Metabolism of epipelagic tropical ctenophores

Measurements of respiration and excretion at 25°C were made for five species of ctenophores collected during five cruises to the Bahamas (1982–1984). The mean element-specific respiration and ammonium excretion rates of freshly collected specimens of all species ranged from 4 to 16% d^-1, the mean atomic O:N ratios were 10 to 16, and ammonium averaged 60 to 90% of the total dissolved nitrogen excreted. For adult ctenophores, the carbon content ranged from 0.6% carbon (as percent of dry weight) for Bolinopsis vitrea to 3.7% carbon for Beroë ovata. There was a marked increase in the organic content (% carbon of dry weight) of small Bolinopsis vitrea with tentacles compared to fully lobate adults. B. vitrea had increasingly higher metabolic rates when held at food concentrations up to 100 copepods 1^-1 (about 250 g C 1^-1). The overall range between starved and well-fed B. vitrea was about two times for respiration and a factor of three for ammonium excretion. B. vitrea decreased from well-fed to a starved metabolic rate in about a day after removal from food. The metabolic rate of Eurhamphaea vexilligera was not measurably affected by short-term starvation or feeding (maximum 25 copepods 1^-1). In feeding experiments, E. vexilligera of 20 to 56 mm length fed at rates equivalent to clearance rates of 250 to 1 800 ml h^-1.     

The feeding behavior of Beroe ovata

Patches of the neritic ctenophores Beroe ovata and Bolinopsis vitrea were observed on the edge of the Great Bahama Bank in order to determine the interaction between the predator Beroe ovata and its prey Bolinopsis vitrea. Laboratory experiments on Beroe ovata showed that it responds chemokinetically to the presence of its prey; as it swims it collides with other etenophores on which it preys. The unique use of macrocilia as cutting implements aids the predator in removing tissue from its prey, yielding maximum gain from each encounter. By means of direct observations while diving, data on horizontal and vertical patchiness, swimming speeds, abundance, and feeding behavior were collected, and used to assess the impact of populations of the predator on its prey. Together, these two animal populations form an ecological feed-back system which affects other portions of the planktonic community.     

Hydrolytic enzymes in the two North Sea ctenophores Pleurobrachia pileus and Beroe gracilis

Hydrolytic enzyme activities were determined in tissue extracts of the two North Sea ctenophores Pleurobrachia pileus and Beroe gracilis. The following activities were measured in field samples and characterized according to their mean specific activities and their pH optima: protease, aminopeptidase, amylase, N-acetylglucosaminidase, -glucosidase, chitinase, carboxylesterase, phosphatase. Many of the enzyme activities are correlated, indicating their control by the same factors (i.e. nutritional conditions). Differences found between the phosphatase activities and the apparent lack of the chitinolytic system in B. gracilis reflect the differences in body composition and food specialization in both ctenophores, respectively. The low amylase and -glucosidase activities in both ctenophores agree with the low carbohydrate content of their prey. In spite of high carboxylesterase activities, no lipolytic activity was detected in either of the ctenophores. In a laboratory feeding experiment with P. pileus, the enzyme activities increased but did not reach the activities found in the field samples. In B. gracilis, seasonal variations of some enzyme activities are discussed in relation to the seasonal abundance of its prey, P. pileus.     

A comparison of strike and prey capture kinematics of three species of piscivorous fishes: Florida gar ( Lepisosteus platyrhincus), redfin needlefish ( Strongylura notata), and great barracuda ( Sphyraena barracuda)

