Effects of intensity and seasonal timing of disturbances on a rocky intertidal benthic community on the southern coast of Korea

The effects of intensity and timing of disturbances on recovery of marine benthic organisms were investigated on a rocky intertidal shore in Gwangyang Bay, Korea. We hypothesized that the recovery pattern of the benthic community structure would be affected by disturbance intensity and season. Twenty-eight permanent plots were set up, with disturbance intensity (cleared plots and sterile plots) and seasonal disturbance (fall 1999 and spring 2000) incorporated into the experimental design. To monitor natural seasonal variation in benthic community abundances, we established seven permanent unmanipulated plots. Turf-forming algae were observed in the unmanipulated plots throughout the experimental period, whereas green algae and invertebrate presence varied with season. In the disturbance-intensity experiment, turf-forming and green algae were dominant in cleared plots. The highest coverage of sessile organisms was observed in sterile plots, which exhibited the highest species richness because of their relatively low macroalgal coverage. Seasonal effects of disturbance were an important factor in the recovery pattern of benthic organisms under high disturbance intensity. Coverage of green algae was higher in sterile spring plots than in sterile fall plots; this result was attributed to low spatial competition, as the disturbances occurred just before green algal blooms. On the other hand, the abundances of barnacles and bivalves were highest on sterile fall plots, as these organisms were suppressed by green algal blooms in other periods. These results indicate that the effects of disturbance intensity on benthic community recovery patterns can be influenced by season of disturbance.     

Trophic cascades in rocky shore tide pools: distinguishing lethal and nonlethal effects

The effects of predators on the density of their prey can have positive indirect effects on the abundance of the preys resource via a trophic cascade. This concept has strongly influenced contemporary views of how communities are structured. However, predators also can transmit indirect effects by inducing changes in prey traits. We show that the mere presence of predator risk cues can initiate a trophic cascade in rocky shore tide pools. In large (mean surface area =9 m²), natural tide pools, we manipulated crab density and their foraging ability to examine the relative importance of lethal (density-mediated) and non-lethal (trait-mediated) predator effects to algal community development. We found that perceived predation risk reduced snail density as much as the direct predation treatment, showing that green crab predation was not an important factor regulating local snail density. Instead, snail emigration away from resident crabs appears to be the most important factor regulating local snail density. As a result, the abundance of ephemeral green algae was similar in the predation risk and direct predation treatments, suggesting that the consumption of snails by crabs plays a minimal role in mediating the trophic cascade. Increased attention to trait-mediated effects that are transmitted by predator-induced changes in prey behavior may change our view of how predators exert their strong influence on community structure.     

Stability and change of phytoplankton communities in a highly dynamic environment—the case of large,shallow Lake Balaton (Hungary)

Time series data of key environmental variables (water temperature, global radiation, vertical light attenuation, internal P load) and biomass of four colour classes of photosynthetically active algae were collected during 2003 and 2004 with daily resolution. Using these data, seasonal patterns of phytoplankton were analyzed as a function of the dynamic environment. Abstraction of the environmental state as a point in multi-dimensional space was used to identify habitat templates of bloom-forming groups and derive an indicator of environmental stability/physical disturbance. These templates were synthesized into a simple threshold model that sufficiently simulated development and collapse of various blooms. Blooms were, however, rare events related to specific environments with strong, unidirectional forcing. Tentative quantification of disturbance and compositional stability/community change allowed discriminating disturbance-driven changes and autogenic succession with reasonable success. The two processes were found to be equally important in shaping the composition and biomass of phytoplankton.     

Long-term trends in algal-dominated rocky subtidal communities on the Swedish west coast — a transitional system?

