Comparison of the Hunting Behavior of Four Piscine Predators Attacking Schooling Prey

Four piscine predator species were observed repeatedly attacking large (> 100,000) schools of flat-iron herring, Harengula thrissina. The predators could be categorized into two groups. Stalking predators (two species) were slow-moving, predominantly solitary hunters attacking from positions beneath the school. Attacks were directed at individual prey and the sequence of events was orient-approach-strike. Although the stalking species were seen most often and were responsible for the majority of the attacks, capture success was low. The remaining two species were fast-moving, pelagic hunters regularly occurring in groups of up to 8 individuals. These predators were extremely proficient at capturing prey, either by orienting on individuals (stragglers) or accelerating into the school and ramming their prey (impact attacks). Group size was positively associated with capture success, but not significantly so. Because stalking predators orient towards individual prey, they may suffer from the effect of confusion when attacking schooling prey. Use of the impact strategy, by comparison, may allow predators to overcome the confusion effect either by attacking prey already separated from the school, or by orienting towards aggregated prey rather than particular individuals.     

Effects of cadmium on foraging behaviour of lake charr, Salvelinus namaycush

Synopsis Foraging behaviour of lake charr, Salvelinus namaycush, exposed to 0.5 and 5.0 g l^–1 cadmium concentrations for 106–112 d, was examined in a flow-through stream channel using fathead minnows, Pimephales promelas, as prey. Five modes of foraging behaviour were observed and quantified (ending at fixate-orient; ending at approach and/or chase; ending at failed attack; ending at capture and loss; successful consumption). Each mode represented an increasingly progressive end-point from initial sighting of the prey to final consumption. Compared to controls., 0.5 g l^–1 Cd-exposed predators captured significantly fewer prey, while 5 g l^–1 Cd-exposed predators made fewer attacks. Both Cd-exposed groups consumed fewer prey than controls.     

Cross frontal variability in hydrological and biological structures observed in a river plume area (Rhône mouth, NW Mediterranean Sea)

The lake herring (Coregonus artedi) is an important coldwater planktivore in the Laurentian Great Lakes and in smaller inland lakes in portions of Canada and the northern United States. Lake herring cruise the pelagia and feed selectively in both gulping and particulate modes. They are visual predators in environments with adequate illumination. Visual predation by fish consists of a series of discrete steps. We studied the first step in the predation sequence, reaction to prey, at light intensities of 2–1500 Lx in a simulated pelagic environment at 10–13°C. We measured lake herring reactive distances, the distance at which a prey item will be detected and attacked, to live Limnocalanus macrurus, a natural prey of lake herring in Lake Superior. We used the reactive distances and associated angles of bearing and elevation, which described the location of the prey relative to the lake herring, to calculate reactive volume. This reactive volume can be envisioned as an irregular sphere surrounding the fish, within which prey are detected and attacked. All of the attacks on prey occurred in the anterior portions of the sagittal and lateral planes of the lake herring, as would be expected for a pelagic, cruising fish. The reactive volume surrounding the lake herring was generally spherical, but was more irregular than the simple spheres, hemispheres, cylinders, cones or other geometries assumed in previous studies. The reactive distances and the reactive volume changed with light intensity and were significantly smaller at 2–10 Lx than at 40–1500 Lx. At 40–1500 Lx, the reactive volume was expanded over that observed at 2–10 Lx laterally and caudally. Collectively our results indicate that lake herring can visually forage most effectively in environments with light levels >10 Lx.     

