Mating system of Bracon hebetor (Hymenoptera: Braconidae)

Abstract.

• 1 We report on the mating system of a field population of the parasitic wasp, Bracon hebetor, on a corn pile infested by the Indian meal moth, Plodia interpunctella. We demonstrate that the mating system is based upon male scramble competition polygyny with male aggregations on high places on the corn.
• 2 The sex ratio among adults was greater than 80% males on the surface of the corn, whereas below the surface the sex ratio was less than 45%. Males actively courted females on the surface, but there were no aggressive interactions among males during courtship or mating.
• 3 Approximately 20% of the females found on the surface of the corn had no sperm in their spermathecae, regardless of age, but the numbers of unmated females decreased later during the day.
• 4 In laboratory studies we showed that females from this population oviposit a female biassed sex ratio, and that only 14% of females were mated before dispersing from their place of emergence.
• 5 Thus sib-mating is unlikely in this gregarious parasitoid. This outcrossing mating system probably arose because of severe inbreeding depression that B.hebetor suffers via a sex locus: diploids that are heterozygous at the sex locus develop into females, but homozygous diploids are male and are generally inviable. The female biassed sex ratio may have evolved in B. hebetor in response to males being the more expensive sex, females dispersing more frequently from the population than males, or a fraction of females remaining unmated in the population.

     

Brood guarding in a bethylid wasp

Abstract.

• 1 Atypical of the parasitoid Hymenoptera, female Goniozus nephantidis Muesebeck remain with offspring until they pupate. This behaviour will only have evolved if the consequent reduction in fecundity is outweighed by fitness returns.
• 2 G.nephantidis competes for hosts with conspecific and non-conspecific parasitoids. The effectiveness of G.nephantidis at deterring superparasitism and multiparasitism is tested. Brood survivorships were compared when G.nephantidis and Bracon hebetor Say intrude on hosts with the mother present and absent and with offspring at different developmental stages.
• 3 Laying by the intruder reduced brood survivorship. Guarding reduced oviposition on unparasitized hosts by intruding females, and prevented superparasitism of hosts with egg broods. Hosts with larval broods were rarely superparasitized. Multiparasitism was common except on hosts with guarded larval broods, but even here survivorship was reduced.
• 4 Competitive asymmetries determined the outcome of contests for possession of host resources.
• 5 The costs of remaining seem at least partially offset by the prevention of oviposition by competing parasitoids.

     

Effect of host availability on reproduction and survival of the parasitoid wasp Trichogramma minutum

Abstract.

• 1 We tested the hypothesis that females of the egg parasitoid, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), could adjust their fecundity schedule according to host availability and that there was a negative correlation between reproduction and survival in these wasps.
• 2 Newly-emerged females were provided with an unlimited or limited number of hosts in the first trial and with either unlimited, limited or zero hosts in the second trial.
• 3 When hosts were unlimited, wasps had the highest rate of reproduction in the first day, which decreased dramatically thereafter. When hosts were limited, wasps from the two trials differed in their response. In Trial I, females with limited hosts had lower first-day fecundity than, and the same subsequent-day fecundity as, those with unlimited hosts. However, in Trial II, females with limited host had a lower first-day but a higher subsequent-day fecundity than those with unlimited hosts. This indicates variation in Trichogramma's ability to shift its fecundity schedule in response to host availability.
• 4 There was a positive (rather than a negative) correlation between reproduction and survival. Wasps that oviposited (in host-unlimited treatment) had greater longevity than those that could not (in host-unavailable treatment).
• 5 The sex ratio of the progeny produced by wasps in both host-unlimited and limited treatments shifted gradually from a female to a male bias as the wasps aged.
• 6 We consider the ability of parasitoids to adjust their fecundity schedule as an adaptation to changing host resources and discuss our findings with regard to theories of life history evolution.

