Selective predation on zooplankton by pelagic Arctic charr, Salvelinus alpinus, in six subarctic lakes

Selective predation by planktivore fish appears to be an important regulatory factor of zooplankton communities, potentially causing large changes in species composition and size distributions within populations. In this study, prey preferences and size-selective predation on zooplankton by Arctic charr were examined in six subarctic lakes with Arctic charr as the dominant pelagic fish species. Most of the lakes had a zooplankton community dominated by copepods (Cyclops scutifer and Eudiaptomus graciloides), but the pelagic charr evidently selected cladoceran species (Bythotrephes longimanus, Daphnia sp. and Bosmina sp.), likely because the copepods have a higher mobility and evasiveness than the cladocerans. Furthermore, a strong size selection was also revealed for both Bosmina sp. and Daphnia sp., as individual prey from Arctic charr stomachs were exclusively larger than individuals sampled in the environment. Additionally, visibility due to size, morphology and pigmentation (egg-carrying females) was also a major factor for the selection of zooplankton prey. In conclusion, Arctic charr was found to be highly selective on zooplankton both in respect to species composition and individual size of Bosmina sp. and Daphnia sp.     

Size-selective predation by the cleaner fish Labroides dimidiatus

A comparison of the size-frequency distribution of parasitic gnathiid isopod larvae in the diet of the cleaner fish Labroides dimidiatus and on six host fish species ( Chlorurus sordidus, Ctenochaetus striatus, Hemigymnus melapterus, Scolopsis bilineatus, Siganus doliatus, Thalassoma lunare ) was made on one occasion. The comparison was repeated with Hemigymnus melapterus on three occasions and between two islands in Australia. L. dimidiatus selected larger gnathiids at all times at Lizard Island but not at Heron Island. Size-selective predation by L. dimidiatus suggests any potential effect of cleaner fish on parasites may vary according to the size of parasite. However, this effect appears to vary spatially.     

Size-selective and sex-selective predation by brown bears on sockeye salmon

Breeding activity increases the vulnerability of many animals to predation, and such predation can affect the subset of animals successfully reproducing. To study the ways in which predation might affect the evolution of Pacific salmon, we measured the intensity and selectivity of predation by bears (primarily brown bears, Ursus arctos) on mature sockeye salmon (Oncorhynchus nerka) breeding in a series of small, spring-fed ponds and creeks near Pedro Bay, Alaska, from 1994 to 1998. Bears killed male salmon more often than females; males constituted 60% of the kills but only 35% of the salmon that died of senescence. The bears also killed fish that were larger, on average, than those dying of senescence (males: 462 vs 452 mm; females: 453 vs 443 mm). The level of predation varied greatly, from 4% (females) and 10% (males) in 1994 to 100% of both sexes in 1996 and 1997. The rate of predation also varied among habitats, being lower in larger ponds than in smaller, shallower ponds and the very small interconnecting creeks. Despite the intense and size-selective predation, the salmon in safer habitats (large ponds) were not larger than those in riskier habitats, and salmon densities were only slightly higher in the safer areas. Compared to a nearby population that experiences no bear predation (Woody Island), the male sockeye salmon from the Pedro Pond system had shallower bodies (i.e., less exposure in shallow water) for a given length, consistent with the hypothesis that selective predation can affect the extent of sexual dimorphism among populations. However, the average length at age for both males and females was greater in the Pedro Pond fish, indicating that selective factors besides predation affect length. Overall, the results indicate that bears can be an agent of natural selection within (and perhaps between) sockeye salmon populations, and predation can greatly affect reproductive success among individuals and years for the population as a whole. Received: 6 April 1999 / Accepted: 1 June 1999     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

Size-selective predation by all-male prawns: implications for sustainable biocontrol of snail invasions

All-male populations of the freshwater prawn Macrobrachium rosenbergii were recently produced by a novel temporal RNA interference (RNAi)-based biotechnology for aquaculture purposes. This biotechnology opens the way to the wide use of all-male prawn populations as sustainable biocontrol agents against invading populations of freshwater snails, for which there is currently no environmentally friendly solution. Among the most damaging of the invasive freshwater snail species are the apple snails (Pomacea spp.), which inflict major damage on natural ecosystems and rice fields. The proposed use of all-male prawn populations as environmentally friendly biocontrol agents against invasive freshwater snails has several advantages: efficient predation by the prawns over a wide range of freshwater snails, the ready availability of the prawns, and the monosex non-reproductive nature of the biocontrol agents. Since the aquatic predators are strongly size selective, we quantified the predation rate as a function of body size of both predator and prey (M. rosenbergii and P. caniculata). Medium-sized and large prawns (~10–30 g) efficiently preyed small and medium-sized snails (up to 15 mm), while small prawns (up to 4 g) immediately and completely eradicated snail hatchlings. Medium-sized prawns (~22 g) exterminated a significant fraction of snail biomass within 24 h (up to 58% of their body mass) after being introduced into a tank of snails. A typical ‘climbing-to-the surface’ anti-predator behavior of the snails was recorded. The potential of all-male prawns as efficient biocontrol agents over hatchling and adult apple snails as part of an integrated pest management program is discussed. Our experiments set the stage for evaluating the ecological and economic implications of this generic solution for a wide variety of habitats.     