Predation rates of nemertean predators: the case of a rocky shore hoplonemertean feeding on amphipods

Estimates of the predation rates of benthic nemerteans are often based on observations of single individuals, and consequently they may not be representative for all members of a population of these predators. Herein we conducted controlled and repeatable laboratory experiments on the predation rate of the hoplonemertean Amphiporus nelsoni Sánchez 1973, which is common at exposed rocky shores along the central Chilean coast. During the austral fall (April, May 2000), nemerteans were observed in relatively high numbers crawling in the intertidal zone during early morning or late-afternoon low tides. When these nemerteans were offered living amphipods held by a forceps, they immediately attacked the amphipods and fed on them. In the laboratory experiments, nemerteans preferred the amphipod Hyale maroubrae Stebbing, 1899, which is also very common in the natural habitat of A. nelsoni. The nemerteans preyed to a higher extent on small males and non-ovigerous females than would have been expected from their abundance. We suggest that these (non-reproductive) stages of H. maroubrae are very mobile and therefore have a high likelihood of encounters with nemerteans. Predation rates reached maxima when nemerteans were provided prey densities of four or more of their preferred prey species, H. maroubrae, furthermore indicating that encounter rates with prey may affect predation rates. In long-term laboratory experiments, A. nelsoni consumed more amphipods during low tide conditions than during high tide conditions. Many nemerteans in the field prefer particular environmental conditions (e.g. nocturnal low tides), which restricts the time available for successful feeding. In the long-term experiment, predation rates of A. nelsoni never exceeded 0.5 amphipods nemertean^–1 d^–1. Maximum feeding events were 3 or 4 amphipods nemertean^–1 d^–1, but this only occurred during 10 out of a possible 2634 occasions. Nemerteans that had consumed 3 or 4 amphipods during 1 day, consumed substantially less prey during the following days. Towards the end of the long-term experiment, average predation rates decreased to 0.2 amphipods nemertean^–1 d^–1, corresponding to predation rates reported for other nemertean species (0.1–0.3 prey items nemertean^–1 d^–1). We suggest that predation rates from laboratory experiments represent maximum estimates that may not be directly transferable to field populations. Additionally, low predator–prey encounter rates with preferred prey in the field may further limit the predation impact of nemertean predators in natural habitats.     

Morphology and defensive structures in the predator-prey interaction: an experimental study of Parabroteas sarsi (Copepoda,Calanoida) with different cladoceran prey

Parabroteas sarsi is a predaceous calanoid copepod commonly found in South Andes lakes. Feeding experiments were carried out in order to estimate the predation rates and attack patterns on different cladoceran prey. Predation rates were related with prey sizes. The smallest prey, Bosmina longirostris, was ingested up to 5 prey pred^–1 h^–1 while the largest, Daphnia middendorffiana, only at 0.12 prey pred^–1 h^–1. The functional response of P. sarsi differed when confronted with different prey although in all cases, the number of kills increased with prey density. A saturation of ingestion rates at high prey densities was only observed for B. longirostris and Ceriodaphnia dubia juveniles. Remains seldom appeared at the end of the experiments, implying that the predator consumed prey totally. Yet, in all experiments carried out with Daphnia ambigua and D. middendorffiana, remains, including soft parts, were found. Direct observations of attack and of number and types of remains showed interference of handling by tail, helmet, and size of the prey.     

The spatial patchiness of Pacific herring larvae

Synopsis The spatial patchiness at age was measured for Pacific herring,Clupea harengus pallasi, larvae of a small coastal inlet. Lloyd's index of patchiness decreased from 3.8 at hatching to a minimum of 1.5 at 25–30 d and then increased to 2.5 at 50 d. This two-cycle pattern resembles that measured by others for the larvae of other species of schooling pelagic fishes. It differs by being several times lower in magnitude. This is attributed to two factors: (1) a rapid reduction in patchiness from a relatively high level in the demersal egg interval to a lower level in the planktonic larval interval as larvae are released into the water column over a period of 2–4 days, and (2) foraging by larval herring for prey that are larger, more dilute, and less patchy than the prey of other, smaller pelagic fish larvae.     

