Odour-mediated foraging by yellowjacket wasps (Hymenoptera: Vespidae): predation on leks of pheromone-calling Mediterranean fruit fly males (Diptera: Tephritidae)

Predation is probably the most important male mortality factor in insect species with courtship displays that render males performing them conspicuous targets of predators. Sexually active Mediterranean fruit fly males, Ceratitis capitata (Wied.), aggregate in leks, where they participate in agonistic encounters and engage in visual, acoustic and pheromone-calling displays to attract receptive females. The objective of this study was to assess: a) whether sexually displaying C. capitata males in leks inside host and non-host foliage are subject to predation by the most prominent predators yellow-jacket wasps, Vespula germanica (F.), and if so, b) whether olfactory, visual or auditive stimuli are used by foraging wasps in locating male C. capitata prey. Studies were carried out in a citrus orchard and surroundings on the island of Chios, Greece. Observations were conducted using perforated containers hung within mulberry, fig or citrus foliage. Living C. capitata flies of different sex and either mature or immature were placed inside. Our results show that the yellowjacket wasps have learned to associate the presence of sexually active medfly males aggregated in leks with their prey's pheromone (kairomone). Foraging wasps, flying through the crowns of host trees, responded to the odour source of C. capitata male pheromone by approaching from downwind. Even inside dense citrus tree foliage, wasps keyed in on aggregations of pheromone-calling males using olfactory stimuli. Stimuli of visual and acoustic male signalling were only used at close range, after having followed the pheromone plume close to its source. Visual cues played a greater role in directing wasp foraging under more open and exposed host foliage conditions. Odour-based foraging of wasps inside host foliage in the mid-morning hours, when medfly male lekking activities peak, shifted gradually to a more visual-based host fruit patrolling in the afternoons to capture ovipositing and feeding medfly females. On ripe fruit, particularly fig, V. germanica visual prey hunting also included the capture of feeding medfly males, other feeding Diptera, as well as medfly larvae extracted from wasp-made perforations in the fruit.     

Host Location and Ovipositional Behavior of Platygaster demades Walker (Hymenoptera: Platygastridae), an Egg Parasitoid of Apple and Pear Leaf Curling Midges

The foraging responses of 1–2-day-old naïve female Platygaster demades to odors of apple and pear foliage and host insect eggs were measured. The host origin of P. demades had no effect on the parasitoids’ longevity, host preference, or foraging behavior. Four distinct behaviors related to oviposition were identified. In choice experiments, more female parasitoids responded to apple foliage with no midge eggs than to midge eggs alone. In a Y-tube olfactometer, parasitoids preferred the plant cues to clean air, and responded equally to both apple and pear odors. The results indicate that P. demades utilizes plant cues to locate the habitat of its host and then searches for host eggs to parasitize.     

Close-Range Host Searching Behavior of the Stemborer Parasitoids <Emphasis Type="Italic">Cotesia sesamiae</Emphasis> and <Emphasis Type="Italic">Dentichasmias busseolae</Emphasis>: Influence of a Non-Host Plant <Emphasis Type="Italic">Melinis minutiflora</Emphasis>

Studies were conducted on the host searching behavior of the larval parasitoid Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) and the pupal parasitoid Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae), both of which attack lepidopteran (Crambidae, Noctuidae) cereal stemborers. The behavior of D. busseolae was observed in a diversified habitat that consisted of stemborer host plants (maize, Zea mays L. and sorghum, Sorghum bicolor (L). Moench (Poaceae)) and a non-host plant (molasses grass, Melinis minutiflora Beauv. (Poaceae)), while C. sesamiae was observed separately on host plants and molasses grass. In previous olfactometer studies, C. sesamiae was attracted to molasses grass volatiles while hboxD. busseolae was repelled. The aim of the present study was to investigate the influence of molasses grass on close-range foraging behavior of the parasitoids in an arena that included infested and uninfested host plants. Dentichasmias busseolae strongly discriminated between host and non-host plants, with female wasps spending most of the time on infested host plants and least time on molasses grass. Likewise, C. sesamiae spent more time on uninfested and infested host plants than it did on molasses grass in single choice bioassays. While on infested plants, the wasps spent more time foraging on the stem, the site of damage, than on other areas of the plant. Overall, the results indicate that presence of the non-host plant does not hinder close range foraging activities of either parasitoid.     

