Amphipod (Gammarus minus) responses to predators and predator impact on amphipod density

Recent theoretical work suggests that predator impact on local prey density will be the result of interactions between prey emigration responses to predators and predator consumption of prey. Whether prey increase or decrease their movement rates in response to predators will greatly influence the impact that predators have on prey density. In stream systems the type of predator, benthic versus water-column, is expected to influence whether prey increase or decrease their movement rates. Experiments were conducted to examine the response of amphipods (Gammarus minus) to benthic and water-column predators and to examine the interplay between amphipod response to predators and predator consumption of prey in determining prey density. Amphipods did not respond to nor were they consumed by the benthic predator. Thus, this predator had no impact on amphipod density. In contrast, amphipods did respond to two species of water-column predators (the predatory fish bluegills, Lepomis macrochirus, and striped shiners, Luxilus chrysocephalus) by decreasing their activity rates. This response led to similar positive effects on amphipod density at night by both species of predatory fish. However, striped shiners did not consume many amphipods, suggesting their impact on the whole amphipod “population” was zero. In contrast, bluegills consumed a significant number of amphipods, and thus had a negative impact on the amphipod “population”. These results lend support to theoretical work which suggests that prey behavioral responses to predators can mask the true impact that predators have on prey populations when experiments are conducted at small scales. Received: 21 March 1997 / Accepted: 15 December 1997     

Identifying gray whale (Eschrichtius robustus) foraging grounds along the Chukotka Peninsula, Russia, using satellite telemetry

The purpose of this study was to evaluate summer and fall residency and habitat selection by gray whales, Eschrichtius robustus, together with the biomass of benthic amphipod prey on the coastal feeding grounds along the Chukotka Peninsula. Thirteen gray whales were instrumented with satellite transmitters in September 2006 near the Chukotka Peninsula, Russia. Nine transmitters provided positions from whales for up to 81?days. The whales travelled within 5?km of the Chukotka coast for most of the period they were tracked with only occasional movements offshore. The average daily travel speeds were 23?km?day^?1 (range 9–53?km?day^?1). Four of the whales had daily average travel speeds <1?km?day^?1 suggesting strong fidelity to the study area. The area containing 95% of the locations for individual whales during biweekly periods was on average 13,027?km² (range 7,097–15,896?km²). More than 65% of all locations were in water <30?m, and between 45 and 70% of biweekly kernel home ranges were located in depths between 31 and 50?m. Benthic density of amphipods within the Bering Strait at depths <50?m was on average ~54?g wet wt m^?2 in 2006. It is likely that the abundant benthic biomass is more than sufficient forage to support the current gray whale population. The use of satellite telemetry in this study quantifies space use and movement patterns of gray whales along the Chukotka coast and identifies key feeding areas.     

A review of Killer Whale interactions with other marine mammals: predation to co-existence

Killer Whales are well-known as predators of other marine mammals, including the large Sperm and baleen whales. Members of all marine mammal families, except the river dolphins and manatees, have been recorded as prey of Killer Whales; attacks have been observed on 20 species of cetaceans, 14 species of pinnipeds, the Sea Otter, and the Dugong. Ecological interactions have not been systematically studied and further work may indicate that the Killer Whale is a more important predator for some populations than previously believed. Not all behavioural interactions between Killer Whales and other marine mammal species result in predation, however. Some involve 'harassment' by the Killer Whales, feeding by both species in the same area, porpoises playing around Killer Whales, both species apparently 'ignoring' each other, and even apparently unprovoked attacks on Killer Whales by sea lions. These non-predatory interactions are relatively common. We conclude that interactions between Killer Whales and marine mammals are complex, involving many different factors that we are just beginning to understand.     

