Cooperative prey capture by young subsocial spiders

Matriphagous young of a subsocial spider Amaurobius ferox exhibit collective predation during their post-maternal social period. In this paper, we examine functional mechanisms of collective predation by sibling groups. Predation efficiency increased with increasing number of individuals within each group. Solitary or paired individuals were generally unable to capture a 20 mg cricket. In larger groups, more individuals participated and captured the prey more quickly. Some siblings did not take part in paralyzing prey, but later consumed it. The proportion of these profiteers within a group increased with the group size. Presented with prey of different sizes (1, 5, or 40 mg), siblings were most aggressive towards each other when predating on 5 mg prey. Prey of this size could be captured by a single individual and yet were sufficiently large for more than one individual to eat. Siblings were much less aggressive towards one another during the capture of 40 mg prey, which require the assistance of other individuals to capture. By providing the same mass of prey in different numbers of individuals (a single cricket of 40 or 40 mg of first-instar crickets), we tested the influence of cooperation on the post-maternal social period. We found no difference in the development of young during the social period nor the timing of dispersal and the body mass of dispersing individuals. We conclude that the young of this subsocial animal increased predation efficiency by cooperative hunting after the mother's death.     

Social spiders catch larger prey: a study of Anelosimus eximius (Araneae: Theridiidae)

Summary During a 1-year-study in tropical Panama, prey of the social theridiid Anelosimus eximius was analysed at two locations and compared with the potential prey spectrum according to sweepnet catches, pitfall traps and bowl traps. Compared with other web-building spiders, A. eximius catch an unusually high number of large insects: about 90% are flying ants, beetles,lepidopterans hemipterans, cockroaches and grasshoppers. This is the result of a communal strategy to overwhelm prey. Webs are maintained commonly, and several spiders attack an entangled insect simultaneously. More spiders participate on insects that are larger and struggle more. The ability to catch large prey insects is discussed as a major driving factor for sociality in spiders.     

Predator foraging behavior in response to perception and learning by its prey: interactions between orb-spinning spiders and stingless bees

Although rewarded bees learn and remember colors and patterns, they have difficulty in learning to avoid negative stimuli such as decorated spider webs spun by Argiope argentata. A. argentata decorates its web with silk patterns that vary unpredictably (Fig. 1) and thus foraging insects that return to sites where spiders are found encounter new visual cues daily. Stingless bees can learn to avoid spider webs but avoidance-learning is slowed or inhibited by daily variation in web decorations (Figs. 3,4; Tables 1,2). In addition, even if bees learn to avoid decorated webs found in one location, they are unable to generalize learned-avoidance responses to similarly decorated webs found at other sites. A. argentata seems to have evolved a foraging behavior that is tied to the ways insects perceive and process information about their environment. Because of the evolutionary importance of bee-flower interdependence, the predatory behavior of web-decorating spiders may be difficult for natural selection to act against.     

Effects of predator visibility on prey encounter: a case study on aerial web weaving spiders

Summary Perhaps the most important factor affecting predator-prey interactions is their encounter probability. Predators must either locate sites where prey are active or attract prey to them, and prey must be able to recognize potential predators and flee before capture. In this study we manipulate and describe three components of the foraging system of predatory, web-weaving spiders, the presence of viscid droplets, silk brightness (achromatic surface reflectance), and visibility of the orb pattern, to determine their effect on insect attraction, recognition, and web avoidance. We found that webs with viscid droplets were more visible to prey at close range, but at greater distances the sparkling droplets lured insects to the web area and hence increased insect capture probability. Although the size of viscid droplets and silk brightness are closely correlated (Table 2, Fig. 3), the relationships among droplet size, spider size, and the visual environments in which webs are found are more complicated (Fig. 2, Tables 2, 3). In environments with predictable light exposure, droplet size and hence silk visibility correlate with spider size, and spiders that forage at night produce relatively more visible silks then spiders that forage during the day (Table 3, Fig. 4). In habitats in which light levels are not predictable, silk surface reflectance and spider size are not closely correlated, suggesting that the complexity of the light environment, as well as the visual and foraging behaviors of insects found there, has played an important role in the evolution of spider-insect interactions.Offprint requests to: C.L. Craig     

Effects of water motion and prey behavior on zooplankton capture by two coral reef fishes

