Learning of host searching cues by the larval parasitoid Microplitis croceipes

We conducted laboratory experiments using a wind tunnel to determine the effects of prior experience on the learning and retention of learned responses in the larval parasitoid Microplitis croceipes (Cresson) (Hymenoptera: Braconidae). Although most females that had either antennated host frass or oviposited in a host in the presence of vanilla odor, made oriented flight to the odor from downwind in the wind tunnel at 30 min after experience, only those that had oviposited in a host with the odor responded to the odor at 24 h after experience. Females that had oviposited in the non‐natural host beet armyworm (BAW) larva with or without the odor did not respond to the odor at 30 min after experience. These results indicate that an oviposition in the host in the presence of odors strongly affects associative learning and the persistence of learned response to the odors. When females were allowed to antennate host frass in the presence of vanilla and to subsequently oviposit in a host within an interval of 5 min or less, their learned response to vanilla also persisted for 24 h. Similarly, when females were conditioned to link vanilla with host frass and then allowed to make an ovipositor contact with host hemolymph, their learned response persisted for 24 h. However, antennal contact with hemolymph after such conditioning of vanilla with host frass did not improve the persistence of learned response to the odor. These results indicate that ovipositor contact with host hemolymph during oviposition is partially responsible for an increased retention of learned response. Females responded to vanilla 48% of the time at 30 min after antennating host frass without the odor, but their response to the odor significantly decreased after oviposition in a BAW larva subsequent to the antennation of host frass. This result indicates that oviposition in a BAW larva decreases subsequent response to general odors. Based on the results, we discuss the foraging behavior of M. croceipes dependent on learning and subsequent experiences.     

Foraging for patchily-distributed leaf miners by the parasitic wasp,Dacnusa sibirica

We studied the rules used by the female parasitoid, Dacnusa sibirica Telenga (Hymenoptera: Braconidae), for deciding when to leave a tomato leaflet on which she is searching for larvae of the leafminer, Liriomyza bryoniae Kalt. (Diptera: Agromyzidae). Females would deposit a marking pheromone on the leaflet and would leave the leaflet when the amount of the pheromone accumulated to the threshold L, which is proportional to the amount of search effort on the leaflet. L appears to increase with host density since it rises after every encounter with a host (or mine).D. sibirica would employ an area-concentrated search, which is advantageous in foraging for hosts showing a clumped distribution.     

Host plant, oviposition behavior and larval ecology of a sawfly leafminer, Profenusa japonica (Hymenoptera: Tenthredinidae)

The host plant, oviposition behavior and larval ecology of Profenusa japonica Togashi are reported for the first time. Adults of P. japonica mated and oviposited on a polyantha rose, Rosa multiflora (Rosaceae), in April. Each female adult laid an egg on the edge of a leaflet. Hatched larvae consumed the parenchymatous layer of leaflets and in so doing created a mine. The larval stage comprised five instars. On average, 70.6% of the total area of a leaflet was consumed by one larva. Female adults of P. japonica laid eggs singly, probably to avoid larval competition for food.     

Reproductive biology of the microtype tachinid fly Zenillia dolosa (Meigen) (Diptera: Tachinidae)

Reproductive biology including mating, adult longevity, fecundity and development of the tachinid fly Zenillia dolosa was investigated for optimizing rearing procedures using Mythimna separata as a host in the laboratory. Females lay microtype eggs containing a first instar larva on food plants of the host and then the eggs must be ingested by the host for parasitization. Mating success was 58.5% with mating duration of 80.7 min. Mating was most successful when day 0–1 females were kept with day 2–4 male flies. Female body size was positively correlated with its fecundity but not with longevity. However, females that survived longer produced more eggs during their lifetime. Parasitoids successfully developed in 4th to 6th instar host larvae. Host instars at the time of parasitoid egg ingestion significantly influenced development time of the immature parasitoid, but did not affect body size of the emerging parasitoid. We suggest that pairing newly emerged females with day 2–4 males should result in higher mating success and using the last instar hosts for parasitization should minimize development time of the parasitoid for rearing.     

