Make love not war: a common arthropod defence compound as sex pheromone in the forest cockchafer Melolontha hippocastani

Mate finding in the forest cockchafer, Melolontha hippocastani, occurs during a spectacular swarming period at dusk. The swarming flights are mainly performed by males whereas most of the females stay within the host trees and continue feeding. Males orientate towards damage-induced green leaf volatiles (GLV) allowing location of mechanically damaged foliage. In order to distinguish between unspecific leaf damage and damage caused by feeding females, male cockchafers orientate by a sex attractant. Here we show this compound to be 1,4-benzoquinone, which has been known for more than 40 years as a highly effective defence compound used by numerous arthropod species to repel enemies. 1,4-benzoquinone synergistically increased the number of males caught in GLV-baited funnel traps during the swarming period. Significantly more males landed on wire cages baited with a combination of 1,4-benzoquinone and GLV than on cages baited with only GLV. The results suggest that the sex pheromone of M. hippocastani might have evolved from a primary role as a defence compound.     

Intimate Rendezvous in a Tritrophic Context? Nothing but the Girls for Male Lysiphlebus testaceipes

In insects, mating often occurs after natal dispersal, and hence relies on a coevolved combination of sexual communication and movement allowing mate encounter. Volatile sex pheromones are widespread, generally emitted by females and triggering in‐flight orientation of conspecific males. In parasitoid wasps, unmated females can start laying unfertilized eggs via parthenogenesis so that host patches could serve as sites of rendezvous for mating. Males could therefore use cues associated with host patches to focus their search on females that have successfully found oviposition sites. We hypothesized that in parasitoids exploiting herbivorous hosts, sex pheromones, and herbivore‐induced plant volatiles (HIPV) should act in synergy, triggering male orientation toward ovipositing females. We tested this hypothesis with the aphid parasitoid Lysiphlebus testaceipes. Results from both field and laboratory experiments show that males are strongly attracted to virgin females, but that volatiles from aphid‐infested plants have no effect on male orientation, neither has a cue, nor in interaction with the female sex pheromone. The absence of synergy between sex pheromones and HIPV contrasts with results on other species and raises interesting questions on mating systems and sexual selection in parasitoid wasps.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

Male load-lifting capacity and mating success in the swarming caddis fly Athripsodes cinereus

Abstract. Males of the caddis fly Athripsodes cinereus (Curtis) (Trichoptera: Leptoceridae) swarm above the water surface of lakes and streams. Females enter swarms and are pursued until grasped by a male. The pair couple their genitalia in the air, and then the male alone flies the pair to the shore where they settle and complete the copulation. About 8% of the pairs (total n = 384 pairs) dipped in the water soon after the coupling manoeuvre and about 25% of those then separated. Males in dipping pairs ( n = 13) were on average smaller and relatively older than the males that successfully carried their mate to the shore ( n = 54). No differences were found for flight muscle ratio (weight of flight muscles/total body weight) or relative load (total load/flight muscle weight). Males were larger than females (wing length), though typically female Trichoptera are the larger sex. Large male body size in A. cinereus may be an adaptation for flight during pairing; i.e. larger males are more likely to be able to carry larger loads.     

Time allocation to the reproductive and feeding behaviors in the male cabbage butterfly

Individually marked males of Pieris rapae crucivora, were observed to determine how they allocate time to reproduction and feeding. Males were found to alternately feed and search for females. This manner of time allocation persisted throughout the day. The total times that males allocated to the two behaviors were positively correlated, i.e. those males that spent longer searching for females, also feed for longer periods. Males, however, tended to allocate more time to the female-searching in the morning than in the afternoon, while time allocated to feeding throughout the day. Older males spent more time searching for females in the morning. The body weight of male butterflies also changed as they aged. The results are discussed in terms of both proximal and ultimate aspects of female-search.     

Sex-role reversed nuptial feeding reduces male kleptoparasitism of females in Zeus bugs (Heteroptera; Veliidae)

Males of a variety of taxa occasionally steal food secured by their mates. In some spiders and insects, males rely entirely on this form of intraspecific kleptoparasitism for their subsistence. However, this male strategy may be costly for females and a variety of different female counteradaptations have been proposed. In Zeus bugs (Phoreticovelia spp.), males ride on the back of their mates for extended periods and females produce a gland secretion that males feed on. By experimentally occluding the dorsal glands in females and varying food availability, we show that nuptial feeding by females reduces the extent to which the males kleptoparasitize their mates. We suggest that females have, at least in part, evolved this unique form of nuptial feeding as a counteradaptation to reduce the rate of kleptoparasitism by males.     

Variation in energy reserves and role of body size in the mating system of Anopheles gambiae

Anopheles gambiae mates in flight. Males gather at stationary places at sunset and compete for incoming females. Factors that account for male mating success are not known but are critical for the future of any genetic control strategy. The current study explored variations in nutritional reserves (sugars, glycogen, lipids, and proteins) in wild‐caught swarming and resting males and evaluated the effect of body size and wing symmetry on male mating success. Our results showed that glycogen and sugar reserves are mobilized for flight. Males consume proportionally 5.9‐fold as much energy derived from sugars in swarming activities than when they are at rest. Mated males were on average bigger than unmated ones (P<0.0001). A strong correlation between the left and right wings in both mated and unmated males was found and additional analysis on fluctuating asymmetry did not show any indication of mated males being more symmetrical than unmated ones. The distribution of wing size of mated males was focused around a central value, suggesting that intermediate size of males is advantageous in the An. gambiae mating system. The results are discussed in the context of sexual selection.     