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.     

Importance of host feeding for parasitoids that attack honeydew-producing hosts

Insect parasitoids lay their eggs in arthropods. Some parasitoid species not only use their arthropod host for oviposition but also for feeding. Host feeding provides nutrients to the adult female parasitoid. However, in many species, host feeding destroys an opportunity to oviposit. For parasitoids that attack Homoptera, honeydew is a nutrient‐rich alternative that can be directly imbibed from the host anus without injuring the host. A recent study showed that feeding on host‐derived honeydew can be an advantageous alternative in terms of egg quantity and longevity. Here we explore the conditions under which destructive host feeding can provide an advantage over feeding on honeydew. For 5 days, Encarsia formosa Gahan (Hymenoptera: Aphelinidae) parasitoids were allowed daily up to 3 h to oviposit until host feeding was attempted. Host feedings were either prevented or allowed and parasitoids had ad libitum access to honeydew between foraging bouts. Even in the presence of honeydew, parasitoids allowed to host feed laid more eggs per hour of foraging per host‐feeding attempt than parasitoids that were prevented from host feeding. The higher egg‐laying rate was not compromised by survival or by change in egg volume over time. In conclusion, host feeding can provide an advantage over feeding on honeydew. This applies most likely under conditions of high host density or low extrinsic mortality of adult parasitoids, when alternative food sources cannot supply enough nutrients to prevent egg limitation. We discuss how to integrate ecological and physiological studies on host‐feeding behavior     

Foraging for solitarily and gregariously feeding caterpillars: A comparison of two related parasitoid species (Hymenoptera: Braconidae)

In the present study we apply a comparative approach, in combination with experimentation, to study behavior of two parasitoid species that attack caterpillar hosts with different feeding strategies (gregarious or solitary). In a semifield setup, consisting of clean cabbage plants and plants infested with one of two host species, the foraging behavior of the specialistCotesia rubecula, on obligate parasitoid of solitarily feedingPieris rapae larvae, was compared to that of the generalistCotesia glomerata, a polyphagous parasitoid of several Pieridae species (mainly the gregariously feedingPieris brassicae).Cotesia glomerata displayed equal propensity to search for and parasitize larvae of both host species. AlthoughC. glomerata exhibited a relatively plastic foraging behavior in that it searched differently under different host distribution conditions, its behavior seems more adapted to search for gregariously feeding hosts. Females exhibited a clear area-restricted search pattern and were more successful in finding the gregariously feeding caterpillars.Cotesia rubecula showed a higher propensity to search forP. rapae than forP. brassicae, i.e., females left the foraging setup significantly earlier when their natural hostP. rapae was not present.C. rubecula showed a more fixed foraging behavior, which seems adapted to foraging for solitarily feeding host larvae. In a setup with onlyP. rapae larvae, the foraging strategies of the two parasitoid species were quite similar. In a choice situationC. glomerata did not show a preference for one of the host species, whileCotesia rubecula showed a clear preference for its natural host species. The latter was shown by several behavioral parameters such as the number of first landings, allocation of search time, and percentage parasitization.     

The parasitoid of a fruit moth caterpillar utilizes fruit components as nutrient source to increase its longevity and fertility

