Host deprivation influence on response of Rhagoletis pomonella to its oviposition deterring pheromone

ABSTRACT. The response of host-depibved Rhagoletis pomonella Walsh (Diptera, Tephritidae) females to host fruit ( Crataegus viridis L.) marked with R.pomonella oviposition deterring pheromone was measured in the laboratory. A positive correlation was found between length of host deprivation (5, 10, 20, 40 or 80 min) and probability of host acceptance (i.e. oviposition) by flies. The results are discussed in relation to current theories on physiological control of insect feeding and oviposition behaviour.     

Influence of host deprivation on egg load and oviposition behaviour of Brachymeria intermedia, a parasitoid of gypsy moth

Abstract. Mated female Brachymeria intermedia (Hymenoptera: Chalcididae) deprived, since emergence, from pupae of their host Lymantria dispar (Lepidoptera: Lymantriidae), accumulated eggs but had a very low rate of hostacceptance. Parasitoids that were host-deprived after encountering pupae early in life also accumulated eggs, but maintained a high acceptance rate. Thus early exposure to hosts promoted active reproductive behaviour. Total egg production depended on the total number of pupae encountered, indicating that B.intermedia adjust their egg production to host availability. Hence, in B. intermedia both the physiological state of the parasitoid (age and egg load) and the informational state (in this case host-availability and experience) interact to shape oviposition behaviour.     

Does fecundity drive the evolution of insect diet?

We investigate whether egg load (a surrogate for fecundity) drives host specificity in a herbivorous insect. In many insects, including our study organism (Edith's checkerspot butterfly), both egg load and tendency to accept low-ranked hosts increase during each search for an oviposition site. Effects on host acceptance of egg load and passage of time are thereby potentially confounded. We conducted two experiments designed to disentangle these effects. In both experiments, we estimated the times of first acceptance of both a high-ranked and a low-ranked host, without allowing the insects to oviposit. In the first experiment, we measured egg load at the time of first acceptance of the low-ranked host. The later the time of first acceptance, the higher was the fecundity. We therefore reject the hypothesis that all insects accepted the low-ranked host at the same predetermined egg load. In the second experiment, we measured egg load 48 h after the high-ranked host was first accepted. We found no relationship between egg load and timing of acceptance of the low-ranked host. Insects with higher rates of egg accumulation did not accept the low-ranked host sooner. Taken together, these results suggest that acceptance of the low-ranked host is not driven directly by egg load. Rather, this acceptance results from some other process that is influenced by time since last oviposition. We conclude that there is no evidence to support the assumption that females with high rates of egg accumulation are more likely to accept low-ranked hosts.     

Behavioral processes and egg-laying decisions of the bean weevil, Callosobruchus maculatus

Empirical studies of behavioral processes were designed in a seed beetle, Callosobruchus maculatus, to distinguish between two alternative tactical models, namely, comparison tactics and threshold tactics of seed choice, and also to deduce the behavioral rule of its oviposition decisions. Search sequences and the oviposition process of the female bruchid beetle were observed and recorded. Analyzing data of the search sequence showed that the female bruchid beetle repeatedly encountered seeds randomly and tended to oviposit on a newly encountered seed in an ovposition bout. These results contradict the usual comparison models, which predict that the female will return to choose any one of the previously encountered seeds after sampling. In addition, the oviposition decision (rejection or acceptance of an encountered seed to oviposit) was analyzed with a generalized linear modeling (GLIM) technique. Modeling showed that the probability of accepting a seed with different numbers of eggs on it changed during her egg-laying process. This evidence supports the idea that the female is using a threshold tactic and that her acceptance threshold is being adjusted by experience gained during the egg-laying process. The analysis of statistical modeling also showed that both the time since the last oviposition and the number of eggs which had been laid by the female had a significant effect on the probability of accepting seeds with different numbers of eggs. Therefore, a time measuring system and the physiological state variable, e.g., eggload of the female, should be included in the behavioral rule to explore how the female makes her decision in the egg-laying process and to examine the importance of choice behavior as a component of selection and adaptation. Received: May 8, 1998 / Accepted: September 20, 1999     

