The roles of adult and larval specialisations in limiting the occurrence of five species of Dacus (Diptera: tephritidae) in cultivated fruits

Summary The relative importance of adult preferences or specialisations of larval physiology in restricting the host range of five species of Dacine fruit flies (Diptera: Tephritidae) was examined, with particular emphasis on their utilization of cultivated fruits. The species; D. tryoni, D. jarvisi, D. cucumis, D. musae and D. cacuminatus differ widely in host range with D. tryoni being highly polyphagous while D. cacuminatus is virtually monophagous. Laboratory experiments showed that larvae of all species survived and developed in many cultivated fruits in which the specialists never occur in the field. By contrast the oviposition preferences and specificity of adult females differed widely between species. Female D. tryoni oviposited in most species of fruit. The specialised species; D. cucumis, D. musae and D. cacuminatus strongly preferred their usual hosts and would not oviposit in novel fruits even in the absence of the preferred host. In contrast, D. jarvisi consistently preferred its main native host but when this was not offered readily accepted cultivated fruits. These differences in preference are consistent with the pattern of infestation displayed by each species in the field. The study indicate that, in general, the occurrence of these species of Dacus in cultivated fruits is constrained more by the behavioural preferences of adult females than by larval specialisations. A genetic change in some aspect of host recognition or acceptance would be necessary for the specialised species to regularly infest cultivated fruits though no change in larval characteristics may be needed. As the types and concentrations of defensive secondary compounds may differ between native and cultivated fruits this conclusion cannot be extended to host shifts among native fruits.     

Host preference and host suitability in an egg-pupal fruit fly parasitoid, Fopius arisanus (Sonan) (Hym., Braconidae)

The relative oviposition rate of the parasitoid Fopius arisanus (Sonan) was investigated across three frugivorous tephritid species, Bactrocera tryoni Froggart, Bactrocera jarvisi (Tryon) and Bactrocera cucumis French. Choice and no-choice tests were both used. The suitability of these three species for sustaining larval development and survival to the adult stage was also assessed. Fopius arisanus parasitized all three tephritid species, regardless of the method of exposure, but showed stronger preference for B. tryoni and B. jarvisi over B. cucumis. Superparasitism was extremely rare. Successful development of F. arisanus varied across host species. Bactrocera tryoni yielded significantly more parasitoids than B. jarvisi, but no wasps emerged from B. cucumis puparia. Tests were set up in replicated trials, but results were not homogeneous across trials. We discuss the host relationships of F. arisanus with reference to this variation and in relation to host suitability for larval development.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

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‐specific parasitism in the Central American striped cuckoo,Tapera naevia

Coevolutionary theories of brood parasite strategy and host defense have been informed by research on egg mimicry and host recognition. However, there is no information on the strategies of New World parasitic cuckoos and their hosts. The striped cuckoo Tapera naevia is a New World cuckoo that uses multiple host species and maintains an egg color polymorphism. To investigate if color‐matching influenced rejection behavior in hosts, I conducted an egg rejection experiment on a host that lays blue‐green eggs, the rufous‐and‐white wren Thryophilus rufalbus and a host that lays white eggs, the plain wren Cantorchilus modestus. I used spectrophotometric analysis of egg color to determine the degree of egg color‐matching. I found that at the field site the striped cuckoo lays highly mimetic eggs for the rufous‐and‐white wren, in both color and brightness. The rufous‐and‐white wren was more likely to accept mimetic artificial eggs than non‐mimetic eggs. The plain wren exhibited low rejection rates for both mimetic and non‐mimetic artificial eggs. The evidence from this study indicates that the striped cuckoo lays eggs that are closely color‐matched to those of its preferred host, the rufous‐and‐white wren, and that this mimicry improves acceptance.     

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.     