Rocky subtidal communities on the Swedish west coast were monitored over approximately 16 years (1970–1985) by means of stereophotographic recordings, at intervals, of fixed sites.Dominating elements of community structure were studied at the 5- and 10 m depth levels of an outer archipelago locality. Excluding seasonal variations, the basic community structure, largely dominated by the perennial phaeophycean Halidrys siliquosa (L.), remained relatively stable over the first 7–8 years of study, but with an increasing trend in cover of filamentous epiphytic red algae. Later, the structure of the studied communities changed radically. The changes initially involved intense settling of the blue mussel Mytilus edulis L. and reduced cover of red algae. The mussels settled largely on Halidrys plants which eventually broke down almost completely. Later, there was intensive predation on Mytilus by the starfish Asterias rubens Linné. After the elimination of Mytilus, the studied test areas were dominated by filamentous ephemeral red algae for the rest of the study period, but with some regeneration of Halidrys at the 5 m level.The observed changes were thought to be indicative of a major disturbance in the coastal ecosystems off the Swedish west coast during the studied period. This notion is discussed in relation to studies of other components of the ecosystem during the same time period and in relation to possible causes. Among the latter may be mentioned a suspected eutrophication process, long-term temperature anomalies and resulting effects on biological interrelationships such as competition, predation and chemical defense.The observed trends included a shift in dominance from slow-growing perennial algae to fast-growing ephemeral algae. If this shift was a general phenomenon, for which there is some circumstantial evidence, it could have important repercussions on the cycling of nutrients and organic matter in the coastal ecosystem. Possible effects are discussed on hypothetical terms in relation to observed anomalies in phytoplankton blooms and benthic oxygenation.     

Effects of interactions between algae and grazing gastropods on the structure of a low-shore intertidal algal community

Summary At low levels on shores in New South Wales, foliose algae are abundant and often occupy all substrata; microalgal grazing gastropods are rare or absent. At higher levels, foliose algae are sparse or absent and grazing gastropods are abundant. Hypotheses for the causes of the lower vertical limits of distribution of these grazers include the effects of increased predation or the deleterious physiological effects of increased period of submergence at lower levels on the shore. Alternatively, the presence of the algae, because they occupy space and deprive the grazers of substratum for feeding, may prevent the downward movement, or survival of the grazers at low levels. Under the first two of these hypotheses, algae are able to colonize and grow in low-shore areas as an indirect result of factors which remove grazers. Under the third hypothesis, the algae are directly responsible for the lack of grazers.Experimental clearings of the low-shore algae and introductions of the mid-shore limpets Cellana tramoserica and Siphonaria denticulata were used to test these hypotheses. C. tramoserica grazes microalgae and removes them from the substratum, preventing colonization. S. denticulata, in contrust, crops the algae, leaving a visible cover of algae on the substratum, which can grow rapidly. Because of its method of feeding, S. denticulata had no measurable impact on the rates of colonization, nor on the dry weights of algae, compared with those of ungrazed areas. C. tramoserica could keep cleared areas tree from foliose algae, but only when the limpets were mainfained in great density (10 per 900 cm²). They were less effective where wave-action was greater.Neither species of limpets could survive when placed onto beds of mature algae, because they had no substratum on which to cling and were swept away by the waves. C. tramoserica did not invade clearings below their lower limit of distribution where they had to move over a bed of foliose algae. Few C. tramoserica moved directly downshore into cleared areas. When placed on bare rock within low-shore beds of algae of different ages, S. denticulata remained amongst the algae and maintained their tissue-weights. Few C. tramoserica remained in areas with well-developed algae, compared with areas having sparse algal growth. Those Cellana which remained amongst well-developed algae lost weight, whereas limpets in areas with less algal growth mammtained their weights. In experimental cages in low-shore beds of algae, where the limpets were inaccessible to potential predators, C. tramoserica lost weight and died. On cleared areas they survived for many weeks, but lost weight and died as algae grew and covered the substratum. In the absence of predation, the micro-algal grazer C. tramoserica could not survive in lowshore areas because algae grew too fast and occupied the substratum, making it inaccessible for the limpets to graze; the algae, once grown beyond small sporelings, are not a suitable food-source for C. tramoserica, and the loss of weight and death of these limpets is attributable to starvation.The lower limit of distribution of C. tramoserica is not due to the direct effects of physical factors associated with prolonged submersion, nor to the impact of predators, but is apparently determined by the presence of rapidly growing, extensive beds of foliose algae at low levels on the shore. The cause of the limit of distribution of S. denticulata is not yet known and predation may prove to be important. Removal of S. denticulata from low-shore algal beds would not, however, affect the domination of substrata by algae. Grazing by S. denticulata at very great density had no effect on algal cover nor weight. In the intertidal community studied, the persistence of a low-shore algal zone, bounded above by abundant grazers is not influenced by the activities of predators, but is a direct result of interactions between the grazers and the algae.     