Persistence of experience effects in the parasitoid Trichogramma nr. brassicae

1. The importance of learning in parasitoids is likely to depend on how long learning effects persist, the number of learned stimuli that can be retained, and whether learning effects are confined to low ranked hosts.
2. The duration of learning of host-specific stimuli was tested in four strains of Trichogramma nr. brassicae (Hymenoptera: Trichogrammatidae). Both the persistence of learning of the rearing host and oviposition experience were tested in separate experiments.
3. The response to a non-preferred host, Sitotroga cerealella , was measured at different times following experience on that host or an alternative host, Heliothis punctigera . Both types of experience persisted for at least 5 days in all four strains.
4. The effect of oviposition in a highly preferred host, H. punctigera , following oviposition in S. cerealella was also tested. The increased acceptance of S. cerealella following oviposition on this host persisted despite a subsequent oviposition on H. punctigera .
5. The implications for host range at both individual and population levels are discussed.     

Conserved oviposition preferences in alpine leaf beetle populations despite host shifts and isolation

Abstract.  1. Choosing the plant on which to lay their eggs is the last act of care that most female herbivorous insects bestow upon their offspring. These decisions play a pivotal role in insect–plant interactions, placing host preference under strong selection and contributing to the diversity of phytophagous insects as one of the first traits to adapt to new hosts.
2. This study presents a test of whether extreme isolation and exposure to different host plants can produce intra-specific divergence in oviposition preference in alpine insects. Geographic variation should impose selection to fine-tune host plant ranking and specificity to the plants normally encountered, to avoid wasting time during the very limited reproductive season experienced at high altitudes.
3. Beetles from five populations of Oreina elongata differing in host availability were offered three natural hosts: Cirsium spinosissimum , Adenostyles alliariae , and Adenostyles glabra . A novel application of a continuation ratio model (logistic regression) was made to sequential no-choice experiments, combined with quasi-likelihood analysis of multiple-choice experiments.
4. The results show little geographic variation in host plant choice: all populations strongly preferred Cirsium in multiple-choice trials, and in no-choice experiments laid around 47% of their remaining eggs during each stage, almost regardless of the host present.
5. Enemy-free space seems to explain the preference for Cirsium , but isolation and exposure to different plants has clearly not caused local adaptation in host plant ranking or specificity. Reasons for this conservatism despite divergence in other characteristics are discussed.     

Adaptive value of host discrimination in parasitoids: when host defences are very costly

Host acceptance decision in parasitic wasps strongly depends on the parasitism status of the encountered host. In solitary species, a host only allows the development of a single parasitic larva and then, any oviposition in an already parasitised host leads to larval competition and loss of offspring. Females of many parasitoid species are able to discriminate between parasitised hosts and healthy ones. However, the host discrimination process may require more time than oviposition, exposing the wasp to high risks when the host has efficient defences. Consequently, depending on the degree of success of the host defence, the cost of host inspection for discrimination can outweigh the benefit of superparasitism avoidance. In the present paper, a theoretical approach was developed for determining how host defences may affect optimal host acceptance behaviour in parasitoids. The present model compares the lifetime reproductive success over the strategy used, discrimination and no-discrimination: a discriminating wasp sets a relatively greater value in its current oviposition, while a non-discriminating female sets a greater value in its own survival and future reproduction. The model predicts that depending on physiological state variables and environmental state variables, the optimal policy is not discriminating. Our results suggest that the low discriminating ability observed in some parasitic wasps could probably be an evolutionary response to host defences pressure.     

Host visitation sequence as a determinant of search persistence in fruit parasitic tephritid flies