Longevity and fecundity of female wasps are two decisive factors for the effectiveness of parasitoid species as biological control agents. Accessibility and suitability of nutrient sources determine parasitoid survival and reproduction. Host, nectar and honeydew feeding are frequent adult parasitoid behaviors to cover nutritional needs. Here we postulate that especially parasitoid species of endophytic herbivores might use plant tissue as a nutrient source that becomes accessible upon herbivory. We investigated the influence of plant consumption and host feeding on longevity and fecundity of Hyssopus pallidus, a gregarious ecto-parasitoid of caterpillars of the codling moth that feed inside apple fruits. Longevity of unmated and mated ovipositing female parasitoids was highest in treatments with fruit pulp. While longevity in this treatment was not significantly different from that with honey, it was significantly higher than in treatments without food, with water or with a host alone.Reproduction was significantly increased by these sugar-rich nutrient sources compared to the control with a host alone. In contrast, host feeding did not yield any significant contribution to longevity and fecundity in a series of bioassays with different host–parasitoid ratios and with differently aged and sized hosts, compared to controls without food.We conclude that in this synovigenic species host feeding does not contribute to longevity and fecundity, but females can increase survival and reproduction in the field relying solely on the plant tissue damaged by their host caterpillar.     

Natural selection in the laboratory for a change in resistance byDrosophila melanogaster to the parasitoid waspAsobara tabida

The selection response of the polymorphic hostD. melanogaster (Meigen) to the braconid waspA. tabida (Nees) is addressed. Cages of flies with and without wasps were initiated with a population ofD. melanogaster that exhibited variation both in larval foraging behavior and in encapsulation ability. Encapsulation ability was measured as the proportion of parasitized larvae that produce a hardened capsule which encapsulates the wasp egg and ultimately kills the wasp larva. We determined whether the host population changed its encapsulation ability and/or its foraging behavior in response to the wasp. Both species were collected from a local orchard whereA. tabida is the only wasp known to parasitizeD. melanogaster larvae. The naturally occurring genetic polymorphism for rover and sitter larval foraging behavior inD. melanogaster is also found in this field population.A. tabida's vibrotactic search behavior enables it to detect rover more frequently than sitter larvae. Rover larvae move significantly more while feeding than do sitter larvae. In this field population, rover larvae also show higher encapsulation abilities than do sitter larvae. Six cage populations, three without wasps and three with wasps, each containing an equal mixture of rover and sitter flies, were established in the laboratory and maintained for 19 fly generations. Selection pressure in the laboratory was similar to that found in the field population from which the flies and wasps were derived. We found that larvae from cages with wasps developed a significantly higher frequency of encapsulation than those reared without wasps. We were, however, unable to detect a change in larval movement (rover or sitter behavior) in larvae from cages subject to selection from wasps compared to larvae from cages containing no wasps. This may have resulted from a balance between two selective forces, selection against rovers by the wasps' use of vibrotaxis, and selection for rovers resulting from their increased encapsulation abilities     

Temperature and forager body size affect carbohydrate collection in German yellowjackets, Vespula germanica (Hymenoptera, Vespidae)

The classic formulation of optimal foraging theory predicts that a central-place forager will gather more food if it is required to travel farther from the nest to find that food. We examined the foraging behavior of German yellowjackets (Vespula germanica) to determine whether carbohydrate foragers follow this pattern. We trained foragers to collect 2 M fructose solution at 5 or 50 m from the nest and measured the time spent feeding, load size, and the rate of delivery. We show that as a forager’s crop fills during a foraging bout, the amount of solution ingested per second decreased. However, load size did not change as wasps collected food up to 50 m from the nest. Instead, temperature and body size were better predictors of the volume of fructose a forager carried. Finally, the rate of fructose delivered to the nest was higher at warmer temperatures. Due to the fact that wasps gather more food but feed for shorter periods of time at warmer temperatures, we found an overall negative relationship between feeding time and load size. We conclude that the strong effects temperature had on the behavior of V. germanica foragers imply that feeding time may not always be an accurate predictor of the size of the load an individual carries back to the nest. Results from this study suggest that in yellowjacket colonies, foragers can collect and bring disproportionately more food back to the nest during the warmest days of the summer, a time of year when this pest species reaches peak population size during its annual colony cycle.     