Repeatability of Foreign Egg Rejection: Testing the Assumptions of Co‐Evolutionary Theory

Most theoretical models of coevolution between brood parasites, whether interspecific or conspecific, and their hosts explicitly assume consistent individual behaviour in host egg‐rejection responses. Accordingly, hosts cast as acceptors always accept, whereas ejectors always reject parasitic eggs when exposed to stable ecological conditions. To date, only few studies have attempted to test this critical assumption of individual repeatability in egg‐rejection responses of hosts. Here, we studied the repeatability of egg rejection in blackbirds (Turdus merula) and song thrush (T. philomelos), species in which females are reported to reject simulated, non‐mimetic foreign eggs at intermediate frequencies at the population level. However, intermediate rates of acceptance and rejection can be consistent with either or both intra‐ and interindividual variability in rejection behaviours. Our experiments revealed generally high individual consistency in these hosts’ responses to experimentally introduced non‐mimetic and mimetic model foreign eggs. Individuals also responded faster on average to second than to first trials within the same breeding attempts, but the difference was statistically significant only in blackbirds. These results are consistent with the critical assumption of co‐evolutionary models, that statistically egg rejection is mostly individually repeatable, but also reveal that some individuals in both species change their responses even within the short time‐window of one breeding attempt. The data suggest that individuals reject foreign eggs faster when perceived parasitism risk is greater because of repeated introductions of experimental parasitic eggs. We provide methodological recommendations to facilitate experimental and meta‐analytical studies of individual egg rejection repeatability and discuss how to reduce technical constraints arising from disparate treatments and variable sample sizes for future studies.     

Behavioural plasticity in the stinging act of female ectoparasitoids

Abstract. The plasticity of the stinging behaviour of female Eupelmidae ( Eupelmus vuilleti Crawford, E. orientalis Crawford Hymenoptera: Eupelmidae) was analysed through the interrelationships of a parasitic community living on larval stage of Callosobruchus maculatus Fabricius (Coleoptera: Bruchidae). In traditional African grainstores, Eupelmus females have at their disposal all the larval stages of their host. Their exploitation is revealed by the sting observed on the host caused by the ovipositor of parasitoid females. This stinging behaviour begins immediately on the adult female's emergence, and thereafter there is a daily increase in the number of hosts stung. Stinging activity is linked to the age of the available hosts, but not all of the hosts available are exploited by the two species studied. This absence of total exploitation could indicate a threshold of activity in the stinging acts which could be in part linked to the secretion activity of the poison glands of parasitoid female.
In females of E. vuilleti and E.orientalis the stinging act is usually associated with egg-laying. The relationship between stinging and egg-laying is stronger in more advanced larval stages of the host.
In the presence of parasitized hosts, females of E. orientalis and E.vuilleti can exhibit hyperparasitism. The majority of larvae (i.e. primary parasitoid) that have been hyperparasitized have been stung and a large number of them are completely inactive. However, E. vuilleti females deposited their eggs in the available space of the pupation chamber of the host. The neonatal larvae from these scattered eggs could exploit any available host without the female having 'prepared the ground' at the time of egg-laying.
In Eupelmidae, all this happens as if there were a complementary action between the females and the neonate which could, if necessary, reinforce and/or compensate for the deficiencies in the host exploitation by the female.     

Host body size as a factor determining the egg complement of Strepsiptera,an insect parasite

The egg complement (total number of eggs produced by a single female) differs greatly among the species of Strepsiptera. The maximum is found in Stichotrema dallatorreanum (750,000 eggs), and the minimum in Triozocera minor (984 eggs). Based on the egg complement of 31 species in 11 genera, the following conclusions were drawn: (1) The egg complement is generally smaller in those species whose hosts gregariously cohabit in a very limited area, or are distinct flower-visitors, compared with those whose hosts display the above two traits weakly; (2) The egg complement is determined by the size of the maternal body. The size of female strepsipterans is reduced when they parasitize smaller host such as males and workers, as compared with those that parasitize larger host such as females and queens; likewise, the size of the strepsipterans becomes larger on increase in size of hosts, showing that their egg complements are principally determined by the size of host species; (3) The increase in the egg complement is compensated for by the reduction in egg size. The relative egg size (length of the firstinstar larva/length of maternal body) is conspicuously reduced according to an increase in the size of the female strepsipterans.     