DIETS OF BAIRD'S BEAKED WHALES, BERARDIUS BAIRDII, IN THE SOUTHERN SEA OF OKHOTSK AND OFF THE PACIFIC COAST OF HONSHU, JAPAN

Stomach contents were analyzed from 127 Baird's beaked whales, Berardius bairdii , taken in coastal waters of Japan. During late July-August of 1985–1987, 1989, and 1991, 107 samples were collected from off the Pacific coast of Honshu. An additional 20 samples were collected from whales taken in the southern Sea of Okhotsk during late August—September of 1988 and 1989. Prey identification using fish otoliths and cephalopod beaks revealed the whales fed primarily on deep-water gadiform fishes and cephalopods in both regions. Prey species diversity and the percentage of cephalopods and fish differed between the two regions. Off the Pacific coast of Honshu the whales fed primarily on benthopelagic fishes (81.8%) and only 18.0% on cephalopods. Eight species of fish representing two families, the codlings (Moridae) and the grenadiers (Macrouridae), collectively made up 81.3% of rhe rotal. Thirty species of cephalopods representing 14 families made up 12.7%. In the southern Sea of Okhotsk, cephalopods accounted for 87.1% of stomach contents. The families Gonatidae and Cranchiidae were the predominant cephalopod prey, accounting for 86.7% of the diet. Gadiform fish accounted for only 12.9% of the diet. Longfin codling, Laemonema longipes , was the dominant fish prey in both regions. Depth distribution of the two commonly consumed fish off the Pacific coast of Honshu indicate the whales in this region fed primarily at depths ranging from 800 to 1,200 m.     

Prey: predator ratio dependence in the functional response of a freshwater amphipod

1. First known for their shredding activity, freshwater amphipods also behave as active predators with consequences for prey population regulation and amphipod coexistence in the context of biological invasions. 2. A way to quantify predation is to determine the average consumption rate per predator, also known as its functional response (FR). 3. Although amphipods are gregarious and can display social interactions that can alter per capita consumption rates, previous studies using the FR approach to investigate amphipod predation ignored such potential mutual interference because they did not consider variations in predator density. 4. We investigated the FR of Echinogammarus berilloni feeding on dipteran larvae with joint variations in prey and predator densities. This bivariate experimental design allowed us to estimate interference and to compare the fits of the three main classes of theoretical FR models, in which the predation rate is a function of prey density alone (prey‐dependent models), of both prey and predator densities (predator‐dependent models) or of the prey‐to‐predator ratio (ratio‐dependent models). 5. The Arditi–Ginzburg ratio‐dependent FR model provided the best representation of the FR of E. berilloni, whose predation rate showed a decelerating rise to a horizontal asymptote as prey abundance increased. 6. Ratio dependence means that mutual interference between amphipods leads to prey sharing. Mutual interference is likely to vary between amphipod species, depending on their level of aggressiveness.     

GRAY WHALE (ESCHRICHTIUS ROBUSTUS) HABITAT UTILIZATION AND PREY SPECIES OFF VANCOUVER ISLAND, B. C.

Habitat utilization and prey species of Vancouver Island gray whales were investigated by (1) summarizing 26 yr of distribution and feeding data and (2) conducting intensive observations in Clayoquot Sound, Vancouver Island, from 1989 to 1996. Whale distribution and movements were monitored from March to November through systematic boat surveys and whale-watch sighting programs. Prey species were collected by suction hose and plankton net or determined through analysis of fecal samples. Gray whales utilized virtually all of the southern west coast of Vancouver Island over the 26-yr observation period. Distribution, prey species, and feeding behavior showed marked variability during any one season and between years. Some feeding areas were used on an annual basis, others with >10-yr intervals between use. Feeding occurred in shallow sand or mud bays, eel grass beds, kelp beds, in the open water column, and at the surface. Young whales appeared to utilize habitat and prey species differently than adults. Main prey species included herring eggs/larvae ( Clupea harengus pallasi ), crab larvae ( Cancer magister megalops, Pachycbeles spp. zoea), mysids ( Holmesimysis sculpta, Neomysis rayii, Acanthomysis spp.), amphipods ( Ampelisca spp., Atylus borealis ), and ghost shrimp ( Callianassa californiensis ). The definition and relative importance of specific feeding grounds and the study of human impacts on this population are complicated by its broad and variable use of habitat and prey species.     