     

Effects of pupal treatment with Proteus® and Metarhizium anisopliae sensu lato on functional response of Habrobracon hebetor parasitising Helicoverpa armigera in an enclosed experiment system

The ectoparasitoid wasp, Habrobracon hebetor Say (Hymenoptera: Braconidae) is a valuable biocontrol agent attacking larval stages of many lepidopteran pests including Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). The possibility of combined application of this parasitoid wasp with other biological control agents and chemical insecticides is necessary for the success of an Integrated Pest Management (IPM) programme. One of the most important behaviours of a parasitoid refers to the number of hosts attacked by per parasitoid as a function of host density. In this research, the single and joint effects of the insecticide, Proteus® and the entomopathogenic fungus, Metarhizium anisopliae sensu lato (Metsch.) Sorokin (Hypocreales: Clavicipitaceae) (isolate M14) were studied on the functional response of H. hebetor. Newly mated females (<24?h) from treated pupae of H. hebetor were used to study the functional response of this parasitoid wasp to different densities of H. armigera larvae. A type II functional response was observed in all treatments. The results revealed that control and Proteus® treatments developed the lowest (0.5599?±?0.0373?h) and the highest (0.5709?±?0.0443?h) handling time, respectively. The most and the least values of attack rate were observed in control (0.0996?±?0.0164?h^?1) and Proteus® (0.0776?±?0.0119?h^?1) treatments, respectively. The maximum theoretical parasitism rate (T/T_h) was estimated to be 42.86 for female control wasps. After conducting field experiments on functional response along with life table parameters, M. anisopliae sensu lato (isolate M14) may be used as a compatible biocontrol agent with H. hebetor in IPM programmes of H. armigera.     

Description of the antennal sensilla of <Emphasis Type="Italic">Habrobracon Hebetor</Emphasis>

The morphology of the antennal sensilla of both male and female Habrobracon hebetor (Say) (Hymenoptera: Braconidae) is described using Scanning Electron Microscopy complemented with Transmission Electron Microscopy. Five types of innervated sensilla as well as uninnervated microtrichia were found. These types are: sensilla trichodea; s. chaetica; s. basiconica; s. coeloconica; and s. placodea. No differences in shape, basic structure, and types of antennal sensilla were found between males and females. The types of sensilla of both sexes of H. hebetor were compared with what has been described in other parasitic Hymenoptera, and their putative functions are discussed with reference to their morphology, distribution and ultrastructure.     

Genetic variation in the behavioural mechanisms involved in the response of the egg parasitoid Trissolcus brochymenae to contact chemical cues left by the pest Murgantia histrionica

1. The ability of parasitoid females to perceive chemical traces left by their hosts is of utmost importance in the host location process. The behaviours involved in such ability have thus most likely been promoted by natural selection in the course of the evolutionary time. For this to happen, however, there must be significant genetic variation in natural populations on which natural selection could act.
2. Using the isofemale line method and motion analysis, we detected significant intra‐population genetic variation for several walking behaviour traits of the egg parasitoid Trissolcus brochymenae (Hymenoptera: Scelionidae) females responding to chemical traces left by its host Murgantia histrionica (Heteroptera: Pentatomidae).
3. Besides opening new avenues of research on the reproductive strategies, behaviour, and biological control potential of parasitoid wasps, these results also have implications for understanding their life‐history evolution in general.

     

Protandry in the parasitoidCardiochiles nigriceps,as related to its mating system

A laboratory study was conducted of the emergence times and mating success ofCardiochiles nigriceps Viereck (Hymenoptera: Braconidae), a larval parasitoid of the tobacco budworm,Heliothis virescens (F.) (Lepidoptera: Noctuidae), and in view of this information, the parasitoid's mating system was explored. Observations on laboratory populations ofC. nigriceps confirmed the occurrence of two types of protandry, i.e. seasonal and diurnal; males emerged 2 days before females in a generation, and emerged about 1 hr before females on a given day. An experiment on mating success showed that newly emerged females are not receptive to males that emerged 1 hr earlier, but that these males and females often mate successfully after 1 more hr. The experiment also showed that 1- to 5-day-old males are more successful than 1-hr-old males in mating with 0- to 1-hr-old females. Thus, it is argued that males emerging on a given day have a disadvantage in competition for mates with males that emerged days earlier, and that this disadvantage may serve as a selection pressure toward diurnal protandry. A monogamous mating system for females ofC. nigriceps is suggested because sexual selection would be expected to be strong in species exhibiting both seasonal and diurnal protandry. A possibility of a sibling mating system inC. nigriceps is questioned partly because newly emerged females are unreceptive to males that emerged 1 hr earlier and partly because this parasitoid is solitary in what is considered highly dispersed hosts in the field.     

Male Mate Choice in Trichogramma Turkestanica

In most animals it is the sex that invests the most in reproduction, generally the female, that expresses mate choice. However, in numerous species, males or both males and females are choosy. We investigated mate choice in males of the egg parasitoid Trichogramma turkestanica Meyer (Hymenoptera: Trichogrammatidae). We tested the impact of age and feeding status of males on their capacity to choose between virgin or mated and kin or non-kin females. As expected, males showed no preference between kin and non-kin mates, but inseminated virgin females over mated ones. No effect of age on the level of choosiness was found, but unfed males were choosier than fed ones. This is the first study to show an effect of feeding status of males on mate choice in insect parasitoids.     

