A re-examination of host relations in the Aphelinidae (Hymenoptera: Ghalcidoidea)

Host relations among the Aphelinidae (Hymenoptera: Chalcidoidea) are highly intriguing from an evolutionary view. Females are usually primary endoparasitoids of whiteflies or scale insects, whereas the development of the male is different and has been used for classification. In heteronomous hyperparasitoids (adelphoparasitoids or autoparasitoids) the male develops as a hyperparasitoid of conspecific females or other endoparasitoid species. We review the consequences of this mode of development which is unique because decisions concerning host selection are inextricably linked with progeny sex ratio. Autoparasitoid field sex ratios can fluctuate dramatically concurrent with changes in the relative availability of male and female hosts. A recent adaptive explanation for these sex ratios involves understanding the reproductive constraints acting on heteronomous parasitoids. Host relations in these parasitoids can show a degree of plasticity. We argue that in many instances autoparasitism may be facultative in nature and should not be used for classification. Heterotrophic parasitism, wherein the male develops as a primary parasitoid of lepidopterous eggs, has been poorly understood in the past due to uncertainties in reports of the biology and taxonomy of heterotrophic parasitoids. The host relations of this group are clarified.     

Mating and oviposition behaviour in the legume pod borer, Maruca testulalis

Experiments were conducted outdoors and in the insect rearing laboratory at the International Institute of Tropical Agriculture (IITA) in Nigeria. Four or five nights pairing duration gave the highest mating percentage and oviposition by the pod borer. Direct observations on adult mating revealed that some males mated more than once. The majority of females mated only once. A one-to-one ratio (with 10 males: 10 females) gave the best results for mating and oviposition. Mating took place between 21.00 h and 05.00 h when temperature ranged between 20–25°C and rh over 80% with peak activity occurring between 02.00 and 03.00 h. Oviposition followed a similar trend to that observed for mating with an equal sex ratio (of 10 or more pairs per cage) giving the highest egg count. The average number of eggs laid was ca. 400 per female. These results show a much higher oviposition potential for Maruca testulalis than has been previously reported. The results are discussed with regard to the rearing of this species under laboratory conditions.

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Accouplement et comportement de ponte de Maruca testulalis

Résumé Tant de l'extérieur qu'au laboratoire (à l'IITA au Nigéria), la durée optimale de maintien des couples de M. testulalis Geyer (Lep. Pyralidae) pour obtenir l'accouplement et la ponte a été de 4 à 5 nuits, toutefois les meilleurs résultats ont été obtenus à l'extérieur. L'observation directe a révélé que contrairement aux femelles, les mâles peuvent copuler plus d'une fois. Le meilleur rapport sexuel pour obtenir l'accouplement et la ponte est 1/1 (10 mâles/10 femelles), suivi de près par un raport avec excédent de mâles. L'accouplement a lieu entre 21 h et 5 h quand la température est entre 20 et 30°C et HR supérieure à 80%; un pic est observé entre 2 et 3 h. La ponte présente la même tendance que l'accouplement, avec une production d'ufs plus élevée pour au moins 10 couples par cage. Le nombre moyen d'ufs par femelle est environ 400. Ces résultats sont supérieurs à ce qui avait été indiqué précédemment. La discussion porte sur les difficultés rencontrées pour élever cette espèce au laboratoire.

     

Oviposition and host-feeding patterns in Aphelinus asychis (Hymenoptera: Aphelinidae) at different aphid densities

Abstract. 1. The patterns of host-feeding and oviposition were examined in Aphelinus asychis Walker, which had been provided with second-instar pea aphids as hosts.
2. Female wasps responded to increasing host density (between one and forty aphids for 24 h) with an increasing tendency to oviposit rather than to feed. Superparasitism occurred at all aphid densities, even when unparasitized aphids were available.
3. Aphids intended for feeding were paralysed and died. Wasps did not feed on and oviposit in the same aphid.
4. Feeding to satiation lasted between 4 min and 42 min. Females that had starved for ≰18 h generally deposited one or more eggs before feeding again, while the reverse was true in wasps that had starved for 21 h.
5. The host-feeding behaviour of A. asychis is determined by a female's nutritional status. At low rates of host encounter, the anhydropic eggs may be resorbed. This reproductive strategy conforms to the destructive non-concurrent type among the Hymenoptera.     

