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1.
Summary Prey species may use many different behaviours to avoid predation. In this study, the antipredator behaviours of juvenile roach (Rutilus rutilus) and juvenile perch (Perca fluviatilis) were studied in wading pools with three kinds of structural complexity: no structure, structure simulating vegetation and structure simulating bottom crevices. Predation experiments with piscivorous perch and habitat choice experiments with the prey were performed, and the foraging success and prey choice of the predators were related to the type of structure. Predator foraging success was lower in the vegetation than in the other treatments. In the absence of structure and with vegetation structure, predators preferred perch over roach, while the preference was reversed in the crevice treatment. Roach and perch differed in their antipredatory behaviours. Roach responded to the presence of predators by schooling, moving fast and remaining at the surface, and escaped from attacks by jumping out of the water. In contrast, perch moved more slowly, dispersed after attacks and tried to hide at the bottom. Perch always preferred the vegetation structure to the non-structured part of the pool, while roach showed preference for the vegetation structure only when predators were present. Roach never occurred in crevices, whereas perch used crevices when predators where present. Predator pursuit speed was lower in the vegetation structure than in the non-structured treatment, but prey escape speed was unaffected. The results suggest that both the quantity and quality of structural complexity interacting with species-specific antipredator behaviours are important for predator-prey dynamics. It is also suggested that the presence of structure can have substantial effects on the structure of North Eurasian fish communities, by affecting relative and absolute predation pressures from piscivorous perch on prey species. Correspondence to: B. Christensen  相似文献   

2.
Summary In a laboratory experiment it was shown that piscivorous predators reversed the outcome of competitive interactions between two fish prey species, juveniles of roach (Rutilus rutilus) and perch (Perca fluviatilis), by behaviorally affecting their use of two available habitats, an open water habitat and a structurally complex refuge. The shift in the competitive relationship was the result of predators forcing the juvenile fishes into a prey refuge with high structural complexity. While roach was competitively superior in the unstructured habitat, perch was superior in the structurally complex prey refuge. The reversal in competitive relationship was demonstrated both with respect to foraging rate and growth rate and resulted from the high structural complexity in the prey refuge interfering with the roach's swimming performance. Because survival and growth patterns through the juvenile stages have profound effects on the population/community dynamics of size-structured populations such as those of fish, behaviorally induced changes in competitive ability should have significant implications also at the population and community levels.  相似文献   

3.
Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.  相似文献   

4.
Field observations and manipulative experiments in a nearshore cobble bed (2 to 3 m below mean low water) at Eagle Head, Nova Scotia, Canada, between 1984 and 1986, showed that small juveniles ofStrongylocentrotus droebachiensis (3 to 6 mm diam) sheltering beneath cobbles had a refuge from predators such as rock crabs, small lobsters, and fish. Sea urchins gradually outgrew these refuges and small adults (25 to 30 mm) required larger rocks as shelter from predators, particularly large cancrid crabs. Small juveniles were usually solitary and well dispersed beneath cobbles, whereas small adults tended to aggregate on the undersides and in the interstices of boulders. These aggregations may develop passively as sea urchins accumulate in suitablysized refuges. Chemotaxis experiments indicate that juvenileS. droebachiensis are repelled by waterborne stimuli from conspecifics. In a factorial experiment, effects of the presence of potential predators (rock crabs and lobsters) and/or food (kelp) on the behaviour of large juvenile (10 to 15 mm) and small adult sea urchins were examined in flowing seawater tanks. Both size classes formed exposed feeding aggregations when kelp was provided as food, irrespective of the presence or absence of predators. In the absence of kelp, each size class responded differently to the presence of a predator: juveniles became more cryptic, whereas adults aggregated on the tank sides. Increased movement to the sides of a tank in the presence of a predator may reflect a flight response, since chemotaxis experiments indicated thatS. droebachiensis is repelled by waterborne chemical stimuli from predators. Observational and experimental data suggest that predation at the late juvenile and early adult stages may influence population structure, distribution and abundance ofS. droebachiensis.  相似文献   

