THE TROPHIC ECOLOGY OF FRESHWATER GAMMARUS SPP. (CRUSTACEA:AMPHIPODA): PROBLEMS AND PERSPECTIVES CONCERNING THE FUNCTIONAL FEEDING GROUP CONCEPT

Gammarus spp. are widespread throughout a diverse range of freshwater habitats and can be the dominant part of many benthic macroinvertebrate assemblages, in terms of both numbers and/or biomass. Although the vast majority of studies have emphasized the herbivorous nature of Gammarus spp. and their ‘shredder’ functional feeding group (FFG) classification, we show that a far wider food base is exploited than has been previously acknowledged. This ‘plasticity’ as herbivore/predator is linked to the success of Gammarus spp. in persisting in and colonizing/invading disturbance-prone ecosystems. Intraguild predation and cannibalism are more common than previously realized. This behaviour appears to be a causal mechanism in many amphipod species replacements. Additionally, Gammarus spp. are major predators of other members of the macroinvertebrate community. Furthermore, while many studies have emphasized fish predation on Gammarus spp., we illustrate how this fish: amphipod, predator: prey interaction may be a two-way process, with Gammarus spp. themselves preying upon juvenile and wounded/trapped fish. We urge that a new realism be adopted towards the trophic ecology of Gammarus spp. and their role as predators and prey and that previously established FFG assumptions of both the food and the feeder be questioned critically.     

The functional role of Gammarus(Crustacea, Amphipoda): shredders, predators, or both?

Gammarus spp. are traditionally viewed under the functional feeding group (FFG) concept as herbivorous `shredders'. Although recent studies suggest that Gammarus should also be viewed as predators, this latter role remains contentious. Here, in a laboratory experiment, we objectively examine the balance between shredder and predator roles in a common freshwater species. Gammarus pulex preyed significantly on mayfly nymph, Baetis rhodani, in both the presence and absence of excess leaf material. There was no significant difference in predation where the alternative food, that is, leaf material, was present as compared to absent. Also, G. pulex shredded leaf material in the presence and absence of B. rhodani. However, shredding was significantly reduced where alternative food, that is, B. rhodani prey, was present as compared to absent. Further, G. pulex had a clear leaf species preference. Our results suggest that Gammarus function as both predators and shredders, with the balance of the two roles perhaps depending on food availability and quality. We discuss implications for the use of the FFG concept in assessing freshwater processes, and the role that Gammarus predation may play in structuring macroinvertebrate communities.     

Prey Selection by the Water Bug Aphelocheirus aestivalis Fabr. (Heteroptera: Aphelocheiridae)

Feeding experiments were conducted in situ in flow through boxes to determine the vulnerability of different prey types (the mayfly larvae Baetis, Ephemerella and Ecdyonurus spp., the caddisfly larva Hydropsyche spp. and the crustacean Gammarus spp.) to the predaceous water bug Aphelocheirus aestivalis and to estimate the predation impact of Aphelocheirus on prey populations. Experiments with equal densities of mixed prey and experiments where each prey was tested individually revealed that Baetis and Ephemerella were most vulnerable to Aphelocheirus predation: Hydropsyche, Ecdyonurus and Gammarus were little/not preyed upon. The present results suggest that vulnerability of prey depends mainly upon prey mobility and microhabitat overlap between predator and prey and that feeding behaviour of Aphelocheirus resembles more that of megalopterans than of stoneflies. Experiments with different prey densities (120–720 prey m⁻²) showed that the predation rate of Aphelocheirus increased with increasing prey density. Experiments with different substrates documented that mortality rates of prey decreased with increasing substrate complexity. When substrate conditions were complex mortality rates in the control and in the experimental boxes were the same which suggest little importance of Aphelocheirus predation on mayflies in the study site.     

Uncoupling strong predator–prey interactions in streams: the role of marginal macrophytes