Male morphology,performance and female mate choice of a swarming insect

1. Females of many species select mates based on important morphological and performance traits. This behaviour likely facilitates reproductive success thus exhibiting sexual selection. Most such studies have evaluated a single morphological variable, and only a minority of them studies the effects of behaviour and performance (functional capacity: Irschick et al., Evolutionary Ecology Research, 10 , 177–196, 2008) at all.
2. This study compares male morphological and performance traits differing in three variables that may correlate with fitness: condition index, flight ascent speed and resistance to torpor in relation to male mating success in the parasitoid wasp species Alabagrus texanus (Braconidae), a species where males swarm about emerging virgin females that display both choosy and non-choosy behavioural phenotypes.
3. Males that successfully mated with choosy females exhibited higher condition indices and somewhat stronger resistance to torpor than other males. Conversely, males that mated with non-choosy females did not differ in any trait from other males we measured. Early-arriving males did not have higher condition indices or greater mating success than other males.
4. Thus, both morphological and performance traits contributed to male success, which acted through female choice, indicating a role for sexual selection. Patterns of choice further differed among females, independent of male traits.

     

Reproductive Isolation and Genetic Variation between Two “Strains” ofBracon hebetor(Hymenoptera: Braconidae)

Bracon hebetorSay(Hymenoptera: Braconidae) is known primarily as a parasitoid of pyralid moth larvae infesting stored grain. In the 1970s, a parasitoid identified asB. hebetorwas released for control ofHeliothis/Helicoverpaspp. (Lepidoptera: Noctuidae) on the island of Barbados. Because life-history traits of this parasitoid differed from those reported forB. hebetorfrom the United States, we conducted a series of laboratory experiments to determine whether this parasitoid was (i) a population ofB. hebetorthat attacks noctuids in the field or (ii) a different species fromB. hebetor.We confirmed thatHeliothis virescens(F.) was a more suitable host for the Barbados strain than forB. hebetor.However, a stored-grain infesting pyralid,Plodia interpunctella(Hübner), was a more suitable host for the Barbados strain than wasH. virescens.Reciprocal crosses between the Barbados strain andB. hebetorshowed that the two populations were reproductively isolated. No mating was observed during a series of 30-min observations of reciprocal crosses, and the crosses produced only male offspring. Examination of each female's spermatheca confirmed that females were not fertilized. Sequence analysis of a 517-bp fragment of the mitochondrial 16S rRNA gene revealed that two populations ofB. hebetorfrom our laboratory were identical but differed in sequence by 2% from the Barbados strain. Collectively, our results indicate that the Barbados strain is a distinct species fromB. hebetor.     

Confirmation of the existence of alloparasitoids in nature – host relationships of an Australian Coccophagus species that parasitizes mealy bugs

Heteronomous hyperparasitoids are parasitic wasps with sex‐related host relationships that are unique to a group of genera in the chalcidoid family Aphelinidae. Females are primary parasitoids of various sedentary bugs (mainly, scale insects, mealy bugs, and whiteflies). Males, in contrast, are hyperparasitic, and they frequently develop at the expense of female conspecifics. Alloparasitoids constitute a special category of heteronomous hyperparasitoids, for their males never develop through female conspecifics. The existence of alloparasitic host relationships and the utility of the category ‘alloparasitoid’ have both been questioned. Here, we present results that confirm the existence of the alloparasitic way of life among heteronomous aphelinids. We investigated an undescribed species of Coccophagus (Hymenoptera: Aphelinidae), an Australian parasitoid that attacks the introduced lantana mealy bug, Phenacoccus parvus Morrison (Homoptera: Pseudococcidae), in Queensland. A year‐long field survey regularly returned large numbers of female Coccophagus spec. near gurneyi individuals from P. parvus (total n = 4212), but only few males (n = 11). Males emerged from samples only when the encyrtid parasitoid Anagyrus diversicornis (Howard) (Hymenoptera: Encyrtidae) was present in samples in relatively high numbers. Laboratory oviposition tests confirmed that A. diversicornis is a male host and showed that males do not develop at the expense of conspecific females. Other studies show that males are attracted in numbers to virgin females held in cages above mealy bug‐infested Lantana montevidensis (Spreng.) Briq. (Verbenaceae) in the field, demonstrating that they are common in the population as a whole. This confirms that the males need hosts other than conspecific females and that their usual hosts are present outside of the lantana/P. parvus system. The implications of these results for developing a realistic classification of heteronomous host relationships are discussed.     

