Metabarcoding and metabolome analyses of copepod grazing reveal feeding preference and linkage to metabolite classes in dynamic microbial plankton communities

In order to characterize copepod feeding in relation to microbial plankton community dynamics, we combined metabarcoding and metabolome analyses during a 22‐day seawater mesocosm experiment. Nutrient amendment of mesocosms promoted the development of haptophyte (Phaeocystis pouchetii)‐ and diatom (Skeletonema marinoi)‐dominated plankton communities in mesocosms, in which Calanus sp. copepods were incubated for 24 h in flow‐through chambers to allow access to prey particles (<500 μm). Copepods and mesocosm water sampled six times spanning the experiment were analysed using metabarcoding, while intracellular metabolite profiles of mesocosm plankton communities were generated for all experimental days. Taxon‐specific metabarcoding ratios (ratio of consumed prey to available prey in the surrounding seawater) revealed diverse and dynamic copepod feeding selection, with positive selection on large diatoms, heterotrophic nanoflagellates and fungi, while smaller phytoplankton, including P. pouchetii, were passively consumed or even negatively selected according to our indicator. Our analysis of the relationship between Calanus grazing ratios and intracellular metabolite profiles indicates the importance of carbohydrates and lipids in plankton succession and copepod–prey interactions. This molecular characterization of Calanus sp. grazing therefore provides new evidence for selective feeding in mixed plankton assemblages and corroborates previous findings that copepod grazing may be coupled to the developmental and metabolic stage of the entire prey community rather than to individual prey abundances.     

Zooplankton swarms: characteristics,proximal cues and proposed advantages

This review focuses on monospecific swarms of four taxonomic groups of small crustaceans: three groups are marine copepods: oithonids (Oithona and Dioithona), Acartia species and Calanus species; and the fourth group includes freshwater cladoceran species in the Order Anomopoda. For each of these groups there is a substantial literature on swarming behavior from field studies and laboratory experiments. Swarming characteristics of each taxonomic group are reviewed, proximal cues for swarming are described, comparisons are made for proposed advantages of swarming, and future research directions are suggested. Swarming characteristics of Calanus spp. are distinctly different from those of the smaller crustaceans: the oithonids, acartiids and cladocerans. In a conceptual model proposed for the smaller crustaceans, swarming behavior is affected by their interactions with light cues, water currents and turbulence, behavior of their predators and prey, and abundance of other trophic levels.     

Distribution of <Emphasis Type="Italic">Calanus</Emphasis> populations in a glaciated fjord in the Arctic (Hornsund,Spitsbergen)—the interplay between biological and physical factors

The distribution and diversity of copepods of the genus Calanus were investigated in Hornsund Fjord (on the southwest coast of Spitsbergen) in summer 2001. The Bhattacharya method was used to sort individuals by species based on their prosome length. The established prosome length boundary values for the Calanus copepodid stages coincided with those defined for the Calanus species from Kongsfjorden (on the northwest coast of Spitsbergen). The predominant species in the main and inner fjord basins was Calanus glacialis, whereas Calanus finmarchicus was the prevailing species outside Hornsund. Younger copepodid stages (CI–CIII) of both species concentrated in the surface water layers (0–50∼70 m), while older copepodids (CIV–CVI females) that were ready for wintering stayed in deep layers (50∼70 m to bottom). Calanus hyperboreus was present in low numbers, predominantly as CIV, and in Hornsund deep water layers. The distribution and diversity of Calanus species complied with the notion that the marine fauna in Hornsund is of a more Arctic character than in Kongsfjorden, a fjord 260 km to the north on the west coast of Spitsbergen.     

Patterns of resource use,food quality,and health status of voles (Microtus pennsylvanicus) trapped from fluctuating populations

Summary Recent studies suggest that diet quality is responsible for differential survivorship of vole cohorts (Boonstra and Boag 1987) and spacing behavior of females (Ims 1987). These phenomena have been related either to a lack of or a deterioration in the quality of the preferred food. To test this hypothesis, we compared foods habits, food quality and health status of meadow voles (Microtus pennsylvanicus) from high and low population density phases. In this study, seven plant species represented the main food items used on a regular basis and biomass values of preferred species decreased with browsing pressure. In addition, food quality of the main dietary items changed between phases forcing females to adjust their feeding strategy accordingly. Health of voles changed also opening the way to speculations on competition for high quality food resources and survivorship.     

