Factors affecting the reproductive biology of Melittobia digitata and failure to meet the sex ratio predictions of Hamilton's local mate competition theory

Melittobia digitata Dahms (Eulophidae, Tetrastichini), a species of parasitic wasp satisfying all of Hamilton's local mate competition requisites, does not exhibit the predicted change in sex ratio with increased foundress number. A multifactorial design was used to test how age, oviposition experience, feeding experience, mating, and foundress number affect host‐acceptance, number of offspring, and sex ratio of this species developing on honey bee pupae, Apis mellifera (L.) (Apidae, Apini). All factors significantly affected the time it took for oviposition to commence. Females oviposited soonest when they were 2 days old, mated, had previous feeding and oviposition experience, and were placed on hosts with multiple foundresses. Although the age difference between 2‐ and 5‐day‐old females is small, it significantly affected reproductive behavior. Age, mating, and foundress number were found to have an effect on sex ratio, however, the foundress effect was found to be a mathematical artifact of the limited host size. After correcting for this variable, females were found to have a constant sex ratio of approximately 0.05. Several 2‐way interactions between factors were revealed: age and experience, age and foundress number, age and mating, foundress number and experience, and foundress number and mating. One 3‐way interaction was found between age, mating, and foundress number. This study demonstrates that the sex ratio of M. digitata is not altered with increased foundresses, as predicted by Hamilton , and that slight changes in preconditioning may modify reproductive behavior.     

Host size,superparasitism and sex ratio in mass-reared <Emphasis Type="Italic">Diachasmimorpha longicaudata</Emphasis>, a fruit fly parasitoid

We analyzed the relationship among host size, superparasitism and sex-ratio in mass reared Diachasmimorpha longicaudata (Hymenoptera: Braconidae). Individual host pupae of Anastrepha ludens (Diptera: Tephritidae) were measured (length and width), and the number of oviposition scars per pupa was used as a reliable indicator of superparasitism. The probability of an emerging parasitoid being a female was positively associated with the number of oviposition scars on the host cuticle, but not with the host size. The number of scars per host pupae from which females emerged was slightly but significantly higher than in those pupae giving raise to males. In D. longicaudata, the influence of host size on sex allocation decisions of individual females seems to be overridden by the level of superparasitism, which itself was positively correlated with pupa length. This suggests that larger pupae could experience a higher number of ovipositions than their smaller counterparts, and that a high level of superparasitism may conduct to a female biased sex ratio. We discuss the relevance of these findings which could provide new elements (e.g., the manipulation of superparasitism) for optimizing the mass rearing of this parasitoid.     

Clutch size in a larval-pupal endoparasitoid

Clutch size decisions by Aphaereta minuta (Nees) (Hymenoptera: Braconidae), a polyphagous, gregarious, larval-pupal endoparasitoid, were studied under laboratory conditions. This parasitoid attacks larvae of Diptera inhabiting ephemeral microhabitats such as decaying plant and animal material. Females oviposit in young larval stages, but the eventual size of the host pupa determines host food availability for competing offspring. The size of the pupa can differ greatly between host species. We questioned how A. minuta females deal with this delay between the moment of oviposition and eventual host food availability, and whether they make clutch size decisions that benefit their fitness. It was shown that females indeed vary their clutch size considerably and in an adaptive way: (1) females lay larger clutches in larvae of host species that produce larger pupae, even when the larvae are the same size at the moment of oviposition, and (2) females lay larger clutches in larger larvae than in smaller larvae of the same host species. The latter seems functional as larvae parasitized at an older stage indeed developed into larger pupae compared to larvae parasitized at a younger stage. Furthermore, mortality of parasitized young host larvae was greater than that of both unparasitized larvae and parasitized older larvae. Under field conditions the risk of mortality of young host larvae is expected to be even higher due to the limited period of microhabitat (host food) availability, strong scramble type competition between the host larvae, and the longer period of being exposed to predation.     