Effects of physiological condition and experience on oviposition behaviour of Trichogramma australicum Girault (Hymenoptera: Trichogrammatidae) on eggs of Helicoverpa armigera Hübner (Lepidoptera: Noctuidae)

We produced an ethogram for and investigated effects of physiological condition and experience on oviposition behaviour for Trichogramma australicum on Helicoverpa armigera eggs. The data, coupled with the ethogram, have enabled the establishment of standard behavioural categories which can be used to assess acceptability of H. armigera eggs to T. australicum reared in vitro or in eggs of other species. Physiological and experiential conditions were investigated using females of three different ages (0–24, 24–48 and 48–72 h old), two types of sexual states (mated and unmated) and two types of ovipositional experience (naive and experienced). Durations of host finding, host examination, and post-ovipositional re-examination were reduced by oviposition experience and were shorter in younger females (0–48 h old). Oviposition experience reduces the duration of the drilling and percentage of females host-feeding. Being mated and young reduces the host examination time in subsequent oviposition bouts. Only ovipositionally inexperienced females host-feed before oviposition and host-fed more frequently than experienced individuals after oviposition. Host-feeding needs consideration for optimal culture.     

Testing multiple hypotheses for the maintenance of male homosexual copulatory behaviour in flour beetles

Diverse animal groups exhibit homosexual interactions, yet the evolutionary maintenance of such behaviours remains enigmatic as they do not directly increase reproductive success by generating progeny. Here, we use Tribolium castaneum flour beetles, which exhibit frequent male homosexual copulations, to empirically test several hypotheses for the maintenance of such behaviours: (1) establishing social dominance; (2) practice for future heterosexual encounters; and (3) indirect sperm translocation. We found no evidence that Tribolium males use homosexual copulations either to establish dominance or to practice behaviours that increase their subsequent heterosexual reproductive performance. Our results provide limited support for the hypothesis of indirect sperm translocation: when males from two genetic strains mated with females immediately following a homosexual copulation, females produced progeny sired not only by the directly mating male, but also by that male’s homosexual partner. However, this phenomenon was detected in only 7% of homosexual pairs, and in each case such indirectly sired progeny accounted for < 0.5% of females’ total progeny. Direct observations indicated that mounting males often released spermatophores during homosexual copulations. These observations suggest that homosexual copulations may be a behavioural mechanism that allows males to expel older, potentially low‐quality sperm. Additional work is needed to test this new hypothesis, and to determine whether sperm release during homosexual copulations occurs in other groups.     

Mating Behavior of Cephalonomia tarsalis (Ashmead) (Hymenoptera: Bethylidae) and the Effect of Female Mating Frequency on Offspring Production

The courtship behavior of Cephalonomia tarsalis, a solitary semiectoparasitoid of Oryzaephilus surinamensis, was investigated in the laboratory. Courtship behavior includes a series of stereotypic movements. Males play the most active role, executing the majority of courtship action, and females respond with relatively limited observable behaviors. Males typically keep antennae still during encounters with females prior to mounting, which may be correlated with recognition of the female's sexual status. After mounting, males display a series of movements on females, such as antennae touching female's antennae, antennae or mouth touching female's head or thorax, and walking around on female, which may serve to stimulate females towards increased receptivity. Females signal receptivity by assuming a stereotypical posture of remaining stationary, with head down, and antennae still in front of the body. The male then inserts his aedeagus and the pair copulates. After an average of 40.4 s of copulation, females signal the end of copulation by waving the antennae and moving away from the copulation site. Males continue copulating for a short time after females start moving but dismount soon thereafter. After dismounting, the two wasps move away from each other immediately, and they typically begin grooming. Neither males nor females exhibit mating preference based on mate's mating status in both choice and no-choice tests. The male is polygynous and the mated female can mate multiple times within the first 3 days after starting oviposition. However, female mating frequency does not affect the production of female progeny.     