Stress and disturbance in the phytoplankton community of a shallow,hypertrophic lake

The validity of Connell's intermediate disturbance hypothesis in phytoplankton communities was tested on data from a hypertrophic, shallow lake, Hjarbæk Fjord, Denmark.The present data from Hjarbæk Fjord demonstrate the difficulties in distinguishing stress from disturbance in a phytoplankton community, and show that great changes in the phytoplankton community can take place within few days.A collapse of blue-green algae in late June 1986 caused remineralization of nutrients and resulted in a rapid increase of fast-growing small chlorococcal green algae and phytoplankton species diversity, without any external disturbances acting on the lake. External disturbances in the form of wind action and brackish water intrusion occurred several days after the onset of these events. Carbon depletion and pH 11.0 were severe stress factors on the phytoplankton community. They were induced by calm, warm weather, but eventually acted as a kind of disturbance to the normally well circulated lake.     

Taxonomy and First Records of Two <Emphasis Type="Italic">Megaselia</Emphasis> Rondani Species (Diptera: Phoridae) Preying upon Eggs of <Emphasis Type="Italic">Phyllomedusa iheringii</Emphasis> Boulenger (Anura: Phyllomedusidae)

Non-aquatic reproductive modes have evolved among frogs possibly favored by some advantages such as the avoidance of aquatic predators. These reproductive modes, however, make the egg clutches susceptible to terrestrial predators, among which Diptera larvae are some of the most harmful. The present work reports the predation by phorid flies of 22 egg clutches of Phyllomedusa iheringii Boulenger in the South of Brazil. Phorid specimens were identified as Megaselia bruchiana (Borgmeier & Schmitz) and Megaselia necrophaga (Enderlein), species that were reported previously to be associated with ants and dead beetles, respectively. Frog-feeding in these species is hypothesized to be use of an alternative seasonal food source. We amend the diagnoses of both Megaselia species and provide new illustrations to facilitate their identification. We also describe the male of M. bruchiana for the first time and associate males with females of both species.     

Facultative mutualism between an herbivorous crab and a coralline alga: advantages of eating noxious seaweeds

Because encrusting coralline algae rely on herbivory or low light levels to prevent being overgrown by competitively superior fleshy algae, corallines are relatively rare in shallow areas with low rates of herbivory. In contrast to this general trend, the branching coralline alga Neogoniolithon strictum occurs primarily in shallow seagrass beds and along the margins of shallow reef flats where herbivory on macrophytes is low. This alga apparently persists in these habitats by providing refuge to the herbivorous crab Mithrax sculptus at mean densities of 1 crab per 75 g of algal wet mass. When crabs were removed from some host corallines, hosts without crabs supported 9 times the epiphytic growth of hosts with crabs after only 30 days. Crabs without access to a coralline alga were rapidly consumed by reef fishes, while most of those tethered near a host alga survived. These results suggest that the crabs clean their algal host of fouling seaweeds and associate with the host to minimize predation. However, to effectively clean the host, the crab must consume the wide array of macroalgae that commonly co-occur with coralline algae in these habitats, including chemically defended species in the genera Halimeda, Dictyota, and Laurencia. Crabs did readily consume these seaweeds, which were avoided by, and are chemically defended from, herbivorous fishes. Even though crabs readily consumed both Halimeda and Dictyota in whole-plant feeding assays, chemical extracts from these species significantly reduced crab feeding, suggesting that factors other than secondary chemistry (e.g., food value, protein, energy content), may determine whole-plant palatability. Having the ability to use a wide variety of foods, and choosing the most profitable rather than the least defended foods, would diminish foraging time, increase site fidelity, and allow the crab to function mutualistically with the host alga. Despite the obvious benefit of associating with N. strictum, M. sculptus did not prefer it over other habitats offering a structurally similar refuge, suggesting that these crabs are not N. strictum specialists, but rather occupy multiple habitats that provide protection from predators. Structurally complex organisms like N. strictum may commonly suppress competitors by harboring protective symbionts like M. sculptus. It is possible that diffuse coevolution has occurred between these two groups; however, this seems unlikely because both herbivore and host appear to respond most strongly to selective pressures from predators and competitors outside this association.     