Oviposition behaviour of two tephritid fruit flies,Dacus tryoni and Dacus jarvisi,as influenced by the presence of larvae in the host fruit

Summary When offered a choice, females of the fruit flies Dacus tryoni (Frogg.) and D. jarvisi (Tryon) strongly preferred to lay in fruits without larvae rather than fruits which already contained larvae. Fruits which contained even low densities of larvae, including newly hatched ones, received many fewer eggs than control fruits. This preference was not influenced by the species of larvae present in the fruits nor by the distance to uninfested fruits. Discrimination occurred when fruits with and without larvae were close together (10 cm apart) and also when they were separated by distances of about one metre. Laboratory assays suggested that the flies detect chemical changes in the fruit associated with the decomposition which accompanies larval feeding, but they do not seem to detect the larvae Perse. This behaviour may be significant when these two species utilise the same host since the species which is able to infest fruits first will reduce the availability of hosts for the other species. In contrast to many other Tephritids (e.g. Rhagoletis, Anastrepha and Ceratitis) female Dacus don't discriminate against fruits which contain eggs nor do they deposit a pheromone to deter oviposition by females that subsequently visit the fruit. An hypothesis is proposed to explain the absence of oviposition-deterring pheromones in Dacus, and their presence in many other species of Tephritidae, on the basis of differences in life history and population structure.     

The effects of physiological factors and host plant experience on the ovipositional activity of the sweet potato whitefly, Bemisia tabaci

Both egg load and levels of vitellogenin and vitellin increased when Bemisia tabaci Gennadius (Homoptera: Aleyrodidae), were deprived of host plants. These increases resulted in significant increases in rates of oviposition. There was no difference in total fecundity between whiteflies temporarily deprived of hosts and a control treatment that had continuous access to hosts. The results of previous exposure to a preferred host, melon (Cucumis melo), and a non-preferred host, cotton (Gossypium hirsutum), were also examined. B. tabaci deposited significantly more eggs during a 6-h period on melon than on cotton, regardless of host species they were previously confined to. In addition, whiteflies previously exposed to cotton, deposited more eggs than females previously exposed to melon. This higher rate of oviposition resulted in a low egg load as well as lower levels of vitellogenin and vitellin. Conversely, a low rate of oviposition led to high egg load and high levels of vitellogenin and vitellin. Individuals from the same treatment exhibited variation in egg load, which was negatively and significantly correlated with previous oviposition activity. We conclude that the effects of previous host plant exposure on oviposition were, in part, due to behavioral differences that are not related to the reproductive physiology.     

Egg Discrimination and Sex-Specific Pecking Behaviour in Parasitic Cowbirds

We studied egg‐pecking behaviour in males and females of three cowbird species: the shiny cowbird (Molothrus bonariensis), a host generalist brood parasite, the screaming cowbird (M. rufoaxillaris), a host specialist brood parasite, and the bay‐winged cowbird (Agelaioides badius), a non‐parasitic species. We conducted three experiments in which we offered each bird an artificial nest with two plaster eggs and recorded whether egg pecking occurred and the number of pecks on each egg. In expt 1, we tested if there were species and sex differences in egg‐pecking behaviour by offering the birds two spotted eggs of similar pattern. Shiny and screaming cowbirds responded in 40.3% and 44% of the trials, respectively, with females and males presenting similar levels of response. In contrast, bay‐winged cowbirds did not show any response. In expt 2, we tested if shiny cowbirds responded differentially when they faced a choice between one host and one shiny cowbird egg, while in expt 3, we tested if screaming cowbirds responded differentially when they faced a choice between one shiny and one screaming cowbird egg. Shiny cowbirds pecked preferentially host eggs while screaming cowbirds pecked more frequently shiny cowbird eggs. Our results show that egg‐pecking behaviour is present in both sexes of parasitic cowbirds, but not in non‐parasitic birds, and that parasitic cowbirds can discriminate between eggs of their own species and the eggs of their hosts or other brood parasites.     