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

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Virgins in the wild: mating status affects the behavior of a parasitoid foraging in the field

In haplodiploid organisms, virgin females can produce offspring, albeit only sons. They may therefore face a trade-off between either: (1) searching for hosts and producing sons immediately; or (2) searching for mates and perhaps producing both sons and daughters later in life. Although this trade-off raises a theoretical interest, it has not been approached experimentally. The objective of this article is thus to document the effect of mating status on the foraging behavior of a haplodiploid parasitoid. For this, we recorded the behavior of virgin and mated female Lysiphlebus testaceipes (Hymenoptera: Braconidae) after being released, in the field, on a colony of their aphid hosts. Half of the virgin females were mated by a wild male after less than 10 min of foraging. Evidently, virgin females attract males while foraging on host patches, so that the two activities are not mutually exclusive. Nonetheless, virgin females stayed motionless more often and for longer periods than mated females. Consequently, they attacked aphids at a lower rate, and in turn, attacked fewer aphids on each patch. Moreover, contrary to mated females, virgins did not aggregate their progeny on large patches. We conclude that in L. testaceipes, the trade-off may not be as hypothesized. By dispersing across patches more than mated females, virgins could promote future mating opportunities for their sons and increase their inclusive fitness. However, by moving too frequently, females may lose immediate mating opportunities for themselves and the immediate advantage of producing offspring of both sexes. The observed behavior of virgin L. testaceipes females on host patches could reflect an optimal solution to such a trade-off.     

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     

Mate Finding Via a Trail Sex Pheromone by <Emphasis Type="Italic">Aphytis melinus</Emphasis> DeBach (Hymenoptera: Aphelinidae) Males

Finding mates is frequently problematic for parasitoid wasps. In some parasitoid species, males rely on volatile, airborne sex pheromones for locating mates, while in others they rely on contact, trail sex pheromones. This study sought to shed light on the mate finding mechanism of males of Aphytis melinus. Specifically, the goal was to determine whether A. melinus males use airborne or contact pheromones, or both, for locating mates. The study showed that A. melinus males rely on a contact, trail sex pheromone for locating mates: A. melinus males responded to substrate-borne cues left by virgin females, while they did not respond to airborne cues from virgin females. Specifically, males more frequently encountered virgin females when the females walked across an arena to a fixed encounter point compared to when they were manually placed at the encounter point, and spent greater than expected time on surfaces previously visited by virgin females compared to control surfaces not visited by females. In contrast, males did not respond to airborne cues from virgin females in an airflow olfactometer nor to traps baited with virgin females in the field, and spent similar lengths of time on surfaces visited by newly-mated or 24-h mated females versus control surfaces not visited by females. The main effect of the trail sex pheromone on the behavior of A. melinus males was to direct their search and, so, increase the likelihood of encountering mates. This effect apparently is not preceded by longer-range attraction of males via an airborne female sex pheromone. Overall, the results of this study support a hypothesis in which A. melinus males searching on substrates on which females may be present rely exclusively on a trail sex pheromone to locate mates.     

A plant pathogen causes extensive mortality in an invasive insect herbivore

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Using hosts of mixed sizes to reduce male-biased sex ratio in the parasitoid wasp, Diglyphus isaea

Overproduction of males in mass rearing of parasitic Hymenoptera contributes to higher costs for biological control because only females directly kill pests. We present a technique, based on manipulating host composition, to generate less male‐biased sex ratios in parasitoid species that adjust their sex allocation in response to relative host size. Our system consisted of chrysanthemum, Dendranthema grandiflora Tzvelev var. ‘Miramar’; a leafminer, Liriomyza langei Frick (Diptera: Agromyzidae); and a commercially available parasitoid, Diglyphus isaea (Walker) (Hymenoptera: Eulophidae). We compared the offspring sex ratios of D. isaea females presented with different compositions of L. langei larvae on chrysanthemum. Presenting individual females with only large hosts increased mean sex ratio from 32 to 67% male over 2 days. However, presenting individual females with progressively larger hosts over 1 or 2 days reduced mean sex ratio from 90 to 100% male to less than 30% male. Groups of females produced sex ratios around 58% male if presented with both plants infested by only small hosts and plants infested by only large hosts. In comparison, groups of females produced sex ratios around 48% male if presented with plants infested by both small hosts and large hosts. We compared the use of both small hosts and large hosts to only large hosts for simulated mass rearing of wasps over 8 weeks. Using both small hosts and large hosts produced similar numbers of wasps as using only large hosts, but reduced mean sex ratio of weekly cohorts from 66% male to 56% male. The two techniques produced females of similar size, but using both large hosts and small hosts produced slightly smaller males than using only large hosts. The use of both small hosts and large hosts for mass rearing of D. isaea could reduce actual costs of females by 23%.     