Mating behaviour of the ‘cosmopolitan’ species Phyllognathopus viguieri (Copepoda: Harpacticoida) and its systematical significance

The mating behaviour was studied and recorded on video with individuals of four cultures of Phyllognathopus viguieri from different populations obtained from the interstitial water of a slow sand filter near the river Ruhr (Germany) (Ruhr population), from a compost heap in Bethesda (Maryland, USA) (Maryland population), from a rain gauge in Windsor Campbell farm (Jamaica) (Jamaica population), and a tree trunk with moss in a forest in the municipality of Rio de Janeiro (Brazil) (Brazil population). The mating behaviour was divided into the well‐known initial phase, copula phase and postcopulatory mate guarding phase. An additional phase prior to the initial phase serves to recognize the female, the recognition phase. The mating behaviour is identical in the males of the Jamaica and Brazil populations of P. viguieri. A postcopulatory mate guarding phase is not found in these two groups. Here, we refute the hypothesis, that a postcopulatory mate guarding phase is found in taxa in which only adult males grasp adult females. The males of the Ruhr and Maryland populations differ from each other in their mating behaviour. Generally, the males of all four populations do not mate with fertilized females which are equally unattractive to the males, i.e., females mate only once in their lifetime to produce offspring. These results corroborate the view that the different populations of P. viguieri do not belong to a single cosmopolitan species.     

Sexual reproduction in Daphnia pulex (Crustacea: Cladocera): observations on male mating behaviour and avoidance of inbreeding

1.  Mating behaviour in Daphnia appears to rely on random contact between males and sexual females rather than diffusible pheromones. Males may be able to discriminate sexually receptive females from females in other developmental stages and increase their mating efficiency. Males may also use chemical signals to avoid mating with females from the same clone and avoid the severe inbreeding depression that has been documented for intraclonal mating. The present study used experiments to test for the avoidance of intraclonal mating and assess male mating efficiency in D. pulex .
2.  Three clones were examined for the avoidance of intraclonal mating by providing males with an opportunity to mate with females of the same or two different clones. The proportion of intraclonal matings did not differ from the proportion of interclonal matings, suggesting that D. pulex males do not use kin discrimination to avoid mating with females from the same clone.
3.  The proportion of mated females decreased with increasing numbers and density of sexual females when exposed to a single male. This observation suggests that a male spends more time pursuing and copulating with sexually receptive females than non-receptive females and there is insufficient time to mate with all sexual females. The decrease in proportion of females mated could also be the result of sperm depletion in the male. Sperm depletion is unlikely to occur in nature because sexually receptive females are much rarer than in the experimental conditions.     

Mating Behavior of Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae): Potential for Outbreeding in a Predominantly Inbreeding Species

The mating behavior of the quasi-gregarious egg parasitoid Trissolcus basalis (Wollaston) was investigated under field conditions. Trissolcus basalis has female-biased sex ratios and is a protandrous species, with males emerging 1–2 days before females. Males competed aggressively for control of the egg mass, with one male assuming dominance and control of the egg mass, although changes in dominance occurred at least once on each egg mass observed. Typical mating behavior involved the dominant male mating his sisters immediately upon their emergence from the egg mass. These behaviors are characteristic of an inbreeding species that manifests local mate competition. However, several aspects of the mating behavior of T. basalis are inconsistent with that of an inbreeding species. Over 18% of emerging females were not mated by the dominant male upon emergence, 13% of females were not observed to be mated at all and may have left their natal site as virgins, 25% of females were mated multiple times and sometimes by multiple males, females remained near the natal site for up to several hours after emergence before emigrating, and males dispersed away from the natal site during female emergence. Trissolcus basalis may be a predominantly inbreeding species but its emergence and mating behavior suggest that low-frequency outbreeding is also likely to occur.     