Water motion is an important factor affecting planktivory on coral reefs. The feeding behavior of two species of tube-dwelling coral reef fish (Chaenopsidae) was studied in still and turbulent water. One species of blenny, Acanthemblemaria spinosa , lives in holes higher above the reef surface and feeds mainly on calanoid copepods, while a second, A. aspera , lives closer to the reef surface, feeds mainly on harpacticoid copepods, and is exposed to less water motion than the first. In the laboratory, these two blenny species were video recorded attacking a calanoid copepod ( Acartia tonsa, evasive prey) and an anostracan branchiopod (nauplii of Artemia sp., passive prey). Whereas A. spinosa attacked with the same vigor in still and turbulent water, A. aspera modulated its attack with a more deliberate strike under still conditions than turbulent conditions. For both fish species combined, mean capture success when feeding on Artemia sp. was 100% in still water and dropped to 78% in turbulent water. In contrast, when feeding on Acartia tonsa, mean capture success was 21% in still water and rose to 56% in turbulent water. We hypothesize that, although turbulence reduces capture success by adding erratic movement to Artemia sp. (passive prey), it increases capture success of Acartia tonsa (evasive prey) by interfering with the hydrodynamic sensing of the approaching predator. These opposite effects of water motion increase the complexity of the predator-prey relationship as water motion varies spatially and temporally on structurally complex coral reefs. Some observations were consistent with A. aspera living in a lower energy benthic boundary layer as compared with A. spinosa: slower initial approach to prey, attack speeds modulated according to water velocity, and lower proportion of approaches that result in strikes in turbulent water.Communicated by P.W. Sammarco, Chauvin     

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.     

Effects of prey escape ability,flow speed,and predator feeding mode on zooplankton capture by barnacles

Experimental studies of feeding on zooplankton often involve the use of non-evasive Artemia spp. to represent zooplanktonic prey. Some zooplankton, however, such as copepods, are potentially evasive due to possession of effective predator-avoidance mechanisms such as high-speed escape swimming. In the present study, we compared the efficiencies with which non-evasive (A. salina) and evasive (copepods) zooplankton were captured by a sessile, suspension feeder, the coral-inhabiting barnacle Nobia grandis (Crustacea, Cirripedia). N. grandis specimens and zooplankton used in the present study were collected near Eilat, Israel in 1993. The effect of different flow speeds (from 0 to 14 cm s^-1) on captures of the two preys was also investigated. Additionally, we examined the effect of a flow-induced barnacle behavioral switch from active to passive suspension feeding, on zooplankton capture. Two video cameras were used to make close-up, three dimensional recordings of predator-prey encounters in a computer-controlled flow tank. Frame-by-frame video analysis revealed a highly significant difference (P< 0.001) in the efficiency with which A. salina and copepods were caught (A. salina being much more readily captured than copepods). After an encounter with cirri of feeding barnacles, copepods were usually able to swim out of the barnacles capture zone within one video frame (40 ms), by accelerating from a slow swimming speed (approximately 1.85 cm s^-1) to a mean escape swimming speed of 18.11 cm s^-1 (ca. 360 body lengths s^-1). This was not the case for A. salina nauplii, which usually remained in contact with cirri before being transferred to the mouth and ingested. Thus, experimental studies addressing the methodology of organisms feeding on zooplankton should consider that slow-swimming prey like Artemia sp. nauplii may only represent the non-evasive fraction of natural mesozooplankton assemblages.     

Salps in the Lazarev Sea, Southern Ocean: II. Biochemical composition and potential prey value

Two species of salps, Salpa thompsoni and Ihlea racovitzai, were sampled during three cruises to the Lazarev Sea, Southern Ocean, in summer (December–January) 2005/2006, Autumn (April–May) 2004 and Winter (July–August) 2006. Dry weight, carbon, nitrogen, protein, lipid and carbohydrate contents were measured to characterize the potential value of salps as a food source for predators in the Antarctic ecosystem. Biochemical composition measurements showed that despite having a high percentage of water (~94% of wet weight), both species had relatively high carbon and protein contents in their remaining dry weight (DW). In particular I. racovitzai showed high carbon (up to 22% of DW) and protein (up to 32% of DW) values during all seasons sampled, compared to lower values for S. thompsoni (carbon content only about 15% of the DW, protein content about 10% of the DW). At the same time, carbohydrates (CH) and lipids (Lip) only accounted for a small portion of salp DW in both species (1.4% CH and 3.6% Lip for I. racovitzai; 2.1% CH and 2.9% Lip for S. thompsoni). There was little variability in the biochemical composition of either salp species between the seasons sampled. Both biochemical composition and life cycle characteristics suggest that Antarctic salps, especially I. racovitzai, may be important prey items for both cold and warm-blooded predators in an environment where food is often very scarce.     