Activity budget and diving behavior of gray whales (Eschrichtius robustus) in feeding grounds off coastal British Columbia

Behavior and diving patterns of summer resident gray whales ( Eschrichtius robustus ) foraging on mysids were studied in coastal bays along the north shore of Queen Charlotte Strait, British Columbia. In this region, gray whales were found to feed primarily on planktonic prey rather than on the benthos as in their primary feeding areas further north. During the summers of 1999 and 2000, whales spent most of their time actively feeding or searching for prey (77%), whereas only 15% of their time was spent traveling and 8% socializing. The majority of the dives were short; the mean dive duration was 2.24 min with approximately three respirations per surfacing and 15 s between blows. Whales dove frequently (26.7 h⁻¹), spending only 17% of their time at the surface with an overall blow rate of 1.14 respirations per minute. Activity states were characterized by significantly different diving and respiratory parameters; feeding whales dove more frequently, with shorter intervals between respirations, thus spending less time at the surface compared to when traveling or searching. This diving pattern differs from benthic-feeding whales and likely optimizes capture of the mobile mysid swarms in shallow waters.     

Patterns of gammaridean amphipod abundance and species composition associated with dominant subtidal macroalgae from the western Antarctic Peninsula

The communities of gammaridean amphipods associated with eight dominant macroalgal species were examined near Palmer Station, Western Antarctic Peninsula. A total of 78,415 individuals belonging to 32 amphipod taxa were identified with mean densities ranging up to 20 individuals/g algal wet wt. The most abundant amphipod taxon, Metaleptamphopus pectinatus, was found to associate predominately with the brown alga Desmarestia menziesii, while the second most common taxon, Jassa spp. occurred primarily on the red alga Gigartina skottsbergii. Non-metric multidimensional scaling analysis demonstrated that the population densities of each amphipod species and amphipod species composition were similar on the same algal species but dissimilar on different species of algae. Comparisons of amphipod communities associated with a given algal species but from different sampling sites indicated that although the structure of species-specific macroalgal-associated amphipod communities can vary across spatial scales of 3 km, 50% of the macroalgal species examined showed no significant inter-site differences in associated amphipod community structure. Spearman rank correlation analyses showed that higher abundances of amphipods occurred on the macroalgae with the highest number of branches. As many Antarctic amphipods are known consumers of macroalgae, their remarkable abundances are likely to play a significant role in mediating energy and nutrient transfer in nearshore Antarctic Peninsular macroalgal communities.     

Influence of natural amphipod (<Emphasis Type="Italic">Victoriopisa</Emphasis><Emphasis Type="Italic">australiensis</Emphasis>) (Chilton, 1923) population densities on benthic metabolism,nutrient fluxes,denitrification and DNRA in sub-tropical estuarine sediment

The influence of natural populations of the sub-surface deposit-feeding amphipod Victoriopisa australiensis on sediment biogeochemistry was assessed by randomly collecting 21 sediment cores in a zone of Coombabah Lake, southern Moreton Bay, Australia, where the benthic infauna was dominated by this species. Cores were incubated sequentially to determine sediment–water column fluxes of oxygen, dissolved inorganic carbon and inorganic N species, followed by incubations to determine rates of denitrification and dissimilatory nitrate reduction to ammonium (DNRA) using the isotope pairing technique. Finally, each core was sieved in order to determine the population and biomass of amphipods present. Whilst all measures of overall benthic metabolism (sediment oxygen demand, and effluxes of inorganic carbon and nitrogen) showed increased with amphipod density, with rates being stimulated 70–220% at the highest categorised density range of 2,500–3,500 ind m⁻², only the correlation with dissolved inorganic carbon was statistically significant. In contrast, there were no discernable trends between amphipod densities and any of the N-cycle processes with the slopes of all correlations being very close to zero. These results highlight the differences in mesocosm simulations of fauna effects, which primarily relate to shifts in rates of organic matter turnover, compared to natural sediments where fauna effects relate more to induced changes in rates of organic matter deposition. Therefore, while mesocosms represent a powerful tool to investigate the mechanisms by which fauna influences microbial metabolism in the sediment, only studies of natural sediments can determine to what extent these mechanisms function in situ. Handling editor: Pierluigi Viaroli     

Patterns of prey use by lesser scaup <Emphasis Type="Italic">Aythya affinis</Emphasis> (Aves) and diet overlap with fishes during spring migration

