Partial prey consumption by antlion larvae

Predictions from two models of partial prey consumption were tested using antilion larvae (third instar Myrmeleon mobilis), a sit-and-wait predator. Griffiths' (1980) ‘Digestion Rate Limitation’ model correctly predicted decreased handling time and increased ingestion rate with increasing encounter rates. The model incorrectly predicted constant percentage extraction; percentage extraction changed significantly with encounter rate. An optimality model appropriate for ambush predators (Lucas & Grafen, in press) qualitatively matched observations, although antlions always discarded prey somewhat earlier than predicted. Thus neither of the models of partial prey consumption quantitatively fits observations. This reduction in handling time has a minor influence on the rate of energy intake, and therefore may be adaptive if other factors are taken into account. I show that discarding prey early is adaptive if prey that arrive when the predator is empty handed are more easily caught than those that arrive when the predator is eating. Preliminary results support this assumption.     

Habitat complexity reduces prey vulnerability: An experimental analysis using aquatic insect predators and immature dipteran prey

The effects of alternative prey and structural complexity of habitat on the selection of mosquito larvae by aquatic insect predators were evaluated in the laboratory. The water bugs Anisops bouvieri, Diplonychus (= Sphaerodema) rusticus, and D. annulatus, and the odonate nymphs, Ceriagrion coromandelianum and Brachydiplax chalybea chalybea, selected mosquito larvae based on their abundance relative to chironomid larvae and on the levels of habitat complexity. The effect of one prey species on the other was asymmetrical, as indicated through prey selectivity values. Compared to open habitat, the presence of macrophytes reduced the vulnerability of mosquito larvae while the effect was reverse in the presence of sediments. When both sediment and macrophytes were present in habitats, all the predators except D. annulatus consumed more mosquito larvae than chironomid larvae. The clearance rate, an indicator of predatory efficiency, varied among the predator species and habitat types. The results suggest that the outcome of the interactions between insect predators and mosquito immatures was context-dependent and that it was mediated by the presence of alternative controphic species and the habitat complexity.     

Partial consumption of prey: the significance of prey water loss on estimates of biomass intake

Summary A number of studies on the feeding behaviour of sucking predators have estimated the weight of biomass the predator extracts from the prey by measuring the weight change occurring in the prey. This method does not consider that a proportion of the prey weight change is lost to the immediate environment. I examined the spider Diaea sp. feeding on the fruit fly Drosophila immigrans and found that the prey lost approximately 28% more weight than the predator gained. This difference was largely explained by water loss from the prey. My results suggest that water loss, which is not available to the predator, is an important part of prey weight loss. To avoid overestimating predator biomass gain it is necessary to measure the predator weight gain directly or take into account water loss as a component of prey weight change.     

Foraging activity of a dominant epigeal predator: molecular evidence for the effect of prey density on consumption

Despite the widely held assumption that ‘generalist’ predators consume most prey available to them, there is a growing body of evidence suggesting otherwise. Generalists are expected to perform well in disturbed areas because they can switch between prey pathways when one food source becomes depleted. Indeed, these predators have the potential to promote diversity by switching to prey in a frequency dependent manner and consume prey groups in relation to local abundance. It is therefore important to understand how predation rates fluctuate as local availability changes. We performed open‐field and mesocosm experiments in a corn and soybean agroecosystem to delineate the role prey density plays in determining predation frequency of a dominant epigeal predator. To track trophic pathways, molecular gut‐content analysis using enzyme‐linked immunosorbent assay (ELISA) was performed to track foraging behavior of the wolf spider Pardosa milvina feeding on dipterans, flies. Extensive monitoring of foraging activity and prey populations revealed that predation varied temporally. Importantly, the frequency of individuals testing positive for flies was lower than predicted when flies were extremely abundant but higher when they were scarce, relative to the prey community as a whole. Furthermore, isolating predators in mesocosms revealed an effect of Diptera density on the likelihood of consumption, as determined by ELISA, only when flies were at low levels (12.5% of prey provided). The molecular results suggest that these spiders do not appear to be consuming flies in a frequency‐dependent manner where the decision to switch between different prey pathways is driven by relative abundance. Rather, selectivity of prey is somewhat independent of variation of other prey groups, which is indicative of their consistent reliance on dipterans and may be related to nutritional requirements and/or capture success.     