The avoidance of superparasitism in four species of parasitic wasp—Mathematical models and experimental results

Experimental results, and interpretation, are presented which relate to oviposition behaviour in four species of parasitic wasp (Hymenoptera: Chalcidoidea). All lay their eggs in pupae of the common house fly (Musca domestica) and of other flies associated with man. Probability models suggested byD. J. Daley for describing the avoidance of superparasitism are used to analyse the data. Certain of the models considered by previous authors appear as specieal cases. A satisfactory fit was obtained either by assuming a fixed probability of oviposition in an already parasitised pupa, or by using a model motivated by supposing that a female will tolerate without oviposition some fixed number of visits to already parasitised pupae. A third model was considered which makes the probability depend on the number of previous ovipositions in the pupae now visited. In all four species the presence of other females (with parasite: host ratio unchanged) increased the probability of oviposition upon encountering an already parasitised pupa. In the presence of other females Spalangia endius laid an increased number of eggs, whereas the other three species laid fewer eggs.     

Influence of experience on the sequential and temporal organization of host-acceptance behavior inBrachymeria intermedia (Chalcididae), an endoparasitoid of gypsy moth

Experienced Brachymeria intermediafemales are almost twice as likely to accept a Lymantria disparpupa as inexperienced parasitoids. The sequence of parasitoid behaviors that leads to host acceptance is highly canalized. Experienced parasitoids, however, have a higher probability of initiating host investigation and making a transition from drumming to grasping, which in turn almost invariably leads to ovipositor insertion. Experienced females found the host more quickly than inexperienced females. Host-handling time did not change with experience but was longer in females that accepted rather than rejected the host. Females exposed to gypsy moth odor but not contacting pupae behaved similarly to females that never experienced host-related stimuli.     

Field superparasitism by Diachasmimorpha longicaudata attacking Anastrepha spp. larvae on mango fruits

The incidence of superparasitism in field populations of Diachasmimorpha longicaudata (Ashmead), a solitary larval–pupal endoparasitoid, attacking Anastrepha fruit fly species was determined. From 14,550 fruit fly larvae infesting mango fruits in the field, 68.9% (10,038) survived to the pupal stage. Of these pupae, 3780 (37.7%) were parasitized, of which 2075 (54.9%) were superparasitized. The number of oviposition scars per pupa ranged between 1 and 29, although high numbers were rare. The presence of one or two scars per host was frequent, but only 8.6% of pupae had more than three scars. Fruit size was positively related to the level of infestation by Anastrepha spp. and was significantly correlated with the numbers of parasitized and superparasitized pupae by D. longicaudata. The length and width of host pupae were positively related to superparasitism, and superparasitism was positively correlated with a sex ratio biased toward females. Our data show that superparasitism is present in natural populations of D. longicaudata at remarkable rates and confers some advantageous features such as a female-biased sex ratio. These findings favor the election of this species as a viable biocontrol agent for augmentative releases because the favorable proportion of adult females emerging from superparasitized pupa should contribute to better pest control.     

Host handling and specificity of the hyperparasitoid wasp, Dendrocerus carpenteri (Curtis) (Hym., Megaspilidae): importance of host age and species

The oviposition behaviour of Dendrocerus carpenteri (Curtis), an ectophagous hyperparasitoid of aphidiine wasps inside mummified aphids was examined. Hyperparasitoids were provided in the laboratory with pea aphids ( Acyrthosiphon pisum ) which had been parasitized by three different species of aphidiine wasps ( Aphidius ervi, Ephedrus californicus and Praon pequodorum ) ranging in physiological age from the late larval stage to the late pupal stage. Females accepted only the hosts inside mummified aphids; they ignored live aphids, and did not accept dead, but not yet mummified aphids, although the latter were sometimes probed with the ovipositor. Female behaviour in handling A. ervi or E. californicus mummies did not change with experience; handling and oviposition times were stereotypic. However, naive females needed experience to locate the cocoon of P. pequodorum and distinguish it from the empty aphid mummy. Host acceptance and specificity were influenced more by the developmental stage than the species of the primary parasitoid. In dichotomous choice tests, hyperparasitoids 'preferred' prepupae over younger pupae of A. ervi , but they did not distinguish between these stages of E. californicus; older pupae were accepted at a low rate. Host preference was not influenced by conditioning on the rearing host. We consider several constraints on the host range of D. carpenteri , and discuss alternative explanations of differential hyperparasitism in the field.     