Long‐term spatial memory in Vespula germanica social wasps: the influence of past experience on foraging behavior

Social insects exhibit complex learning and memory mechanisms while foraging. Vespula germanica (Fab.) (Hymenoptera: Vespidae) is an invasive social wasp that frequently forages on undepleted food sources, making several flights between the resource and the nest. Previous studies have shown that during this relocating behavior, wasps learn to associate food with a certain site, and can recall this association 1 h later. In this work, we evaluated whether this wasp species is capable of retrieving an established association after 24 h. For this purpose, we trained free flying individuals to collect proteinaceous food from an experimental plate (feeder) located in an experimental array. A total of 150 individuals were allowed 2, 4, or 8 visits. After the training phase, the array was removed and set up again 24 h later, but this time a second baited plate was placed opposite to the first. After 24 h we recorded the rate of wasps that returned to the experimental area and those which collected food from the previously learned feeding station or the nonlearned one. During the testing phase, we observed that a low rate of wasps trained with 2 collecting visits returned to the experimental area (22%), whereas the rate of returning wasps trained with 4 or 8 collecting visits was higher (51% and 41%, respectively). Moreover, wasps trained with 8 feeding visits collected food from the previously learned feeding station at a higher rate than those that did from the nonlearned one. In contrast, wasps trained 2 or 4 times chose both feeding stations at a similar rate. Thus, significantly more wasps returned to the previously learned feeding station after 8 repeated foraging flights but not after only 2 or 4 visits. This is the first report that demonstrates the existence of long‐term spatial memory in V. germanica wasps.     

Feeding time strategies of the fringe-toed lizard,Uma inornata,during breeding and non-breeding seasons

Summary I examined the foraging behavior during the breeding and non-breeding seasons, May and July 1986, of the fringe-toed lizard Uma inornata (Iguanidae). During the breeding season males differ from females in their diet and in their foraging time strategy, males exhibiting time minimization and females energy maximization. In May, plant associated foods were selectively eaten. Males concentrated on flowers, a readily available quick energy food, which reduced foraging time and increased time for reproductive activities. Time budgets indicate that males spend over twice as much time in the open and in movement in May than do females. Females at this time restrict their activities to the cover of perennial bushes, and feed primarily on plant foods (flowers and arthropods). Energy maximization appears to be maintained by both sexes in the non-breeding season when food resources diminished to one-half of those in the breeding season. The lizards were less selective in their July feeding habits, broadening their diets to include ground-dwelling arthropods and foliage. Predation by these lizards follows a wait-ambush mode of foraging.     

Host-handling behavior of the parasitoid,Diglyphus minoeus [Hym.: Eulophidae], parasitizing the honeysuckle leaf-miner,Phytomyza lonicerae [Dipt.: Agromyzidae]

The host-handling behavior of individual female waspsDiglyphys minoeus which attack the larvae of the honeysuckle leaf-miner,Phytomyza lonicerae, include 4 types of behavior: probing, ovipositor insertion, host-feeding, and resting. Ovipositor insertion may either be injection of venom, probing of the host by the sensilla on the ovipositor's tip or egg laying. Three types of attacks were distinguished: oviposition attack for unparasitized larvae, host-feeding attack for unparasitized host larvae and host rejection for previously parasitized host larvae. Oviposition attack was characterized by frequent alternation between probing and ovipositor insertion, long duration of ovipositor insertion and resting, and the long duration of host-handling. Resting behavior is thought to protect the progeny against superparasitism, host-feeding by other wasps or hosts' recovery from paralysis. Host-feeding attack was characterized by frequent alternation between host-feeding and ovipositor insertion and long duration of host-feeding. Host rejection was composed mainly of probing and ovipositor insertion and short handling time.      

