Parasitism by Pteromalus cerealellae (Hymenoptera: Pteromalidae) on the Cowpea weevil, Callosbruchus maculatus (Coleoptera: Bruchidae): Host density, temperature effects, and host finding ability

Studies on interactions between a larval parasitoid, Pteromalus cerealellae (Boucek) and one of its hosts, Callosobruchus maculatus (F.) were carried out in the laboratory. The number of host larvae parasitized by P. cerealellae increased with host larvae at low densities and tended to a plateau at a density of 25 larvae per female parasitoid. Each parasitoid was able to parasitize more hosts and produced more offspring at 20 and 25 °C than at 30 °C. The number of non-infested seeds mixed with seeds infested with the last instar of C. maculatus did not preclude P. cerealellae from identifying infested seeds and attacking larvae inside them. When infested seeds were tightly packed, several host larvae escaped parasitism. P. cerealellae may be a useful biological control agent in newly harvested cowpea with low C. maculatus infestation, and lowering the temperature of the storage system may enhance the effectiveness of this parasitoid.     

Factors affecting the parasitism rate and the number and sex ratio of offspring of Asecodes hispinarum Bouček (Hymenoptera: Eulophidae), a biological control agent of Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae)

To optimise the production of Asecodes hispinarum Bou?ek (Hymenoptera: Eulophidae), a parasitoid of coconut leaf beetle, Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae), some of the factors affecting rates of parasitism, number of offspring produced per host and sex ratio of A. hispinarum were investigated. The numbers and sex ratio of A. hispinarum offspring per host reduced significantly at extreme low humidity (30% relative humidity [RH]), but there was no significant effect on parasitism. Photoperiod had no significant effects on any of the life traits tested. A. hispinarum was able to reproduce via arrhenotoky, and while increasing the proportion of female parents increased the number of parasitoids produced, the proportion of female offspring decreased. Older females showed a lower rate of parasitism than young females, however, maternal age did not affect the number or the sex ratio of offspring. Increasing the number of hosts offered to a pair of parasitoids significantly increased the number of parasitised hosts but decreased the parasitism rate while the sex ratio of progeny was not affected. Present work showed that to maximise the production of female parasitoids, a parasitoid/host ratio of 1:1, using one-day old A. hispinarum at a female/male ratio of 3:1 and RH of at least 55% is recommended.     

Offspring production and self-superparasitism in the solitary ectoparasitoid Spalangia cameroni (Hymenoptera: Pteromalidae) in relation to host abundance

Parasitoid fitness strongly depends on the availability and quality of hosts, which provide all resources required for larval development. Several factors, such as host size and previous parasitation, may affect host quality. Because self-superparasitism induces competition among a female's offspring, it should only occur if there is an imperfect recognition of self-parasitized hosts or if there is a fitness advantage to self-superparasitism. Against this background, we investigated self-superparasitism and offspring production in Spalangia cameroni (Hymenoptera: Pteromalidae) in relation to the abundance of a novel host, Ceratitis capitata (Diptera: Tephritidae). Individual pairs of parasitoids were provided with either two (low host abundance) or ten (high host abundance) pupae per day. Under high host abundance, lifetime fecundity (number of eggs laid), offspring number, number of pupae parasitized and hosts killed were greater than under low host abundance, whereas the number of eggs per host was lower; and the proportion of hosts that did not produce offspring tended to be lower. The latter suggests the occurrence of ovicide, when hosts are scarce due to an at least imperfect recognition of previously self-parasitized hosts. Offspring production per parasitized pupa was higher when hosts were scarce and levels of self-superparasitism high, suggesting the existence of beneficial effects of self-superparasitism.     