More than a barrier: The complex effects of ecotone vegetation type on terrestrial consumer consumption of an aquatic prey resource

Community structure and dynamics can be influenced by resource transfers between ecosystems, yet little is known about how boundary structure determines both the magnitude of exchanges and their effects on recipient and donor communities. Aquatic and terrestrial ecosystems are often linked by resource fluxes and riparian vegetation is commonly affected by anthropogenic alterations to land use or river hydrological regime. I investigated whether shrubs at the freshwater–terrestrial interface alter the supply, distribution and importance of aquatic prey resources to terrestrial consumers. Shrubs were predicted to alter the larval community composition of aquatic insects and the emergence of winged adults, thus affecting aquatic prey subsidies to terrestrial consumers. In addition, shrubs were hypothesized to alter the microclimatic suitability of the riparian zone for adult aquatic insects, act as a physical barrier to their dispersal and affect terrestrial community composition, particularly the abundance and type of predators that could benefit from the aquatic prey resource. Stable isotope dietary analyses and a survey of shrub‐dominated and open grassland riparian habitats revealed that larval densities of aquatic insects (EPTM: Ephemeroptera, Plecoptera, Trichoptera and Megaloptera) were higher in shrub than grassland habitats; however, reduced emergence and lateral dispersal in shrub areas led to lower densities of adults. The temperature and relative humidity of the riparian zone did not differ between the habitats. Ground‐active terrestrial invertebrate communities had a higher proportion of cursorial spiders in grassland, coinciding with greater abundances of aquatic prey. Aquatic prey contribution to cursorial spider diet matched adult aquatic insect abundances. Overall, riparian shrubs reduced the magnitude, or at least altered the timing, of cross‐ecosystem subsidy supply, distribution and use by consumers through mechanisms operating in both the aquatic and terrestrial ecosystems. Thus, the structure of ecosystem boundaries has complex effects on the strength of biological interactions between adjacent systems.     

Foraging tactics of a terrestrial salamander: Sustained yield in territories

Territorial red-backed salamanders (Plethodon cinereus) were given a high density of two prey types differing in size and caloric profitability. They chose a diet that approximated optimal foraging when they foraged in areas previously marked with advertisement pheromones. However, when foraging in previously unmarked or conspecific-marked areas they did not forage optimally, in a short time frame. In the latter two situations they evinced a lower rate of net energy due to their failure to specialize on the more profitable prey type and their longer intercapture intervals while time was devoted either to marking behaviour or submissive posturing. The salamanders appeared to opt for a strategy of sacrificing initial caloric yield until they had established marked territories and then switching to a higher sustained caloric yield. The data show that experimental handling of a predator per se can alter its foraging behaviour. Also a predator just initiating a territory may forage quite differently from one that has already established ownership, making the time frame during which observations are made important to understanding the predator's foraging strategy.     

Foraging responses of predators to novel toxic prey: effects of predator learning and relative prey abundance

Learning to avoid toxic prey items may aid native predators to survive the invasion of highly toxic species, such as cane toads Bufo marinus in tropical Australia. If the predators’ initial aversion is generalized, native prey that resemble the toxic invader may receive a benefit through accidental mimicry. What ecological factors influence the acquisition of learned avoidance (and hence, the impact of invasion on both predators and native prey)? We conducted laboratory experiments to evaluate how the relative abundance of toad tadpoles compared to palatable native tadpoles (Litoria caerulea and L. rubella) affected the ability of native aquatic predators to discriminate between these two prey types. Both fish (northern trout gudgeon, Mogurnda mogurnda) and frogs (Dahl's aquatic frog, Litoria dahlii) learned to discriminate between toads and frogs within an eight‐day period. Higher abundance of toad tadpoles relative to frog tadpoles enhanced rates of predator learning, and thus reduced predation on toads and increased predation on native tadpoles. In the field, spatial and temporal variation in the relative abundance of cane toads compared to native frogs may influence the rates at which these novel toxic items are deleted from predator diets, and the duration of predator protection afforded to natives that resemble the invader.     