Top-down effects of crustacean zooplankton on pelagic microorganisms in a mesotrophic lake

A series of single-factor in situ experiments was conductedin a mesotrophic lake in Brandenburg, North Germany, to studythe predatory impact of Eudiaptomus graciloides (adults, copepodites,nauplii), cyclopoid copepods (adult Diacyclops bicuspidatus,Thermocyclops oithonoides) and daphnids (adult Daphnia hyalina,Daphnia cucullata) on the microbial community (bacteria, autotrophicpicoplankton, flagellates, ciliates). All zooplankton speciestested reduced the ciliate community significantly and ingestionrates were always higher for ciliates in the 20–55 µmsize category as compared to smaller ciliates (10–20 µm).Adult E.graciloides, which exhibited the highest predatory impacton ciliates, differed from cyclopoids and daphnids by theirability to decimate ciliates to very low abundances. Ingestionrates of ciliates by the crustacean zooplankton followed thesequence E.graciloides > daphnids = cyclopoids = copepodites.While top-down control was evident for ciliates, top-down effectsdown to the autotrophic picoplankton and flagellates were mostlyrestricted to Daphnia-dominated treatments. Top-down effectswere never strong enough to produce negative bacterial growthrates. For all zooplankton tested, clearance rates for ciliatesexceeded those for phytoplankton. Besides the potential of thecrustacean zooplankton to influence the structure of ciliatecommunities, ciliates may contribute to the energy demands ofcopepods and daphnids, especially when phytoplankton resourcesare limited.     

Top-down control in pelagic systems: a role for invertebrate predation

Limnologists have long recognized the importance of predation in freshwater communities. The majority of study of predator effects has involved vertebrate predators, with emphasis on planktivorous fish. Documented effects of planktivorous fish have been so dramatic that manipulations of their populations are seen by many as potential tools in lake management. However, the success of such manipulations is often less than desired due to the ubiquitous complexity of food webs and the pervasiveness of compensatory responses to food web manipulation. Recently, enormous effort has been applied to the Lake Kinneret pelagic food web in effort to reduced the abundance of the planktivorous Kinneret bleak Acanthobrama terraesanctae and thereby increase the biomass of herbivorous zooplankton in the hopes of increasing water clarity. We compared potential predation pressure on Lake Kinneret herbivorous zooplankton by bleak and the other major zooplankton predators in the lake, the cyclopoid copepods Mesocyclops ogunnus and Thermocyclops dybowskii. We found that, despite having much lower biomass, cyclopoid copepods accounted for a greater portion of the predation mortality on herbivorous zooplankton than bleak. Our results suggest that reductions in predation pressure by bleak will not yield subsequent increases in herbivorous zooplankton biomass. Rather, reductions in bleak predation pressure may allow for increases in cyclopoid copepod abundance and thereby a net increase in predation pressure on herbivorous zooplankton.     

Size-selective predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha): field stuides

Summary Studies of predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha) in a large, eutrophic lake showed that there was a clearly marked size threshold ( 160 mm SL) above which roach began to feed on mussels. Roach preying on various sizes of mussels selected them in proportions different from their abundance and accessibility in the habitat. The mean size of mussels ingested by roach of 220 mm and larger, which fed predominantly on Dreissena, closely followed the pattern expected for a constant ratio of mean prey size to mean predator mouth size = 0.59. To explain the size selection we applied an optimal foraging approach, based on the ability of different-sized fish to crush (cost) mussels of different sizes, and hence crushing resistance, and energy contents (benefit). We found that fish smaller than 160 mm, which showed no inclination to eat Dreissena, would only be able to take small mussels with a very high cost/benefit ratio. The real switch to Dreissena would be expected in fish of 230–240 mm that could take most of their prey from highly profitable, numerous, and easily accessible size classes while keeping the mean prey size at the optimal level relative to mean predator mouth size.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Diel vertical migration of a pelagic amphipod in the absence of fish predation

The pelagic amphipod, Hyalella montezuma, migrates vertically into the surface waters at twilight in Montezuma Well, Arizona, USA despite the absence of fish predators or thermal stratification. We suggest that a persistent, dense, neustonic algal assemblage may provide a food resource incentive for the twilight ascent.     