Effect of host density on offspring sex ratios and behavioral interactions between females in the parasitoid waspNasonia vitripennis (Hymenoptera: Pteromalidae)

Pairs of females of the parasitoid waspNasonia vitripennis were videotaped with one or two hosts. The presence of an additional host decreased the number of interactions between females but had no measured effect on the nature of the interactions, i.e., on whether the interaction involved physical contact or occurred while one of the females was parasitizing a host. The number of hosts did not itself affect offspring sex ratios but did influence which other factors were correlated with sex ratio. When there was one host, the proportion of sons was more positively correlated with utilization of previously drilled holes than with female-female interactions, whereas when there were two hosts, the reverse was true. Parasitizing an already parasitized host appeared to affect a female's sex ratio beyond any effects of the physical presence of another female: When two hosts were present, the proportion of sons was greater from hosts parasitized by both females than from hosts parasitized by only one female. The observation that parasitizations in previously drilled holes and female-female interactions are correlated with sex ratios is consistent with previous studies; however, that these relationships are host density dependent is a new result and remains unexplained.     

Benefits of self-superparasitism in a polyembryonic parasitoid

Macrocentrus grandii is a polyembryonic parasitoid, with embryos that divide asexually within the host (European corn borer, Ostrinia nubilalis) to produce broods of clonal offspring. From a biological control standpoint, polyembryony seems advantageous because each parasitized host yields multiple parasitoids with minimal time and egg investment. When we observed M. grandii in the field, however, we found that the parasitoid virtually always invested additional time and, if possible, stings into hosts that it had already stung, apparently reducing some of the advantages of polyembryony. We therefore investigated and found support for two potential benefits that can be gained by self-superparasitism in this system. First, multiple stings allowed production of mixed-sex broods: 27% of multiply-stung versus 0% of singly-stung hosts produced mixed-sex broods. Second, multiple stings increased mean parasitoid progeny produced per host, primarily by reducing the chance of complete brood failure. Our results indicate substantial benefit for a second sting, but little benefit for three or more stings, even though M. grandii was sometimes observed to invest more than two stings within a single host. However, we also found that within-host larval competition is prevalent, suggesting that supernumerary stings may pay off in competition against conspecific larvae. Such additional investment within a single host would be particularly beneficial when hosts, rather than eggs, are limiting, but would decrease the overall efficacy of M. grandii as a biological control agent.     

Interactions between the egg and larval parasitoids of a gall-forming midge and their impact on the host

1. The gall‐forming midge Rhopalomyia californica was exposed experimentally to parasitism and predation during only the egg stage, during only the larval stage, during neither stage, or during both stages. 2. The combined action of natural enemies that attack during both the egg stage and the larval stage led to the lowest number of midges and total insects (midges + parasitoids) in the next generation, and the highest percentage parasitism. 3. The larval parasitoid killed a large fraction of hosts without producing new parasitoid offspring, while there is some indication that the egg parasitoid on its own tended to produce the most parasitoid offspring. The contrasting implications of host mortality versus parasitoid production for biological control are discussed. 4. Exposure to larval parasitoids resulted in a reduction in the number of egg parasitoid offspring produced, but exposure to the egg parasitoid did not affect the number of larval parasitoid offspring produced significantly.     

Sex Allocation Decision Under Superparasitism by the Parasitoid Wasp Eupelmus vuilleti

Superparasitism is a widespread phenomenon in parasitoids and may be advantageous in some circumstances. In this study, offspring sex ratio was analysed in three superparasitism situations: when the second egg was laid by a random Eupelmus vuilleti (Hymenoptera: Eupelmidae) female from a group, when an isolated female was allowed to lay two eggs on the same host (self-superparasitism) or laid one egg on a host already parasitized by a conspecific (conspecific superparasitism). Females produced a different offspring sex ratio according to the different superparasitism situations tested. These sex ratios are in line with the local mate competition theory. The results further suggest that females can discriminate between hosts parasitized by a conspecific or by themselves and adapt the sex of the eggs they lay accordingly.     