Predation refugia can facilitate the coexistence of predators and prey within an ecosystem by weakening trophic links between the two. The marginal macrophytes of shallow lakes are used facultatively by zooplankton to escape fish predation, leading to the stabilisation of lentic food webs. Little is known about such a role for the marginal macrophytes of lotic systems. In this paper, we examine whether the marginal macrophytes of chalk streams help buffer the interaction between the freshwater shrimp, Gammarus pulex and a benthic fish predator, Cottus gobio, both characteristically abundant in such systems. Quantitative surveys were taken of Gammarus and bullheads in winter and summer in a southern English chalk stream. These indicated that Gammarus seasonally switched their habitat preference, from marginal macrophytes in summer, to midchannel habitats in winter. Bullheads exhibited an opposite trend, preferring midchannel habitats in summer and all habitats, particularly marginal macrophytes, in winter. Large Gammarus and precopula pairs were found almost exclusively in the margins in summer. This spatial separation between Gammarus and bullheads in summer, though not in winter, was reflected in bullhead diets, as determined by gut analysis. In field manipulation experiments, bullhead presence was the strongest factor explaining the between‐habitat distribution of Gammarus in both summer and winter, indicating that the habitat shift of Gammarus was driven by the distribution of fish. Other abundant invertebrate taxa, including Asellus, mayflies and chironomids, exhibited little avoidance of habitat patches of high bullhead density. We conclude that marginal macrophytes in chalk streams can potentially facilitate the coexistence of high densities of both Gammarus and bullheads by spatially separating predator and prey in summer. They may further allow large populations of Gammarus to persist in the presence of high bullhead density at stretch‐wide spatial scales, by reducing the predation by bullheads of large breeding and newly born individuals of Gammarus in summer.     

Differential physico-chemical tolerances of amphipod species revealed by field transplantations

Physico-chemical regimes of river systems are major determinants of the distributions and relative abundances of macroinvertebrate taxa. Other factors, however, such as biotic interactions, may co-vary with changes in physico-chemistry and concomitant changes in community composition. Thus, direct cause and effect relationships may not always be established from field surveys. Equally, however, laboratory studies may suffer from lack of realism in extrapolation to the field. Here, we use balanced field transplantation experiments to elucidate the role of physico-chemical regime in determining the generally mutually exclusive distributions of two amphipod taxa, Gammarus (two species) and Crangonyx pseudogracilis. Within two river systems in Ireland, the former species dominate stretches of well oxygenated, high-quality water, whereas the latter dominates stretches of poorly oxygenated, low-quality water. G. pulex and G. duebeni celticus did not survive in bioassay tubes in areas dominated by C. pseudogracilis, which itself survived in tubes in such areas. However, both C. pseudogracilis and Gammarus spp. survived equally well in tubes in areas dominated by Gammarus spp. Physico-chemical regime thus limits the movement of Gammarus spp. into C. pseudogracilis areas, but some other factor excludes C. pseudogracilis from Gammarus spp. areas. Since previous laboratory experiments showed high predation rates of Gammarus spp. on C. pseudogracilis, we propose that predation by the former causes exclusion of the latter. Hence, presumed effects of physico-chemical regime on macroinvertebrate presence/abundance may often require experimental field testing and appreciation of alternative explanations. Received: 14 June 1999 / Accepted: 20 January 2000     

Different Predation Impacts of Two Cyclopoid Species on a Small-sized Zooplankton Community: An Experimental Analysis with Mesocosms

We used mesocosms to analyze predation impacts on the prey populations and prey community structures by two cyclopoid copepod species, the larger Mesocyclops pehpeiensis and the smaller Thermocyclops taihokuensis, who coexist with small-sized herbivorous zooplankton species in a fish-abundant lake. The overall predation impact on the prey populations was stronger for Mesocyclops than for Thermocyclops. Mesocyclops had a strong and less selective impact on the rotifer community but a selective impact on the crustaceans. In contrast, Thermocyclops had a selective predation impact on rotifers but a weak and less selective impact on the crustacean community. As a result, the former predator reduced the diversity of the crustacean community but not the rotifer community, while the latter had an opposite impact on the diversities of the two communities. It has been suggested that fish induce development of a zooplankton community dominated by the small-sized zooplankton species in fish-abundant lakes. Our results demonstrated that cyclopoid copepods altered species composition and diversity of the small-sized zooplankton community in such lakes. Thus, the results have given an important suggestion on the role of the invertebrate predator cyclopoid copepods, which often coexist with fish, that they determine population dynamics and community structures of small-sized zooplankton in fish-abundant lakes.     