A second examination of predation on pelagic copepods by the brittle star Astrotoma agassizii

Euchaeta antarctica and Calanoides acutus, common pelagic copepodsof the Southern Ocean, dominate stomach contents of the brittlestar Astrotoma agassizii from the South Georgia shelf in November–December.This finding does not support our earlier hypothesis of restrictedfeeding by Astrotoma in austral fall on these copepods duringtheir seasonal migration.     

Epipelic and pelagic primary production in Alaskan Arctic lakes of varying depth

We compared on eight dates during the ice-free period physicochemical properties and rates of phytoplankton and epipelic primary production in six arctic lakes dominated by soft bottom substrate. Lakes were classified as shallow ( < 2.5 m), intermediate in depth (2.5 m <  < 4.5 m), and deep ( > 4.5 m), with each depth category represented by two lakes. Although shallow lakes circulated freely and intermediate and deep lakes stratified thermally for the entire summer, dissolved oxygen concentrations were always >70% of saturation values. Soluble reactive phosphorus and dissolved inorganic nitrogen (DIN = NO₃ ⁻–N + NH₄ ⁺–N) were consistently below the detection limit (0.05 μmol l⁻¹) in five lakes. However, one lake shallow lake (GTH 99) periodically showed elevated values of DIN (17 μmol l⁻¹), total-P (0.29 μmol l⁻¹), and total-N (33 μmol l⁻¹), suggesting wind-generated sediment resuspension. Due to increased nutrient availability or entrainment of microphytobenthos, GTH 99 showed the highest average volume-based values of phytoplankton chlorophyll a (chl a) and primary production, which for the six lakes ranged from 1.0 to 2.9 μg l⁻¹ and 0.7–3.8 μmol C l⁻¹ day⁻¹. Overall, however, increased resulted in increased area-based values of phytoplankton chl a and primary production, with mean values for the three lake classes ranging from 3.6 to 6.1 mg chl a m⁻² and 3.2–5.8 mmol C m⁻² day⁻¹. Average values of epipelic chl a ranged from 131 to 549 mg m⁻² for the three depth classes, but levels were not significantly different due to high spatial variability. However, average epipelic primary production was significantly higher in shallow lakes (12.2 mmol C m⁻² day⁻¹) than in intermediate and deep lakes (3.4 and 2.4 mmol C m⁻² day⁻¹). Total primary production (6.7–15.4 mmol C m⁻² day⁻¹) and percent contribution of the epipelon (31–66%) were inversely related to mean depth, such that values for both variables were significantly higher in shallow lakes than in intermediate or deep lakes. Handling editor: L. Naselli-Flores     

Behavioural responses of benthic and pelagic Arctic charr to substratum heterogeneity

Responses to environmental heterogeneity were studied in laboratory-reared offspring of two morphs of Arctic charr Salvelinus alpinus from Loch Rannoch, Scotland, one occupying the pelagic zone and feeding predominantly on zooplankton and the other being benthic in habit and feeding mainly on macroinvertebrates. When housed in groups in tanks with a black-and white striped base, benthic charr demonstrated a clear preference for dark areas, whereas pelagic fish positioned themselves at random with respect to substratum colour. In general, pelagic charr were much less aggressive than benthic charr. In pelagic fish, neither spacing nor aggression was affected by the visual heterogeneity of the substrate. In contrast, benthic charr swam closer together and fought more when housed over a uniform as opposed to a non-uniform substratum. The results are discussed in the context of habitat-specific visual requirements and of an interaction between visual complexity and territoriality previously described for Atlantic salmon Salmo salar .     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

The effects of selective fish predation on the horizontal distribution of pelagic Cladocera in a southern French reservoir

A four-year study showed a clear seasonal succession of species within the cladoceran community of the large, oligotrophic Sainte-Croix reservoir (S.E. France). Diaphanosoma brachyurum and Ceriodaphnia pulchella were strictly limited to the warm stratified period (July to October), whereas Bosmina longirostris and Bosmina coregoni were dominant during spring and autumn. Daphnia longispina was the only species to occur throughout the year with higher densities in spring.In spring and late autumn, the discharge of the inflowing river Verdon was high and the abundance of all species showed a gradient over the whole lake with lower densities close to the inflow. During the stratified period, water inflow was very low and species showed different patterns. Densities of the small form Ceriodaphnia pulchella were similar all along the long axis of the lake, whereas Daphnia densities were significantly higher near the dam. The distribution pattern of Diaphanosoma, an intermediate-sized species, showed similar trends to that of Daphnia. The only planktivorous fish in the pelagic zone, the bleak (Alburnus alburnus), fed mostly on large-bodied species (> 1.0 mm) and was more abundant close to the inflow current. A comparison between the length frequency distributions of cladocera upstream and downstream provided a clear demonstration of the effects of size-selective predation on prey populations. Finally, the interactions between spatial heterogeneity and long-term development of the zooplankton community and the indirect effects of predation are discussed.     