Egg mimicry by the Pacific koel: mimicry of one host facilitates exploitation of other hosts with similar egg types

When brood parasites exploit multiple host species, egg rejection by hosts may select for the evolution of host‐specific races, where each race mimics a particular host's egg type. However, some brood parasites that exploit multiple hosts with the ability to reject foreign eggs appear to have only a single egg type. In these cases, it is unclear how the parasite egg escapes detection by its hosts. Three possible explanations are: 1) host‐specific races are present, but differences in egg morphology are difficult for the human eye to detect; 2) the brood parasite evolves a single egg type that is intermediate in appearance between the eggs of its hosts; 3) or the parasite evolves mimicry of one of its hosts, which subsequently allows it to exploit other species with similar egg morphology. Here we test these possibilities by quantifying parameters of egg appearance of the brood‐parasitic Pacific koel Eudynamys orientalis and seven of its hosts. Koel eggs laid in the nests of different hosts did not show significant differences in colour or pattern, suggesting that koels have not evolved host‐specific races. Koel eggs were similar in colour, luminance and pattern to the majority of hosts, but were significantly more similar in colour and luminance to one of the major hosts than to two other major hosts, supporting hypothesis 3. Our findings suggest that mimicry of one host can allow a brood parasite to exploit new hosts with similar egg morphologies, which could inhibit the evolution of host defences in naïve hosts.     

A new family of Strepsiptera parasitizing fruit flies (Tephritidae) in Australia

ABSTRACT. A new family of Strepsiptera (Dipterophagidae) from Australia containing a new genus and species ( Dipterophagus daci ) is described and figured. This is the first species of Strepsiptera to be described as a parasite of Diptera. D.daci is recorded from ten fruit fly species, viz Dacus aquilonis (May), D.bellulus Drew, D.cacuminatus (Hering), D.decurtans (May), D.mayi Hardy, D.neohumeralis Hardy, D.peninsularis Drew, D.tenuifascia (May), D.tryoni (Froggatt) and Dacus sp. n. ( near musae ). Gregarious parasitism (two to six adult parasites per fly) is common.     

Egg dumping and life history strategy of Callosobruchus maculatus

Abstract.  Previous studies have demonstrated that female Callosobruchus maculatus adjust oviposition rates to cope with changes in host availability. A female lays fewer eggs when host availability is low, and hence decreases larval competition. However, females will also dump eggs on unsuitable substrates under conditions of host deprivation. Because the female does not feed as an adult, egg dumping possibly wastes energy and may thus be a maladaptive behaviour. In this study, the effect of mating and age on the egg-dumping behaviour and the life history strategy of the female are explored. Under host-deprived conditions, mating is seen to trigger egg-dumping behaviour. Also, females mated at 6 days dump significantly fewer eggs and live longer than females mated at 0 or 3 days. Thus, a trade-off between fecundity and longevity is seen among females subjected to different manipulations. In addition, 6-day-old virgin females contain more mature eggs than females mated at 6 days can produce when deprived of hosts. This finding indicates that the female reallocates internal energy resources by oosorption in a resource-limited environment. To test the maladaptive hypothesis, mated females were deprived of a host for 6 days and then given sufficient hosts each day. The results show that the more eggs dumped by a female in the first 6 days, the more eggs are laid later on beans. Egg dumping is thus not maladaptive.     

Oviposition acceptance and fecundity schedule in the cactophilic sibling species Drosophila buzzatii and D. koepferae on their natural hosts

We tested for the occurrence of oviposition acceptance for different media prepared with cactus tissues of three alternative cactus hosts: Opuntia sulphurea, O. quimilo and Trichocereus terschekii for 4 consecutive days in lines of two Drosophila buzzatii populations and one population of D. koepferae. Our results showed that the former laid significantly more eggs on both Opuntia cacti than on T. terschekii, whereas D. koepferae preferred T. terschekii. In addition, fecundity schedules differed between species: D. buzzatii laid similar numbers of eggs along the four-day sampling period, whereas D. koepferae showed an oviposition peak on the second day of egg collection on T. terschekii. We suggest that the between-species disparities observed in oviposition acceptance and fecundity schedule may be related to the temporal and spatial predictability of Opuntia versus T. terschekii (cardón) as part of the different adaptive strategies that have evolved after the split of D. koepferae and D. buzzatii from their recent common ancestor. Therefore, the willingness to accept hosts would be an important factor in the habitat selection and in the maintenance of species diversity.     