Environmental variation and the predator-specific responses of tropical stream insects: effects of temperature and predation on survival and development of Australian Chironomidae (Diptera)

The threat posed by predation varies among predator species and with environmental context, and prey species often adjust their responses accordingly. We investigated such effects within an insect assemblage from a tropical Australian stream. These systems are frequently subjected to catastrophic floods, often suggested to reduce the importance of predation in streams, and invertebrate faunas are characterised by relatively broad environmental tolerances. Impacts of the hunting predator Australopelopia prionoptera (Diptera: Chironomidae) and an undescribed ambush predator from the Polycentropodidae (Trichoptera) on survival and development of two species of tubicolous Chironomidae, Echinocladius martini (Orthocladiinae) and Polypedilum australotropicus (Chironominae), were assessed in laboratory microcosms. A further experiment investigated how impacts of Australopelopia varied over a broad range of temperatures, exceeding that experienced annually by the studied populations. Neither predator impacted survivorship for E. martini, but the presence of the polycentropodid caused E. martini to spend longer as larvae and reduced adult longevity, and adult females were smaller-sized and had smaller oocytes. In contrast, both predators reduced survivorship of P. australotropicus, but only Australopelopia affected its development, causing reductions in pupal duration and oocyte size. The observed non-lethal impacts of predation reflect the threat each predator is known to pose to each prey species in situ. Impacts of predation varied little with temperature, reflecting the broad thermal tolerances of all study species. The predator-specific responses of the prey species imply that predation is a significant selective force in tropical Australian streams, although fluctuation in intensity of predation associated with flooding may limit its importance for community structure and prey diversity at larger scales. Our results indicate a more limited scope for environmental modification of predator–prey relationships in faunas characterised by broad physiological tolerances.     

The effects of spate-induced disturbance, predation and environmental complexity on macroinvertebrates in a tropical stream

• 1 The combined effects of fish predation, substrate complexity and flow on benthic macroinvertebrates inhabiting riffles was investigated in a Hong Kong stream. Predation was manipulated using fish inclusion/exclusion cages containing complex (= many refuges) or simple (= few refuges) substrates. Experiments were undertaken during the winter dry season, when disturbance due to flow events was minimal, and repeated during the summer monsoon, when the stream experienced spates of varying intensity and duration.
• 2 Predation by the fish, Oreonectes platycephalus, significantly reduced the abundance of macroinvertebrates, especially chironomids and mayflies. Because chironomids (mostly Chironominae) were the dominant cage colonists, there was also a reduction in total macroinvertebrate density.
• 3 Predator impacts were significantly lessened during the wet season, when macroinvertebrate densities increased considerably, but significant reductions in the densities of vulnerable taxa and total macroinvertebrate abundance were nevertheless apparent.
• 4 Substrate complexity (the presence of prey refuges) had no significant effect on the ability of predators to reduce prey abundance.
• 5 Detritus accumulated in cages during the latter part of the study, and densities of most taxa were correlated with detrital standing stocks.
• 6 The results of this experiment indicate that biotic interactions such as predation may be suppressed during periods of spate-induced disturbance, although they can still influence benthic communities significantly. However, the effects of predation are highly taxon specific and may vary among streams in response to changes in predator and prey species composition, or the severity and duration of spates.

     

Invasive pike establishment in Cook Inlet Basin lakes,Alaska: diet,native fish abundance and lake environment

Northern pike (Esox lucius) were introduced to the northern Susitna Basin of south-central Alaska in the 1950’s, and have since spread throughout the upper Cook Inlet Basin. Extirpations of several native fish populations have been documented in this area. It is hypothesized here that invasive pike remodel the ecology of lakes by removing vulnerable prey types and that these changes are reflected in the diet of invasive pike. Trends in pike diet suggest that pike switch to less desirable but more abundant macroinvertebrate prey as preferred fish prey are eliminated. The impacts of pike introduction were studied in detail for one species of resident fish, the threespine stickleback (Gasterosteus aculeatus). Stickleback abundance decreases as pike invasion progresses. Conductivity is a significant environmental predictor of stickleback abundance, with higher conductivity apparently mitigating population reduction. Higher conductivity water may lessen the physiological costs of developing more robust armor, which reduces vulnerability to predation. Maximum lake depth also appears to predict stickleback abundance, though this trend was only marginally significant. Deeper lakes may provide an open-water refuge from pike predation by allowing stickleback to exist outside of the pike inhabited littoral zone. These findings indicate the importance of diverse habitat types and certain chemical and physical characteristics to the outcome of invasion by fish predators.     