Foraging behaviour and prey size of the painted stork

The foraging behaviour of painted stork Mycteria leucocephala was studied during 2004–2006 at 14 different sites in the Delhi region, India. Observations were recorded on 131 individuals, including 29 juvenile birds using a video camera. Recordings were also made at the nesting colony in Delhi zoo to study the prey sizes regurgitated to nestlings. The results confirm that the painted stork is a tactile forager and exclusively piscivorous. Foraging group size ranged from 1 to 18 individuals. Per 5 min foot stirring rates in the vegetated habitats were significantly higher than in non-vegetated habitats. The attempt rate and feeding rate in the breeding season were significantly higher than that in the non-breeding season. Prey sizes taken in the breeding season were significantly smaller than those taken in the non-breeding season. About 80% fish fed to the chicks were smaller than 10 cm. Young chicks were offered smaller prey compared with older chicks. The variations in foraging parameters are discussed in relation to habitats and their conservation in the Delhi region.     

Induced defense mechanisms in an aquatic angiosperm to insect herbivory

In terrestrial angiosperms, defense and resistance mechanisms against herbivores have been studied extensively; yet this topic is poorly understood in aquatic angiosperms. We investigated induced response mechanisms in Myriophyllum spicatum to the generalist insect herbivore Acentria ephemerella in three independent experiments. Various morphological and chemical response variables were examined in grazed apical shoots and compared to undamaged controls. We further estimated plant palatability of induced and non-induced apices in choice assays, and assessed the growth response of Acentria larvae in no-choice feeding assays. Leaves of induced apices were splayed out horizontally and changed in color from green to red. The dry matter content and thus plant toughness increased by up to 19 %, but silica levels stayed constant. Induced apices exhibited a decline in chlorophyll content of up to 34 %, reflected also by a 10 % decrease in nitrogen levels, while nitrogen increased by 14 % in lower parts of grazed shoots. Also, herbivore-deterring total phenolic compounds increased by up to 20 % in apices. In choice trials, Acentria larvae strongly avoided grazed tips, and growth was reduced by 25 % on induced apices. In total, we observed five different induced resistance and defensive traits in grazed apices: changes in appearance, increased plant toughness, delocalization of N-containing metabolites, increased polyphenols, and reduced nutritional value. The observed changes prevent herbivore damage and loss of apical tissue, which are most valuable for plant fitness. Our study presents the first evidence of multiple, parallel defense strategies including constitutive and induced defense mechanisms in a freshwater angiosperm.     

Flow increases tolerance of heat and hypoxia of an aquatic insect

Recent experiments support the idea that upper thermal limits of aquatic insects arise, at least in part, from a lack of sufficient oxygen: rising temperatures typically stimulate metabolic demand for oxygen more than they increase rates of oxygen supply from the environment. Consequently, factors influencing oxygen supply, like water flow, should also affect thermal and hypoxia tolerance. We tested this hypothesis by measuring the effects of experimentally manipulated flows on the heat and hypoxia tolerance of aquatic nymphs of the giant salmonfly (Plecoptera: Pteronarcys californica), a common stonefly in western North America. As predicted, stoneflies in flowing water (10 cm s⁻¹) tolerated water that was approximately 4°C warmer and that contained approximately 15% less oxygen than did those in standing water. Our results imply that the impacts of climate change on streamflow, such as changes in patterns of precipitation and decreased snowpack, will magnify the threats to aquatic insects from warmer water temperatures and lower oxygen levels.     