Diversity,heterogeneity and consumer pressure in a tropical rocky intertidal community

Summary Previous studies indicated that at Taboguilla Island (Gulf of Panama), persistence of many intertidal organisms depended on holes and crevices in the rock as refuges from both vertebrate (fishes) and invertebrate (crabs, gastropods, chitons) consumers. Here, we evaluate the influences of substratum heterogeneity and consumers on patterns of diversity of sessile organisms in this habitat. Local substratum topography is highly variable, ranging from smooth to irregular surfaces. Algal crusts typically dominate all low zone rock surfaces, and most other sessile spegies (invertebrates and foliose algae) occur in holes and crevices. Number (S) and diversity (H) of sessile species is lower on homogeneous surfaces than on heterogeneous surfaces. Rate of increase in S with area sampled is positively correlated with substratum heterogeneity; number of species sampled per transect at a homogeneous site would be about 10 vs 30 to 60 on a heterogeneous site. Large fishes and crabs forage intensively over both substratum types, but cannot enter holes and crevices to eat prey. Gastropods, chitons, limpets, and small crabs feed on both substrata but vary in abundance from hole to hole. Prey mortality is thus intense and constant on open surfaces, but variable in space and time in holes and crevices. When consumers are excluded from the general rock surface, algal crusts are settled upon and overgrown by foliose algae, hydrozoans, and sessile invertebrates, particularly bivalves. Both S and H first increase, as sessile species invade and become more abundant, and then decrease as the rock oyster Chama echinata begins to outcompete other species and dominate primary space. Hence, consumers normally keep local diversity low by removing most sessile prey from open surfaces.In these experiments, a consumer pressure gradient was established by removing 0, 1, 2, 3, and all of 4 distinct groups of consumers. As predicted by the intermediate disturbance hypothesis, lowest diversity occurred at lowest (total exclusion) and highest consumer pressure (normal condition). Highest diversity occurred at intermediate consumer pressure. Unexplained variation in this relationship is probably due to quantitative and qualitative differences in consumer regime, variation among plots in substratum heterogeneity, and insufficient time for competitive dominance by Chama to be fully expressed. On a small (0.25 m²) spatial scale, consumers maintain low diversity by keeping prey scarce and causing local extinctions. On larger spatial scales, they may maintain and even produce high diversity through their interaction with substratum heterogeneity and possibly low dispersal rates of sessile species.     

The Structure and Density of Aggregations of the Hermit Crab <Emphasis Type="Italic">Pagurus proximus</Emphasis> Komai, 2000 in Epibiosis of the Brown Alga <Emphasis Type="Italic">Costaria costata</Emphasis> (C. Agardh) De A. Saunders, 1895 in the Sublittoral Zone of Vostok Bay,Sea of Japan

A study of epibiosis of the brown alga Costaria costata in the sublittoral zone of Vostok Bay, Sea of Japan during the spring and summer of 2016 showed that hermit crabs of the following four species accounted for approximately 85% of the total biomass: Pagurus proximus, P. minutus, P. brachiomastus, and P. middendorffii. Of these, the most abundant one (96.5% of the total number of hermit crabs) was P. proximus, the mean aggregation density of which on C. costata (429 ± 221 ind./m²) was higher by an order of magnitude than that in the adjacent area of the bouldery bottom. The size ranges of males and females of this species in aggregations on boulders and those on C. costata were similar, with a predominance of larger individuals on algae. In the size–frequency distribution, three groups of individuals corresponding to three size–age cohorts were differentiated. The male-to-female ratio was 1 : 1.7. Hermit crab aggregations on C. costata are ephemeral; their existence is determined by the duration of the period of algae development, that is, from late spring to the middle of the summer.     