Egg resorption behaviour by the solitary egg parasitoid Anaphes nitens under natural conditions

We investigated under outdoor conditions and inside a climate chamber: (i) whether Anaphes nitens Girault (Hymenoptera: Mymaridae), an egg parasitoid of the Eucalyptus snout beetle, Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae), experienced egg resorption, and (ii) how various treatments (location, food, and/or host supply) and body size influenced egg load, egg resorption, fecundity, and longevity. One‐day‐old females were mated and randomly assigned to five groups: (A) honey + hosts, (B) water + hosts, (C) only honey, (D) only water, and (E) control females killed at emergence. We compared the egg load of the newly emerged females, which represent the control group (egg load = ovarian eggs present at emergence), with the lifetime egg load (i.e., ovarian eggs + emerged offspring + not emerged offspring) of the females with various host and diet treatments, by dissection of the ovaries to find evidence of egg resorption. All groups reared outdoors had fewer eggs than the control, while indoors there was no significant difference. Outdoors, starved but host‐provided wasps (B) experienced the highest reduction of the lifetime egg load (51%). Groups without access to food (B + D) resorbed more eggs than groups provided with honey (A + C). Females with honey and hosts (A) had the highest lifetime fecundity, but those with water and hosts (B) showed a higher daily realized fecundity. Host‐deprived females with access to food (C) attained the longest lifespan. Our results suggest that under stress conditions, such as low temperature and food shortage, A. nitens females practice egg resorption, probably to save energy.     

Seasonal monitoring of Drosophila suzukii and its non-crop hosts in Wuhu,Eastern China

Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) is a serious pest that prefers fresh fruits and is native to Southeast Asia. In our study, apple cider vinegar bait traps were used to capture and monitor the population dynamics of this native pest in Wuhu City, China, from May/June 2017 to May 2018. The research was conducted at 15 locations in two fruit orchards in Wuhu. Traps caught more adults in general in a Meiling blueberry orchard than in a Xicun mixed orchard, and the highest trap counts occurred near harvest time (October). Females had more mature eggs from September to November, and the number of mature eggs declined thereafter. We found several non-crop hosts, which can provide food and reproductive resources for D. suzukii and are common in forests and field margins. By comparing the number of captured adults in the Meiling and Xicun orchards, we found that blueberry was preferred by D. suzukii among the fruits in our search. Fruit ripening times differed among crops; therefore, fly populations moved between crop and non-crop habitats during the year or had varying population dynamics on different crops in different seasons. The D. suzukii population and the number of mature eggs decreased in summer and winter but increased in spring and autumn. Drosophila suzukii had higher survival rates with blueberry than those with other fruits, and D. suzukii could use four non-crop species growing around the orchards as host plants.     

Effect of host diet on acceptance of eastern spruce budworm eggs byTrichogramma minutum

The acceptance behaviour of the parasitic wasp, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), was studied on two types of egg masses of eastern spruce budworm (SBW), Choristoneura fumiferana Clemens. Host eggs were obtained from moths that had been fed balsam fir or artificial diet during their larval stages. Parasitoids were reared from both types of hosts. Host acceptance assays were conducted where females were provided with a single egg mass and with a choice between egg masses. Regardless of the parasitoid's rearing host, females parasitised hosts reared from artificial diet more often than those from balsam fir. Response was consistent in both no-choice and choice assays. Rejection of hosts tended to occur during the examining stage of behaviour, suggesting that acceptability is judged externally. If a host was accepted, the same number of eggs were laid per egg mass regardless of treatment.