Foraging among cannibals and kleptoparasites: effects of prey size on pike behavior

The northern pike (Esox lucius) is an important and selective piscivorethat chooses smaller prey than predicted from energy / timebudgets. In a laboratory experiment, we investigated pike predatorybehavior to explain this selectivity. Northern pike feedingon different prey sizes in aquaria were observed when foragingalone, when in the presence of chemical cues from similar-sizedor larger conspecifics, and when in the presence of conspecifics thatwere allowed to interact with the focal pike. The results showthat prey handling time increases with prey size and that theduration of manipulating and handling prey inflicts a risk ofexposure to cannibals and kleptoparasites on the pike. Therefore,the risk of falling victim to cannibals or kleptoparasites increaseswith prey size. Attracting and experiencing intraspecific interactorscan be regarded as major fitness costs. Chemical cues from foragingconspecifics have only minor effects on pike foraging behavior.Furthermore, the ability to strike and swallow prey head first improvespike predatory performance because failing to do so increases handlingtime. Our findings emphasize the increasing potential costswith large prey and explain previous contradictory suggestionson the underlying mechanisms of behavior, selectivity, and trophiceffects of northern pike predation.     

Foraging for intermittently refuged prey: theory and field observations of a parasitoid

Many insect herbivores feed in concealed locations but become accessible intermittently, creating windows of greater vulnerability to attack, and generating a proportion of the prey population that is readily accessible to foraging natural enemies. We incorporated accessible prey into an extant optimal foraging model, and found that this addition allowed opportunistic exploitation of prey that have already emerged from refugia (the leaving strategy) as a viable strategy, in addition to waiting at refugia for prey to emerge (the waiting strategy). We parameterized the model empirically for the parasitoid Macrocentrus grandii and its host, Ostrinia nubilalis, under field conditions. The model predicted that M. grandii should adopt a leaving strategy when host patch density is high (travel time between patches is short), but a waiting strategy when host patch density is low (travel time between patches is long). Field observations of M. grandii patch tenure were consistent with model predictions, indicating that M. grandii exhibited flexible behaviour based on experience within a foraging bout, and that these behavioural shifts improved foraging efficiency. Behaviour of M. grandii was responsive to heterogeneity in host emergence rates, and appeared to be driven by the relatively small proportion of the host population that became accessible at a fast rate. Therefore understanding forager responses to intermittently refuged prey may require characterization of the behaviour of a subset of the prey population, rather than the average prey individual. The model can potentially be used as a framework for comparative studies across forager taxa, to understand when foragers on intermittently accessible prey should adopt fixed waiting or leaving strategies vs. a flexible strategy that is responsive to the current environment.     

Relationship between prey consumption and colony size in an orb spider

Summary I studied the relationship between prey consumption and colony size in the orb spiderPhiloponella semiplumosa. Observations of unmanipulated colonies showed that prey biomass per juvenile spider was positively correlated with colony size, indicating that prey consumption was highest in the largest colonies observed. In contrast, the relationship between prey biomass per adult female and colony size was curvilinear; prey consumption tended to be highest in intermediatesized colonies. Adult female cephalothorax width was positively correlated with colony size. Number of egg sacs per adult female tended to be highest in intermediate-sized colonies. Prey biomass per juvenile was lower in experimentally reduced colonies than in large control colonies. Aerial-arthropod abundance was not correlated with colony size, and experimental prey supplementation did not affect colony size. Thus, the relationship between prey consumption and colony size was influenced by coloniality directly, rather than by a correlation between prey abundance at a site and colony size.     

Seasonal diversity in aquatic insect communities in an all-year stream system

Summary The general pattern of the data in this study shows that a riffle zone carries more species and, usually, more organisms/m² than a slower current zone. The slower current zone, however, is still apparently able to support a larger number of organisms and species than a pool of standing water. In the West Fork of the Oak Creek system the dominant insects controlling diversity are the Trichoptera, particularly one species of Helicopsyche. Diversity in the fast current zone is high in the summer, and low during other seasons. Diversity in the slow current zone is highest in spring, but is also fairly high in summer. This reflects to a large extent the seasonal fluctuations in abundance of the dominant Trichopterans in the two habitats.     