Behavioral games involving a clever prey avoiding a clever predator: An individual-based model of dusky dolphins and killer whales

Faced with an intermittent but potent threat, animals exhibit behavior that allows them to balance foraging needs and avoid predators and over time, these behaviors can become hard-wired adaptations with both species trying to maximize their own fitness. In systems where both predator and prey share similar sensory modalities and cognitive abilities, such as with marine mammals, the dynamic nature of predator-prey interactions is poorly understood. The costs and benefits of these anti-predator adaptations need to be evaluated and quantified based on the dynamic engagement of predator and prey. Many theoretic models have addressed the complexity of predator-prey relationships, but few have translated into testable mechanistic models. In this study, we developed a spatially-explicit, geo-referenced, individual-based model of a prototypical adult dusky dolphin off Kaikoura, New Zealand facing a more powerful, yet infrequent predator, the killer whale. We were interested in two primary objectives, (1) to capture the varying behavioral game between a clever prey and clever predator based on our current understanding of the Kaikoura system, (2) to compare evolutionary costs vs. benefits (foraging time and number of predator encounters) for an adult non-maternal dusky dolphin at various levels of killer whale-avoidance behaviors and no avoidance rules. We conducted Monte Carlo simulations to address model performance and parametric uncertainty. Mantel tests revealed an 88% correlation (426 × 426 distance matrix, km²) between observed field sightings of dusky dolphins with model generated sightings for non-maternal adult dusky dolphin groups. Simulation results indicated that dusky dolphins incur a 2.7% loss in feeding time by evolving the anti-predator behavior of moving to and from the feeding grounds. Further, each evolutionary strategy we explored resulted in dolphins incurring an additional loss of foraging time. At low killer whale densities (appearing less than once every 3 days), each evolutionary strategy simulated converged towards the evolutionary cost of foraging, that is, the loss in foraging time approached the 2.7% loss experienced by evolving near shore-offshore movement behavior. However, the highest level of killer whale presence resulted in 38% decreases in foraging time. The biological significance of these losses potentially incurred by a dusky dolphin is dependent on various factors from dolphin group foraging behavior and individual energy needs to dolphin prey availability and behavior.     

Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.     

Clam predation by whelks (Busycon spp.): Experimental tests of the importance of prey size,prey density,and seagrass cover

In 57 l-m² samples within a meadow of Halodule wrightii in Bogue Sound, North Carolina, USA, densities of the clams Mercenaria mercenaria and Chione cancellata were positively associated with seagrass cover. Where seagrass was experimentally removed, marked individuals of both clam species exhibited high rates of mortality in fine sand sediments during two successive experiments spanning 13 months. In the unaltered (control) seagrass meadow, M. mercenaria density remained constant over 13 months and C. cancellata density declined at a slower rate than in the unvegetated plots. Seagrass provides these clams with a refuge from whelk (Busycon carica, B. contrarium, and B. canaliculatum) predation, the major cause of mortality and population decline in experimentally unvegetated plots. In 2 factorial field experiments in unvegetated substratum in which densities of M. mercenaria and C. cancellata were varied independently, first over 5 levels (0 X, 1/2X, 1 X, 2 X, 4 X) and subsequently over 4 levels (0 X, 1/4 X, 1 X, 4 X), there was no repeatable intra- or interspecific effect of density on percent survival, or on the rate of any mortality type. Whelk predation fell preferentially on larger size classes of both species, whereas factors which contribute to clam disappearance usually acted more intensely on smaller sizes. Experimental exclusion of large predators by caging demonstrated that even in unvegetated substratum survivorship of both clam species was high in the absence of whelks and other predators. Individuals of C. cancellata live closer to the sediment surface than those of M. mercenaria, which may explain why seagrass does not serve as effectively to protect them from whelk predation. The mechanism of whelk inhibition may depend upon sediment binding by the H. wrightii root mat, which produces a demonstrable decrease in the physical penetrability of surface sediments.     

Risk vs. reward: how predators and prey respond to aging olfactory cues

Many animals use olfaction to find food and avoid predators, and must negotiate environments containing odors of varying compositions, strengths, and ages to distinguish useful cues from background noise. Temporal variation in odor cues (i.e., “freshness”) seems an obvious way that animals could distinguish cues, yet there is little experimental evidence for this phenomenon. Fresh cues provide a more reliable indicator of donor presence than aged cues, but we hypothesize that the benefits of responding to aged cues depend on whether the cue indicates the proximity of a predator or a potential meal. As prey cannot remain eternally risk averse in response to predator odor, we predict that antipredator responses should diminish as predator cues age. In contrast, animals searching for food should investigate aged prey cues if investigation costs are sufficiently low and the potential benefit (a meal) sufficiently high; thus, we predict that predators will maintain interest in aged prey cues. We tested these ideas using free-ranging rats (Rattus spp.) in two separate experiments; firstly assessing giving-up densities in the presence of predator odor, and secondly examining investigation rates of prey odors. As predicted, giving-up densities dropped once predator odor had aged, but investigation rates remained similar for aged and fresh prey odor. Thus, rats used temporal variation in odor cues to evaluate the cost–benefit relationship of responding to predator and prey odors. We suggest that the ecological significance of variable cue age needs more research and should be considered when interpreting behavioral responses to olfactory information.     