A review of predation as a limiting factor for bird populations in mesopredator‐rich landscapes: a case study of the UK

The impact of increasing vertebrate predator numbers on bird populations is widely debated among the general public, game managers and conservationists across Europe. However, there are few systematic reviews of whether predation limits the population sizes of European bird species. Views on the impacts of predation are particularly polarised in the UK, probably because the UK has a globally exceptional culture of intensive, high‐yield gamebird management where predator removal is the norm. In addition, most apex predators have been exterminated or much depleted in numbers, contributing to a widely held perception that the UK has high numbers of mesopredators. This has resulted in many high‐quality studies of mesopredator impacts over several decades. Here we present results from a systematic review of predator trends and abundance, and assess whether predation limits the population sizes of 90 bird species in the UK. Our results confirm that the generalist predators Red Fox (Vulpes vulpes) and Crows (Corvus corone and C. cornix) occur at high densities in the UK compared with other European countries. In addition, some avian and mammalian predators have increased numerically in the UK during recent decades. Despite these high and increasing densities of predators, we found little evidence that predation limits populations of pigeons, woodpeckers and passerines, whereas evidence suggests that ground‐nesting seabirds, waders and gamebirds can be limited by predation. Using life‐history characteristics of prey species, we found that mainly long‐lived species with high adult survival and late onset of breeding were limited by predation. Single‐brooded species were also more likely to be limited by predation than multi‐brooded species. Predators that depredate prey species during all life stages (i.e. from nest to adult stages) limited prey numbers more than predators that depredated only specific life stages (e.g. solely during the nest phase). The Red Fox and non‐native mammals (e.g. the American Mink Neovison vison) were frequently identified as numerically limiting their prey species. Our review has identified predator–prey interactions that are particularly likely to result in population declines of prey species. In the short term, traditional predator‐management techniques (e.g. lethal control or fencing to reduce predation by a small number of predator species) could be used to protect these vulnerable species. However, as these techniques are costly and time‐consuming, we advocate that future research should identify land‐use practices and landscape configurations that would reduce predator numbers and predation rates.     

Decomposing Predation: Testing for Parameters that Correlate with Predatory Performance by a Social Bacterium

Predator–prey interactions presumably play major roles in shaping the composition and dynamics of microbial communities. However, little is understood about the population biology of such interactions or how predation-related parameters vary or correlate across prey environments. Myxococcus xanthus is a motile soil bacterium that feeds on a broad range of other soil microbes that vary greatly in the degree to which they support M. xanthus growth. In order to decompose predator–prey interactions at the population level, we quantified five predation-related parameters during M. xanthus growth on nine phylogenetically diverse bacterial prey species. The horizontal expansion rate of swarming predator colonies fueled by prey lawns served as our measure of overall predatory performance, as it incorporates both the searching (motility) and handling (killing and consumption of prey) components of predation. Four other parameters—predator population growth rate, maximum predator yield, maximum prey kill, and overall rate of prey death—were measured from homogeneously mixed predator–prey lawns from which predator populations were not allowed to expand horizontally by swarming motility. All prey species fueled predator population growth. For some prey, predator-specific prey death was detected contemporaneously with predator population growth, whereas killing of other prey species was detected only after cessation of predator growth. All four of the alternative parameters were found to correlate significantly with predator swarm expansion rate to varying degrees, suggesting causal interrelationships among these diverse predation measures. More broadly, our results highlight the importance of examining multiple parameters for thoroughly understanding the population biology of microbial predation.     

Performance as a fish predator of largemouth bass [<Emphasis Type="Italic">Micropterus salmoides</Emphasis> (Lacepède)] invading Japanese freshwaters: a review

This review focuses on how predator performance of the invasive largemouth bass [Micropterus salmoides (Lacepède)] has been, or will be, formed in Japanese freshwaters. Predation impacts of largemouth bass on fish communities appear pervasive in both Japanese as well as North American freshwaters. Factors affecting performance as a piscivorous predator are (1) light intensity and water clarity, (2) oxygen depletion, (3) prey size and gape size, (4) behavioral refuge of prey, (5) weed beds as refuge for prey fish, (6) interaction with bluegill. Size and behavioral refuges requirements are so rigorous that they may have evolved only in some North American prey fish species like bluegill; therefore, most Japanese native fish species are unlikely to be equipped with such refuges. However, refuge habitats like aquatic weed beds could develop in Japanese freshwaters, allowing prey fish species to survive under predation pressure. The density, architecture, and species composition of aquatic plants may affect their suitability as refuges. Studies in Japanese waters have suggested that the presence of rich aquatic vegetation or invasive bluegill in bass-introduced waters have suppressed the predation impact of largemouth bass on fish communities. In addition to these environmental factors, original genotypic and phenotypic traits of the introduced largemouth bass, and hybridization between different lineages of largemouth bass or with Florida bass [Micropterus floridanus (Lesueur)] may be involved in further adaptation of invasive largemouth bass to Japanese freshwaters.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Identifying inter‐ and intra‐guild feeding activity of an arthropod predator assemblage

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Is coarse woody debris in lakes a refuge or a trap for benthic invertebrates exposed to fish predation?