Summary We tested, through field experiments and simulation models, the hypothesis that fruit-searching tephritid fruit flies adjust their within-tree search persistence according to the sequence and timing of encounters with parasitized (i.e. egg-infested) and unparasitized Crataegus sp. host fruit. In the field, we presented flies with 4 different sequences of unparasitized [=C] and parasitized [=M] fruit: 5M+0C; 1C+5M; 5M+1C, 1C+0M. Following fruit presentation, flies were permitted to forage freely within trees, which harboured no fruit, until emigration occurred. Under these conditions, flies that encountered the aforementioned different sequences of hosts, displayed differences in Giving Up Time, measured as active foraging time and number of leaf visits, in a manner predicted prior to testing. These differences were, however, not statistically significant. Based upon the results described above, we then built 3 simulation models that predicted within-tree Giving Up Times for individual flies: Model 1-Giving Up Time is incremented and decremented by fixed amounts following encounters by the fly with suitable (i.e. for oviposition) and unsuitable hosts, respectively; Model 2-similar to Model 1, but increment and decrement values are variable and are dependent upon the time since previous encounters by the fly with suitable and unsuitable hosts; Model 3-Giving Up Time is fixed. Comparison with previously reported field data for tephritid flies showed that Model 2 predicted rather well, and significantly better than Models 1 and 3, Giving Up Time for wild type tephritid flies under seminatural field conditions. We discuss our results in light of contemporary foraging theory.     

Motivation models fail to explain oviposition behaviour in the diamondback moth

Abstract. It is widely accepted that previous experience and internal physiological factors (such as egg‐load) affect host‐plant discrimination during oviposition by phytophagous insects. However, there is some debate as to how these factors interact in a mechanistic sense to control acceptance. The role of learning and host deprivation in host acceptance by adult diamondback moths (Plutella xylostella L.) was investigated. In the first experiment, we tested whether experience of a lower ranked host cabbage increased acceptance of a higher ranked host as predicted by a hierarchy threshold model. Moths trained on cabbage were over three times more likely to accept cabbage during testing than untrained moths. There was no effect of cabbage training on acceptance of cress, indicating that the effect of training was species‐specific. In a second experiment, designed to test the prediction of motivation models that insects become less discriminating when deprived of oviposition opportunities, depriving females of host plants for 2 nights significantly increased female egg‐load (×2.3). Host deprivation did not decrease discrimination between the preferred host cress and cabbage. Cabbage and cress plants were equally likely to have been accepted by nondeprived moths after 1 night of exposure, yet moths deprived of hosts for 2 nights strongly preferred cress when tested during the first 20 min of the scotophase. During this 20‐min period, previous host deprivation increased acceptance of host plants generally but did not decrease discrimination between hosts. These data contradict the expectation that there is an inverse relationship between host species discrimination and the failure of an insect to find hosts as found in existing oviposition acceptance models. As an alternative, the Incremental Acceptance Model of host acceptance behaviour is presented, in which responsiveness to a host is a function of the recent encounter rate with host‐specific stimuli, and the oviposition reflex is regulated by nonspecific cues such as egg‐load.     

Host-feeding and acceptance by a parasitic wasp (Hymenoptera: Ichneumonidae) as influenced by egg load and experience in a patch

Optimal host selection models based on dynamic programming predict that the physiological state of a foraging insect, i.e. egg load, energy reserves etc., influences behavioral decisions. To test this prediction, the effect of physiological state on host acceptance of the ectoparasitic wasp Agrothereutes lanceolatus was investigated. Female wasps in plastic cups (regarded as patches) were presented with hosts, and their responses to the hosts were continuously observed. After observations, the wasps were dissected and the number of mature and immature eggs they carried were counted. The results showed that behavioral decisions by the female wasps were influenced by mature egg load, but not by wasp size or immature egg load. Hence the wasps with higher egg loads were more likely to oviposit. The number of hosts previously encountered in a patch (i.e. wasp experience) also had an independent effect on females' host acceptance, indicating that female informational state was updated during foraging in that patch. Female wasps host-fed only when mature egg load approached zero. Concurrent host-feeding was not observed. Parasitoid survival was almost zero when parasitoid eggs were transferred onto hosts that were fed upon, indicating that concurrent host-feeding could cause a high degree of offspring mortality. These three results supported the assumption and prediction of optimal host-feeding models. Parasitoid host selection and host-feeding are discussed in the context of recent models.     