Mating Behavior and Colony Founding of the Slave-Making Ant Formica sanguinea (Hymenoptera: Formicidae)

The mating and postmating behavior of reproductives belonging to two sympatric dulotic colonies of the facultative slave-making ant Formica sanguinea was analyzed in the field. Our observations showed that the European blood-red ant adopts a reproductive behavior similar to the male aggregation syndrome. Newly mated females return to a dulotic colony and often wait for a raid. Following a slave raid is an advantageous strategy to locate and invade host nests and to establish a new dulotic colony. In the laboratory, the following modes of colony founding were studied: independent, adoption, alliance, usurpation, and brood raiding. Independent foundation was possible only when several females were kept together. Alliance was obtained with females of two potential slave species (F. cunicularia, F. rufibarbis). Usurpation and adoption were more frequent in the incipient than in the mature host colonies. Mixed colonies were always obtained after the sack of the host pupae. It seems likely that, rather than conspecific adoption followed by budding, F. sanguinea relies on temporary parasitism to start new colonies.     

Variable Female Mating Positions and Offspring Sex Ratio in the Spider Pityohyphantes phrygianus (Araneae: Linyphiidae)

Chromosomal sex determination and male heterogamety have been thought to seriously impede direct sex ratio control. However, in Pityohyphantes phrygianus, a solitary sheetweb spider with a skewed sex ratio, earlier experimental studies suggested that there are options for female control of offspring sex ratio, if females change their position during the normal mating sequence. Here we show that under natural conditions there is considerable between-female variation in positions, especially after termination of mating. Computer simulations of the orientation of female inner genitalia suggest that sperm are placed in different storage sites depending on the positions adopted. This means that a specific position after mating might potentially influence offspring sex ratio. The variance in offspring sex ratio among females in earlier experiments was binomially distributed, which leads us to conclude that females control the mean sex ratio but do not exercise direct control of the sex of individual offspring.     

Subsocial behaviour in Oniticellus cinctus (Coleoptera, Scarabaeidae): effect of the brood on parental care and oviposition

ABSTRACT. In Oniticellus cinctus (F.) the nest chambers each contain about twenty brood balls. Females enlarge the brood balls during the egg and larval stages and remain in the chamber for the whole period of brood development (1 month); they then make a new nest after 1 week. The presence of the brood releases parental care and ensures that the mother remains in the nest: she repairs defects in the brood balls and the nest, and expels other O. cinctus females. A new ball is formed around a naked O. cinctus larva, but larvae of other species are killed. In addition, the brood inhibits oviposition: removal (or addition) of brood balls stimulates (or inhibits) egg laying. In inhibited ovarioles, existing follicles are resorbed and production of new ones ceases. Control of clutch size by the brood is an adaptation to the nest structure and life history of O. cinctus. It may have an important role in the reproductive strategy of other insects with parental care.     

The Race for Females: The Mating System of the Red Mason Bee, Osmia rufa (L.) (Hymenoptera: Megachilidae)

The ecology of a species strongly influences the strategies with which males and females maximize their lifetime reproductive success. When males and females do not invest equally in offspring, the sex with the higher parental investment becomes a rare resource for the other. The spatial and temporal distribution of the limiting sex forms the basis of the mating system. In nest-constructing Aculeata such as the red mason bee, Osmia rufa, females perform intensive brood care, whereas males do not invest in their offspring but instead compete for access to mates. Receptive females of this species are widely distributed and do not assemble at certain places. Therefore, territorial behavior is not an advantageous mating tactic for males, which instead search for females within individual home ranges usually centered around food plants. The unmarked or defended home ranges of different males may completely overlap. Competitive searching leads to a random distribution of matings among males that is largely independent of body size. The mating system of O. rufa can be described as scramble competition polygyny.     