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Long-term responses of zooplankton to invasion by a planktivorous fish in a subarctic watercourse

1. Introduced or invading predators may have strong impacts on prey populations of the recipient community mediated by direct and indirect interactions. The long-term progression of predation effects, covering the invasion and establishment phase of alien predators, however, has rarely been documented.
2. This paper documents the impact of an invasive, specialized planktivorous fish on its prey in a subarctic watercourse. Potential predation effects on the crustacean plankton, at the community, population and individual levels, were explored in a long-term study following the invasion by vendace ( Coregonus albula ).
3. Over the 12-year period, the density and species richness of zooplankton decreased, smaller species became more abundant and Daphnia longispina , one of the largest cladocerans, was eliminated from the zooplankton community.
4. Within the dominant cladocerans, including Daphnia spp., Bosmina longispina and Bosmina longirostris , the body size of ovigerous females and the size at first reproduction decreased after the arrival of the new predator. The clutch sizes of Daphnia spp. and B. longirostris also increased.
5. Increased predation pressure following the vendace invasion induced many effects on the crustacean zooplankton, and we document comprehensive and strong direct and indirect long-term impacts of an introduced non-native predator on the native prey community.     

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.     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.     

Is prey predation risk influenced more by increasing predator density or predator species richness in stream enclosures?

The direct lethal impacts and the indirect effects predators have on prey characteristics, such as behavior, have fitness consequences for the prey. Whether the level of predation risk that prey face in the presence of multiple predator species can be predicted from a null model that sums the risk from each predator species in isolation is unclear. In field enclosures, we tested whether the predation risk experienced by Stenonema mayfly larvae from a dragonfly larva (Boyeria vinosa) and a hellgrammite (Corydalus cornutus) together matched the predictions of the multiplicative risk model. We then compared whether any deviations from the models predictions were larger in the presence of two predator species than in the presence of an equivalent density of individuals from either predator species alone, to determine if unique effects arise for the prey in the presence of multiple predator species. We also determined if prey moved preferentially into predator-free refuge spaces or decreased their movement in the presence of predators. Stenonemas risk of predation was reduced compared to the models prediction, but no unique multiple predator species effects were present because this risk reduction was comparable in magnitude to the level exhibited in the presence of each predator species alone. The prey did not move into predator-free refuge spaces in the presence of predators in the field enclosures. Thus, these predators appear to interfere interspecifically and intraspecifically, which may facilitate the coexistence of the predators and the prey.     

Can non-native smallmouth bass, <Emphasis Type="Italic">Micropterus dolomieu</Emphasis>, be swamped by hatchery fish releases to increase juvenile Chinook salmon, <Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>, survival?

One of the strategies that can be used to reduce predation impacts to valued fish species is by swamping predators with more prey than they can eat. We examined whether this approach was viable by calculating the maximum bioenergetic consumption potential of non-native smallmouth bass Micropterus dolomieu on fall Chinook salmon Oncorhynchus tshawytscha juveniles in the Yakima River throughout the spring between 1998 and 2002 and comparing those estimates to previously published estimates of fall Chinook salmon consumption. We found that the smallmouth bass population consumed fall Chinook salmon well below their bioenergetic potential. However, individual smallmouth bass that were piscivorous were eating other food items at a level near satiation. Furthermore, the maximum consumption potential was relatively low prior to mid-April, and then increased substantially to a peak in May. Predation mortality to hatchery fall Chinook salmon could be reduced within a year by releasing hatchery fall Chinook salmon that will emigrate quickly prior to mid-April, when predation potential is still very low. However, attempting to swamp predators with hatchery Chinook salmon to benefit naturally produced Chinook salmon poses uncertain benefits to natural origin fish and likely unacceptable costs to hatchery fish. Considerable swamping is occurring by other naturally produced fish species in the Yakima River such as dace Rhinichthys spp., mountain whitefish Prosopium williamsoni, and crayfish Pacificastus spp. Therefore, it is important to consider impacts to these non-target species because they could have indirect predation impacts on Chinook salmon.