Ram feeding is the process by which a predatory fish uses a high-velocity lunge or chase to overtake its prey. This study compares the strike and prey capture behaviors and kinematics of three species of ram-feeding fishes: Florida gar Lepisosteus platyrhincus, redfin needlefish Strongylura notata, and great barracuda Sphyraena barracuda. These ram-feeding piscivores are morphologically similar with fusiform bodies, posteriorly placed dorsal and anal fins, and large, conical teeth. Strike and prey capture kinematics for five individuals of each species were recorded with high-speed video. Pre-strike behavior in L. platyrhincus consists of a slow stalk, resulting in the close, lateral positioning of the predators head relative to the prey. Lepisosteus employ a sideways lunge of the head during the strike, which lasts only 25–40 ms and is the most rapid strike among these three species. Strongylura notata and Sphyraena barracuda exhibit longitudinal orientation to the prey before the strike, followed by a high velocity, head-on lunge, initiated by an s-start in Sphyraena barracuda. Prey capture in adult L. platyrhincus and Strongylura notata is characterized by the jaws closing on the prey, with the prey held orthogonal to the jaws. This is followed by manipulation using the inertia of the prey to reposition the prey head first, and then suction transport into the buccal cavity. Prey capture in juvenile Sphyraena is accomplished by closing the jaws after the prey has entered the buccal cavity, resulting in possible ram transport of the prey with no oral manipulation under these experimental conditions. Although these three species all employ ram feeding for prey capture of elusive prey, each species has a unique repertoire that appears to minimize hydrodynamic constraints and prey response, utilize locomotory capabilities, and may be suited to each species specific habitat.Communicated by P.W. Sammarco, Chauvin     

Ontogeny of diet changes in a tropical benthic carnivorous fish, Parupeneus barberinus (Mullidae): relationship between foraging behaviour, habitat use, jaw size, and prey selection

Parupeneus barberinus forages on benthic invertebrates using a wide range of foraging modes, including vigorous digging in the substratum, resulting in considerable disturbance to the benthos. Polychaetes were the most important prey item for all size classes, but fishes less than 120 mm total length consumed more small ostracods and nematodes than did larger fishes. Fishes greater than 120 mm total length consumed mostly bivalves, and fishes over 240 mm total length consumed mostly bivalves and crabs. A morphological examination of the feeding apparatus suggested that the size of important prey items consumed was determined by gape height and jaw width. Prey available to different size classes of fishes was determined by combining information on microhabitat use, foraging behaviours, and prey volumes in the substratum. Small fishes spent more time foraging on the reef flat and slope, compared with larger fishes that foraged mostly on the reef edge and base. In addition smaller fishes foraged mostly in the upper 2 cm of sediment, whereas larger fishes often foraged to depths of 10 cm. Selection ratios showed that different size classes of fishes selectively extracted different prey items from the substratum. Small fishes showed a preference for ostracods whereas large fishes selected for bivalves and crabs. Although polychaetes were the dominant prey item for all size classes, they were consistently selected against.     

Not all ctenophores are hermaphrodites. Studies on the systematics,distribution, sexuality and development of two species of Ocyropsis

The lobate ctenophores Ocyropsis maculata and O. crystallina are not simultaneous hermaphrodites, based on morphological, histological and experimental evidence. Sex ratios in populations, sex ratios of size classes within populations, and average sizes of males and females support the hypothesis that both species are dioecious, rather than sequential hermaphrodites. We have divided each species into two subspecies, based on morphology and geographic distribution. Preliminary evidence suggests that the subspecies also differ in reproductive behavior. One subspecies, O. crystallina guttata, spawns on a daily cycle in the laboratory, and spawning becomes more synchronous when males and females are placed together. Species of Ocyropsis, all of which are oceanic in distribution, are probably descended from a common ancestor that was a simultaneous hermaphrodite. That this group of oceanic ctenophores evolved dioecy directly contradicts the assertion that there is a selective advantage to hermaphroditism in environments where the probability of finding a mate is reduced.     