Studying <Emphasis Type="Italic">Daphnia</Emphasis> feeding behavior as a black box: a novel electrochemical approach

We present a novel approach for examining the complex feeding behavior of a filter feeder at a previously unexploited scale. A Daphnia lives in a viscous environment and thus creates a feeding current with a distinct laminar inflow and a repetitive pulsed outflow. We propose that by treating the feeding apparatus as a black box, and using the pulsed outflow current as a surrogate to the inside working of the apparatus, we can calculate feeding rate in near real time. The structure of the outflow is interpreted as a direct representation of the organism’s response to its environment. Therefore, we examine how the work performed by an organism’s feeding apparatus is altered according to environmental factors and metabolic demands. Our approach is an integration of optical (Schlieren system) and electrochemical (chronoamperometry) techniques that allow for real time visualization and temporal analysis of flow systems, respectively. As electrochemistry requires a tracer chemical, we employed low dopamine concentrations (≤ 1mM), and tested the effect of dopamine on the heart rate and swimming of Daphnia. It appears that dopamine free in solution at concentrations below 10 mM has no adverse effects on the organism, and all observed differences in Daphnia feeding behavior were due to environmental or metabolic factors. The feeding nature of daphnids in the presence or absence of food, and differences between the sexes is reported. Our results indicate that in the absence of food a Daphnia has a strict and repetitive feeding behavior with short delays between pumping actions. However, in the presence of food this behavior becomes complex, with increased delays between pumps, perhaps designed to maximize feeding efficiency. Our observations demonstrate that males have a higher appendage beat frequency than females under identical conditions. We hypothesize that the difference may be dictated by metabolic demand, as a male spends more time actively seeking a mate. The application of electrochemistry to the study of Daphnia feeding behavior is an improvement over current methods for its near real time quantification of behavioral response, its versatile application under varying environmental conditions and its extreme sensitivity to changes in the organism’s feeding behavior. This technique is a valuable addition to the current tools available for studying Daphnia feeding behavior and will allow us to learn more about the interactions of an organism with its environment. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Protein polymorphisms in six species of the genus Calanus

Isoelectric focusing was used to study total protein patterns and allozyme variations of six species of the genus Calanus. Each species could be characterized by total protein patterns. The results of the allozyme study indicated, in agreement with previous morphological studies, that the six Calanus species belong to three different groups: the C. finmarchicus group C. finmarchicus, C. glacialis and C. marshallae the C. helgolandicus group (C. helgolandicus, C. pacificus), and C. hyperboreus, which stands apart. There is no indication that there are more loci coding for the proteins studied in species with larger genome sizes. Nor is the degree of enzyme polymorphism related to genome size in these species.     

Advances in the Study of Feeding Behaviors, Mechanisms, and Mechanics of Sharks

Sharks as a group have a long history as highly successful predatory fishes. Although, the number of recent studies on their diet, feeding behavior, feeding mechanism, and mechanics have increased, many areas still require additional investigation. Dietary studies of sharks are generally more abundant than those on feeding activity patterns, and most of the studies are confined to relatively few species, many being carcharhiniform sharks. These studies reveal that sharks are generally asynchronous opportunistic feeders on the most abundant prey item, which are primarily other fishes. Studies of natural feeding behavior are few and many observations of feeding behavior are based on anecdotal reports. To capture their prey sharks either ram, suction, bite, filter, or use a combination of these behaviors. Foraging may be solitary or aggregate, and while cooperative foraging has been hypothesized it has not been conclusively demonstrated. Studies on the anatomy of the feeding mechanism are abundant and thorough, and far exceed the number of functional studies. Many of these studies have investigated the functional role of morphological features such as the protrusible upper jaw, but only recently have we begun to interpret the mechanics of the feeding apparatus and how it affects feeding behavior. Teeth are represented in the fossil record and are readily available in extant sharks. Therefore much is known about their morphology but again functional studies are primarily theoretical and await experimental analysis. Recent mechanistic approaches to the study of prey capture have revealed that kinematic and motor patterns are conserved in many species and that the ability to modulate feeding behavior varies greatly among taxa. In addition, the relationship of jaw suspension to feeding behavior is not as clear as was once believed, and contrary to previous interpretations upper jaw protrusibility appears to be related to the morphology of the upper jaw-chondrocranial articulation rather than the type of jaw suspension. Finally, we propose a set of specific hypotheses including: (1) The functional specialization for suction feeding hypothesis that morphological and functional specialization for suction feeding has repeatedly arisen in numerous elasmobranch lineages, (2) The aquatic suction feeding functional convergence hypothesis that similar hydrodynamic constraints in bony fishes and sharks result in convergent morphological and functional specializations for suction feeding in both groups, (3) The feeding modulation hypothesis that suction capture events in sharks are more stereotyped and therefore less modulated compared to ram and bite capture events, and (4) The independence of jaw suspension and feeding behavior hypothesis whereby the traditional categorization of jaw suspension types in sharks is not a good predictor of jaw mobility and prey capture behavior. Together with a set of questions these hypotheses help to guide future research on the feeding biology of sharks.     