Prey susceptibilities,prey utilization and variable attack efficiencies of Ural owls

Summary To investigate the factors that influence prey utilization among predators with active prey, three series of experiments were performed in which Ural owls (Strix uralensis) searched for and attacked three prey species of wild mice, Microtus montebelli, Apodemus speciosus, and A. argenteus, in a large flight cage. Over the whole study, owls attacked mice about ten times a night. The number of attacks on each prey species did not differ from that predicted by a random attack model. M. montebelli was taken more than either Apodemus species. Prey utilization appears to be influenced by prey susceptibilities only and it is unlikely that prey selection by the owls affected prey utilization patterns. Under the experimental conditions, random attack is predicted by optimal foraging theory. However, random attack may be explained just as well by the inability of the owl to discriminate prey type. The owls, energy gain was adjusted not by alteration in the number of attacks on a prey species but rather by alteration in the capture success between experiments. Capture success increased in poor food conditions for the same prey species. This flexibility in capture success has not been considered in the assumptions of optimal foraging theory. In conventional optimal foraging theory, the probability of capture success is implicitly assumed as constant and unity. We suggest that this assumption is inadequate to understand the foraging behavior of owls.     

Effect of natural zooplankton feeding on the tissue and skeletal growth of the scleractinian coral<Emphasis Type="Italic"> Stylophora pistillata</Emphasis>

Laboratory experiments were designed to estimate the ingestion rates of the scleractinian coral Stylophora pistillata under varying prey concentrations and feeding regimes and to assess the effect of feeding on the tissue and skeletal growth. Six sets of corals were incubated under two light (80 and 300 µmol photons m^–2 s^–1) and three feeding levels (none, fed twice, and fed six times per week) using freshly collected zooplankton. Results showed that the number of prey ingested was proportional to prey density, and no saturation of feeding capability was reached. Capture rates varied between 0.5 and 8 prey items 200 polyp^–1 h^–1. Corals starved for several days ingested more plankton than did fed corals. Fed colonies exhibited significantly higher levels of protein, chlorophyll a, and chlorophyll c₂ per unit surface area than starved colonies. Feeding had a strong effect on tissue growth, increasing it by two to eight times. Calcification rates were also 30% higher in fed than in starved corals. Even moderate levels of feeding enhanced both tissue and skeletal growth, although the processes involved in this enhancement remain to be determined.     

Stable isotopes show food web changes after invasion by the predatory cladoceran <Emphasis Type="Italic">Cercopagis pengoi</Emphasis> in a Baltic Sea bay

Cercopagis pengoi, a recent invader to the Baltic Sea and the Laurentian Great Lakes, is a potential competitor with fish for zooplankton prey. We used stable C and N isotope ratios to elucidate trophic relationships between C. pengoi, zooplankton (microzooplankton, 90–200 m, mostly copepod nauplii and rotifers; mesozooplankton, >200 m, mostly copepods), and zooplanktivorous fish (herring, size range 5–15 cm and sprat, 9–11 cm) in a coastal area of the northern Baltic Sea. The isotope ratios in C. pengoi and fish were much higher than those of zooplankton, showing general trends of enrichment with trophic level. Young-of-the-year (YOY) herring had a significantly higher ¹⁵N/¹⁴N ratio than C. pengoi, suggesting of a trophic linkage between the two species. To evaluate the possible relative importance of different food sources for C. pengoi and YOY herring, two-source isotope-mixing models for N were used, with micro- and mesozooplankton as prey for C. pengoi and mesozooplankton and C. pengoi as prey for YOY herring. These models indicate that mesozooplankton was the major food source of both species. However, microzooplankton may be important prey for young stages of C. pengoi. Comparative analyses of the herring trophic position before and after the invasion by C. pengoi showed a trophic level shift from 2.6 to 3.4, indicating substantial alterations in the food web structure. Our findings contribute to a growing body of evidence, showing that C. pengoi can modify food webs and trophic interactions in invaded ecosystems.     