Prey-capture benefits in a mixed-species group of Amazonian tamarins,Saguinus fuscicollis and S. mystax

Summary Tamarins of the genus Saguinus feed on a wide range of arthropods and small vertebrates, which compose a critical component of their diet. This paper examines the foraging patterns and capture success of the Avila-Peres saddle-back (S. fuscicollis avilapiresi) and the red-capped moustached tamarin (S. mystax pileatus) in very stable mixed-species groups, and whether and how any foraging benefits for either species resulted from their association. Moustached tamarins actively searched for prey items which were mainly well exposed on the midstorey foliage. Saddle-back tamarins, on the other hand, foraged at lower heights, largely by manipulating a variety of microhabitats potentially concealing embedded prey. The foraging activity of the numerically dominant and larger-bodied moustached tamarins often resulted in prey items escaping to lower substrates, usually the forest leaf-litter. The beating effect of this species substantially facilitated captures of large, mobile prey items by saddle-backs, which were highly adept at locating and retrieving flushed prey. It is estimated that, while saddle-backs obtained 66–73% of their prey biomass from flushed items, this proportion was substantially lower (2–9%) for moustached tamarins. Commensal insectivory appears to involve a highly asymmetric benefit to saddle-backs, and a low cost to moustached tamarins, which partly explains the stability of mixed-species groups. Correspondence to the present address     

Feeding mechanism of Pelagia noctiluca (Scyphozoa: Semaeostomeae); laboratory and open sea observations

The behaviour of the oral arms of Pelagia noctiluca (Forsskål) has been investigated in open sea predation and in predation induced both in the laboratory and the natural environment. Specimens were first studied in the field and then collected from coastal aggregations in the Gulf of Trieste, North Adriatic Sea, in December 1985 (Lat. 13°40 E, Long. 45°42 N) and in June 1986 (Lat. 13°39 E, Long. 45°43 N). The results of laboratory experiments and in situ observations, recorded on videotapes and photographs, show that the marginal tentacles are utilized to: (1) paralyze the prey; (2) contract and bend inward towards the nearest oral arm. Occasionally the tentacle does not contract and the prey is released; thus prey selection may occur. The oral arms are therefore involved in the: (1) transport of prey from the tentacle to the gastric cavity; (2) catching of motionless prey; (3) anchoring the medusa to the substratum. A similar feeding pattern can explain the survival of several specimens of P. noctiluca near the bottom during the winter of 1985/1986 in the Gulf of Trieste (North Adriatic Sea) in spite of the severe climatic conditions.     

Temporal variability in zooplankton prey capture rate of the passive suspension feeder<Emphasis Type="Italic"> Leptogorgia sarmentosa</Emphasis> (Cnidaria: Octocorallia), a case study

There is increasing evidence that suspension feeders play a significant role in plankton–benthos coupling. However, to date, active suspension feeders have been the main focus of research, while passive suspension feeders have received less attention. To increase our understanding of energy fluxes in temperate marine ecosystems, we have examined the temporal variability in zooplankton prey capture of the ubiquitous Mediterranean gorgonian Leptogorgia sarmentosa. Prey capture was assessed on the basis of gut content from colonies collected every 2 weeks over a year. The digestion time of zooplankton prey was examined over the temperature range of the species at the study site. The main prey items captured were small (80–200 µm), low-motile zooplankton (i.e. eggs and invertebrate larvae). The digestion time of zooplankton prey increased when temperature decreased (about 150% from 21°C to 13°C; 15 h at 13°C, 9 h at 17°C, and 6 h at 21°C), a pattern which has not previously been documented in anthozoans. Zooplankton capture rate (prey polyp^–1 h^–1) varied among seasons, with the greatest rates observed in spring (0.16±0.02 prey polyp^–1 h^–1). Ingestion rate in terms of biomass (g C polyp^–1 h^–1) showed a similar trend, but the differences among the seasons were attenuated by seasonal differences in prey size. Therefore, ingestion rate did not significantly vary over the annual cycle and averaged 0.019±0.002 g C polyp^–1 h^–1. At the estimated ingestion rates, the population of L. sarmentosa removed between 2.3 and 16.8 mg C m^–2 day^–1 from the adjacent water column. This observation indicates that predation by macroinvertebrates on seston should be considered in energy transfer processes in littoral areas, since even species with a low abundance may have a detectable impact.Communicated by S.A. Poulet, Roscoff     