Host-marking behaviour as a quantitative signal of competition in the walnut fly Rhagoletis juglandis

Abstract.  1. Walnut-infesting flies in the Rhagoletis suavis species group actively re-use hosts for oviposition despite engaging in a genus-typical host-marking behaviour which, in other Rhagoletis groups, deters oviposition. In a study of the walnut fly, R. juglandis (Cresson), alternative hypotheses for the putative marking behaviour were evaluated.
2. The oviposition site attraction hypothesis proposes that the host mark guides females to oviposition sites on occupied fruit. The competition intensity signal hypothesis proposes that the host mark is an indicator of the level of competition to be incurred if fruit are re-used.
3. In a field cage, females were presented simultaneously with fruit previously exposed to 25 females that were also allowed to oviposit and engage in the putative marking behaviour, and control fruit on which females were allowed only to oviposit. The occurrence of host marking reduced a female's propensity to oviposit from 46% to just over 10%, consistent with the competition intensity signal hypothesis only.
4. In a laboratory assay, the duration of host marking was correlated positively with the size of a female's clutch. This result, also consistent with the competition intensity signal hypothesis, suggests that the amount of marking pheromone on a fruit is a reliable indicator of the number of eggs already deposited within.
5. In a second field-cage experiment, females were allowed to mark on fruit for 0, 10, 20, or 30 min and fruit were presented to test females. Whether or not females alighted on a particular host was not affected by the duration of marking; however, the frequency of both ovipositor probing and egg deposition decreased with increasing duration of marking. Consistent with the competition intensity signal hypothesis, this result suggests that the host mark permits females to assess the level of competition that a clutch will incur within re-used fruit.     

Insect oviposition plasticity in response to host availability: the case of the tephritid fruit fly Bactrocera dorsalis

1. Insect oviposition behaviour is ecologically and physiologically plastic. For tephritid fruit flies, Bactrocera dorsalis Hendel, host availability varies spatially and temporally. Females are expected to adopt adaptive oviposition strategies to maximise lifetime reproductive fitness, including survival. Bactrocera dorsalis oviposition tactics in response to different host availabilities were investigated. 2. This study includes three treatments: (i) variable host densities (host density varied according to a fixed cycle from day to day over values of 1, 5, 10 and 20 hosts per cage), (ii) a fixed high host density (20 hosts per cage), and (iii) a fixed low host density (1 host per cage). 3. Daily egg‐laying number per female over the course of 27 days was entirely independent of host density and highly dependent on female age. As host availability increased, females accepted significantly more hosts, generally laid small egg clutches, and more broadly distributed the eggs. 4. Tephritid fruit flies adaptively adjusted egg clutches in ways that reflected the variability of host availability. Egg‐ and time‐limitation constraints appeared to drive these adjustments. Female egg maturation was triggered by oviposition activity and reflected marked lifetime trade‐offs. Such strategies involved specific time schedules for egg laying. 5.This study defined the oviposition plasticity of the tephritid fruit fly. These results have general implications for the behavioural ecology of insect herbivores and parasitoids.     

Influence of intertree distance on foraging behaviour of Rhagoletis pomonella in the field

Abstract. 1. We tested a prediction from contemporary foraging theory that animals should decrease their allocation of energy to the searching of individual patches when interpatch travel costs decrease.
2. We used individual Rhagoletis pomonella Walsh (Diptera: Tephritidae) females foraging for oviposition sites (= Crataegus fruit) in a host tree which was surrounded by four other trees at varying distances.
3. We found that flies generally invested less search, measured as time spent searching a tree or number of leaves visited on a tree, when neighbouring trees were nearby than when farther away.
4. Under our test conditions, flies appeared to have difficulty locating neighbouring trees at a distance of more than 1.6 m.
5. Our study calls into question the interpretation of search effort by insects within resource patches in the absence of information on interpatch distances.     

Effects of egg load on finding and acceptance of host fruit in Ceratitis capitata flies

Abstract. In studies conducted on potted host trees in field cages and in the laboratory, we examined the influence of egg load on the finding and acceptance of high-ranking (kumquat) and lower-ranking (grapefruit) hosts for oviposition by wild-origin Mediterranean fruit fly females, Ceratitis capitata (Wiedemann). By prescribing the periods during which females had access to protein prior to testing, we generated four classes of females having progressively increasing egg loads but not differing in population origin, age, degree of protein hunger at testing, or amount of prior experience with host fruit (none). Egg load had no discernible effect on behaviour associated with finding either type of fruit but did have a significant effect on several behaviours associated with oviposition after alighting on fruit. Increasing egg load led to increasing propensity to engage in ovipositional-type behaviour on both kumquats and grapefruits. There was no evidence, however, to support a hypothesis that medflies would become less discriminating against grapefruits relative to kumquats as egg load increased. Relative to kumquats, grapefruits were accepted for oviposition by intermediate and high egg load females to a substantially greater degree in laboratory cages than on trees, suggesting that results of laboratory cage experiments on host discrimination by tephritid flies may poorly reflect differences in behavioural responses expressed under less constrained conditions.     