Recent decline in the lesser scaup Aythya affinis population has been linked to changes in wetland conditions along their spring migration routes. In particular, the use of amphipod prey by lesser scaup has declined in many regions of the upper Midwest U.S.A. and has been linked to expanded fisheries, although empirical data on diet overlap are lacking. To explore patterns of prey use by lesser scaup and diet overlap with fishes, we quantified diets of scaup and fishes during the 2003 and 2004 spring migration in eastern South Dakota, U.S.A. We compared diet overlap between lesser scaup and fishes collected from Twin Lakes, South Dakota—an important stopover location for spring-migrating scaup. Plant seeds occurred in >95% of lesser scaup diets (n = 118) and represented an appreciable amount of consumed biomass (>70%). Gastropods, amphipods, and chironomids were the most abundant invertebrate prey taxa and occurred in 29–34% of lesser scaup diets. Although relatively frequent, these taxa each contributed only 4–27% of the diet by weight. Percent dry mass of amphipods, a preferred prey by lesser scaup, was low (4%) indicating that amphipod availability may be reduced during spring migration. Analysis of fish diets showed that black bullhead Ameiurus melas and yellow perch Perca flavescens had the highest diet overlap with lesser scaup at 94% and 92%, respectively. Moreover, mean size of amphipods and chironomids found in fish diets were significantly larger than that consumed by lesser scaup. Our findings support the notion that amphipod use by spring-migrating lesser scaup has declined and that size-selective predation by fishes may influence prey availability for lesser scaup. Handling editor: K. Martens     

Partitioning mechanisms of Predator Interference in different Habitats

Prey are often consumed by multiple predator species. Predation rates on shared prey species measured in isolation often do not combine additively due to interference or facilitation among the predator species. Furthermore, the strength of predator interactions and resulting prey mortality may change with habitat type. We experimentally examined predation on amphipods in rock and algal habitats by two species of intertidal crabs, Hemigrapsus sanguineus (top predators) and Carcinus maenas (intermediate predators). Algae provided a safer habitat for amphipods when they were exposed to only a single predator species. When both predator species were present, mortality of amphipods was less than additive in both habitats. However, amphipod mortality was reduced more in rock than algal habitat because intermediate predators were less protected in rock habitat and were increasingly targeted by omnivorous top predators. We found that prey mortality in general was reduced by (1) altered foraging behavior of intermediate predators in the presence of top predators, (2) top predators switching to foraging on intermediate predators rather than shared prey, and (3) density reduction of intermediate predators. The relative importance of these three mechanisms was the same in both habitats; however, the magnitude of each was greater in rock habitat. Our study demonstrates that the strength of specific mechanisms of interference between top and intermediate predators can be quantified but cautions that these results may be habitat specific. An erratum to this article can be found at     

Seasonal changes of macrozooplankton and Benthic Boundary Layer macrofauna from the Bay of Saint-Brieuc (western English Channel)

The macrozooplankton and Benthic Boundary Layer (BBL) macrofauna over a coarse sand and pebble community in the Bay of Saint-Brieuc (western English Channel) were sampled with a WP2 zooplankton net and with a modified MACER-GIROQ suprabenthic sledge, respectively, from February 1994 to November 1995. One hundred and sixty-four species were collected in 44 suprabenthic sledge hauls and 19 taxa in 30 zooplankton net hauls. In the water column, appendicularians and cnidarians dominated, while, in the BBL, holoplanktonic amphipods, chaetognaths, amphipods and mysids dominated the fauna; among them Apherusa spp., Sagitta setosa Muller, Anchialina agilis (Sars), Sirella clausii Sars and Eusirus longipes Boeck were the dominant species. The density and biomass of the BBL macrozooplankton were lower than those of the macrozooplankton in the water column. The density and biomass of suprabenthos remained a low throughout the year. In the water column, density and biomass of macrozooplankton showed a maximum in spring and remained low from autumn to winter; conversely, in the BBL, the density and biomass of both macrozooplankton and suprabenthos were higher from summer to autumn. The change in abundance of both BBL and pelagic taxa was seasonal. Some species were primarily sampled in the water column (appendicularians, cladocerans and cnidarians), while others were preferentially found at the BBL (suprabenthic species, holoplanktonic amphipods and cephalopods). A third group was collected throughout the water column (chaetognaths and fish larvae).      