The rostral micro-tooth morphology of blue marlin,Makaira nigricans

Billfish rostra potentially have several functions; however, their role in feeding is unequivocal in some species. Recent work linked morphological variation in rostral micro-teeth to differences in feeding behavior in two billfish species, the striped marlin (Kajikia audax) and the sailfish (Istiophorus platypterus). Here, we present the rostral micro-tooth morphology for a third billfish species, the blue marlin (Makaira nigricans), for which the use of the rostrum in feeding behavior is still undocumented from systematic observations in the wild. We measured the micro-teeth on rostrum tips of blue marlin, striped marlin, and sailfish using a micro–computed tomography approach and compared the tooth morphology among the three species. This was done after an analysis of video-recorded hunting behavior of striped marlin and sailfish revealed that both species strike prey predominantly with the first third of the rostrum, which provided the justification to focus our analysis on the rostrum tips. In blue marlin, intact micro-teeth were longer compared to striped marlin but not to sailfish. Blue marlin had a higher fraction of broken teeth than both striped marlin and sailfish, and broken teeth were distributed more evenly on the rostrum. Micro-tooth regrowth was equally low in both marlin species but higher in sailfish. Based on the differences and similarities in the micro-tooth morphology between the billfish species, we discuss potential feeding-related rostrum use in blue marlin. We put forward the hypothesis that blue marlin might use their rostra in high-speed dashes as observed in striped marlin, rather than in the high-precision rostral strikes described for sailfish, possibly focusing on larger prey organisms.     

On birth and death in the sea

We present the first comparative study of the stage-specific patterns of mortality of Calanus and Pseudocalanus, two widely distributed genera that are representative of a relatively large-bodied, broadcast spawning calanoid copepod and a relatively small-bodied, egg-brooding calanoid. The study site is Georges Bank, a continental shelf locality in the Northwestern Atlantic with retentive circulation that renders it suitable for studies of population dynamics. Based on extensive mortality estimates from 30 cruises, we find that co-occurring Calanus finmarchicus and Pseudocalanus spp. have markedly different patterns of stage-specific mortality, the former bimodal and the latter relatively uniform with respect to developmental stage. Neither taxon exhibits a monotonic decline in mortality with developmental stage, nor are rates of mortality predictable in a useful manner by copepod body size or by ambient temperature. Young stages of the broadcast-spawning C. finmarchicus show conditional density-dependence of mortality rates, i.e. mortality rates are independent of population density when adult females are low in abundance but positively related to population density at high female abundances. This density-dependence, which is probably attributable to egg cannibalism, introduces a quadratic mortality term into population dynamic models. The egg-brooding Pseudocalanus spp., in contrast, show no evidence of density-dependent mortality. The two taxa illustrate a life history trade-off: the broadcast-spawning Calanus exhibits birth rates that are greatly elevated with respect to those of Pseudocalanus, but there is a compensatory cost in very low survivorship of the freely spawned eggs. Both the high fecundity, high mortality life history of Calanus and the low fecundity, low mortality life history of Pseudocalanus appear to have approximately equal fitness in this study site.     

Pristane in the marine environment

Summary 1. Methods are described for the sensitive determination of pristane and similar hydrocarbons in individual planktonic organisms and for the isolation of pristane from copepod oil.2. Pristane (2, 6, 10, 14-tetramethylpentadecane) occurs in unusually high concentrations (1–3% of the body lipid) inCalanus finmarchicus, C. glacialis, andC. hyperboreus, and at lower concentrations in a wide range of planktonic animals from the Gulf of Maine and continental slope waters.3. A predator,Paraeuchaeta norvegica, contains pristane at an intermediate level between that ofCalanus and the other herbivores, probably as a result of feeding onCalanus.4. On the basis of relative abundance and structural similarity, phytol is suggested as the precursor of pristane in herbivorous zooplankton.5. Pristane content is unaltered or increased during the metabolism of the major deposit lipids ofCalanus hyperboreus during starvation in the laboratory or in nature. Because of its low density, pristane may contribute to the bouyancy ofCalanus, especially when the other lipids are metabolized.6. Pristane may prove useful as a biochemical integrator for the total assimilation of phytoplankton byCalanus.7. Some planktonic organisms may be characterized by the presence of specific compounds, e. g. pristane inCalanus and several unknown compounds inRhincalanus.8. Specific products of organisms of limited geographical occurence may prove useful as biochemical tags of water masses.