Factors that affect the selection of tomato leaflets by two whiteflies,Trialeurodes vaporariorum and Bemisia tabaci (Homoptera: Aleyrodidae)

In Japan, although greenhouse whitefly, Trialeurodes vaporariorum (Westwood), and sweet potato whitefly, Bemisia tabaci (Gennadius), co-occur on tomato plants under greenhouse conditions, the two whiteflies are distributed differently with regard to leaf position. To elucidate the factors that determine the leaf position of these whiteflies, we investigated traits for leaflets collected from three positions on tomato plants. Furthermore, we examined leaflet selection by and fertility of the two whiteflies under choice and non-choice conditions. In addition, the effect on whitefly behavior of volatile compounds released from leaflets was evaluated by use of a Y-tube olfactometer test. Nitrogen and carbon content were highest for upper leaflets. In choice tests, more T. vaporariorum and B. tabaci adults selected upper and middle leaflets, respectively. Similarly, they oviposited more eggs on upper and middle leaflets. In non-choice tests, T. vaporariorum oviposited more eggs on upper leaflets, but B. tabaci oviposited equally on each leaflet. In Y-tube olfactometer tests, more T. vaporariorum adults moved to upper leaflets whereas more B. tabaci adults moved to middle leaflets. These results suggest that different leaflet selection by adults of these two whiteflies is likely to be associated with the different volatile compounds emitted by tomato leaflets at each position.     

A biological assessment of Apophua simplicipes (Hymenoptera: Ichneumonidae) as a parasitoid of the oblique banded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

Apophua simplicipes (Cresson) (Hymenoptera: Ichneumonidae) is a common parasitoid of the oblique banded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) in organically managed apple orchards in the southern interior of British Columbia, Canada. The biological characteristics of a laboratory colony of A. simplicipes were studied. When held at 15 and 25°C and provided with honey water, individual females survived an average of 60.6±6.1 and 29.8±4.7 days, oviposited 196.7±50.7 and 326.6±51.3 eggs and parasitized a total of 163.4±40.4 and 229.4±35.8 hosts, respectively. Females oviposited into first through fourth instar oblique banded leafrollers, with significantly more parasitism occurring in the first two instars compared to the third and fourth instars. No parasitoid larvae survived past the first larval stage in parasitized fourth instar hosts. Apophua simplicipes did not parasitize larvae of three-lined leafroller, Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) which is sympatric with oblique banded leafrollers in orchards in the southern interior of British Columbia. Female predation and host feeding from wounds on early instars of both leafroller species was observed under laboratory conditions. In addition, early instar hosts exited diet feeding sites in response to the probing activity of the ovipositing wasps. A similar escape reaction in the orchard may cause a leafroller larva to move away from its feeding site, making it more vulnerable to predation or movement off the tree. Apophua simplicipes larvae emerged from fifth and sixth instar hosts. Parasitized oblique banded leafroller hosts consumed significantly less meridic diet than unparasitized female larvae from fifth instar through to parasitoid emergence or leafroller pupation. Our laboratory results suggest that A. simplicipes may reduce field populations of oblique banded leafroller and decrease pest feeding damage.     

INTRAPOPULATIONAL VARIATION IN LEAFLET MORPHOLOGY OF ZAMIA PUMILA L. IN RELATION TO MICROENVIRONMENT AND SEX

The cycad Zamia pumila L. was examined for differences in leaflet morphology between sun and shade plants and between males and females within a single population in northern Puerto Rico. Sun leaflets are significantly smaller than shade leaflets in length, width, and surface area; and sun leaflets have a higher length: width ratio than shade leaflets. Also, average densitythickness is greater for sun leaflets than for shade leaflets. Males and females are similar in leaflet size; but the females had a larger number of leaflets per leaf than males.     

Butterfly Host Plant Choice in the Face of Possible Confusion

We tested predictions from the theory that ovipositing females of phytophagous insects are limited by their neural capacity for information processing. Previous studies have found that relatively specialized insects make faster and/or more accurate identifications of host plants compared to generalists. The study species was the polyphagous comma butterfly, Polygonia c-album (Nymphalidae). We compared females originating from two populations (Sweden and England) which differ in degree of specialization on the preferred host Urtica dioica (Urticaceae). Females were given a choice between this plant and a very similar nonhost, white dead nettle, Laminum album (Lamiacease), or a choice between a relatively poor host, Betuala pubescens, and the nonhost Betula pendula (Betulaceae). Oviposition rate was lower in cages with Betula compared to cages with Urtica, demonstrating that P. c-album females will withhold eggs when preferred hosts are not available. As predicted, females originating from the Swedish generalist population oviposited more often on the nonhost Lamium. However, females of both populations discriminated very strongly against oviposition on B. pendula. We found that newly hatched larvae have some ability to move from herbaceous nonhost to hosts. Although alternative interpretations are possible, the results give further support to the hypothesis that there are trade-offs between diet breadth and the ability to discriminate among plants.     