Parent-offspring aggression in moorhens

Summary Colonial orb-weaving spiders from Mexico were studied to test predictions of risksensitive foraging theory: 1. group foraging increases prey capture/individual, and reduces prey variance; 2. spiders should be expected to exhibit risk-averse behavior (forage in groups) when the average level of prey exceeds individual needs, and exhibit risk-prone behavior (forage solitarily) when prey are searce. Laboratory and field studies show that group foraging increases capture efficiency and reduces variability in prey captured per spider. In desert/mesquite grassland habitat, where prey availability is low, M. atascadero forage solitarily in most cases. In tropical rainforest/agriculture sites, M. increassata forage in large colonies of thousands of webs. In intermediate habitats, M. spinipes forages solitarily or in groups, depending on prey availability. Over a range of sites with varying levels of prey, M. spinipes shifts from a risk-prone to a risk-averse group foraging strategy as prey increases.Group foraging behavior observed in colonial Metepeira fits the predictions of risk-sensitive foraging models. These findings explain why spiders tend to group webs together only in areas of superabundant prey. The role of risk-sensitivity in the evolution of coloniality in spiders is discussed.     

Foraging, prey capture, and gut contents of the mesopelagic narcomedusa Solmissus spp. (Cnidaria: Hydrozoa)

Narcomedusae are the most common group of medusae in the mesopelagic depths of Monterey Bay, California. Numerous capture events of various prey taxa were recorded in situ and analyzed using the Monterey Bay Aquarium Research Institute's remotely operated vehicle "Ventana". In situ video analysis of the stomach contents of 82 Solmissus incisa and S. marshalli revealed 88 identifiable prey items. Most (88%) were gelatinous animals. Of these gelatinous prey, 60.3% were ctenophores, 20.5% were cnidarians, 12.8% were salps, 3.8% were chaetognaths, and 2.6% were polychaetes. Euphausiids accounted for 11.4% of the diet, but they were probably captured adventitiously, as the gut contents of ctenophore prey. The tentacle-first foraging behavior of the narcomedusae is an effective way to capture large, comparatively fast-moving prey, because the fluid disturbance caused by swimming is decoupled from the area of prey capture. This behavior contrasts with the prevailing models of feeding behavior in medusae. Stealth predation may be the dominant mode of capturing prey by medusae in the mesopelagic depths of the oceans.     

Food and daily ration of the rock soleLepidopsetta bilineata (Pleuronectidae) in the eastern Bering Sea

A total of 129 specimens ofLepidopsetta bilineata (Ayres) were collected in the eastern Bering Sea (57°02–58°11N; 161°58–164°38W) between 28 August and 7 September 1984. Examination of its stomach and intestine contents revealed that polychaetes, gammarid amphipods and echiuroids constitute the bulk of its diet. Prey type is related to fish size. Juveniles consume more mobile prey, such as small crustaceans (gammarid amphipods, cumaceans, carideans), while adults, although still continuing to feed on gammarid amphipods, preferentially feed on infaunal organisms such as polychaetes, echiuroids and mollusks. Some prey, such as echinoderms, fishes and urochordates, were encountered only in stomachs, which resulted in a higher index of feeding diversity for this organ. Rock sole feeds primarily during daylight hours, peaking at dusk. Its daily ration was estimated as 0.49, 1.12, and 1.61% of fish body weight from contents of stomach, intestine, and stomach plus intestine, respectively.     