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Pristan in der marinen Umwelt

Kurzfassung Pristan (2, 6, 10, 14-Tetramethylpentadecan) kommt in niedriger Konzentration in manchen planktonischen Tieren des Golfs von Maine, USA, und der Gewässer des kontinentalen Schelfes vor. Dieser Kohlenwasserstoff leitet sich vermutlich vom Phytol in der Nahrung der planktonischen Herbivoren ab. Seine Konzentration ist ungewöhnlich hoch inCalanus finmarchius, C. glacialis undC. hyperboreus, wo er möglicherweise als auftriebsregulierende Substanz in Hungerzeiten wichtig ist. Arten, wieParaeuchaeta norvegica, welcheCalanus verzehren, stellen sekundäre Pristanquellen innerhalb der marinen Nahrungskette dar. Die proCalanus-Individuum angetroffene Pristanmenge könnte als Indikator für die gesamte Nahrungsassimilation dienen. Pristan und andere artspezifische Stoffe können in das die Individuen umgebende Wasser gelangen und dann als biochemische Markierungs-Substanzen dienen.

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Contribution No. 1438 of the Woods Hole Oceanographic Institution.     

Differences in food delivered to chicks by males and females of little auks ( Alle alle ) on South Spitsbergen

Sex differences in the quality and quantity of food loads brought to little auk (Alle alle) chicks were investigated in a large colony in Hornsund (South Spitsbergen). Adults returning to the colony were caught in mist-nets and food loads were taken from their gular pouch. The sex of each bird was determined by means of molecular methods. Females brought significantly more food per single load than males in terms of wet weight (30% more on average), number of prey items (46%) and energy contents (39%). However, there was no difference between the sexes in the size of prey taken. Energy-rich Calanus glacialis, originating from cold Arctic waters, was the most frequent prey item and made up the majority of food loads brought by both males and females (75 and 72%, respectively). This indicates that both sexes forage mainly in Arctic waters. However, differences in the proportion of cold water Calanus species (C. glacialis and C. hyperboreus), warm water Calanus species (C. finmarchicus), as well as other taxa, between males and females may suggest different ways of exploiting the feeding area.     

Plant-feeding and non-plant feeding phytoseiids: differences in behavior and cheliceral morphology

In previous studies plant feeding behavior of plant- and non-plant feeding phytoseiids was never examined directly. Moreover, in these studies the cheliceral morphology of phytoseiids was not associated with their ability to feed on plants. In the present study, we monitored the plant-feeding behavior of Euseius scutalis and Amblyseius swirskii. Only E. scutalis was observed penetrating the leaf surface with the movable digit and feeding. Second, using a dye and coloring the gut as an indicator for feeding, we found that E. scutalis pierced an artificial membrane and fed whereas A. swirskii did not. Finally, to identify morphological characteristics typical of plant feeders versus non-plant feeders, we used scanning electron microscopy to examine the adaxial (inner) profile of the chelicerae in 13 phytoseiid species. The only parameter that distinguished between plant- and non-plant feeders was the ratio of the dorsal perimeter length of the fixed digit to the ventral perimeter length of the movable digit. Plant-feeders were characterized by ratio values greater than one whereas the values for non plant-feeders were lower than one. We suggest that a shorter and less curved movable digit, expressed by a high ratio, will facilitate the penetration of the leaf surface. Cheliceral traits proposed here as typical of plant feeders, were observed for five genera, indicating that plant-feeding may be more common in the Phytoseiidae than previously reported. We propose that the ability to feed on plants be added as a cross type trait of phytoseiid life-style types.     