Predation on Ceriodaphnia cornuta and Brachionus calyciflorus by two Mesocyclops species coexisting in Barra Bonita reservoir (SP,Brazil)

Feeding experiments with two species of carnivorous copepod, Mesocyclops longisetus (Thiebaud) and Mesocyclops kieferi Van de Velde from Barra Bonita, a eutrophic reservoir in São Paulo, Brasil, were performed using two common types of prey: Ceriodaphnia cornuta, a cladoceran, with a mean body length of 464 µm (including spines) or 393 µm (without spines), and Brachionus calyciflorus, a rotifer with a mean body length of 350 µm (including spines) or 279 µm (without spines).Both species showed higher consumption rates on Brachionus than on Ceriodaphnia. For Mesocyclops longisetus, the average rates were: 2.19 prey ind^–1 h^–1 (Brachionus), and 1.30 prey ind^–1 h^–1 (Ceriodaphnia). For Mesocyclops kieferi, the rates were 1.85 prey ind^–1 h^–1 (Brachionus) and 0.60 prey ind^–1 h^–1 (Ceriodaphnia). These experimental data are discussed with reference to the dynamics of the predator and prey populations in the reservoir.Laboratorio de Limnologia, Departamento de Ciencias Biologicas, Universidade Federal de São CarlosCentro de Recursos Hidricos e Ecologia Aplicada Lab. de Limnologia, Departamento de Hidraulica e Saneamento, Escola de Engenharia de São Carlos, Universidade de São Paulo     

Effects of successive predator attacks on prey aggregations

We study the cumulative effect of successive predator attacks on the disturbance of a prey aggregation using a modelling approach. Our model intends to represent fish schools attacked by both aerial and underwater predators. This individual-based model uses long-distance attraction and short-distance repulsion between prey, which leads to prey aggregation and swarming in the absence of predators. When intermediate-distance alignment is added to the model, the prey aggregation displays a cohesive displacement, i.e., schooling, instead of swarming. Including predators, i.e. with repulsion behaviour for prey to predators in the model, leads to flash expansion of the prey aggregation after a predator attack. When several predators attack successively, the prey aggregation dynamics is a succession of expanding-grouping-swarming/schooling phases. We quantify this dynamics by recording the changes in the simulated prey aggregation radius over time. This radius is computed as the longest distance of individual prey to the aggregation centroid, and it is assumed to increase along with prey disturbance. The prey aggregation radius generally increases during flash expansion, then decreases during grouping until reaching a constant lowest level during swarming/schooling. This general dynamics is modulated by several parameters: the frequency, direction (vertical vs. horizontal) and target (centroid of the prey aggregation vs. random prey) of predator attacks; the distance at which prey detect predators; the number of prey and predators. Our results suggest that both aerial and underwater predators are more efficient at disturbing fish schools by increasing their attack frequency at such level that the fish cannot return to swarming/schooling. We find that a mix between aerial and underwater predators is more efficient at disturbing a fish school than a single type of attack, suggesting that aerial and underwater foragers may gain mutual benefits in forming foraging groups.     

Age and growth of Pacific herring larvae based on length-frequency analysis and otolith ring number

Synopsis Age and growth in length and dry weight of cohorts of wild Pacific herring larvae, Clupea harengus pallasi, were measured using successive modes in the length-frequencies of the catches and the number of rings in the otoliths. Average linear rates of growth in length ranged from 0.36–0.41 mm d^–1, and average exponential rates of growth in dry weight ranged from 0.063–0.084 d^–1. Length-date and dry weight-age curves were best described using one-cycle and two-cycle Gompertz functions, respectively. Weight-length relationship were, therefore, curvilinear on double logarithmic plots and were best described by a non-linear allometric function. Average rates of otolith ring deposition were 0.90, 1.09 and 0.73–1.26d^–1. Rings were deposited daily from the day of complete yolk absorption in the first two cohorts, but interrupted ring deposition was observed over the first 27 d of the third cohort. Relatively low water temperatures, <9° C, coincided with the interruption and may have caused it.     