Foraging for patchily-distributed leaf-miners by the parasitoid,Dapsilarthra rufiventris (Hymenoptera: Braconidae). II. Stopping rule for host search

We studied the stopping rule which the female parasitoid,Dapsilarthra rufiventris, uses for deciding when to leave the leaflet on which she is searching for leaf-mining larvae,Phytomyza ranunculi. She is unlikely to employ some current stopping rules, such as fixed-number and fixed-time rules and others. The searching female appears to deposit a marking pheromone on the leaflet. We formulated a model for predicting the amount of pheromone accumulated on the leaflet. The model assumes that she will deposit the pheromone on the leaflet at a given rate (a) per unit time which is proportional to search speed, and will leave it when the amount of pheromone reaches the thresholdL. In this modelL denotes the amount of the search effort spent on the leaflet. The model was fitted fairly well to the data. A comparison of the observed results with the predictions of the model suggests thatL increases markedly at the first encounter with the mine and at a lower rate for the subsequent encounters. This appears to be a kind of area-concentrated search, that is, searching for hosts for a while in the immediate vicinity after finding one host, and would be adaptive in foraging forP. ranunculi larvae, which exhibit clumped distributions among leaflets in the field.     

Effect of body size and sugar meals on oviposition of the yellow fever mosquito,Aedes aegypti (Diptera: Culicidae)

The effects of dietary sugar and body size on the oviposition of Ae. aegypti were studied under laboratory conditions. In female mosquitoes provided with sugar, the start of maximum fecundity was significantly delayed and the oviposition period was longer than in females provided with water. The peak of oviposition was also delayed in sugar‐fed females. Large females oviposited more eggs per day than small females at maximum fecundity and during eight days of observations. Large females also visited significantly more water‐containing cups in their cages per day than small females at maximum fecundity. During the eight days of observations, large females and sugar‐fed females visited more water‐containing cups in their cages than water‐fed small females. Both large females and sugar‐fed females oviposited their eggs at sites higher above the water line than water‐fed small females. These results suggested that large and sugar‐fed female Ae. aegypti mosquitoes had more energy reserves and oviposited their eggs at higher sites, which would lead to a time lag in hatching.     

Prostephanus truncatus mate choice on contact: does pheromone signalling by males affect their mating success?

Prostephanus truncatus is an economically important beetle pest of stored maize and cassava in the tropics. Male beetles signal using an aggregation pheromone that attracts both female and male beetles over large distances. Females preferentially orientate towards the pheromone signals of particular males when given a choice. The influence of pheromone signalling on courtship and mating success was investigated using pheromone biossays and mating trials in both of which a female made a choice between two males. Signalling was manipulated by exposing males to a Female Factor that inhibits pheromone production. The relative attractiveness of males to females based on pheromone bioassays was found not to influence short-range courtship behaviour or mating success.     

Oviposition site selection of Episyrphus balteatus

The oviposition behaviour of Episyrphus balteatus DeGeer (Diptera: Syrphidae) was investigated in four-choice experiments in screen cages and in experiments with direct behavioural observations. Females laid hardly any eggs in the absence of aphids and preferred larger aphid colonies to small ones. Honeydew attracted the females via olfaction and elicited oviposition. Females were able to perceive the presence of conspecific eggs and oviposited less often in aphid colonies in which eggs were present. The oviposition deterring stimulus was also active when the eggs were removed, hence, the deterrent did not act visually, but probably olfactorily.     

Responses to aphid sex pheromones by the pea aphid parasitoids Aphidius ervi and Aphidius eadyi

Aphidius ervi and Aphidius eadyi, two parasitoids of the pea aphid Acyrthosiphon pisum, were attracted to components of the aphid sex pheromone in laboratory bioassays. Pre-test experience with host aphids in the presence of aphid sex pheromone did not affect the response of A. ervi to pheromone in a 4-way olfactometer, compared with that of naive parasitoids. Aphidius ervi females exposed only to the pheromone prior to testing did not respond in the olfactometer, suggesting habituation to the foraging cue by the parasitoid. In a wind tunnel, aphid sex pheromone increased the attraction of A. ervi to the plant-host complex (Vicia faba/A. pisum), suggesting an additive effect when two different foraging cues are present simultaneously.     