Foraging behavior related to habitat characteristics in the invasive wasp Vespula germanica

When foraging on carrion resources, the wasp Vespula germanica usually makes repeated visits to the feeding site until depleting the resource. In the present study we analyzed how environmental cues affect wasps' behavior when re‐locating a protein food source. We studied this behavior in two different natural habitats: closed and open habitats. As closed habitats have more references to orient wasps to the feeding site than open habitats, we hypothesized that they would return to the foraging site more frequently in closed habitats than in open ones. We tested this hypothesis by studying wasp behavior in three different natural habitat conditions: (i) closed habitats, (ii) open habitats, and (iii) open habitats artificially modified by adding five sticks with flagging. Experiments consisted of training individual wasps to feed from a certain array, and at the testing phase we removed food and displaced the array by 60 cm. Therefore, we recorded wasps’choices when returning to the training area, by counting both the wasps’first approaches and the number of visits to the original feeding site and the displaced array. Wasps' behavior while re‐locating a protein food source was different if foraging at open or closed habitats. Wasps more frequently revisited a previous feeding location when foraging in closed habitats than when foraging in open ones. Furthermore, wasps more frequently visited the displaced array than the original feeding site in all three treatments. Nevertheless, when wasps were trained in closed habitats, they returned to the original feeding site more frequently than if trained in open ones. Interestingly, when five sticks with flagging were added in open habitats, wasps responded similarly as in closed habitats without these references. The results show that foraging behavior in V. germanica seems to be different in closed and open habitats, probably associated with the existence of references that guide foragers when re‐locating undepleted resources.     

Is the poleward expansion by Atlantic cod and haddock threatening native polar cod, <Emphasis Type="Italic">Boreogadus saida</Emphasis>?

During a recent period of increased influx of warm Atlantic water to the western coast of Svalbard, we have observed a northward expansion of boreal Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) into areas dominated by the native polar cod (Boreogadus saida). To determine the potential impact of new ecological interactions, we studied the diet of co-occurring juvenile gadoids in fjords, open water, and sea ice around Svalbard. We also reviewed the available literature on polar cod feeding in different habitats across the Arctic to determine whether region, habitat, or fish size may influence diet. Feeding by polar cod in the pelagic zone was size dependent, with small fish primarily consuming Calanus spp. and smaller copepods, with an increasing ration of Themisto spp. at larger sizes. In benthic habitats, diets were more varied and included considerably more unidentified material and sediment. Less than 40% dietary overlap was detected among the three species when they were found together. Stable isotope analyses indicated these patterns were representative of longer-term assimilation. The low interspecific dietary overlap suggests little direct competition. Future increases in abundance and the high predation potential of the boreal taxa, however, may impact the persistence of polar cod on some Arctic shelves.     

Arctic seabird food chains explored by fatty acid composition and stable isotopes in Kongsfjorden, Svalbard

Marine birds are important predators in the marine ecosystem, and dietary studies can give useful information about their feeding ecology, food webs and oceanographic variability. The aim of this study was to increase our understanding of the diet and trophic level of the seabirds breeding in Kongsfjorden, Svalbard. We have used fatty acids and stable isotopes, both of which integrate diet information over space and time, to determine trophic relationships in marine food webs. Fatty acid compositions of muscle from Little auk (Alle alle), Brünnich’s guillemot (Uria lomvia), Black-legged kittiwake (Rissa tridactyla), Northern fulmar (Fulmarus glacialis) and Glaucous gull (Larus hyperboreus) were determined and compared with their prey species. Canonical analysis (CA) showed that fatty acid composition differed among the five seabird species. Little auk, Black-legged kittiwake and Northern fulmar had high levels of the Calanus markers 20:1n9 and 22:1, indicating that these seabirds are a part of the Calanus food chain. Brünnich’s guillemot differed from the other species with much lower levels of 20:1n9 and 22:1. Brünnich’s guillemot is a pursuit diver feeding on fish and amphipods deeper in the water column, below 30 m. Glaucous gull also differed from the other seabird species, with a larger variation in the fatty acid composition indicating a more diverse diet. Trophic level analysis placed Little auk at the lowest trophic level, Brünnich’s guillemot and Black-legged kittiwake at intermediate levels and Glaucous gull and Northern fulmar at the highest trophic level.     

Feeding by an omnivorous planktonic copepod aetideus divergens Bradford

Aetideus divergens Bradford is representative of a large group of marine planktonic calanoid copepods which are omnivores or mixed feeders. Because very little quantitative information is available on the feeding behavior of these copepods, laboratory feeding experiments have been carried out with adult female A. divergens presented with various sizes and concentrations of diatoms and freshly hatched nauplii of Artemia. The copepod fed most efficiently on the largest size of diatom and on Artemia nauplii, but was peculiarly inefficient at feeding on small diatoms, even when they were available at very high concentrations. In this respect, the copepod differs from filterfeeding copepods, such as Calanus pacificus Brodsky. A possible explanation of this difference depends upon Aetideus being less capable of handling very small food particles than Calanus. Aetideus divergens and its congeners usually occur at subsurface depths not far below the mixed layer and seem to be adapted for feeding on large particles, possibly large phytoplanktonic organisms and fecal pellets, which sink out of the mixed layer.     