5.
Four decades of observations on the limnology and fishes of Oneida Lake, New York, USA, provided an opportunity to investigate causes of mortality during winter, a period of resource scarcity for most juvenile fishes, in age-0 yellow perch (Perca flavescens) and age-0 white perch (Morone americana). This time series contains several environmental (e.g., winter severity) and biological (e.g., predator abundance) signals that can be used to disentangle multiple effects on overwinter mortality of these fishes. A multiple regression analysis indicated that age-0 yellow perch winter mortality was inversely related to fish length in autumn and to the abundance of alternative prey (gizzard shad [Dorosoma cepedianum] and white perch). However, no length-selective predation of yellow perch by one of the main predators, adult walleye (Sander vitreus), was detected. In contrast, white perch mortality was directly associated with total predator biomass and abundance of white perch in autumn, and inversely related to yellow perch abundance as a potential buffer species, but not to the abundance of gizzard shad. Winter severity was not a significant predictor of mortality for either perch species. Predicted winter starvation mortality, from a model described in the literature, was much lower than observed mortality for yellow perch. Similar models for white perch were correlated with observed mortality. These results collectively suggest that predation is the main mechanism shaping winter mortality of yellow perch, while both predation and starvation may be important for white perch. This analysis also revealed that gizzard shad buffer winter mortality of yellow perch. Although winter duration determines the northern limit of fish distributions, in mid-latitude Oneida Lake and for these species, predator-prey interactions seem to exert a greater influence on winter mortality than starvation.  相似文献   

6.
Human disturbances, such as overfishing, may disrupt predator–prey interactions and modify food webs. Underwater surveys were carried out at six shallow-water reef barrens in temperate waters of northern-central Chile from October to December 2010 to describe the effects of predation, habitat complexity (low, medium and high) and refuge availability on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Rhynchocinetidae), an important mesoconsumer on subtidal hard substrata. Three sites were within managed (restricted access) areas for fishermen, and three were unmanaged (open-access). Field observations and tethering experiments were conducted to examine the relationship between fish and shrimp abundances, and the relative predation rates on shrimps. Direct effects of predation on R. typus body-size distribution were examined from shrimps collected in the field and fish stomachs. The presence and the abundance of R. typus increased with habitat reef complexity and refuge availability. Shrimp abundance was negatively related to fish abundance in managed areas, but not in open-access areas, where shrimp densities were the highest. Also, predation rates and body-size distribution of shrimps were unrelated, although fish consumed more large shrimps than should be expected from their distribution in the field. R. typus occurred most often in shelters with wide openings, offering limited protection against predators, but providing potential aggregation sites for shrimps. Overall, direct effects of predation on shrimp densities and population structure were weak, but indirect effects on shrimp distribution within reefs appear to have been mediated through behavioural responses. Our study highlights the need to assess both numerical and behavioural responses of prey to determine the effects of predator loss on mesoconsumer populations.  相似文献   

7.
Predators such as crabs, whelks, and sea stars attack their bivalve prey in different ways, and predator-induced defenses are an important means of protection. The degree to which induced defenses are specific to different predators, however, remains largely unknown. In laboratory experiments (June to August 1998), we raised mussels (Mytilus edulis L.) in the presence of a drilling predator [the whelk Nucella lapillus (L.)] or a crushing predator [the crab Carcinus maenas (L.)] to determine whether induced changes in prey shell thickness, size, or shape occurred and whether changes were predator-specific. Over a 2 month period, juvenile mussels were exposed to waterborne cues from actively feeding crabs or whelks. Mussels produced thicker shell lips in response to both predators relative to control mussels raised in their absence, and the difference was significantly greater in response to whelks than to crabs. Mussels exposed to whelks showed significantly smaller increases in shell length and width and total wet weight than did mussels exposed to crabs. Thus, there may be a trade-off between shell thickness and linear shell growth and a potential delay in attaining a size refuge from predation. Received: 4 August 1999 / Accepted: 31 January 2000  相似文献   

8.
Predation is often thought of as an unforgiving and strong selective force, quickly selecting against maladaptive behaviour in the prey. It is argued that experience is likely to have low influence on the phenotypic response to predation, as failing to react correctly to a predator may mean death to the prey and no second chance to learn and correct the behaviour. Individuals from different populations of Eurasian perch are known to differ in risk-taking behaviour. Variation in predation pressure has been suggested as a key factor causing these differences, but little is known about the underlying mechanism by which predation generates risk-taking phenotypes in perch. We compared the degree of boldness between two natural populations of Eurasian perch, living under different predation regimes, and the same populations hatched and reared under identical conditions, free from predation. By this common-garden approach, we sought to investigate patterns in the influence of inheritance and experience on boldness phenotype. The wild fish differed in risk taking, with fish from the low predation-risk population acting bolder than fish from the high-risk environment. In the reared fish, both populations behaved equally bold. Only the fish originating from the high predation population showed different behaviour when comparing wild and reared ecotypes. Our results suggest that experience has an important impact on the response to predators and that geographic variation in risk taking between populations of Eurasian perch to a high degree is shaped by adjustments to the current environment. Habituation had an effect of risk-taking behaviour over the experimental period, but consistent differences between individuals were also found. Furthermore, we also show, by the estimation of variance components, that the behaviour we observe is affected by a range of random effects, such as aquaria and group membership, that in concert shapes the behaviour of an individual perch.  相似文献   