Two different life-history strategies determine the competitive outcome between Dirhinus giffardii (Chalcididae) and Pachycrepoideus vindemmiae (Pteromalidae), ectoparasitoids of cyclorrhaphous Diptera

Dirhinus giffardii Silvestri and Pachycrepoideus vindemmiae Rondani are solitary parasitoids attacking puparia of many cyclorrhaphous flies. They are not typical ectoparasitoids, as they feed on host pupae within puparia that develop from the exoskeleton of host larvae. Dirhinus giffardii did not kill its host until the parasitoid egg developed into a larva, while P. vindemmiae permanently paralysed its host at the time of oviposition. As a result, ovipositing into a young host puparium (< 1 day old) in which the host pupa has not yet fully formed resulted in complete death of offspring in P. vindemmiae, but D. giffardii, although suffering higher mortality than in older host puparia, still showed a level of successful development. In a choice experiment, both parasitoids preferred to attack 2- to 3-day-old puparia in which the host pupae had fully formed, rather than 1-day-old host puparia. Pachycrepoideus vindemmiae always prevailed in competition because it injected venom that not only paralysed the host, but also caused the death of D. giffardii larvae in multi-parasitized hosts. Dirhinus giffardii preferred to attack unparasitized hosts rather than hosts previously parasitized by P. vindemmiae, while P. vindemmiae did not show a preference between unparasitized hosts and hosts previously parasitized by D. giffardii.     

The role of physical cues in the regulation of host recognition and acceptance behavior ofAphidius ervi Haliday (Hymenoptera: Braconidae)

The role of color and shape in the host recognition and acceptance behavior ofAphidius ervi Haliday was studied. A quantitative analysis of the oviposition behavior ofA. ervi was carried out with a computer-aided analysis of 150 video-recorded oviposition sequences on its natural host,Acyrthosiphon pisum (Harris). The importance of visual stimuli was assessed in a choice condition bioassay, observing the behavioral reaction of female parasitoids to various test materials flame-sealed into glass capillaries. Glass beads 2 and 6 mm in diameter and a flat arena were coated with cornicle secretion ofA. pisum, and their acceptance rates by both naive and experienced female parasitoids were assessed under no-choice conditions. In most cases,A. ervi females switched from random searching to attack position when the host was within a range of 1 cm, suggesting that host recognition is regulated in part by cues acting before physical contact. The glass capillary bioassay indicated that visual cues are important factors in the host recognition and acceptance phases. Pea aphid color alone can elicit the oviposition response of naiveA. ervi females, and this response is enhanced when color is combined with aphid shape. The cornicle secretion ofA. pisum stimulated an oviposition response which was stronger in naive females ofA. ervi than in experienced ones and was not significantly affected by the glass bead size or flat surface. These results, along with those from previous studies, suggest that manipulation of the oviposition behavior ofA. ervi is feasible under laboratory conditions.     

Female's preference for oviposition site and larval performance in the water-lily beetle,Galerucella nymphaeae (Coleoptera: Chrysomelidae)

Water-lily beetles prefer younger rather than older water-lily leaves as oviposition sites. By the time of hatching, however, young leaves have aged consieerably. Larval performance of the water-lily beetle was measured on different types of leaves of the yellow water-lily and compared with oviposition preference of females. The leaf types used in the experiments were categorized as (i) young, (ii) natal (medium-aged) and (iii) old. The natal leaves were the ones on which larvae from a particular egg-batch had hatched. There were two sets of experiments. First, larvae were raised from eggs to pupae on young and on old leaves. Second, the growth of the 1st-instar larvae was measured on young, natal, and old leaves. The development time from egg to pupa did not differ between young and old leaves, but larvae growing on young leaves attained a higher pupal weight. In the second experiment the 1st-instar larvae grew fastest on their natal leaves, but there was also variation in the growth rate of progeny from different egg-batches. Larval growth on young and old leaves did not differ significantly. Larvae tried to emigrate much less from natal than other types of leaves. Females tended to lay eggs on leaves where larval growth was fastest. It seems that medium-aged leaves are best for larval growth, but the leaf characteristics responsible for this remain unresolved.     