Oviposition decisions in an endoparasitoid under self-superparasitism conditions

Superparasitism is a widespread phenomenon. Having accepted superparasitism, mated female parasitoids must decide on the sex of each egg they subsequently lay into the same host. Theory predicts that this decision is either based on host quality, when more male eggs are laid in hosts that are already parasitized because they are perceived to be of poorer quality; or more eggs are laid of the sex that is most likely to be a strong larval competitor, i.e. generally females.Anastatus disparis is a facultative endoparasitic egg parasitoid. We used ‘artificial’ hosts to explore outcomes of decision making by A. disparis during superparasitism under a manipulated absence of larval competition. When only one egg was laid it was always female. As the number of eggs laid increased, so more of them were male. This supports the theory that oviposition decisions are based on host quality; more male eggs were laid in hosts that were already parasitized and thus of poorer quality.In a second experiment, eggs were exposed to parasitoids for different periods of time. Half the eggs were dissected to determine the number of parasitoid eggs that had been laid. The remaining eggs were incubated and the number and sex of offspring that ultimately emerged, following larval competition, were recorded. Under superparasitism conditions fierce larval competition ensued; only one offspring survived and they were predominantly female.In conclusion, oviposition decisions by female A. disparis accepting self-superparasitism were made based on host quality.     

Incidence of superparasitism in the egg parasitoid,Ooencyrtus kuvanae Howard (Hymenoptera: Encyrtidae)

The encyrtid Ooencyrtus kuvanae is a solitary parasitoid of the gypsy moth, Lymantria dispar L. (Lepidoptera-Lymantridae) that is used in biological control programmes and whose mass rearing is influenced by superparasitism. The purpose of this study was to investigate the effect of the self-superparasitism of O. kuvanae at different host densities (5, 10, 15 and 20), female ages (3 and 5 days) and durations of exposure (1 and 5 days) under various laboratory conditions (25°C?±?1°C, RH 60?±?5% and a 16:8?h light:dark photoperiod) as well as in a new laboratory host, Philosamia ricini (Danovan) (Lepidoptera: Saturniidae) of O. kuvanae. In this study, we determined the rate of egg superparasitism and adult emergence and recorded development time, longevity and body weight. Superparasitism increased with female age and the duration of exposure to parasitoids when females had access to five host eggs. Superparasitism increased the number of parasitoid offspring, but it resulted in male-biased (56.90%) progeny. Furthermore, superparasitism caused deleterious effects to the fitness of the progeny by prolonging the developmental process, and decreasing longevity. For example, we found that when four adults can emerge from one superparasitised host egg, the body size of the parasitoid offspring decreases significantly. Hence, superparasitism should be avoided when mass rearing O. kuvanae.     

Adaptive self- and conspecific superparasitism in the solitary parasitoid Leptopilina heterotoma (Hymenoptera: Eucoilidae)

One of the foraging decisions facing parasitoids is whetherto accept (superparasitize) or reject hosts that have alreadybeen parasitized. An important distinction is whether the hosthas been parasitized by the female parasitoid herself or bya conspecific. In solitary parasitoids, the pay-off from anegg laid in the latter host type (conspecific superparasitism)is the probability that the second egg wins the competitionfor the host and results in an offspring. The pay-off from anegg laid in the former type (self-superparasitism) increaseswith an increasing probability that another female will superparasitizethe host in the near future. When this probability equals one,self-superparasitism and conspecific superparasitism have thesame payoff. However, conspecific superparasitism will generallyhave a higher pay-off than self-superparasitism. It will thereforebe beneficial for a female parasitoid to be able to distinguishbetween a host she parasitized and one parasitized by a conspecific.The degree of benefit depends on the probability of conspecificsuperparasitism in the near future. Using an optimal diet model,I show that when a parasitoid encounters a patch containinga mixture of unparasitized and already-parasitized hosts, afemale that can distinguish between the two types of parasitizedhosts gains more offspring than a female without this ability.However, when parasitoids search a patch together with conspecifics,it is adaptive to self-superparasitize, and the pay-off fromthis ability may be negligible. It is therefore predicted thatwhen a female parasitoid searches a partially depleted patchalone, it will reject the hosts parasitized by itself more frequentlythan hosts parasitized by conspecifics. In contrast, femaleparasitoids searching together are predicted to accept hoststhat they parasitized themselves much more often. The resultsshow that the solitary parasitoid Leptopilina heterotoma (Hymenoptera:Eucoilidae) is able to distinguish between hosts that it parasitizedand hosts parasitized by conspecifics. The predictions of themodel are met in a second experiment that shows that L. heterotomaself-superparasitizes when the probability of conspecific superparasitismis high.     