Evidence for acoustic communication in the parasitoid wasp Glyptapanteles flavicoxis

Females of the parasitoid wasp Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae) deposit sex pheromone on substrate that elicits attraction and wing fanning in conspecific males. We tested the hypothesis that wing fanning sound induces a behavioral response from females which, in turn, affects the males’ orientation toward them. Females exposed to playback of the males’ wing fanning sound engaged in short flights, with sound characteristics different from those of the males’ wing fanning sound. In two‐choice bioassays, playback of the females’ flight sound attracted significantly more males than a silent control stimulus, and in combination with pheromone‐containing body extract of females it attracted more males than female body extract alone. Our data support the conclusion that the males’ wing fanning induces sound and visual reply signals from females that help males orient toward them.     

Floral resources impact longevity and oviposition rate of a parasitoid in the field

1. The use of floral resource subsidies to improve herbivore suppression by parasitoids requires certain trophic interactions and physiological changes to occur. While the longevity and fecundity of parasitoids are positively affected by nectar subsidies in laboratory studies, the impacts of floral subsidies on the fecundity and longevity of freely foraging parasitoids have not been studied. 2. We studied the longevity and per capita fecundity of naturally occurring Diadegma insulare foraging in cabbage plots with and without borders of flowering buckwheat, Fagopyrum esculentum, as well as relationships between longevity, fecundity, sugar feeding and parasitism rates on larvae of the diamondback moth, Plutella xylostella. 3. Relative longevity was estimated by counting broken setae on the fringe of the forewing. Floral borders increased the longevity of males and females in adjacent cabbage plots. 4. The egg maturation rate of D. insulare was estimated by comparing egg loads of females collected early in the day with egg loads of females held without hosts in field cages throughout the day. Females in buckwheat cages matured 2.7 eggs per hour while females in control cages resorbed 0.27 eggs over the same time period. 5. The fecundity of females collected in the afternoon was estimated by comparing their actual egg load to the estimated egg load in the absence of oviposition for females in a given plot. Females foraging in buckwheat plots had marginally fewer eggs remaining in their ovaries, and laid marginally more eggs than females in control plots. Females from both treatments carried 30-60 eggs by the afternoon and therefore were time-limited rather than egg-limited. 6. Plots where a greater proportion of females had fed on sugar had longer-lived females. This suggests that feeding enhanced longevity of D. insulare. However, plots with longer-lived and more fecund females did not exhibit higher parasitism rates, although the power of these tests were low.     

Sex differences in diet and foraging behavior in white-faced capuchins (Cebus capucinus)

I examined sex differences in diet and foraging behavior in two groups of white-faced capuchin monkeys (Cebus capucinus)in a tropical dry forest at Santa Rosa National Park, Costa Rica. I tested three hypotheses: sex differences in diet and foraging behavior are best explained by (1) sexual dimorphism, (2) the energy demands of pregnancy and lactation for females;and (3) avoidance of competition between the sexes. Sexual dimorphism offered the best explanation of sex differences in the diet and foraging behavior of C. capucinus,accurately predicting that males do more strenuous foraging activity, make less use of small foraging supports, and spend more time on or near the ground. Females spent more time foraging than males did but probably obtained a lower protein yield per unit foraging time. Females exploited more small and embedded invertebrates, while males ate more large invertebrates and vertebrate prey. Pregnant and lactating females spent more time resting and less time foraging than other females did, increasing their foraging return by focusing on foods requiring little handling. There was little evidence of competition avoidance between the sexes.     

Stopping rule of host search by the parasitoid,Chrysocharis pentheus (Hymenoptera: Eulophidae), in host patches

We studied the stopping rule obeyed by the female parasitoid, Chrysocharis pentheus, in deciding when to leave the leaflet on which she is searching for larvae of Phytomyza ranunculi. She seemed not to employ some stopping rules that have been suggested; i.e., a fixed-number rule and a fixed-time rule and others. The stopping model formulated for Dapsilarthra rufiventris parasitic on the same host species fitted well to the results. The model assumes that the searching female will deposit a marking pheromone on the leaflet at a rate proportional to the search speed and will leave the leaflet when the amount of the pheromone that has accumulated on the leaflet reaches the threshold, L. In this model, L denotes the amount of search effort spent on the leaflet. A comparison of the observed results with the predictions from the model suggested that L increased markedly at the first encounter with the mine (host), but less at later encounters. C. pentheus appears to employ a mixed strategy of a fixed search-effort and an area-concentrated search. This would confer an adaptive advantage in foraging for P. ranunculi larvae, which are distributed in clumps among leaflets in the field.     