The oddity effect drives prey choice but not necessarily attack time

The tendency of predators to preferentially attack phenotypically odd prey in groups (the oddity effect) is a clear example of how predator cognition can impact behaviour and morphology in prey. Through targeting phenotypically odd prey, predators are thought to avoid the cognitive constraints that delay and limit the success of attacks on homogenous prey groups (the confusion effect). In addition to influencing which prey a predator will attack, the confusion and oddity effects would also predict that attacks on odd prey can occur more rapidly than attacking the majority prey type, as odd prey are more easily targeted, but this prediction has yet to be tested. Here, we used kerri tetra fish, Inpaichthys kerri, presented with mixed phenotypic groups of Daphnia dyed red or black to investigate whether odd prey in groups are preferentially attacked and whether these attacks were faster than those on the majority prey type. In agreement with previous work, odd prey were targeted and attacked more often than expected from their frequency in the prey groups, regardless of whether the odd prey was red in a group of black prey or vice versa. However, no difference was found in the time taken to attack odd vs. majority prey items, contrary to our predictions. Our results suggest that the time taken to make an attack is determined by a wider range of factors or is subject to greater variance than the choice of which prey is selectively targeted in a group.     

Density-dependent effects of prey defences

In this study, we show that the protective advantage of a defence depends on prey density. For our investigations, we used the predator-prey model system Chaoborus-Daphnia pulex. The prey, D. pulex, forms neckteeth as an inducible defence against chaoborid predators. This morphological response effectively reduces predator attack efficiency, i.e. number of successful attacks divided by total number of attacks. We found that neckteeth-defended prey suffered a distinctly lower predation rate (prey uptake per unit time) at low prey densities. The advantage of this defence decreased with increasing prey density. We expect this pattern to be general when a defence reduces predator success rate, i.e. when a defence reduces encounter rate, probability of detection, probability of attack, or efficiency of attack. In addition, we experimentally simulated the effects of defences which increase predator digestion time by using different sizes of Daphnia with equal vulnerabilities. This type of defence had opposite density-dependent effects: here, the relative advantage of defended prey increased with prey density. We expect this pattern to be general for defences which increase predator handling time, i.e. defences which increase attacking time, eating time, or digestion time. Many defences will have effects on both predator success rate and handling time. For these defences, the predator’s functional response should be decreased over the whole range of prey densities. Received: 15 September 1999 / Accepted: 23 December 1999     

Predation by rock bass on other stream fishes: experimental effects of depth and cover

Synopsis Effects of water depth and cover availability on predation rates by adult rock bass, Ambloplites rupestris, on juvenile central stoneroller, Campostoma anomalum, pumpkinseed, Lepomis gibbosa, and fantail darter, Etheostoma flabellare, were measured in a laboratory stream. A predation rate experiment and a prey activity experiment were conducted. In the first experiment, each prey type shared experimental chambers with predatory rock bass under all combinations of two depths (shallow and deep) and two cover levels (absent and present). Predation rates after 72 h were greater in deep (35–38 cm) than in shallow (7–10 cm) water for fantail darter and two sizes of central stoneroller, but not for pumpkinseed. Presence of cover (opaque plastic tubes) on the stream bottom had no direct effect on predation rates, which were higher for pumpkinseed and small stoneroller (35–64 mm total length) than for fantail darter and large stoneroller (70–89 mm total length). In the second experiment, diel patterns of small stoneroller activity were monitored under all combinations of two water depths and three levels of predation risk. Small stonerollers were more active during the day and in the absence of rock bass, but were not affected directly by water depth. My results suggest that effects of habitat features (e.g., depth, cover) on predator-prey interactions vary according to the natural history and behavior of particular prey and predators. Future research should integrate habitat-specific responses of prey to predation risk into models that predict the distribution of prey among available habitats.The unit is jointly supported by the U.S. Fish and Wildlife Service, the Virginia Department of Game and Inland Fisheries, the Wildlife Management Institute, and Virginia Polutechnic Institute and State University.     