Prey capture by Pisaster brevispinus (Asteroidea: Echinodermata) on soft substrate

Observations on the subtidal seastar Pisaster brevispinus (Stimpson) indicate that this predator can extend the central tube feet into sand-mud substrate for a distance roughly equal to the radius of the seastar. Field and laboratory evidence demonstrates the use of the elongated tube feet by the asteroid for the capture of burrowed prey items. A relationship between predator size and the size of juveniles of the deep-burrowing clam Tresus nuttallii taken as prey is a product of the ability of larger seastars to extend the tube feet further into the substrate and capture deeper prey. The role of P. brevispinus as an important subtidal predator along the Pacific Coast of North America is discussed.     

The effect of prey consumption on territorial defense by harriers: differential responses to neighbors versus floaters

Summary Food consumption may reduce fighting intensity of territory owners by decreasing resource value (additional food cannot be consumed) and/or increasing fighting costs (food in the digestive tract may raise injury risks). A territorial harrier's (Circus cyaneus, adult females) decision to reduce its level of aggression should depend upon whether or not the intruder was a competitor for individual prey items (as are smaller male floaters) or for the territory proper (as are female floaters and especially female neighbors). Accordingly, following meals, aggressive intensity of owners was strongly reduced towards male floaters (more were ignored), slightly reduced towards female floaters (more were called at than chased), and remained unchanged towards neighbors (virtually all were chased). Hence, alterations in aggressive behavior of territory owners following food consumption may depend upon the type of intruder and the resource under contest (a food item or a territory).     

Teams in social insects: group retrieval of prey by army ants (Eciton burchelli,Hymenoptera: Formicidae)

Summary Eciton burchelli workers frequently form groups to retrieve large prey items. Such groups have a definite structure. There is a constant relationship between total ant dry weight and prey item dry weight for both individual porters and groups, and this relation is such that a larger weight of ant or ants can carry disproportionately heavy items. Furthermore, all prey items are carried at a standard retrieval speed. This means that groups are superefficient; they can carry items that are so large that if they were fragmented the original members of the group would be unable to carry all the fragments. Groups also have a distinct caste distribution. There is a significant tendency for each group to have a single submajor, the specialist porter caste in Eciton burchelli. These findings which were verified by experiments suggest that groups fulfill the criteria of efficient teams. The biomechanics are proposed to explain the superefficiency of groups. The organization of co-operation is considered as is the role of teams in the economy of these societies. Colony caste profiles can only be understood by examining the role of teams which form a plastic supercaste.     

Use of pelagic prey subsidies by demersal predators in rocky reefs: insight from movement patterns of lingcod

Allochthonous subsidies of energy and nutrients can affect community structure in patchy marine habitats, including rocky reefs, and their ecological consequences may depend on the mechanism of energy transfer. Lingcod (Ophiodon elongatus) are demersal predators that trophically link nearshore rocky reefs with offshore pelagic habitats through consumption of pelagic fishes. We quantified lingcod habitat use and movement patterns to make inferences about the temporal and spatial conditions under which lingcod may acquire pelagic prey. Lingcod maintained small home ranges (21,272 ± 13,630 m²) within a rocky reef in the San Juan Archipelago, Washington; eight of nine individuals used rocky habitat exclusively. Depths occupied by lingcod (0–50 m) coincided with pelagic fish distribution on the rocky reef; however, diel patterns in lingcod activity varied inversely with occurrence of pelagic fishes on the reef. Our findings suggest that the pelagic subsidy to lingcod is not strongly mediated through directed off-reef foraging by lingcod.     

Oviposition by butterflies on young leaves: Investigation of leaf volatiles

Summary Various butterflies select young foliage on which to lay their eggs; volatiles emitted by young and old leaves have been compared (by sorption enrichment, followed by GC-MS) to gauge possible qualitative and quantitative differences between the two age groups. The plants investigated are cabbage (Brassica oleracea), two milkweeds (Asclepias syriaca andA. curassavica), the bitter orange (Citrus aurantium) and the lime (C. aurantiifolia). The chemical compounds identified belong to three classes, isoprenoids, fatty acid derivatives and benzenoids. Quantitative differences were found between young and old leaves, of which a few may be characteristic of young leaves only. Thirty-four single trials withDanaus plexippus exposed to volatiles from young and old leaves are recorded.