Effect of Mating Disruption on Reproductive Behavior in the European Pine Sawfly, Neodiprion sertifer (Hymenoptera: Diprionidae)

Females of the European pine sawfly, Neodiprion sertifer (Geoffr.) (Hymenoptera: Diprionidae), were released and observed inside and outside (=control) of areas treated with their sex pheromone for the purpose of mating disruption. In 1992 and 1993, respectively, 0 and 2.5% of the females were observed mating in the pheromone treated area during the first day, compared with 28 and 26% in the control area. Of the females that mated, approximately 50% stayed on their twigs and oviposited, while the remaining 50% disappeared. Significantly more females disappeared from their twigs inside the treated area compared to the control area in both years. No difference was found in the proportion of unmated females ovipositing (2–12%) between the treated and the control area. Predation was responsible for much of the disappearance, but deliberate dispersal was also noted. Mating and dispersal in female N. sertifer are discussed in relation to population density and overall fitness.     

Microhabitat use by larvae and females of a rare barrens butterfly,frosted elfin (<Emphasis Type="Italic">Callophrys irus</Emphasis>)

The frosted elfin (Callophrys irus) is a localized and declining butterfly found in xeric open habitats maintained by disturbance. We described the effects of woody plant canopy cover, topography and host plant size and density on the quality of microhabitat of wild indigo (Baptisia tinctoria) host plants containing late instar frosted elfin larvae at four study sites in southeastern Massachusetts, United States. We also assessed whether females preferentially depositing eggs on host plants within specific microhabitats, therefore conferring greater survivorship to the larvae through the late-instar stage. We found that moderate amounts of canopy cover and large plant size characterized larvae-occupied host plants. In the absence of tree canopy cover, late instar larvae density remained low even when host plant density was high. However, females oviposited on wild indigo plants without regard to any of the vegetative or environmental variables we measured. These results indicate that canopy cover was an important characteristic of microhabitats containing late instar larvae, and late instar larvae occupancy was determined by suitable microhabitat conditions, and not female oviposition selection. Managing for canopy cover and microhabitat heterogeneity within relatively open habitats is recommended for the maintenance of frosted elfin populations.     

Avoidance of occupied hosts by the Hessian fly: oviposition behaviour and consequences for larval survival

Abstract 1. Unoccupied wheat plants and wheat plants occupied by conspecific eggs or larvae were presented to ovipositing female Hessian flies in choice tests.
2. The presence of conspecific eggs on the leaf surfaces of wheat plants did not appear to have any effect on the responses of ovipositing Hessian fly females.
3. The presence of conspecific larvae at the base and nodes of wheat plants for 1, 6, or 11 days had significant effects on Hessian fly oviposition. Eggs oviposited on plants were inversely proportional to larval densities and days of larval occupation.
4. Feeding by Hessian fly larvae is associated with several changes in wheat plants. One of these changes, the growth arrestment of the plant, was measured by recording the heights of plants used in oviposition tests. Plant heights were inversely proportional to both larval densities and days of occupation. Plant heights were directly proportional to eggs oviposited on plants.
5. The consequences of adult female avoidance of plants occupied by conspecific larvae were investigated by allowing females to oviposit on unoccupied plants and 1-day, 6-day, and 11-day larval occupied plants, then scoring at the end of the first larval instar the survival of the offspring that resulted from this oviposition.
6. Survival during the first larval instar was 88% for the offspring of females that oviposited on unoccupied plants, decreasing to 82, 31, and 4% on the 1-day, 6-day, and 11-day occupied plant treatments. On these four plant treatments, a positive correlation was found between larval performance (i.e. survival) and the preferences of ovipositing females.
7. On the four plant treatments, relationships between first-instar larval density and first-instar larval survival varied significantly. On unoccupied plants, survival was inversely proportional to density. On plants oviposited on at 6 days of larval occupation, survival was directly proportional to density.