Intra- and interspecific host discrimination in two closely related egg parasitoids

Intraspecific host discrimination is frequently found in solitary parasitoids, but interspecific host discrimination, where female parasitoids recognize hosts already parasitized by females of other species, is rare. This particular behaviour appears to be adaptive only under specific circumstances. In this paper, we quantified intraspecific host discrimination in Anaphes n. sp. (Hymenoptera: Mymaridae), an endoparasitoid of the eggs of Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) and interspecific host discrimination toward eggs parasitized by Anaphes sordidatus (Girault), a sympatric species competing for the same resource in similar habitats. To examine host discrimination, choice experiments were used where the females had to choose between different categories of eggs (unparasitized, parasitized by Anaphes n. sp. or A. sordidatus). Superparasitism and multiparasitism were avoided in experiments where the female had a choice between unparasitized hosts and hosts parasitized by the same female, by a conspecific or by a female A. sordidatus. When all hosts available were parasitized, conspecific superparasitism occurred more often than self-superparasitism or multiparasitism. These results indicated that females Anaphes n. sp. were capable of self-, conspecific and interspecific discrimination. Self-discrimination followed recognition of an external marking while interspecific discrimination occurred mostly after insertion of the ovipositor. Interspecific discrimination could result from the recent speciation of these species and could be associated with a genotypic discrimination. This behavior appears to be adaptive because of the competition for common hosts between the two parasitoid species.     

寄主大小及寄生顺序对蝇蛹佣小蜂寄生策略的影响

寄主大小模型认为寄生蜂后代性比与寄主大小相关,寄生蜂倾向于在大寄主上产出更多雌性后代,在小寄主上产出更多雄性后代.探讨了以家蝇蛹为寄主时,蝇蛹佣小蜂后代产量和性比变化;单次寄生情况下,寄主大小及寄生顺序对寄生蜂后代性比等影响.结果表明,蝇蛹佣小蜂的产卵期为(8.93±3.34)d,单头雌蜂能产雌性后代(34.11±16.34)头和雄性后代(11.04±8.87)头,且雄性百分比为0.24±0.11.随成蜂日龄的增大,寄生蜂产生雄性后代的比率显著增加.蝇蛹佣小蜂在寄生家蝇蛹时,会优先选择寄生个体较大的蛹;在单次寄生的情况下,蝇蛹佣小蜂倾向于在较大的家蝇蛹内产出更多的雌性后代.     

Parasitisation of Tetrastichus brontispae (Hymenoptera: Eulophidae), a biological control agent of the coconut hispine beetle Brontispa longissima (Coleoptera: Chrysomelidae)

The gregarious endoparasitoids Tetrastichus brontispae Ferrière is one of the important natural enemies of the coconut hispine beetle Brontispa longissima (Gestro), a serious invasive pest on coconut palm plants (Cocos nucifera L.) in Southeast Asia. Development at different temperatures, effect of host and female ages, effect of food and oviposition frequency and superparasitism were investigated in the laboratory. Females were allowed only one attack against one host in all experiments. The wasp developed in a host between 19 and 30°C, whilst no wasp completed its immature development at 16 and 31°C. Host and female ages affected parasitisation. Parasitoid emergence was high on day 0 and day 1 pupal hosts, and younger females produced more offspring than older females. The longevity of the female was affected not only by food supply, but also by oviposition frequency. The female survived longer when oviposition frequency was low. However, the total number of hosts parasitised by the female during her lifetime did not differ at different oviposition frequencies. In superparasitism, although the percentage of adult emergence and body size of offspring decreased with an increasing number of attacks per host, a host parasitised by up to four females could produce parasitoid offspring.     