Prey capture and structure of the visual space of an insect that hunts by sight on the litter layer (Notiophilus biguttatus F., Carabidae,Coleoptera)

Summary The carabid beetle Notiophilus biguttatus hunts springtails and mites by visual cues. The preycapture behaviour of the beetle and the escape behaviour of the springtails were analysed by means of highspeed films. N. biguttatus has between 900 and 1250 ommatidia in each compound eye. The visual space covers ca. 200° in the horizontal plane, with a binocular overlap of no more than 74°. The fovea, the part of the eye where the pseudopupil is largest, points straight ahead of a beetle in its normal posture.The structure of the visual space was determined from measurements of the optical axes in the horizontal plane (plane of fixation) over the middle of the eye. Because of the slanted position of the ommatidia under the cornea, the optical axes point more towards the front or the back of the animal than do the anatomical axes.The optical axes were used to construct the binocular visual space in the horizontal plane. The point E , to which an estimation of distance is possible, lies on the midline 42.6 mm away from the front edges of the eyes. Resolution rapidly decreases with increasing distance, particularly depth resolution.At a distance corresponding to that from which the beetle attacks its prey, depth and width resolution correspond roughly to the dimensions of the smallest prey animals. The smallest measured directional corrections made by the beetle prior to attack (2°–3°) correspond approximately to the divergence angles in the fovea ( _h=2.2°), and the smallest measured distance correction prior to attack (0.2 mm) corresponds approximately to the depth resolution at attack distance.Supported by the Deutsche Forschungsgemeinschaft (SFB 4)     

Daily ration and specific daily ration of the chaetognath Sagitta enflata

Digestion times, obtained directly from laboratory feeding, were used in conjunction with a day/night series of field samples to estimate the daily ration and specific daily ration of Sagitta enflata Grassi, the most abundant chaetognath in the Gulf Stream near Miami, Florida (USA), during winter and early spring. Feeding was independent of time of day and increased with increasing chaetognath size from 1.7 prey day^-1 for the 11.5 mm size class to 2.9 prey day^-1 for the 21.5 mm class. The diet of s. enflata consisted of 94.8% copepods having a mean weight of 21.8 g, and 5.2% prey chaetognaths estimated to be two-thirds the length of the predators. The daily ration of S. enflata from 12.5 to 20.5 mm in length increased from 48.3 to 143.3 g of prey dry weight and from 18.5 to 48.1 g of prey carbon. The specific daily ration declined from 0.12 to 0.08 dry weight basis, and from 0.26 to 0.14, carbon basis, over the same size range. Prey chaetognaths contributed as much as 51% of the ration on a carbon basis.     

Going deep: common murres dive into frigid water for aggregated,persistent and slow-moving capelin

Owing to the necessity of delivering food to offspring at colonies, breeding seabirds are highly constrained in their foraging options. To minimize constraints imposed by central-place foraging and to optimize foraging behavior, many species exhibit flexible foraging tactics. Here we document the behavioral flexibility of pursuit-diving common murres Uria aalge when foraging on female capelin Mallotus villosus in the northwest Atlantic. Quite unexpectedly, being visual foragers, we found that common murres dived throughout the day and night. Twenty-one percent of recorded dives (n = 272 of 1,308 dives) were deep (≥50 m; maximum depth = 152 m, maximum duration = 212 s), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; 40–180 m) of the Labrador Current. Deep dives occurred almost exclusively during the day when murres would have encountered spatially predictable aggregations of capelin between 100 and 150 m in the water column. Temperatures within the CIL shaped trophic interactions and involved trade-offs for both predators and prey. Sub-0°C temperatures limit a fish’s ability to escape from endothermic predators by reducing burst/escape speeds and also lengthening the time needed to recover from burst-type activity. Thus, while deep diving may be energetically costly, it likely increases certainty of prey capture. Decreased murre foraging efficiency at night (indicated by an increase in the number of dives per bout) reflects both lower light conditions and changing prey behavior, as capelin migrate to warmer surface waters at night where their potential to escape from avian predators could increase.     