Does Host Value Influence Female Aggressiveness,Contest Outcome and Fitness Gain in Parasitoids?

Intraspecific competition for resources is common in animals and may lead to physical contests. Contest outcomes and aggressiveness can be influenced by the resource holding potential of contestants but also by their perception of the resource value (RV). Competitors may assess resource quality directly (real RV) but may also estimate it according to their physiological status and their experience of the habitat quality (subjective RV). In this article, we studied contests between females of the solitary parasitoid Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae) when exploiting simultaneously a host, a Delia radicum L. (Diptera: Anthomyiidae) pupa. We tested the effect of factors modifying host value on the occurrence of agonistic behaviours, contest outcomes and host exploitation. The factors tested were: the quality of the previous habitat experienced by females, female egg load, host parasitism status and the stage reached by the owner female in her behavioural oviposition sequence. Females successfully protected their host against intruders during its exploitation, but not after oviposition, and their aggressiveness did not seem to be influenced by their perception of the RV. The fact that the host is subsequently parasitized by the opponent females appears to mainly depend on the host selectiveness of females.     

Superparasitism strategies by a native and an exotic parasitoid species attacking the Mexican fruit fly,Anastrepha ludens (Diptera: Tephritidae)

Superparasitism refers to the action of parasitoids ovipositing eggs in hosts that are already parasitized; this inevitably results in the elimination of supernumerary larvae in solitary parasitoids. Here, we investigated superparasitism performed by two species of solitary parasitoids on the larvae of Anastrepha ludens (Loew; Diptera: Tephritidae): a native species, Doryctobracon crawfordi (Viereck; Hymenoptera: Braconidae), and an exotic species, Diachasmimorpha tryoni (Cameron; Hymenoptera: Braconidae). Tests were conducted under laboratory conditions evaluating the behaviour of females acting alone (self-superparasitism) or in groups (conspecific superparasitism). Parasitism strategies were different between these two species. In D. crawfordi, the number of first instar larvae found in each dissected host pupa was never greater than two, regardless of the number of oviposition scars observed per pupa. In contrast, there was a positive correlation between the number of oviposition scars and the number of first instar larvae in D. tryoni. The survival and fecundity of D. crawfordi females emerging from pupae with one scar was higher than in females emerging from pupae with more scars. In D. tryoni, the number of oviposition scars did not show deleterious effects on life history traits and was positively correlated with the proportion of emerging females. An understanding of the superparasitism strategy adopted by parasitoid species could be of great interest to augmentative biological control programmes because the mass rearing of natural enemies could be negatively or positively affected by this condition.     

Communal use of integumental wounds in honey bee (Apis mellifera) pupae multiply infested by the ectoparasitic mite Varroa destructor

The ectoparasitic bee mite, Varroa destructor, is highly adapted to its natural and adopted honey bee hosts, Apis cerana and Apis mellifera. Adult females perforate the integument of bee pupae in such a way that they and their progeny can feed. We examined the wounds that founder females made, and usually found one, and rarely up to three, integumental wounds on pupae of A. mellifera multiply infested by V. destructor. The punctures were mainly on the 2nd abdominal sternite of the host. These perforations are used repeatedly as feeding sites by these hemolymph-sucking mites and by their progeny. The diameter of the wounds increased during pupal development. In brood cells containing 4-5 invading female mites and their progeny, healing of the wound is delayed, normally occurring just before the imaginal moult of the bee pupa. These wounds are subject to microbial infections, and they are relevant to the evolution of behavioral traits in these parasitic mites and their relations to host bees.     