Functional response and egg distribution among hosts by Pseudapanteles dignus,a larval endoparasitoid of Tuta absoluta

The functional response of a biocontrol agent has been classically pointed out as a quantitative evaluation criterion to understand its killing capacity to an arthropod pest. In this paper, we revisited the functional response of the internal larval parasitoid Pseudapanteles dignus (Muesebeck) (Hymenoptera: Braconidae), a candidate for biocontrol of the South American tomato leafminer, Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae), measured as the daily egg oviposition of single parasitoid females at increasing densities of T. absoluta larvae through dissection of hosts. The functional response curve of the number of parasitized hosts was fitted out taking into account the number of eggs laid and their distribution among hosts at each T. absoluta larval density. The data also allowed us to discuss the self-superparasitism strategy of this parasitoid as an adaptive trait. Pseudapanteles dignus showed a sigmoid shape functional response and a contagious distribution of eggs among hosts, favouring self-superparasitism and laying a similar number of eggs in each superparasitized host at each T. absoluta density. This research is firstly intended to ascertain about the oviposition behaviour of P. dignus and additionally to provide information on its reproduction to be applied in mass rearing procedures and augmentative releases against T. absoluta.     

Performance of Sclerodermus brevicornis,a parasitoid of invasive longhorn beetles,when reared on rice moth larvae

Biological control efficiency can be improved by developing effective mass‐rearing systems to produce large numbers of high‐quality parasitoids. This study explored an alternative host for rearing Sclerodermus brevicornis (Kieffer) (Hymenoptera: Bethylidae), a potential biocontrol agent for the suppression of exotic and invasive wood‐boring longhorn beetle (Coleoptera: Cerambycidae) populations in the European agroforestry ecosystems. We tested larvae of the rice moth, Corcyra cephalonica Stainton (Lepidoptera: Pyralidae), as host for the parasitoid. We quantified the probability and timing of host attack and parasitism as well as reproductive success, offspring production, and the characteristics of adult offspring. As S. brevicornis is a quasi‐social species (multiple females, communally produced offspring broods), we also explored the effects of varying the number of females to which individual hosts were presented, with the aim of determining the optimal female‐to‐host ratio. As time to host attack can be a limiting factor in S. brevicornis rearing protocols, we tested the use of adult females of another bethylid species, Goniozus legneri Gordh, to paralyse C. cephalonica larvae prior to presentation. We identified the conditions within our experiment that maximized offspring production per host and offspring production per adult female parasitoid. We found that C. cephalonica is suitable as a factitious host and, as it is considerably more straightforward for laboratory rearing than cerambycid species, it is a good candidate for adoption by future S. brevicornis mass‐rearing and release programmes.     

Bionomics of Opius bellus (Hymenoptera: Braconidae), an endoparasitoid of Anastrepha fraterculus (Diptera: Tephritidae) in fruit-growing areas of Northwestern Argentina

Opius bellus is a neotropical larval-prepupal parasitoid known to attack the pestiferous fruit fly, Anastrepha fraterculus. Due to interest in the use of native parasitoids in forthcoming fruit fly biocontrol programmes in Argentina, O. bellus was colonised for the first time using laboratory-reared A. fraterculus larvae. A series of experiments were conducted to (1) best achieve an efficient parasitoid rearing by determining optimal larval host age, host:parasitoid ratio and host exposure time and (2) assess their potential as biological control agents by determining reproductive parameters. The most productive exposure regimen was: 7–9 d-old (early and middle third-instars) A. fraterculus larvae for 4 h at a 4:1 host:parasitoid ratio; this array of factors was sufficient to achieve the highest average adult emergence (48%) and an offspring sex ratio at equitable proportion. Increasing both host:parasitoid ratio further than 4:1 and the host exposure time beyond 4 h did not significantly enhance parasitoid female offspring yield. Females produced eggs for 29.5 ± 1.4 days. At 32 days of age, 50% of the females were still alive. The majority of the progeny were produced by females between 20 and 24 d-old. At 26°C, gross fecundity rate, net reproductive rate, intrinsic rate of increase and mean generation time were 20.7 ± 4.2 offspring/female, 9.6 ± 2.5 females/newborn females, 0.06 ± 0.01 females/female/day and 8.4 ± 0.2 days, respectively. The long lifespan and reproductive parameters suggest that this parasitoid species has suitable attributes for mass-rearing.     