Status of the Nemertea as predators in marine ecosystems

The ecology of nemertean predators in marine ecosystems is reviewed. Nemerteans occur in most marine environments although usually in low abundances. Some species, particularly in intertidal habitats, may reach locally high densities. During specific time periods appropriate for hunting, nemerteans roam about in search of prey. Upon receiving a stimulus (usually chemical cues), many nemertean species actively pursue their prey and follow them into their dwellings or in their tracks. Other species (many hoplonemerteans) adopt a sit-and-wait strategy, awaiting prey items in strategic locations. Nemerteans possess potent neurotoxins, killing even highly mobile prey species within a few seconds and within the activity range of its attacker. Most nemertean species prey on live marine invertebrates, but some also gather on recently dead organisms to feed on them. Heteronemerteans preferentially feed on polychaetes, while most hoplonemerteans prey on small crustaceans. The species examined to date show strong preferences for selected prey species, but will attack a variety of alternative prey organisms when deprived of their favourite species. Ontogenetic changes in prey selection appear to occur, but no further information about, e.g. size selection, is available. Feeding rates as revealed from short-term laboratory experiments range on the order of 1–5 prey items d^–1. These values apparently are overestimates, since long-term experiments report substantially lower values (0.05–0.3 prey items d^–1). Nemerteans have been reported to exert a strong impact on the population size of their prey organisms through their predation activity. Considering low predation rates, these effects may primarily be a result of indirect and additive interactions. We propose future investigations on these interactive effects in combination with other predators. Another main avenue of nemertean ecological research appears to be the examination of their role in highly structured habitats such as intertidal rocky shore and coral reef environments.     

The guild structure of a community of predatory vertebrates in central Chile

Summary The trophic ecology of eleven predator species (Falconiforms: Buteo polyosoma, Elanus leucurus, Falco sparverius, Geranoaetus melanoleucus, Parabuteo unicinctus; Strigiforms: Athene cunicularia, Bubo virginianus, Tyto alba; Carnivores: Dusicyon culpaeus; Snakes: Philodryas chamissonis, Tachymenis peruviana) in two nearby localities of central Chile is analyzed. The localities exhibit the typical climate (hot-dry summers, coldrainy winters), and vegetation (chaparral), of mediterranean ecosystems. Densities of the staple prey (small mammals) were estimated by seasonal trapping during two years in both open and dense patches of chaparral.The trophic parameters examined are: 1) proportion of diurnal, crepuscular, or nocturnal prey found in the predators' diet; 2) relationship between abundance of different mammalian prey in the predators' diet, and in both open and densely vegetated habitat patches; 3) mean weight and variance of weight of small mammal prey consumed; 4) average weight of the predators; 5) food-niche breadth of the predators; 6) relationship between average weight of predators and mean weight of mammalian prey taken, its variance, and food-niche breadth; 7) overlap in food-niche between all the predator species; 8) guild packing of the predators. Parameters 1) and 2) are used to assess the importance of temporal and habitat segregation of the predators, respectively; parameters 3), 4), 5), and 6) provide information on the possibilities of partitioning the prey resources among the predators; parameters 1), 2), 7) and 8) are used to investigate the organization of the community in terms of guilds.Three niche dimensions seem to be important in determining the structure of the predator community: 1) hunting activity period (diurno-crepuscular, nocturno-crepuscular), 2) hunting habitat (open, or both open and dense patches), and 3) mean prey size taken. Segregation along these three axes results in generally low food niche overlaps (<54% in 47 of the 55 pairwise comparisons) among the predators, but it is not possible to determine whether this was produced by competitive interactions or stochastic differences. Three guilds (niche overlap >90% in pair-wise comparisons) can be recognized: a) the carnivorous-insectivorous guild formed by the diurnal raptors A. cunicularia and F. sparverius, which tend to hunt in open habitat patches; b) the herpetophagous guild formed by the diurnal snakes P. chamissonis and T. peruviana, which presumably hunt in open habitat patches; c) the carnivorous guild (highly specialized in the capture of two rodent species) formed by the diurnal raptors B. polyosoma, G. melanoleucus, P. unicinctus, and the carnivore D. culpaeus, which hunt in open habitat patches. The diurnal raptor E. leucurus is not clearly associated with any guild, and the only two nocturnal raptors in the community (B. virginianus and T. alba) exhibit marked differences in their trophic ecology.     