Body size and alternative mating tactics in the beewolf Philanthus zebratus (Hymenoptera; Sphecidae)

A study of the mating behaviour of males of the beewolf Philanthus zebratus revealed that in one population males display variability in mating tactics and that this variability is related to male body size. There was a tendency for large males to patrol the airspace above the nesting area while smaller males were territorial adjacent to it. The mean sizes of the two groups of males were significantly different, although the size ranges of the two groups overlapped. Only 2.5% of the males were observed to undertake both mating tactics, at different times. Observations are presented on daily and seasonal activity patterns and on the relative location of nests, territories, and patrolling males. A second population, with lower nest density, was observed for several days, revealing only territorial males. It is suggested that the presence of patrolling males is related to the higher nest density of the one population. The fact that patrolling males tend to be relatively large is possibly related to flight energetics or simply to the ability of large males to seize females, which are usually larger than males, in mid-air.     

Male Behavior and Regulation of Worker Mating in a Ponerine Ant, Pachycondyla (Bothroponera) sp. (Hymenoptera: Formicidae)

Male behavior and regulation of worker mating were observed in Indonesian Pachycondyla sp. The colonies had some mated workers but only one gamergate (mated and egg-laying workers). Males frequently tried to mate with workers in their natal nests irrespective of worker age and dominance rank, however, workers never accepted mounting of nestmate males. Worker mating occurred only under the gamergateless condition with alien males.     

Costs and Benefits of Mating in the Grasshopper Chorthippus parallelus (Caelifera: Acrididae)

Fitness consequences of different levels of mating benefits, viz., (1) virgin, (2) seldom mated, (3) free mating chances with the same male throughout the female's life span, and (4) free mating chances with three males always offered, were investigated in female meadow grasshoppers Chorthippus parallelus. No differences among the treatments were apparent in egg number per pod, total egg number per female, and hatching success of larvae. Seldom-mated females had offspring with the heaviest average larval dry weight. We did not find costs of mating in terms of reduced female longevity and residual dry weight between the treatments.     

The role of the androconia in the mating behaviour of the European skipper, Thymelicus lineola, and evidence for a male sex pheromone

Abstract. Laboratory experiments were carried out to determine the role and characteristics of male scent scales (androconia), located in patches (stigmata) on the forewings of Thymelicus lineola (Ochsenheimer) (Lepidoptera: Hesperiidae). In behavioural tests, there were 30—40% fewer matings of virgin females by males with the stigmata removed or where females had their antennal sensilla covered with nail polish, when compared to sham-treated controls. These reductions occurred despite a large increase in male courtship activity. No physical contact was observed between male wings and female antennae during mating. A scanning electron microscope study of male wings and female antennae showed that: (1) lysis zones form over time on the androconia and pieces (osmo-phores) distal to these zones break off; (2) while few of these zones are present at emergence they are numerous in-day-old males; (3) during a single episode of courtship and mating approximately two-thirds of the available osmophores break off, yet the commonly observed courtship between males does not appear to result in osmophore release; and (4) osmophores were never observed on the female antennae. Electroantennogram recordings (EAGs) indicated that: (1) female antennae responded strongly to odours from male forewings but not from other parts of either males or females; (2) female antennae responded more strongly to young field-collected males than to newly-emerged or old, field-collected males; (3) female antennal response to male forewings was reduced if the androconia were scraped off, and was eliminated if the stigmata were removed; (4) male antennae gave weak EAG responses to both male and female forewings. We conclude that males release pheromone via the breakage of osmophores during courtship. We propose that the frequently observed refusals by females of courting males are at least partially dependent on the quantity or quality of male pheromone released, which are in turn correlated with the male's age and mating status.