Costs and consequences of superparasitism in the polyembryonic parasitoid Copidosoma koehleri (Hymenoptera: Encyrtidae)

Abstract.  1. Polyembryonic wasps provide dramatic examples of intra-specific developmental conflict. In these parasitoids, each egg proliferates into a clonal lineage of genetically identical larvae. If more than one egg is laid in a host (superparasitism), individuals of different clones may compete for food resources.
2. In the polyembryonic encyrtid Copidosoma koehleri , one larva per clone can differentiate into a sterile soldier. It is shown that soldiers are always females, and that they attack intra-specific competitors.
3. Research hypotheses were that (a) clones that develop in superparasitised hosts suffer heavier mortality than clones that develop in singly parasitised hosts, and (b) female clones cause higher mortality to their competitors than male clones, hence larval survival is lower in superparasitised hosts that contain females than in male-only broods.
4. The potential frequency of superparasitism in C. koehleri was manipulated by varying parasitoid–host ratios and exposure durations.
5. As parasitoid densities and exposure durations increased, the frequency of superparasitism rose, brood sizes increased, but the number of hosts that completed development was reduced. The number of offspring per parasitoid female decreased with increasing parasitoid–host ratios. Offspring size and longevity were inversely correlated with brood size. As superparasitism rates increased, fewer all-male broods were produced. Male–female broods were female-biased, suggesting selective killing of males by female soldiers. All-female broods were significantly smaller than all-male broods at high parasitoid densities only, possibly reflecting aggression among soldiers of competing clones.
6. The results support the working hypotheses, and suggest that female larvae outcompete males in superparasitised hosts.     

Inter-larval Competition and its Subsequent Effect on Pediobius furvus (Hym.: Eulophidae) Broods for the Management of Graminaceous Stem Borers

The effects of superparasitism on the size, rate of development, progeny production, sex ratio and percentage parasitism of the parasitoid Pediobius furvus (Gah.) and the number of parasitized hosts producing mature parasitoids were investigated. High parasitoid egg densities in hosts were obtained using two host densities (1 and 2) and three arbitrary parasitoid densities (1, 2 and 5) under two exposure times (12 and 24 h). Superparasitism in P. furvus resulted in a reduction in the number of adults that matured and a reduction in the size and preponderance of male production. Dissections of the parasitized host pupae of the stem borer Chilo partellus (Swinhoe) showed no combat among developing wasps. Immature supernumeraries were eliminated by starvation and/or suffocation. The results showed that P. furvus larvae are not harmed by surplus food material in their hosts and that the female parasitoid preferred to lay more eggs into unparasitized than into parasitized host pupae.     

The developmental strategies and related profitability of an idiobiont ectoparasitoid Sclerodermus pupariae vary with host size

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Effect of expected offspring survival probability on host selection in a solitary parasitoid

Optimal Foraging Theory predicts that parasitoid females should optimize their host selection to maximize their lifetime fitness gain and parasitize the most profitable hosts. In particular, in solitary parasitoids, females should avoid superparasitism, at least when sufficient unparasitized hosts are available. However, when unparasitized hosts are scarce, they should prefer, among already parasitized hosts, those that provide the best survival probability to their progeny, which depends on the age and the developmental stage of the first parasitoid. To test this hypothesis in a solitary ectoparasitoid, Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae), we first assessed the survival probability of a second parasitoid according to the time elapsed since initial parasitism. We then analyzed the female selection behavior in patches containing a mixture of hosts parasitized over various time intervals. Our results showed that the older the opponent larva was, the lower the survival probability of the second parasitoid was. However, when the first individual had reached the prepupal stage, both individuals could complete their development. At this stage, the survival probability of the second parasitoid was surprisingly high but such individuals were reduced in size. Our study also showed that host acceptance by females was strongly correlated with the survival probability of their progeny when the first parasitoid was from 0 to less than 10 days‐old. When the first parasitoid had reached the prepupal stage, females usually rejected these hosts, although the survival probability of the offspring was quite high. This discordance between female host selection behavior and progeny survival probability is discussed.