9.
The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.  相似文献   

10.
We evaluated the effects of potential predators from intertidal habitats on Strongylocentrotus purpuratus survival using laboratory experiments and assessed abundances of main predatory species along the Pacific coast of North America. The interactive effects of urchins’ and predators’ sizes in mediating predation were quantified. Habitat complexity (substrate pits, adult spine canopy) was manipulated to examine its effects on predation of most susceptible individuals (<14 mm). Pachygrapsus crassipes was identified as a major predator of urchins up to ≈30 mm. A positive effect of predator size on consumption of progressively larger urchins was detected, probably due to a mechanical limitation on crabs’ ability to consume large prey. Larger claws of males with respect to females of comparable sizes facilitated the handling of larger prey. Substrate refuges significantly reduced mortality on juvenile urchins. These results show that crab predation may be important in organizing intertidal communities, despite multiple ecological mechanisms promoting sea urchin survival.  相似文献   

11.
Prey often adopt antipredator strategies to reduce the likelihood of predation. In the presence of predators, prey may use antipredator strategies that are effective against a single predator (specific) or that are effective against several predators (nonspecific). Most studies have been confined to single predator environments although prey are often faced with multiple predators. When more than one predator is present, specific antipredator behaviours can conflict and avoidance of one predator may increase vulnerability to another. To test how prey cope with this dilemma, I recorded the behaviours of lizards responding to the nonlethal cues of a bird and snake presented singly and simultaneously. Lizards use specific and conflicting antipredator tactics when confronted with each predator, as evidenced by refuge use. However, when both predators were present, lizards refuge use was the same as in the predator-free environment, indicating that they abandoned refuge use as a primary mechanism for predator avoidance. In the presence of both predators, they reduced their overall movement and time spent thermoregulating. This shift in behaviour may represent a compromise to minimize overall risk, following a change in predator exposure. This provides evidence of plasticity in lizard antipredator behaviour and shows that prey responses to two predators cannot be accurately predicted from what is observed when only one predator is present.Communicated by W. Cooper  相似文献   

12.
Habitat structure affects intraguild predation   总被引:4,自引:0,他引:4  
Intraguild predation is thought to be ubiquitous in natural food webs. Yet, theory on intraguild predation predicts the intraguild prey to persist only under limited conditions. This gap between theory and empirical observations needs scrutiny. One reason might be that theory has focused on equilibrium dynamics and a limited set of species (usually three) that interact in well-mixed populations in unstructured habitats, and these assumptions will often not hold in natural systems. In this review, we focus on the effects of habitat structure on intraguild predation. Habitat structure could reduce encounter rates between predators and prey and could create refuges for prey. In both cases, habitat structure could reduce the strength of intraguild interactions, thereby facilitating species coexistence. A meta-analysis of studies on manipulation of habitat structure shows that intraguild prey indeed suffer less from intraguild predation in structured habitats. This was further confirmed by a meta-analysis in which studies on intraguild predation were classified according to habitat structure. Intraguild predation reduced densities of the intraguild prey significantly more in habitats with little structure than in habitats rich in structure. The effect of intraguild predation on the shared prey was negative, and not significantly affected by habitat structure. We conclude that habitat structure may increase persistence of the intraguild prey by decreasing the strength of the interaction between intraguild predator and intraguild prey.  相似文献   

13.
Predation risk and foraging behavior of the hoary marmot in Alaska   总被引:2,自引:0,他引:2  
Summary I observed hoary marmots for three field seasons to determine how the distribution of food and the risk of predation influenced marmots' foraging behavior. I quantified the amount of time Marmota caligata foraged in different patches of alpine meadows and assessed the distribution and abundance of vegetation eaten by marmots in these meadows. Because marmots dig burrows and run to them when attacked by predators, marmot-toburrow distance provided an index of predation risk that could be specified for different meadow patches.Patch use correlated positively with food abundance and negatively with predation risk. However, these significant relationships disappeared when partial correlations were calculated because food abundance and risk were intercorrelated. Using multiple regression, 77.0% of the variance in patch use was explained by a combination of food abundance, refuge burrow density, and a patch's distance from the talus where sleeping burrows were located. Variations in vigilance behavior (look-ups to search for predators while feeding) according to marmots' ages, the presence of other conspecifics, and animals' proximity to their sleeping burrows all indicated that predation risk influenced foraging.In a forage-manipulation experiment, the use of forage-enhanced patches increased six-fold, verifying directly the role of food availability on patch used. Concomitant with increased feeding, however, was the intense construction of refuge burrows in experimental patches that presumably reduced the risk of feeding. Thus, I suggest that food and predation risk jointly influence patch use by hoary marmots and that both factors must be considered when modeling the foraging behavior of species that can be predator and prey simultaneously.  相似文献   