The temporal selfish herd: predation risk while aggregations form

The hypothesis of the selfish herd has been highly influential to our understanding of animal aggregation. Various movement strategies have been proposed by which individuals might aggregate to form a selfish herd as a defence against predation, but although the spatial benefits of these strategies have been extensively studied, little attention has been paid to the importance of predator attacks that occur while the aggregation is forming. We investigate the success of mutant aggregation strategies invading populations of individuals using alternative strategies and find that the invasion dynamics depend critically on the time scale of movement. If predation occurs early in the movement sequence, simpler strategies are likely to prevail. If predators attack later, more complex strategies invade. If there is variation in the timing of predator attacks (through variation within or between individual predators), we hypothesize that groups will consist of a mixture of strategies, dependent upon the distribution of predator attack times. Thus, behavioural diversity can evolve and be maintained in populations of animals experiencing a diverse range of predators differing solely in their attack behaviour. This has implications for our understanding of predator–prey dynamics, as the timing of predator attacks will exert selection pressure on prey behavioural responses, to which predators must respond.     

Broken pieces: can variable ecological interactions be deduced from the remains of crab attacks on bivalve shells?

Shell fragmentation patterns that result from attacks by durophagous predators on hard‐shelled marine invertebrates are a rich source of indirect evidence that have proved useful in interpreting predation pressure in the fossil record and recent ecology. The behaviour and effectiveness of predators are known to be variable with respect to prey size. It is less well understood if variable predator–prey interactions are reflected in shell fragmentation patterns. Therefore, we conducted experimental trials to test the behavioural response of a living crab, Carcinus maenas, during successful predatory attacks on the blue mussel Mytilus edulis on two prey size categories. Further, we examined resultant shell fragments to determine whether specific attack behaviours by C. maenas could be successfully deduced from remaining mussel shells. In contrast to previous studies, we observed no significant differences in attack behaviour by the predators attributable to prey size. In most experimental predation events, crabs employed an ad hoc combination of five mechanisms of predation previously described for this species. We identified seven categories of shell breakage in predated mussels, but none of these were unambiguously correlated with specific attack behaviour. Combined attack behaviours may produce shell breakage patterns that have previously been assumed to be attributable to a single behaviour. While specific patterns of shell breakage are clearly attributable to durophagy, the results of this study provide important insights into the limitations of indirect evidence to interpret ecological interactions.     

Distribution, abundance, behavior and metabolism of Periphylla periphylla, a mesopelagic coronate medusa in a Norwegian fjord

The distribution, behavior and metabolism of the mesopelagic jellyfish, Periphylla periphylla (Péron & Lesueur), were investigated in Lurefjorden, Norway. Field studies, conducted in 1998–1999 with plankton nets and a remotely operated vehicle, indicated that 80-90% of the dense (up to 2.5 m^–3) population migrated 200–400 m vertically each day throughout the year. In situ observations with red light revealed that swimming rates and feeding activity varied with age and time of day. Detection of turbulence and contact with surfaces caused this medusa to conceal one or all of its tentacles in the stomach or to shed nematocyst-laden tissue from the tentacles. Stomachs of medusae collected with nets were often full of prey entangled with the sloughed tissue. Stomachs of medusae captured individually with ROV samplers were empty or contained only a few prey in their stomachs (typically, 1–4 copepods Calanus spp. or chaetognaths Eukrohnia hamata Möbius per medusa). Low rates (0.4–5.6 l O₂ mg C^–1 h^–1) of oxygen consumption of P. periphylla suggested that this species was sustained by relatively few (1–34) prey d^–1.