Scaling the effects of predation and disturbance in a patchy environment

The effects of hydraulic disturbances on the impact of two predatory benthic invertebrates on their prey were examined in a stream at two distinct spatial scales. At the scale of small habitat patches (0.0625 m²), hydraulic patch type was an important determinant of the microdistribution of prey and predators. Prey abundances were similar across all patch types at baseflow, but local densities were higher in patches identified as low-flow refugia after periods of high and fluctuating flow. The microdistribution pattern of predatory larvae of a caddisfly, Plectrocnemia conspersa, was similar to that of its prey, whereas predatory larvae of an alderfly, Sialis fuliginosa, did not shift their microdistribution significantly with discharge and were always most abundant in lowflow refugia. There was little evidence of an aggregative response of predators with prey, even though both predators and prey are mobile. Both predator species showed similar patch-specific patterns of per capita consumption rates: uniform consumption rates across hydraulic patch types at low and moderate flows, but highest in flow refugia during high flows. Species-specific patterns, however, were apparent in the magnitude and direction of differences between consumption rates during disturbance events, and in comparable patches at base flow: At high flow, consumption rates for P. conspersa were exaggerated (3.9 times higher) in flow refugia but at par in other patches; for S. fuliginosa they were at par in flow refugia but reduced in other patches (up to 3.3. times lower). These differences may be related to species-specific foraging behaviours (search vs ambush predators) and the influence of prey movements on feeding success. Using the patch-scale results only, it is difficult to predict the effects of physical disturbance on predation intensity at the larger scales of whole habitats, populations or communities. At the large scale (>200 m²), net predator impacts were estimated over the stream reach, using a spatially explicit model that accounts, in an additive way, for habitat heterogeneity and patch-specific responses of predators and prey. The relationship between predator impact over the whole reach and hydraulic disturbance differed for the two predators. The predator impact of S. fuliginosa decreased with increasing hydraulic disturbance, as predicted by the harsh-benign hypothesis. There was no directional trend for P. conspersa, however, and maximum predator impact may occur at intermediate disturbance levels. For the prey community in this stream, predation pressure from S. fuliginosa appears to fluctuate directly with the discharge hydrograph, whereas predation from P. conspersa may be more persistent. Flow refugia may play a dual role in the sructure of stream communities by preventing catastrophic mortality of animals (predators and prey) from physical forces during disturbances, and by maintaining (or perhaps increasing) predation pressure. Summing the effects of species interactions in small habitat patches to the larger scale of a whole stream reach indicates that the scale of approach influences the observed patterns and their implied underlying process.     

Differential seed and seedling predation by crabs: impacts on tropical coastal forest composition

Recently, the importance of seed predation by crabs on mangrove species distributions and densities has been established by several studies. In a tropical coastal terrestrial forest in Costa Rica, we investigated the relative importance of predation by land crabs, Gecarcinus quadratus, and hermit crabs, Coenobita compressus, on measured forest composition through a series of seed removal and seedling establishment experiments. We also used natural light-gaps and adjacent non-gap sites to test how canopy cover affects crab predation (seed removal) and seedling establishment. We found fewer tree species (S=18) and lower densities (seedlings, saplings, and adults) in the coastal zone within 100 m of coastline, than in the inland zone (S=59). Land crab densities were higher in the coastal zone (3.03±1.44 crabs m^–2) than in the inland zone (0.76±0.78 crabs m^–2), and hermit crabs were not present in the inland zone. Seed removal and seedling mortality also were higher in the coastal zone than in the inland zone, and in the open controls than in the crab exclosures. Mortality of seeds and seedlings was two to six times higher in the controls than exclosures for four of the five experiments. Crabs preferred seeds and younger seedlings over older seedlings but showed no species preferences in the seed (Anacardium excelsum, Enterolobium cyclocarpum, and Terminalia oblonga) and seedling (Pachira quinata and E. cyclocarpum) stages. We conclude that the observed differences in tree densities were caused by differential crab predation pressure along the coastal gradient, while the differences in species composition were due to predator escape (satiation) by seed quantity. Canopy cover did not affect seed removal rates, but did affect seedling survival with higher mortality in the non-gap versus gap environments. In summary, crab predation of seeds and seedlings, and secondarily canopy cover, are important factors affecting tree establishment in terrestrial coastal forests.     

The impacts of overwintered green frog larvae on wood frog embryo mortality under a wetter climate

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红树林湿地相手蟹科动物摄食生态研究进展

相手蟹科物种是红树林湿地的主要底栖动物类群,在生态系统中起着重要的作用。我国大陆地区目前已记录的相手蟹科物种数量为12种,低于其它红树林地区(国内常用的采样方法会造成螃蟹物种数量和密度的低估),其中褶痕相手蟹(Sesarma plicata)、无齿相手蟹(S.deaani)和双齿相手蟹(S.bidens)等是常见种。红树植物叶片是相手蟹科动物的主要食物来源,相手蟹科动物通过地表摄食和洞穴贮存的形为消耗了大量的红树植物凋落叶,使这些凋落叶的有机质和营养元素得以保留在生态系统内,在凋落叶的周转和物质归还方面起到重要的作用。它们同时也摄食红树植物的繁殖体并且对不同物种的繁殖体具有摄食偏好,这可能对一些红树物种的植被更新能力和红树植被群落结构产生影响。相手蟹科动物对不同物种和不同状态的红树叶片也存在摄食偏好,通常对腐烂的叶片摄食偏好较强;螃蟹的摄食偏好与叶片的营养成份、粗纤维和单宁含量以及C/N比等性质有关;但在恶劣的野外环境下,螃蟹则会表现出随机性摄食。目前关于相手蟹科动物生态学作用的认识仍不充分,例如它们的种群大小和对凋落物的转化作用等,有待于进一步研究。     