Lifetime Reproductive Success in the Solitary Endoparasitoid, Venturia canescens

Parasitoid wasps have long been considered excellent organisms in studies examining the evolution of reproductive and life-history strategies. In examining the lifetime reproductive success of parasitoids in the laboratory, most investigations have provided the insects with excess hosts and food, where they exist in a relatively constraint-free environment. Importantly, these conditions may not accurately reflect the true heterogeneity of natural systems, where suitable hosts and food sources are likely to be limiting. This study examines the influence of differences in host and food availability on reproductive and life-history parameters in an asexual strain of the solitary endoparasitoid, Venturia canescens (Hymenoptera: Ichneumonidae). Lifetime reproductive success in V. canescens was measured in response to temporal variations in host and food (honey solution) access. Cohorts of parasitoids were provided with 200 fifth-instar larvae of the Indian meal moth, Plodia interpunctella (Lepidoptera: Pyralidae), and food for variable periods daily after eclosion. V. canescens is synovigenic, and host-deprived wasps continued to mature eggs over the first few days after eclosion until the egg storage capacity was reached in the oviducts. When these wasps were subsequently provided with hosts, oogenesis resumed and continued until later in adult life. Constantly fed wasps lived longer and produced more progeny than wasps from cohorts which were alternately fed and starved or were starved from eclosion. Moreover, wasps with constant host and food access produced most progeny early in life and usually experienced prolonged periods of postreproductive survival. In contrast, the reproductive period of wasps with limited host access was more evenly distributed throughout the adult life. Consequently, the cumulative progeny production by V. canescens with constant food access was fairly uniform irrespective of host availability. Longevity and fecundity in V. canescens were positively correlated with adult size. However, variable host access had little effect on the longevity of wasps which were constantly supplied with honey. Over the first 2 days of adult life, variation in food access also had no effect on progeny production by V. canescens. We argue that manipulating temporal host and food access to parasitoids in the laboratory more closely approximates natural conditions, where these resources are likely to be spatially separated. Moreover, our findings suggest that many highly synovigenic parasitoids like V. canescens, which produce microtype (=hydropic) eggs, have a considerably higher reproductive potential than ovary dissections have revealed. Our findings are discussed in relation to life-history evolution in the parasitic Hymenoptera.     

Differential energy allocation as an adaptation to different habitats in the parasitic wasp <Emphasis Type="Italic">Venturia canescens</Emphasis>

Environmental pressures are expected to favour organisms that optimally allocate metabolic resources to reproduction and survival. We studied the resource allocation strategies and the associated tradeoffs in the parasitoid wasp Venturia canescens, and their adaptation to the characteristics of the environment. In this species, individuals of two reproductive modes coexist in the same geographical locations, but they mainly occur in distinct habitats. Thelytokous (asexual) wasps are mostly found in anthropogenic habitats, where hosts tend to aggregate and food is absent. Arrhenotokous (sexual) wasps are exclusively found in natural habitats, where hosts are scattered and food is present. We analysed (1) the quantity of energy stored during ontogeny, (2) the tradeoff between reproduction and survival, by measuring egg load and longevity and (3) the host patch exploitation behaviour of the wasps at emergence. Arrhenotokous wasps emerged with more metabolic resources than thelytokous ones, especially glycogen, a nutrient that could be used for flying in search of hosts and/or food. Thelytokous wasps allocated more energy than arrhenotokous wasps to egg production: this would allow them to parasitize more hosts. The tradeoff between egg production and longevity was not revealed within reproductive modes, but when comparing them. At emergence, arrhenotokous wasps tended to exploit host patches less thoroughly than thelytokous wasps, suggesting that by leaving the host patch, they search for food. The results clearly showed adaptations to the characteristics of habitats preferentially inhabited by the two reproductive modes, and suggested a mechanism that facilitates their coexistence in natural conditions.     

Host Stage Selection and Sex Allocation by Gyranusoidea tebygi (Hymenoptera: Encyrtidae), a Parasitoid of the Mango Mealybug, Rastrococcus invadens (Homoptera: Pseudococcidae)