Analysis of Tiphia parasitoids preovipositional behaviors and of their scarab host defensive responses

Tiphia vernalis Rohwer and Tiphia popilliavora Rohwer were introduced as biocontrol agents against Japanese beetles (Popillia japonica Newman) and oriental beetles (Anomala orientalis Waterhouse). Studies have shown that under field conditions, T. vernalis parasitize Japanese beetles more often than oriental beetles. This study was done to understand how tiphiid wasps handle the two different host species and the influence of host defensive behaviors on the oviposition process of tiphiid wasps. The preovipositional behaviors performed by Tiphia wasps included: stinging, examining, moving soil, kneading, host-feeding, and host scraping. The frequency, sequence, and total time spent on each behavior before oviposition were scored and compared between two host species. The sequence and frequency of preovipositional behaviors performed by both Tiphia wasps did not show a difference between the two host species. However, female T. vernalis spent significantly longer time trying to sting oriental beetles than Japanese beetles in order to paralyze them. The time T. popilliavora spent on prestinging behaviors did not show a difference between Japanese and oriental beetles. The defensive behaviors performed by Japanese and oriental beetle grubs included: vigorous movements, rubbing their abdomen or head against the wasp’s abdomen, and biting at the attacking wasp. The frequency and total time spent on each defensive behavior was scored and compared between two host species. Overwintered, third instar oriental beetle grubs spent significantly longer time on defensive behaviors when they were attacked by T. vernalis which likely cause wasps to spend longer time trying to sting oriental beetle grubs. The active host resistance gained through behavioral defenses could make oriental beetle grubs less susceptible to T. vernalis attack than Japanese beetle grubs, especially under field conditions. Younger grubs attacked by T. popilliavora did not exhibit these differences.     

Preference and performance of peach fruit fly (Bactrocera Zonata) and Melon fruit fly (Bactrocera Cucurbitae) under laboratory conditions

In Pakistan and all over the world, the Peach Fruit Fly (PFF), Bactrocera zonata (Saunders.) and the Melon Fruit Fly (MFF), Bactrocera cucurbitae (Coquillett.) are considered severe and polyphagous insect pests for various fruits and vegetables. The current study was conducted to check the Laboratory preference and performance of B. cucurbitae and B. zonata on selected Fruits Citrus (Citrus sinensis), Apple (Malus domestica), Banana (Musa acuminate), and vegetable, Sponge gourd (luffa aegyptiaca), Bitter gourd (Momordica charantia) Pumpkin (Cucurbita moschata) under laboratory conditions. The study showed that Sponge Gourd was the preferable host with the mean pupae resurgence of (242.33), followed by Bitter Gourd (78.333) among selected vegetables. At the same time, among fruits, a banana was the preferable host with mean pupae resurgence (204.33), followed by orange (158.33). The pumpkin and apple was the least preferable host for both B. cucurbitae and B. zonata, with mean pupae resurgence (35.667) and (79.000), respectively. Furthermore, the study showed that Banana was the preferable host for B. Zonata among intact and infested fruits, whereas B. cucurbitaee showed the most preference to Bitter gourd among intact and infested vegetables showing significantly different results among intact and infested fruits and vegetables. Maximum number of eggs, pupa, female flies, male flies, adult emergence from pupa (flies) and period of pupa of B. zonata and B. cucurbitae on banana and bitter gourd. While, other fruits and vegetables showed the minimum number of eggs, pupa, female flies, male flies, adult emergence from pupa (flies) and period of the pupa. The current study concluded there is a need to evaluate other host plants against these fruit fly species for effective control.     