Behavioral responses of female oriental fruit flies to the odor of papayas at three ripeness stages in a laboratory flight tunnel (Diptera: Tephritidae)

Behavioral responses of adult female oriental fruit flies, Dacus dorsalisHendel, to the odor of papayas from three ripeness classes were studied using a threechoice flight tunnel bioassay. Laboratoryreared flies were allowed to respond freely to any of three papaya odors (mature green, colorbreak to one-fourth ripe, and one-half to full ripe) emanating from identical (spherical) fruit models. Five behaviors were measured in assessing the fly's relative attraction to the odors (number of landings), arrestment (total fly seconds on sphere), fly-fly interactions on the fruit model (maximum and modal fly density), and acceptance for oviposition (total eggs laid). Females showed no significant difference in total fly landings based on all age classes combined. Significant differences were noted among age classes. Females spent more total time on the sphere and showed a higher maximum density and modal fly density to ripe fruit than to green fruit odors. Ovipositional acceptance of fruit models based on the total number of eggs laid in a sphere was greater in response to the ripefruit odor than to the other two odor classes. Olfactorystimulated behavioral responses of females to the odor of ripe papayas were significantly different from the other ripeness classes for all behaviors at 8 days postemergence and then declined in 11-day-old flies. Behavioral responses were greater during the afternoon than in the morning. Observations of wild oriental fruit flies to papayas in the field indicated a preference for residing on riper fruit. The results of this study are discussed with regard to the role of olfactory inputs generated by the odor of ripening fruit on female attraction and oviposition behavior resulting in infestation of papayas by oriental fruit fly.     

Pre-oviposition mate-guarding and mating behaviour of Argia vivida (Odonata: Coenagrionidae)

Abstract. 1. At Halcyon Hotsprings, British Columbia, Canada, male and female Argia vivida Hagen encountered to mate in two different ways.
2. In the morning (before 12.30 hours solar time), males basked at sunspots in the forest and darted out at passing females, attempting to take them in tandem (the first method of encounter).
3. If a male was successful, the pair engaged in a 31.3±4.8 min copulation followed by an hour of tandem flight before beginning oviposition.
4. As the day progressed, unmated males moved slowly toward the water and arrived at the water at about the same time as the earliest ovipositing pairs (1131±27.5 min solar time).
5. Males retained their grasp on their mates during oviposition (contact-guarding) but since some tandems separated during oviposition, non-tandem males at the water could capture recently released, gravid females (the second method of encounter).
6. The new pairs performed a brief copulation (10.2±3.38 min) and began ovipositing immediately thereafter.
7. Some females that avoided recapture attempted to oviposit unguarded.
8. We believe the long duration of morning copulations and period of tandem constitute a male strategy, which we call 'pre-oviposition guarding', to guard females until it is warm enough at the oviposition site for the females to begin ovipositing.
9. Separation of tandems during oviposition may be initiated by either member of the pair and we suggest that one benefit to a female of leaving a guarding mate is increased efficiency of oviposition when the intensity of male harassment is low.
10. The mating system of A. vivida thus comprises a series of complementary male and female mating behaviours.     

Physical and chemical cues affect oviposition by Neoleucinodes elegantalis

Recognition and acceptance of a suitable host plant by phytophagous insects requires the integration of visual, physical and chemical cues. The present study investigates the host cues that a specialist insect integrates to optimize oviposition decisions and whether these cues are weighted in a specific way. The study also determines whether the tomato fruit borer Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae), an important pest on Solanaceae in Brazil, shows a preference for oviposition sites that differ in physical and/or chemical cues. When styrofoam balls are provided as artificial fruits, N. elegantalis deposit significantly more eggs on rough artificial fruits than on smooth ones. Hexane fruit extracts applied to the artificial fruits stimulate female oviposition strongly. Physical and chemical cues also affect the oviposition of females when offered together. Furthermore, certain parts of the artificial fruits are prefered, irrespective of the presence of chemical cues. Both physical and chemical cues affect oviposition decisions; hence, the fruit borer relies on cues of different sensory modalities.     