Structure and distribution of pelagic ostracods (Ostracoda: Myodocopa) in the Arctic Ocean

The study of the pelagic ostracod fauna of the Arctic Ocean based on materials collected by numerous Russian expeditions (1929–1993) and data from the literature showed the extreme poorness of the Arctic pelagic ostracod fauna, its mainly North Atlantic genesis and complete isolation from the Pacific fauna. Maximum ostracod abundance was observed in the epipelagic zone, and the greatest species diversity occurred in the relatively warm deep Atlantic layer throughout the year. To the north, east, and west of Franz Josef Land and Spitsbergen, the number of species and abundance indices of pelagic ostracods were decreased. In superficial water layers of the Central Arctic, maximum ostracod density and biomass were recorded in June and September. The best bioindicator of warm Atlantic water in the Arctic basin is Obtusoecia obtusata; and of cold polar water in the North Atlantic, Boroecia maxima.     

The influence of hunger and predation risk on group cohesion in a pelagic fish, walleye pollock Theragra chalcogramma

Variation in the intensity of schooling behavior in fishes suggests that the benefits of aggregation are balanced by certain costs. We examined the proximity of group members to each other in juvenile walleye pollock, Theragra chalcogramma, under different environmental conditions. Food availability, simulated by a gradient of six ration treatments, had a major influence on group cohesion, with increasing dispersion as food level decreased. Group cohesion also decreased at night relative to daytime levels. Small juveniles (x=53 mm TL) maintained on high rations were highly responsive to the potential threat of a predator, with groups becoming more cohesive and remaining so for up to an hour after the initial threat. A chronic threat (continual presence of predators) resulted in tighter group cohesion than an acute threat (single simulated attack). Small juveniles maintained on low rations were less responsive to predation threats and recovered quickly, supporting the hypothesis that hunger induces risk-taking behavior. Large juveniles (x=149 mm TL) did not change their degree of aggregation in response to either type of predation threat. An overall plasticity in the degree of cohesiveness among group members indicates that walleye pollock are capable of gradually modifying their schooling behavior according to the environmental context.     

Heterotrophic bacterial and viral dynamics in Arctic freshwaters: results from a field study and nutrient-temperature manipulation experiments

Heterotrophic bacterial and viral concentrations (range, 0.7 × 10⁴ to 206.2 × 10⁴ ml⁻¹ and 0.05 × 10⁶ to 128.9 × 10⁶ ml⁻¹, respectively) were determined in several Arctic freshwater environments, including lakes and glacial ecosystems (78.55°N, 11.56°E). Our bacteria and virus results mirrored trends seen in temperate lakes, with an average virus-to-bacteria ratio (VBR) of 13 (range, 7.3–25.2) and viral concentrations and DOC positively correlated with bacterial concentrations (R = 0.964, P < 0.01 and R = 0.813, P < 0.05, respectively). Lysogenic bacteria, determined by induction with Mitomycin C, were not detected in any of the investigated Arctic freshwater environments. Nutrient-addition experiments at in situ and at elevated temperatures were performed to elucidate the factors which influenced the bacterial growth and the virus–bacteria interactions in Arctic freshwaters. Our results suggest that multiple limiting factors interacted and constrained bacterial growth. Bacterial concentrations and doubling times increased at elevated temperatures and appeared to be co-stimulated by phosphorus and carbon. However, viral concentrations showed a lack of response to nutrient addition thus indicating an uncoupling between bacteria and viruses in the experiment.     

Size-selective foraging behaviour of blue crabs,Callinectes sapidus (Rathbun), when feeding on mobile prey: Active and passive components of predation

Size-selective predation by small (30-50 mm carapace width) and large (90-110 mm) Callinectes sapidus when feeding on three different size classes of Litopenaeus setiferus was investigated. Laboratory experiments using no-choice and multiple-choice designs were carried out. Active and passive components of predation were evaluated by comparing the probability of attacking (active) and capturing (passive) shrimp. Small C. sapidus consumed shrimp of all size classes in similar frequencies during multiple and no-choice experiments, and the probability of attacking and capturing shrimp of all size classes was similar. Large crabs consumed large shrimp with a significantly higher frequency during multiple than during no-choice experiments, and the probability of capturing large shrimp was significantly higher than medium and small ones. Results in the present study suggest that size-specific consumption rates of C. sapidus are mainly the result of a passive mechanism associated with the salience of shrimp to predation. However, an active component of foraging behaviour related to the decision to attack an alternative prey whilst consuming one that has already been captured, helps to explain the differences in shrimp consumption between multiple and no-choice experiments. The complex interaction between active, passive components and satiation is discussed.