Host handling behavior of the egg parasitoid Telenomus triptus to the egg mass of the stink bug Piezodorus hybneri

Host handling behavior of Telenomus triptus Nixon (Hymenoptera: Scelionidae) to an egg mass of Piezodorus hybneri Gmelin (Heteroptera: Pentatomidae) was studied in the laboratory. Five distinct behavioral events could be distinguished in the host handling behavior: drumming, ovipositor-insertion, marking, walking, and resting. Female wasps showed two types of behavioral sequence in an ovipositional bout. One consisted of drumming, ovipositor-insertion, and marking, and the other drumming, ovipositor-insertion, walking, and resting. Females did not seem to lay an egg when ovipositor-insertion was not followed by marking. This was observed frequently in the early oviposition bout, on average 2.9 times per female. Females finally succeeded in parasitizing all the eggs in a host egg mass in most cases. The durations of drumming, ovipositor-insertion, and marking on an egg mass were nearly constant, while the total time spent by a female on and beside a host egg mass varied considerably as a result of variable durations of walking and resting. Self-superparasitism occurred when almost all the eggs in a host egg mass were parasitized. Females laid the first male egg within the first four eggs; this could be an adaptation to small egg masses or single egg.     

The effect of host acceptability on oviposition and egg accumulation by the small white butterfly, Pieris rapae

The influence of host‐plant acceptability on oviposition rate, egg load, internal fat storage and longevity was studied in the small white butterfly, Pieris rapae L. (Lepidoptera: Pieridae). Newly emerged females and males were presented with either cabbage (Brassica oleracea L. var. gemmifera), which is a highly acceptable host for the small white, or wallflower (Cheiranthus cheiri L.), which is much less acceptable for oviposition. Individuals were dissected when 3, 6 and 9‐days‐old. Females kept on cabbage contained fewer mature eggs than those kept on wallflower and there was no change in the load of mature eggs over the time course studied. The number of immature eggs carried did not vary with host plant species, but did fall significantly between days 6 and 9. The body fat content of individuals declined with increasing age, but the decline was slower for individuals kept on the host of low acceptability. Individuals that were allowed to spend their natural lifespan on cabbage showed similar oviposition patterns over time, where the oviposition rate started high and gradually reduced until death. However, considerable differences in oviposition pattern were found in individuals kept on wallflower, varying from that found on cabbage to no oviposition at all. The implications of these findings are discussed in the light of existing oviposition theories. This leads to the conclusion that in one species the premises of existing theories on optimization of oviposition are not mutually exclusive but rather play in concert.     

Dependence of juvenile hormone activity on haemolymph factor in Tenebrio molitor pupae

Morphogenetic response of Tenebrio molitor pupae to juvenile hormone (JH) varies with age, it being low during the first 6 hr after pupal ecdysis and reaching a peak at about 18 hr after ecdysis. Parabiosis of 6-hr pupa to a 18-hr-old pupa causes a higher morphogenetic response in 6-hr pupa to JH. A similar increase in morphogenetic response was not noticeable when a 0·45 μm Millipore filter was placed between the parabiotic partners, suggesting that the factor(s) may be macromolecular. This view is further supported by the observation that the Millipore filter did not prevent transport of ecdysterone another agent that enhances morphogenetic response of pupae to JH.     

Natural History and Comparative Morphology of Immatures of <Emphasis Type="Italic">Gamelia anableps</Emphasis> (C. Felder &amp; R. Felder) (Lepidoptera: Saturniidae,Hemileucinae)

Egg, larva, and pupa of Gamelia anableps (C. Felder & R. Felder, 1874) are described for the first time and the chaetotaxy of the first instar provided. Eggs and selected structures of larvae and pupae were also investigated by means of scanning electron microscopy. Eggs are laid in clusters, are ellipsoidal and laterally flat. Larvae passed through eight instars in about 78 days. Pupation occurs inside a poorly organized cocoon of yellowish silk. The dark brown pupa is cylindrical in shape with appendages firmly fused together and to the body wall. The chaeotaxy is compared with other Saturniidae species belongining to Hemileucinae, Ceratocampinae, and Saturniinae and the natural history of G. anableps compared with other Saturniidae.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999