Learning pays off: influence of experience on host finding and parasitism in Lariophagus distinguendus

Abstract. 1. The influence of experience on egg maturation, parasitism rate, and behaviour during host searching was investigated for Lariophagus distinguendus (Först.) (Hymenoptera: Pteromalidae) parasitizing larvae of the granary weevil Sitophilus granarius (L.) in grains of wheat Triticum aestivum L.
2. Dissection of female parasitoids and parasitism bioassays at high host density revealed that experience with hosts (e.g. by oviposition or by host feeding) is not required either for triggering oogenesis or for oviposition.
3. In parasitism experiments at low host density, when single host-infested grains were offered within a bulk of healthy grains, host finding and parasitism rate were increased by experience.
4. Behavioural observations revealed that searching time required for finding an infested grain was shorter for experienced parasitoids than for naive parasitoids, because travel time from grain to grain is shorter for experienced parasitoids, and because experienced parasitoids spend less time than naive parasitoids on non-infested grains.
5. In conclusion, experience due to host exposure increases parasitism and thereby the fitness of the parasitoids. It is discussed that this increase is more likely due to learning than to different egg load dynamics of experienced parasitoids.     

Host-plant use in the range expansion of the pine processionary moth, Thaumetopoea pityocampa

Abstract.  1. During range expansions of phytophagous insects, secondary or novel hosts may allow colonisation of areas without primary hosts. Because plant species often differ in their relative attractiveness and suitability for insects, insect preference for, and performance on, these hosts can determine recruitment potential in the current and future expansion areas.
2. This study explores the relative roles of female preference and larval performance in an important pine defoliator, Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera, Notodontidae), which colonises three Pinus species at its current range margin in the Italian Alps: P. nigra (primary host), P. sylvestris (secondary host), and P. mugo (novel host).
3. Host use patterns in multiple insect populations were studied through choice and no-choice oviposition experiments in cages, field surveys of mixed stands, and laboratory and field monitoring of larval growth and mortality. It was predicted that a specific life-history trait – time limitation of short-lived females to lay a single batch of eggs – would act as a component of female performance, and lead to similar rates of host acceptance in no-choice settings.
4. In the choice experiment, P. nigra was accepted the most frequently while P. sylvestris was accepted the least frequently, confirming nest density patterns in the field. Contrary to prediction, females remained discriminating in no-choice settings in spite of time limitation. In contrast, relative growth rate (RGR) and mortality of larvae did not differ significantly among the three hosts, highlighting a discrepancy between female preference and larval performance.
5. Recruitment potential of T. pityocampa in future expansion into stands of P. sylvestris and P. mugo is evaluated by combining host quality, conservatism in oviposition behaviour, habitat suitability, and the opportunity for local adaptation.     

Low oviposition stimuli reduce egg production in the pollen beetle Meligethes aeneus

Abstract. Gravid female pollen beetles, Meligethes aeneus (F.) (Coleoptera: Nitidulidae), were exposed at different intervals to oviposition sites that varied in acceptability.The egg load of dissected individuals which had been deprived of oviposition sites was not greater than individuals which had been frequently exposed to oviposition sites.However, the egg load of individuals which had been exposed to highly acceptable host plants was greater than those exposed to a host plant of low acceptability.Over the experimental period the total number of eggs which an individual produced was lower for those that were either deprived of an oviposition site or frequently exposed to a low-quality host plant.There was no evidence of either increased oviposition probability or clutch size as the period since the last oviposition increased.It is concluded that low host quality and low host encounter rate reduce the egg production of M.aeneus. It is further concluded that the suppressed egg production reduces the accumulation of eggs, so that physiological motivation for the insect to oviposit is not increased.If low oviposition site quality acts to reduce oogenesis, as found here, then the probability of egg deposition on low-quality species may not increase over time and the effect of antixenotic resistance may be enhanced.