14.
Summary During their first 1–2 years of life, juvenile coho salmon (Oncorhynchus kisutch) are stream-dwelling, and feed upon drifting invertebrates. They move upstream from a holding position to intercept individual prey items; the distance moved (attack distance) is an increasing, but decelerating, function of prey size. Since the fish are presumably more visible to predators during such feeding excursions, prey size and risk are associated variables.The effect on attack distance of the presentation of a model predator (a photograph of a rainbow trout) was examined in the laboratory. Attack distances are shortened following presentation of a predator; this is particularly true when the prey are large (Fig. 1). The extent of the reduction of attack distance is directly related to predator presentation frequency, although there appears to be a minimum level to which it will decline (Fig. 2). Hungry fish and fish in the presence of a competitor (simulated by a mirror) are less responsive to the predator, suggesting a trade-off of energetic requirements and risk (Fig. 3 and Table 3). The effect of predation risk should be to reduce the relative proportion of large prey in a juvenile coho's diet, and its net rate of energy intake.  相似文献   

15.
Although scyphomedusae have received increased attention in recent years as important predators in coastal and estuarine environments, the factors affecting zooplankton prey vulnerability to these jellyfish remain poorly understood. Current models predicting feeding patterns of cruising entangling predators, such as Chrysaora quinquecirrha (Desor, 1948), fail to account for the selection of fast-escaping prey such as copepods. Nevertheless, our analysis of gastric contents of field-collected medusae showed that this scyphomedusa fed selectively on the calanoid copepod Acartia tonsa (Dana, 1846) and preferentially ingested adult over copepodite stages. We measured feeding rates in a planktonkreisel while simultaneously videotaping predator–prey interactions. C. quinquecirrha consumed adult A. tonsa ten times faster than copepodites. Differences in prey behavior, in the form of predator–prey encounter rates or post-encounter escape responses, could not account for the elevated feeding rates on adults. Prey size, however, had a dramatic impact on the vulnerability of copepods. In experiments using heat-killed prey, feeding rates on adults (1.5 times longer than copepodites) were 11 times higher than on copepodites. In comparison, medusae ingested heat-killed prey at only two to three times the rate of live prey. These results suggest that during scyphomedusan–copepod interactions, prey escape ability is important, but ultimately small size is a more effective refuge from predation. Received: 26 September 1997 / Accepted: 20 May 1998  相似文献   

16.
Predator–prey relationships provide an excellent opportunity to study coevolved adaptations. Decades of theoretical and empirical research have illuminated the various behavioral adaptations exhibited by prey animals to avoid detection and capture, and recent work has begun to characterize physiological adaptations, such as immune reactions, metabolic changes, and hormonal responses to predators or their cues. A 2-year study quantified the activity budgets and antipredator responses of adult Belding’s ground squirrels (Spermophilus beldingi) living in three different California habitats and likely experiencing different predation pressures. At one of these sites, which is visually closed and predators and escape burrows are difficult to see, animals responding to alarm calls remain alert longer and show more exaggerated responses than adults living in two populations that likely experience less intense predation pressure. They also spend more time alert and less time foraging than adults at the other two sites. A 4-year study using noninvasive fecal sampling of cortisol metabolites revealed that S. beldingi living in the closed site also have lower corticoid levels than adults at the other two sites. The lower corticoids likely reflect that predation risk at this closed site is predictable, and might allow animals to mount large acute cortisol responses, facilitating escape from predators and enhanced vigilance while also promoting glucose storage for the approaching hibernation. Collectively, these data demonstrate that local environments and perceived predation risk influence not only foraging, vigilance, and antipredator behaviors, but adrenal functioning as well, which may be especially important for obligate hibernators that face competing demands on glucose storage and mobilization.  相似文献   