Erratum to: Prey selection by resident common bottlenose dolphins (Tursiops truncatus) in Sarasota Bay, Florida

Prey selection was investigated in wild, resident common bottlenose dolphins, Tursiops truncatus, during the summer months in Sarasota Bay, Florida, USA. Stomach content analyses of 15 dolphins with extensive sighting histories and well-documented distributions were used to determine prey use. Prey availability was assessed by purse seine surveys. We compared the relative abundances of prey available to estimates of prey use at closely matching spatial and temporal scales. G-tests determined that dolphins in this study significantly selected for prey at the species, family, and soniferous/non-soniferous prey levels (G _adj  = 753.98–1,775.93, df = 1–21, p ≤ 0.01). While comprising only 6.3% of the total available prey, soniferous fishes accounted for 51.9% of the total prey consumed. Manly’s standardized forage ratios and 95% Bonferroni confidence intervals determined significant positive selection for soniferous prey and against non-soniferous prey (β^S = 0.9461 vs. β^NS = 0.0539). Dolphins selected against Gerridae, Clupeidae, and Sparidae (β ≤ 0.0014), as well as against all the species within those families (β ≤ 0.0190). It is likely that passive listening for soniferous prey provides an ecological or energetic advantage to cetaceans utilizing this specific foraging technique.     

Prey selection in <Emphasis Type="Italic">Octopus rubescens</Emphasis>: possible roles of energy budgeting and prey nutritional composition

This study explores the relationship between energy budgeting and prey choice of Octopus rubescens. Seventeen male Octopus rubescens were collected between June 2006 and August 2007 from Admiralty Bay, Washington. Prey choices made by individuals in the laboratory deviated widely from those expected from a simple optimal foraging model. O. rubescens chose the crab Hemigrapsus nudus over the clam Nuttallia obscurata as prey by a ratio of 3:1, even though prey energy content and handling times suggested that this octopus could obtain 10 times more energy intake per unit time when choosing the latter compared to the former prey species. Octopus energy budgets were similar when consuming either of the prey species except for lipid extraction efficiency that was significantly higher in octopuses consuming H. nudus. This suggests that lipid digestibility may play an important role in the prey choice of O. rubescens.     

Forager specialization and the control of nest repair in Polybia occidentalis Olivier (Hymenoptera : Vespidae)

We measured patterns of individual forager specialization and colony-wide rates of material input during periods of response to experimental nest damage and during control periods in three colonies of the tropical social wasp Polybia occidentalis.

Species-specific antipredatory behaviours: effects on prey choice in different habitats

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

Diet of young Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in eastern and western Atlantic foraging grounds

Atlantic bluefin tuna (Thunnus thynnus) are highly migratory predators whose abundance, distribution, and somatic condition have changed over the past decades. Prey community composition and abundance have also varied in several foraging grounds. To better understand underlying food webs and regional energy sources, we performed stomach content and stable isotope analyses on mainly juvenile (60–150 cm curved fork length) bluefin tuna captured in foraging grounds in the western (Mid-Atlantic Bight) and eastern (Bay of Biscay) Atlantic Ocean. In the Mid-Atlantic Bight, bluefin tuna diet was mainly sand lance (Ammodytes spp., 29% prey weight), consistent with historic findings. In the Bay of Biscay, krill (Meganyctiphanes norvegica) and anchovy (Engraulis encrasicolus) made up 39% prey weight, with relative consumption of each reflecting annual changes in prey abundance. Consumption of anchovies apparently declined after the local collapse of this prey resource. In both regions, stable isotope analysis results showed that juvenile bluefin tuna fed at a lower trophic position than indicated by stomach content analysis. In the Mid-Atlantic Bight, stable isotope analyses suggested that >30% of the diet was prey from lower trophic levels that composed <10% of the prey weights based upon traditional stomach content analyses. Trophic position was similar to juvenile fish sampled in the NW Atlantic but lower than juveniles sampled in the Mediterranean Sea in previous studies. Our findings indicate that juvenile bluefin tuna targeted a relatively small range of prey species and regional foraging patterns remained consistent over time in the Mid-Atlantic Bight but changed in relation to local prey availability in the Bay of Biscay.