Population genetics of drifting (Calanus spp.) and resident (Acartia clausi) plankton in Norwegian fjords

Kaartvedt distinguished between drifting and resident planktonand hypothesized that the latter were distinguished by theirability to maintain their horizontal position in desired habitats(Kaartvedt, 1993). In this study, we examined the populationgenetic consequences of these two lifestyles for copepods infour fjords of western Norway (Lurefjorden, Masfjorden, Sognefjordenand Sørfjorden) and one fjord in eastern Norway (Oslofjorden).Based on DNA sequence variation of a region of mitochondrial16S rRNA, we contrasted population genetic diversity and structurein drifting populations of Calanus spp. with that of residentpopulations of Acartia clausi. With the exception of Sørfjorden(where Calanus spp. were rare), two or three species of Calanusco-occurred in significantly different proportions in the fjords.Based on a 350 base-pair region of mitochondrial 16S rRNA, Calanusspp. varied in molecular genetic diversity, with the highestvalues for C.helgolandicus. There was no evidence of significantgenetic structure of fjord populations for either C.finmarchicusor C.helgolandicus; the population structure of C.glacialiscould not be evaluated as the species was only abundant in Lurefjorden.Acartia clausi was abundant in all five fjords sampled for thisstudy. Molecular genetic diversity of A.clausi, based on a 220bp region of mt 16S rRNA, was within the range of Calanus spp.values. Populations of A.clausi showed significant genetic structure(i.e. haplotype frequencies differed markedly) among the fjords.The results of this study indicated that little exchange (geneflow) occurs between populations of A.clausi in different fjords,and suggested that the populations are long-term residents ofa fjord. In contrast, most Calanus spp. fjord populations maybe replaced periodically, as they drift with currents flowingto and from coastal and fjord environments.     

Behaviour of <Emphasis Type="Italic">Odontomachus chelifer</Emphasis> (Latreille) (Formicidae: Ponerinae) Feeding on Sugary Liquids

Sugary solution intake behavior by Odontomachus chelifer (Latreille), is studied. The feeding mechanism involved is described. Effects of sucrose concentration and solution viscosity on ingested load, feeding time and rate of liquid intake are assessed. Suction was always the feeding mechanism, regardless of concentration or viscosity of the solution. There were no differences in loads ingested for concentrations of up to 30% w/w, but feeding took longer at this concentration. Liquid intake rates were higher at the lowest concentrations. Above 40% w/w, values of all three variables were smaller. O. chelifer’s ability to ingest mildly viscous fluids could be related to its capacity to ingest other viscous fluids present in its diet.     

Selection of Bemisia nymphal stages for oviposition or feeding,and host-handling times of arrhenotokous and thelytokous <Emphasis Type="Italic">Eretmocerus mundus</Emphasis> and arrhenotokous <Emphasis Type="Italic">E. eremicus</Emphasis>

Host-handling behavior is an important aspect of parasitoid foraging behavior. When a parasitoid encounters a potential host, the handling behavior starts with the evaluation of the host and continues if the host has been judged acceptable. Host handling is usually terminated after egg laying or host feeding and host marking. Host-handling behavior of an arrhenotokous population of two Eretmocerus species, E. mundus Mercet and E. eremicus Rose and Zolnerowich, along with a thelytokous population of E. mundus were compared under laboratory conditions. Several elements of host-handling behavior, including encountering, ascending, turning on host, descending, preening, egg laying, and host feeding were recorded. There were no correlations among the durations of these phases across parasitoid populations/species or host nymphal instars. Duration of different phases of host-handling behavior showed only slight and sometimes significant differences between different Eretmoceruspopulations/species. The actual laying of the egg had the longest duration of all host-handling behaviors, and was longer on third nymphal instars than on younger ones. Females of the three populations/species accepted the first three nymphal stages either for egg laying or for host feeding. Females spent a lot of time to make wounds in the host when preparing for host feeding, and eventually killed the host. The implications of these findings for the use of the different Eretmoceruspopulations/species in biological control are discussed.