17.
Levenbach S 《Ecology》2008,89(10):2819-2828
Little attention has been given to associational refuges in ecology, despite their potential for maintaining species diversity and supporting higher trophic levels. Here I show how the colonial anemone, Corynactis californica, creates a refuge for benthic macroalgae and invertebrate fish prey on intensively grazed shallow rocky reefs in the Santa Barbara Channel off southern California, U.S.A. On reefs heavily grazed by sea urchins, benthic macroalgae and invertebrate fish prey were relatively more abundant among Corynactis colonies than adjacent areas lacking the anemone. Results from field experiments showed that Corynactis facilitated the recruitment of macroalgae and tubicolous amphipods in "urchin-barren" areas subjected to intensive grazing. In areas forested by giant kelp (Macrocystis pyrifera), where grazing intensity from urchins was low, Corynactis suppressed algal recruitment but facilitated tubicolous amphipods. A manipulation of fish and sea urchins suggested that grazing by urchins, as opposed to predation from fish (primarily surfperch Embiotocidae), suppressed tubicolous amphipods, and this activity was hindered by the presence of Corynactis. In systems where human activity has intensified herbivory, associational refuges may maintain species diversity and support higher trophic levels.  相似文献   

18.
The trophic ecology of the stomiid assemblage (Pisces, Stomiiformes, Stomiidae) in the eastern Gulf of Mexico, a region with physical and biological characteristics typical of oligotrophic low-latitude regimes, was investigated. Over 1400 specimens representing 69 species and 17 genera were examined. Four patterns of feeding were evident among the abundant stomiids: (1) myctophid predation; (2) zooplankton/small micronekton predation; (3) penaeidean shrimp predation; and (4) copepod/micronekton predation. One rare species preyed on cephalopods. Il was concluded that stomiids exhibited a high level of prey-selectivity, particularly considering the broad range of prey types available in the eastern Gulf of Mexico. The absence of numerically dominant potential prey (e.g.Cyclothone spp., sternoptychids) in the diets of piscivorous stomiids is possibly a function of feeding periodicity coupled with stomiid vertical migration. Stomiids may feed at night in the upper 200 m on vertically migrating myctophids while disregarding co-occurring nonmigrating prey during the daytime. Integration of stomiid abundance and diet data suggests that: (1) stomiids are the dominant upper trophic-level predators of the Gulf of Mexico mesopelagial, (2) stomiids inflict the highest predation impact on myctophids in low-latitude midwater ecosystems, and (3) the historic use of predation-avoidance arguments to explain certain mesopelagic phenomena (e.g. vertical migration, ventral photophores) appears to be substantiated by estimates of stomiid predation-impact. The stomiids may serve as key trophic mediators in the transfer of energy from the mesopelagial to the bathyand benthopelagial.  相似文献   

19.
Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining in refuge. In the Iberian rock lizard, Lacerta monticola, we studied the effects of sex, reproductive season, speed of predator approach, and potential loss of mating opportunities on time spent in refuge following simulated predatory attacks. Lizards of both sexes adjusted refuge use to the level of risk by spending more time in refuge when approached rapidly than slowly. Females remained in refuge for equal times in the mating and postreproductive seasons, but males emerged sooner during the mating season, suggesting adjustment to a cost of lost opportunity to search for mates during the mating season. When a tethered female was nearby, males emerged from refuge earlier than if no female was present, indicating a trade-off between risk and mating opportunity. Approach speed affected emergence time when females were absent, but not when a female was present. Approach speed did not affect the probability that, after emerging, a male would return to court the female. For males that courted females intensely (bit them) before entering refuge, approach speed did not affect latency to emerge, but males that courted less intensely emerged sooner if approached slowly than rapidly. These findings show that males adjust the length of time spent in refuge to both risk of predation and reproductive cost of refuge use.Communicated by A. Mathis  相似文献   

20.
Suspension-feeding bivalves are organisms of major functional importance in several aquatic environments around the world. They are also important food items for many fish and benthivorous seabirds. It has commonly been thought that predation pressure on blue mussel (Mytilus edulis) populations is negligible in the Baltic Sea, owing to the scarcity of major invertebrate predators such as starfish and crabs. It has recently been shown, however, that the blue mussel is the main food item for roach (Rutilus rutilus) in the archipelago areas of the western Gulf of Finland, where this freshwater fish species has become increasingly abundant, mainly due to increased eutrophication. To quantify the influence of roach predation on blue mussel populations we measured the standing biomass and size structure of the local blue mussel population and used a bioenergetic model to estimate mussel consumption by individual roach during two consecutive summers, 1997 and 1998. The results of the model were combined with existing data on roach abundance, giving annual consumption estimates of 75–105 kg blue mussel dry weight ha–1 in the study area, approximately two-thirds of these consumed mussels being >10 mm. This corresponds to approximately one-third of the standing population of mussels >10 mm in the area. Our results suggest that the predation effects of vertebrates on Baltic blue mussel populations are not insignificant, as previously believed. Predation by roach and other predators may have an important structuring effect on unstable blue mussel communities within the Gulf of Finland, where the species lives at the edge of its range.Communicated by M. Kühl, Helsingør  相似文献   

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