The composition and distribution of benthic amphipods (Crustacea: Amphipoda) in the southern Barents Sea

The composition and distribution of near-bottom amphipods in the southern part of the central Barents Sea were studied based on quantitative materials collected by expeditions of the Murmansk Marine Biological Institute in 1996–1997. A total of 144 amphipod species (on average 13 ± 1 species per station) were found; the average biomass was 0.49 ± 0.07 g/m², average abundance 106 ± 15 specimens/m². Boreal-Arctic (46%), Arctic (17%), and boreal (21%) species were predominant among the amphipod fauna. The present results were compared with the data for 1968–1970.     

Predicting Cetacean Habitats from Their Energetic Needs and the Distribution of Their Prey in Two Contrasted Tropical Regions

To date, most habitat models of cetaceans have relied on static and oceanographic covariates, and very few have related cetaceans directly to the distribution of their prey, as a result of the limited availability of prey data. By simulating the distribution of six functional micronekton groups between the surface and ≃1,000 m deep, the SEAPODYM model provides valuable insights into prey distributions. We used SEAPODYM outputs to investigate the habitat of three cetacean guilds with increasing energy requirements: sperm and beaked whales, Globicephalinae and Delphininae. We expected High Energy Requirements cetaceans to preferentially forage in habitats of high prey biomass and/or production, where they might easily meet their high energetic needs, and Low Energy Requirements cetaceans to forage in habitats of either high or low prey biomass and/or production. Cetacean sightings were collected from dedicated aerial surveys in the South West Indian Ocean (SWIO) and French Polynesia (FP). We examined cetacean densities in relation to simulated distributions of their potential prey using Generalised Additive Models and predicted their habitats in both regions. Results supported their known diving abilities, with Delphininae mostly related to prey present in the upper layers of the water column, and Globicephalinae and sperm and beaked whales also related to prey present in deeper layers. Explained deviances ranged from 9% for sperm and beaked whales in the SWIO to 47% for Globicephalinae in FP. Delphininae and Globicephalinae appeared to select areas where high prey biomass and/or production were available at shallow depths. In contrast, sperm and beaked whales showed less clear habitat selection. Using simulated prey distributions as predictors in cetacean habitat models is crucial to understand their strategies of habitat selection in the three dimensions of the ocean.     

Prey selection by an intertidal beetle: field test of an optimal diet model

Summary Adult beetles Thinopinus pictus LeConte (Staphlyinidae) live on sand beaches in temporary burrows from which they emerge at night to prey on amphipods Orchestoidea calforniana (Brandt). I constructed models of amphipod size selection by beetles, using the size distributions of amphipods measured on the beach, and the results of laboratory experiments on capture success, reaction distance and feeding rates. Capture success decreased and the probability that an amphipod was detected increased with increasing amphipod size. Beetles observed during beach searches selected larger sizes of amphipods than predicted from availability and vulnerability of different sizes. To apply an optimal foraging model, I estimated the profitability of different sizes of amphipods from the number of amphipods of a given size required to satiate a beetle in the laboratory. Profitability was highest for large amphipods and lowest for small amphipods and isopods. However, amphipod abundance on the beach was always below the threshold at which specialization on larger sizes was predicted to occur.     

The specialized diet of Harpagifer bispinis:

The diet of Harpagifer bispinis (Pisces: Nototheniidae) from two localities of the South Shetland Archipelago was studied. Simultaneous to the capture of H. bispinis and at the same sites the availability of food was considered, and amphipod diversity was compared with the density of Harpagifer. It was found that three quarters of the fish fed only on amphipods (mainly Gondogeneia antarctica) and for the rest amphipods were also the main component, even when other prey species were available. The high selectivity of G. antarctica is due to its high mobility and to the fact that Harpagifer is an ambush feeder. At different predator densities the amphipod fraction of the community appears to be highly modified by the predator both numerically and in species evenness. We postulate that Harpagifer can be a key species in structuring the mobile epibenthic community, even when this environment is subject to strong physical stress.     