Foraging tactics and prey-selection patterns of omnivorous and carnivorous calanoid copepods

The foraging tactics and prey-selection patterns of omnivorous and carnivorous calanoid copepods are reviewed. Calanoid foraging tactics are envisioned as falling along several closely coupled continua reflecting swimming behavior, feeding behavior, and dietary habit. The consequences of these foraging tactics on prey-selection patterns are explored in the context of a graphical model. It is hypothesized that the prey-selection patterns of calanoid copepods are determined, to a large extent, by calanoid foraging tactics and the size relationships of predator and prey.     

Fluctuations in prey density: effects on the foraging tactics of scolopendrid centipedes

Foraging animals are faced with the problem of acquiring information about prey populations and utilizing that information in making foraging decisions. In this paper the effect of variation in prey density on the search tactics and mechanisms of prey density assessment in the centipede Scolopendra polymorpha are examined. Centipedes exhibited a prey density-dependent repertoire of search tactics. After 50 min of exposure to high prey density, centipedes switched from active search to ambush-like tactics, while maintaining a high rate of search at the lowest prey density. An initial period of sampling of prey density was involved in the switch in search behaviour and it is suggested that the encounter rate with prey was the key element in density assessment. When the prey density was changed from low to high, centipedes switched from active search to ambush tactics and when prey density was changed from high to low centipedes switched from ambush to active search within 40 min. Such behaviour may decrease the unreliability of sampling information and the risk involved in foraging decisions in variable environments.     

Feeding economy of larvae of a flower fly,Metasyrphus corollae [Dip.: Syrphidae]: Partial consumption of prey

Metasyrphus corollae (F.) larvae ingested aphid contents at a decreasing rate over time because the contents of the aphid became increasingly difficult to obtain as the body of the prey was emptied. Starved larvae usually handled prey longer than wellfed larvae. Younger starved larvae fed longer than older larvae. Larvae starved for 24 h ate the most of a prey. Larvae held prey, on average, until 71% of dry mass was extracted after which the prey carcass was discarded. Prey handling time and amount of each prey consumed were determined by size and hunger of larvae and degree of depletion of prey contents.

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Résumé Les larves deMetasyrphus corollae (F.) ingèrent de plus en plus lentement le contenu des pucerons en fonction du temps car ce contenu est de plus en plus difficile à extraire puisque la proie se vide. Les larves qui ont été privées de nourriture se nourrissent généralement plus longuement que les larves bien nourries, et les jeunes larves affamées plus longuement que les plus agées. Les larves qui n'ont pas mangé depuis 24 h sont celles qui consomment le plus. Les larves retiennent la proie, en moyenne, jusqu'à ce que 71% de son poids sec ait été extrait, après quoi la carcasse de la proie est jetée. Le temps de manipulation et la quantité consommée de la proie sont déterminés par la taille et la faim de la larve, ainsi que par l'épuisement du contenu de la proie. Il n'y a aucune preuve consistante que les larves deM. corollae répondent directement au nombre de proies.

     

Digestive constraints on an aquatic carnivore: effects of feeding frequency and prey composition on harbor seals

We hypothesized that increased feeding frequency in captive harbor seals would increase nutrient loads and thus reduce retention time and the digestive efficiency of natural prey. We measured daily feed intake and excretion during 6 feeding trials and fed herring (49% lipid), pollock (22% lipid) or an equal mix of each diet over 24 months. Animals were accustomed to feeding at either high or low frequency. Body mass and intake did not vary with season. Although mean retention times were similar between diets and feeding frequencies, solute and particulate digesta markers separated at high feeding frequency. Consistent dry matter digestibility resulted in greater gut fill from pollock than from herring. Digestible energy intakes from pollock were approximately 25% greater than from either herring or the mixed diet. Lipid digestibility of herring declined from 90% to 50% when lipid intake exceeded 60 g kg^–0.75 day^–1. Our hypothesis of a trade-off between intake and digestion was not supported for protein but was supported for lipid. Results of this study imply that a flexible digestive system for harbor seals can compensate for ingesting prey of lower energy density by increasing gut fill and enhancing protein and lipid assimilation, to sustain digestible energy intake.Abbreviations DM dry matter - DEI digestible energy intake - DIT diet-induced thermogenesis - FF feeding frequency - MRT mean retention time Communicated by: G. Heldmaier