Observations of Toxic Blue-Green Algae (Cyanobacteria) in Some Scandinavian Lakes

Blooms of blue-green algae from 51 eutrophic Scandinavian lakes were investigated during the period 1978–1984, to ascertain the occurrence of toxinogenic species. Toxicity assays were performed by intraperitoneal injection of suspensions of freeze-dried algal material in mice. Toxin-producing blue-green algae were found in 30 lakes. They belonged to 11 different species of the six genera Anabaena, Aphanizomenon, Gomphosphaeria, Microcystis, Nodularia and Oscillatoria. The presence of toxinogenic strains of blue-green algae seemed quite constant in several of the localities studied. In some lakes, more than one toxic species were found to develop simultaneously. The level of toxicity showed large variation (MLD100, 6 to > 2500 mg/kg), but clinical and pathological changes were quite uniform. The results indicate that water-blooms of toxin-producing blue-green algae, in the geographical area in question, are regionally widespread. In some localities, blooms of blue-green algae are apparently always toxic. Several aspects of the toxic blue-green algae problem are discussed.     

13C and 15N depletion in components of a foodweb from an ephemeral boreal wetland compared to boreal lakes: putative evidence for microbial processes

Stable carbon and nitrogen isotope ratios were used to posit the relative importance of microbial processes on energy pathways in an ephemeral, humic boreal wetland compared to four clearwater lakes in northwestern Ontario, Canada. In addition to algae and dipteran larvae, odonate larvae were sampled as these latter organisms are known to predate indiscriminately on smaller invertebrates and are thus likely to have average isotope ratios reflective of their habitats. Similarities in ¹³C and ¹⁵N values between lake insect larvae and emerged adults suggested that littoral foodwebs in these oligotrophic lakes may rely to a considerable degree upon terrestrial carbon. Wetland insect larvae and algae were depleted in both ¹³C and ¹⁵N compared to biota in lakes. Carbon isotope analysis implied a substantial presence of microbial respiration from decomposition in the humic wetland, whereas nitrogen isotope analysis suggested the prevalence of microbially modified nitrogen dynamics, including the possibilty of N-fixation.     

Experimental study of indirect effects of fish on the demographic parameters of cladoceran species under eutrophic conditions

Indirect effects of fish on the demographic parameters of cladoceran species were studied under eutrophic conditions. Laboratory experiments were performed with water from control and fish mesocosms to avoid the direct impact of fish predation. In the experiments with the water from the fish mesocosms, fish indirectly negatively affected the demographic parameters of large cladocerans (Daphnia magna and D. pulicaria) due to the enhanced abundance of blue-green algae in the phytoplankton. However, small Ceriodaphnia quadrangula and littoral species Simocephalus vetulus did not respond to the presence of blue-greens. Due to this mechanism, the total abundance of cladoceran species can be sustained during the development of blue-green algae because large and small bodied species differ in their resistence to high concentratons of blue-green algae. Fish chemical signals (kairomones) did not influence the demographic parameters of any cladoceran species.     

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.     

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.     

Effect of acanthocephalan parasites on hiding behaviour in two species of shore crabs

The effect of acanthocephalan parasites (Profilicollis spp.) on the hiding behaviour during low tide of two species of shore crabs (intermediate hosts), Macrophthalmus hirtipes (Brachyura: Ocypodidae) and Hemigrapsus crenulatus (Brachyura: Grapsidae), was examined at Blueskin Bay, South Island, New Zealand. Exposed M. hirtipes were found to have significantly higher infection levels than did hidden conspecifics. This pattern was not observed for H. crenulatus. Mean cystacanth numbers were found to be considerably higher in M. hirtipes than H. crenulatus. Crabs exposed at low tide are at a greater risk of predation by definitive shorebird hosts than are hidden conspecifics. Preferential manipulation of one intermediate host species over another could influence diversity within ecosystems.