Host stage selection and sex allocation by Gyranusoidea tebygi Noyes (Hym,, Encyrtidae) were studied in choice and no-choice experiments in the laboratory. The parasitoid reproduced on first, second, and third instars of the mango mealybug, Rastrococcus invadens Williams (Hem., Pseudococcidae), and it avoided hosts that were already parasitized. Host feeding was occasionally observed. Sex ratios of the offspring produced by individual wasps were highly biased in favor of females, whereas the sex ratio of groups of wasps foraging under crowded conditions varied from male biased in smaller hosts to female biased in larger hosts. Females had longer developmental times than males, developed faster in larger mealybugs than in smaller ones, and were always larger than males emerging from the same host instar. Their size increased with the instar of the host at oviposition. About 90% of all ovipositions in second and third instar nymphs resulted from an attack with multiple stings, starting with a sting in the head of the host for the most part. The function of these head stings is either to assess quality of the host or to subdue hosts prior to oviposition. Encounter rates, number of attacks, and number of stings during one attack increased, while ovipositions decreased with host instar. Time investment per oviposition and time spent preening increased with increasing host age because older hosts defended themselves more vigorously than younger ones. Thus, while fitness of the parasitoid increased with host size, fitness returns per time decreased. The implications of this host selection behavior for the biological control of the mango mealybug are discussed.     

Intra-plant host selection, oviposition preference and larval survival of Chrysophtharta agricola (Chapuis) (Coleoptera: Chrysomelidae: Paropsini) between foliage types of a heterophyllous host

Abstract 1 Paropsine chrysomelid beetles defoliate commercial eucalypt plantations in Australia. Adults and larvae feed on the same host, with the larval food source determined by the oviposition choice of females. Most eucalypt species are heterophyllous, with their foliage undergoing distinct morphological and chemical changes between adult and juvenile growth. 2 The intra‐plant foliage feeding and oviposition preference adults and the larval development of Chrysophtharta agricola were examined using adult and juvenile foliage of a heterophyllous plantation species, Eucalyptus nitens. The foliage types differ in chemistry, toughness, waxiness and timing of production. 3 In the field, feeding damage caused by adult beetles was 15% more frequent on adult foliage than on juvenile foliage; however, egg batches were three times more common on juvenile than on adult foliage. 4 Oviposition preference for juvenile foliage over adult foliage was confirmed in choice trials in the laboratory, with adult fecundity and longevity not significantly different between foliage types. 5 Larval survival, development time and subsequent pupal weight were also unaffected by foliage type, suggesting that neither foliage type is nutritionally superior for adults or for larvae. However, adult foliage was significantly thicker than juvenile foliage and this may prove a physical constraint to larval establishment. Biotic and abiotic factors (including interactions with natural enemies, competition, microclimate and mate location) that may affect patterns of host plant utilization are discussed.     

Glandular Trichomes on Alfalfa Impede Searching Behavior of the Potato Leafhopper Parasitoid

New cultivars of alfalfa, Medicago sativa L., have been released with glandular trichomes for resistance to potato leafhopper, Empoasca fabae (Harris). Yet, the impact of the glandular trichomes on the primary natural enemy of the leafhopper, Anagrus nigriventris Girault, is unknown. We compared the host searching behavior of the egg parasitoid on four alfalfa clones varying in trichome characters. Female wasps were videotaped on Ranger, a susceptible clone with relatively sparse trichomes, B14, a resistant clone with dense but nonglandular trichomes, and FG12 and FG18, two resistant clones with glandular trichomes. Although the number of leafhopper eggs per stem exposed to wasps did not significantly differ among the four clones, the frequency of foraging and total foraging time were less on the two clones with glandular trichomes than on the two clones with nonglandular trichomes. In addition, an analysis of covariance demonstrated that, although the number of ovipositional probes increased with egg density on a stem, the number of probes on stems with glandular trichomes was significantly less than that on stems without glandular trichomes. The allocation of time by wasps among drumming, probing, and grooming behaviors was similar among the clones. Wasps tended to fly off of clones with glandular trichomes more often than off of clones with nonglandular trichomes. This study suggests that cultivars with glandular trichomes may interfere with host searching by A. nigriventris.     

Vertical stratification of feeding by Japanese beetles within linden tree canopies: selective foraging or height per se?