A KINEMATIC STUDY OF SUCTION FEEDING AND ASSOCIATED BEHAVIOR IN THE LONG-FINNED PILOT WHALE, GLOBICEPHALA MELAS (TRAILL)

Analysis of videotaped feeding sequences provides novel documentation of suction feeding in captive juvenile long-finned pilot whales ( Globicephala melas ). Swimming and stationary whales were videotaped while feeding at the surface, mid-water, and bottom. The ingestion sequence includes a preparatory phase with partial gape followed by jaw opening and rapid hyoid depression to suck in prey at a mean distance of 14 cm (duration 90 msec), although prey were taken from much greater distances. Depression and retraction of the large, piston-like tongue generate negative intraoral pressures for prey capture and ingestion. Food was normally ingested without grasping by teeth yet was manipulated with lingual, hyoid, and mandibular movement for realignment; suction was then used to transport prey into the oropharynx. Whales frequently rolled or inverted before taking prey, presumably to avoid grasping and repositioning. Prey were sucked off the bottom or sides of the pool without direct contact; lateral suction was used to ingest items from the sides of the mouth.     

Observations on feeding in a South African suctorial hoplonemertean, Nipponnemertes sp. (Family Cratenemertidae)

Marine hoplonemerteans were collected intertidally inAlgoa Bay, Port Elizabeth, South Africa in 1983.Algoa Living worms appear to belong to the genus Nipponnemertes, perhaps N. africanus (Wheeler, 1940). The external morphology and thestylet apparatus are described and illustrated. Laboratory observations showed that this speciesattacked and consumed two species of sympatricamphipods, Elasmopus pectenicrus and Hyalegrandicornis (the only two species tested). Thefeeding behavior was similar to that documented forother suctorial nemerteans that feed on amphipods. Onecomplete feeding sequence, from the initial strike tocompletion of feeding on H. grandicornis,took approximately 12 min: the proboscis struck theventral side of the amphipod, which was immobilized in<1 min; the head eventually wedged between thesternal plates, and the internal organs were evacuatedby suctorial action. The addition of E. pectenicrus andH. grandicornis as potentialprey for suctorial hoplonemerteans brings the totalknown number of amphipod species to 25, involving atleast ten families. A summary of all species ofAmphipoda known to be potential prey for suctorialhoplonemerteans is presented.     

CONSEQUENCES OF ALTERNATIVE FUNCTIONAL RESPONSE FORMULATIONS IN MODELS EXPLORING WHALE-FISHERY INTERACTIONS

We evaluated the utility of Ecosim for exploring interactions between cetacean predators, their prey, and fisheries. We formulated six Ecosim parameterizations, representing alternative hypotheses of feeding interactions (functional response) between cetaceans and their main fish prey, and examined differences in the predicted responses to simulated harvesting regimes for minke whales and their prey. Regardless of the type of function response formulated, intense fishing on the main fish prey of minke whales had a longer-lasting negative impact on minke whales than when minke whale biomass was removed directly by harvesting. Consumption rate, biomass, feeding time and mortality of minke whales were all sensitive to the type of functional response specified. Inclusion of "handling time" limited minke whales consumption at high prey densities and predicted higher consumption at low prey densities; features characteristic of a type II functional response. Predicted decline and recovery rates of minke whales were slower than when consumption rates were not limited. Addition of "foraging time" adjustments resulted in more conservative estimates of decline and recovery. However, when "other mortality" was linked to time spent foraging, exposure to higher mortality at low prey densities, and reduced mortality at high prey densities resulted in dramatic differences in predicted biomass trajectory. Sensitivity to the "other mortality" assumption is important for cetaceans whose predation mortality is only a small proportion of total mortality. Differences in the feeding and biomass dynamics were also observed when prey availability to predators was represented by changes in prey vulnerability, confirming earlier reports that Ecosim predictions are sensitive to this parameter.     

Abundance, biomass and small-scale distribution of cryopelagic amphipods in the Franz Josef Land area (Arctic)

Arctic sea ice is inhabited by several amphipod species. Abundance, biomass and small-scale distribution of these cryopelagic (=ice associated) amphipods were investigated near Franz Josef Land in summer 1994. The mean abundance of all species was 420 ind./m²; the mean biomass was 10.61 g ww/m². Gammarus wilkitzkii was the dominant species, whereas Apherusa glacialis, Onisimus nanseni and O. glacialis were only scarcely found. Amphipods were concentrated at the edges of ice floes and were less frequent in areas further away under the ice. The relationship between the distribution and ecological/physiological requirements of cryopelagic amphipods, as well as the small-scale morphology of Arctic sea ice, are discussed. Received: 14 January 1998 / Accepted 14 April 1998