Superparasitism and mutual interference in the egg parasitoid Anagrus delicatus (Hymenoptera: Mymaridae)

Abstract.

• 1 In nature, interference among Anagrus delicatus (Hymenoptera: Mymaridae) parasitoids reduced the per-capita number of hosts parasitized. Interference increased with parasitoid density.
• 2 Anagrus delicatus did not avoid parasitizing hosts that had recently been parasitized by conspecific wasps. Evidence indicated that this superparasitism was largely a random process, increasing with the ratio of parasitized to unparasitized hosts.
• 3 Individual parasitoid efficiency, the number of hosts killed per wasp per unit time, decreased with increasing wasp density. This occurred whether wasps searched the patch together (simultaneously) or one by one (sequentially), and was the result of an increase in time spent superparasitizing hosts at higher wasp density. This is known as indirect mutual interference.
• 4 Increasing numbers of parasitoids together on the same patch caused a significant decline in the rate and per-capita number of hosts parasitized. However, there was not a correspondent decline in searching efficiency with increasing wasp density (i.e. no direct mutual interference).
• 5 These forms of parasitoid density dependence should contribute to the stability of the host—parasitoid interaction.

     

Flexible patch time allocation by the leafminer parasitoid, Opius dimidiatus

Abstract.

• 1 The ability to use flexible decision rules can be an advantage to parasitoid females searching for patchily-distributed hosts. In a series of laboratory experiments the hypothesis that Opius dimidiatus, a solitary parasitoid of the chrysanthemum leafminer (Liriomyza trifolii), adjusts the time she allocates to searching for her larval hosts in response to both patch qualities and experiences with hosts was tested by varying such patch parameters as area, presence of host mines and density of host mines, and by allowing ovipositions and encounters with parasitized hosts.
• 2 Though leaf area was not a factor, the presence of host mines in a leaf did increase the time a female O.dimidiatus spent searching, over time spent on unmined leaves.
• 3 When host mine density was increased, females responded by increasing their search period in a density-dependent manner, suggesting a perception of patch quality.
• 4 Ovipositions in hosts caused females to reset their‘giving-up time’(GUT), or increase search intensity, by adding an amount of search time that increased with each successive oviposition. Conversely, encounters with parasitized (unsuitable) hosts incremented the GUT, but by an amount that decreased with each successive encounter.

     

Putative Host Volatiles Used by <Emphasis Type="Italic">Habrobracon hebetor</Emphasis> (Hymenoptera: Braconidae) to Locate Larvae of <Emphasis Type="Italic">Plodia interpunctella</Emphasis> (Lepidoptera: Pyralidae)

Indianmeal moth, Plodia interpunctella (Hubner) (Lepidoptera: Pyralidae), is a post-harvest pest of grains, milled and processed food, processing plants, warehouses and bakeries. The parasitoid, Habrobracon hebetor (Say) is among the most important natural enemies of Pyralidae infesting stored grains and grain products. Many parasitoids use semiochemicals originating from their hosts, or host’s habitat as cues to locate hosts, hosts’ food or habitat. The authors used Y-tube and four-way olfactometers to assay responses to stimuli with the moth host and thereby understand the role of host-associated semiochemicals in host location by H. hebetor. Responses of mated parasitoid females were assayed to the following stimuli: P. interpunctella sex pheromone, female adults, larvae, or hexane extracts of residue of the rearing medium. Generally, host-related odor sources generated stimuli that elicited better responses than those to blank controls. Previous exposure to odor sources from the host shortened latency periods and response times compared to naïve females. Odors emanating from live moth larvae elicited the strongest responses. When responses from the four odor sources were compared in a four-way olfactometer, it was confirmed that volatiles from larval moths elicited the strongest attraction to the parasitoid. The involvement of host-specific chemical cues in both long and short range host location by female parasitoid is discussed.