Adult Japanese beetles, Popillia japonica Newman (Coleoptera: Scarabaeidae), tend to aggregate and feed most heavily in the upper canopy of host plants, defoliating them from the top down. We examined characteristics of linden (Tilia cordata L.) foliage from different canopy zones and tested two hypotheses that might account for vertical stratification of feeding by this vagile, polyphagous folivore. In the field, P. japonica caused 4–12 times more damage to upper canopy leaves in full sunlight than to lower canopy leaves in sun or shade. However, this within-tree pattern apparently cannot be explained by differences in nutritional parameters (protein, water, and sugar content) or defensive properties (toughness, tannins) of leaves. Furthermore, beetles did not discriminate between foliage from different canopy zones in laboratory choice tests, nor were fecundity or longevity higher for beetles fed upper canopy, sun-exposed leaves. Clonal grape plants suspended from vertical posts in the field at 3.65, 1.83 or 0 m above ground showed a top-down defoliation pattern identical to that seen in linden trees. This suggests that the height of foliage per se strongly affects initial orientation and attack by P. japonica. Recent related studies have shown that both sexes of P. japonica are strongly attracted to host volatiles induced by feeding of conspecific beetles. We suggest that Japanese beetles begin to feed in the upper canopy for reasons unrelated to host nutritional variation (e.g., behavioral thermoregulation, visual orientation to the host silhouette), and that top-down defoliation follows as additional beetles are attracted to feeding-induced volatiles acting as aggregation kairomones.     

Feeding rate and attack specialization: the roles of predator experience and energetic tradeoffs

Synopsis Individual mosquitofish, Gambusia affinis, can adopt a broad range of attack selectivities. In part, this variation can be explained by the past experiences of a fish. Individuals selected the more profitable Ceriodaphnia dubia (Cladocera) over less profitable cyclopoid copepods to a greater degree after being exposed to both prey types than did individuals experienced with only one of the prey types. Feeding rate (biomass ingested per unit time) declined with increased attack specialization on the profitable prey (Ceriodaphnia) when such prey were scarce, a result in agreement with assumptions of optimal diet theory. When profitable prey were abundant feeding rate was a bimodal function of the intensity of specialization on profitable prey; fish that specialized on cyclopoid copepods (the less profitable prey type) fed at higher rates than did generalists. This may be the result of antagonistic learning that precluded feeding efficiently on more than one type of prey at a time. The data are consistent with the hypothesis that rejection of unsuitable prey involves a time cost. The two preceeding aspects of foraging behavior, which are absent from most optimal diet models, could lead to failure in predicting the attack specialization of some predators, An additional aspect of the results was the generally weak relationship between feeding efficiency and specialization behavior. This suggests that feeding rate may not have been as tightly linked to the specialization behavior a predator adopts as is assumed by current foraging theory.     

Larval feeding experience influences adult predator acceptance of chemically defended prey

Generalist predatory paper wasps, Polistes dominulus, experience plant secondary defensive compounds as developing larvae through their herbivorous lepidopteran caterpillar prey and as adults through attacking caterpillars while foraging. We evaluated the role that larval and early adult experience with unpalatable prey plays in subsequent foraging choices by adult wasps. For periods of two or four weeks, caged wasps were raised exclusively on caterpillars of either unpalatable Buckeye, Junonia coenia, that sequester substantial levels of iridoid glycosides (IG) or on Painted Lady, Vanessa cardui, that contain very low levels of IG. Wasps were then allowed to forage on both caterpillar species simultaneously. Patterns of prey capture differed significantly based on previous prey experience. Regardless of previous feeding experience, adult wasps overwhelmingly preferred to take Vanessa. Yet Junonia-experienced wasps continued to attack and take back to the nest over 50% more Junonia than did Vanessa-experienced wasps. The longer the wasps' larval experience with Junonia, the more likely they were to capture Junonia caterpillars. However, the life stage at which the wasps experienced Junonia was also influential as young adult experience with the unpalatable prey was more of a deterrent than was experience strictly as larvae for Junonia-experienced wasps. The results demonstrate that, in these predators, previous experience with deterrent chemicals during their larval development alters patterns of prey acceptability to the adult insects.