Acoustic orientation of male Aedes diantaeus during mating

The reaction of males of Aedes diantaeus to different auditory stimuli was studied in natural environment during swarming and under laboratory conditions. The males were attracted by 250-420 Hz that corresponds to the main frequencies of flight tones of sympatric females and the range of sensitivity of male Johnston's organs. Behavioural data show that in natural environment swarming males are attracted by flying conspecific and heterospecific females rather than by dead or immobilized females. Contact with heterospecific females always ended in parting of mosquitoes while contact with cospecific females resulted in pairing. Thus, swarming males of Aedes diantaeus localize females by their flight tones but recognize the females of their own species in contact.     

Sexual dimorphism in relation to swarming and pair formation patterns in leptocerid caddisflies (Trichoptera: Leptoceridae)

North European Leptoceridae (Trichoptera) perform three types of swarming flight patterns: (1) swarming males of Athripsodesand Ceracleafly in horizontal zigzag patterns over the water surface, (2) the Mystacidesspp. perform vertical zigzag movements, and (3) the flight of males of Triaenodes unanimisMcLach. is a mixture of the horizontal and vertical zigzagging. Also three groups of pair formation behavior can be distinguished. In the first group, of Athripsodesand Ceraclea,the females fly into the male swarms, where they are grasped and carried to the riparian vegetation by the flying males with the females hanging upside-down in genitalia coupling. In the second group, a Mystacidesfemale is caught by a male, when approaching a swarm and both use their wings to fly in tandem to the shore where they copulate. In the third group, of Triaenodes bicolor(Curt.) and Oecetis lacustris(Curt.), the males fly searching for females sitting on aquatic plants and when a female is found the male lands and they copulate immediately while clinging to the plant. The different swarming and mating behaviors might have favored selection for three types of sexual dimorphism: (1) longer forewings in males than females in species which fly in copula, (2) larger eyes in males of the vertically zigzagging species, and (3) much smaller males in the group where males search for females sitting on aquatic plants. In the second group approaching females are detected by males before reaching the swarm and in the third group the female almost always mates with the male which is the first to find her. In conclusion, we suggest that females of Athripsodesand Ceracleahave a greater choice among swarming males than do females of Mystacides, T. bicolor,and O. Lacustris.     

Pre-copulatory isolation in sympatric Melolontha species (Coleoptera: Scarabaeidae)

1 The two most abundant cockchafer species in Europe, the forest cockchafer Melolontha hippocastani Fabr. and the European cockchafer Melolontha melolontha L., tend to form calamitous mass breedings with casual reports on sympatric and simultaneous occurrence. 2 Both species are known to use feeding‐induced green leaf volatiles (GLV) as primary attractants (sexual kairomones) for mate finding. The attractiveness of GLV is enhanced by the sex pheromones 1,4‐benzoquinone in M. hippocastani and toluquinone in M. melolontha. Phenol attracts males from both species. All three compounds are present in females of both species. 3 In the present study, it is confirmed that only male M. melolontha perform the typical swarming flight at dusk, as has already been shown for M. hippocastani. Furthermore, whether swarming Melolontha males were cross‐attracted to heterospecific females, and whether males could discriminate olfactorily between conspecific and heterospecific females, was tested in the field. 4 Males of both species preferred females when given the choice between females and males of the other species. However, they preferred conspecific females when females from both species were offered simultaneously. 5 The results suggest that species‐specific pheromone blends contribute to precopulatory reproductive isolation in sympatric populations of M. melolontha and M. hippocastani, but are not mutually exclusive or indispensable prerequisites for mate finding as in other insects.     

Behavioural responses to ectoparasites: time-budget adjustments and what matters to Blue Tits Parus caeruleus infested by fleas

Blue Tit nests are often heavily infested by fleas, which feed on the incubating female and the nestlings. Depending on habitat quality, the drawing of blood by fleas reduces offspring quality, or it is compensated by an increase in food provisioning by the adults and may reduce their future reproduction. Given these fitness costs, tits are expected to have evolved behavioural responses enabling them to remove, destroy or minimize the contact with fleas. To identify these traits, we video-recorded the changes in frequency and duration of the hosts' potential anti-flea behavioural defences in nests experimentally infested with low and high flea densities. We also investigated whether flea load affected the number of male feeds delivered to incubating females, and whether the parents increased their rate of food provisioning to the nestlings equally at high flea density. Flea density significantly affected the nest sanitation and sleeping behaviour of Blue Tit females but had no significant effect on grooming. Female Blue Tits increased the frequency but decreased the duration of bouts of these behavioural traits, and hence their time-budgets, based on per hour duration of behaviour, were not significantly affected by flea density. High flea density reduced nestling weight at the early nestling stage but these costs were fully compensated by an increase in female feeding effort. Males did not increase their frequency of food provisioning to incubating females nor to nestlings in heavily infested nests. The results are discussed in the light of parasite-mediated selection on host behaviour and the reciprocal host selection on flea life-history and behavioural traits.     

Female feeding priority in bonobos, Pan paniscus, and the question of female dominance

The question of whether bonobos show feeding priority and female dominance has been proposed and examined, both in the wild and in captive studies, with differing results. The relationship between female dominance and female feeding priority has been best studied in prosimian primates. These studies use established criteria of females consistently evoking submissive behavior from males in dyadic encounters for determining female dominance. Although the relationship is complex, female dominance in prosimians is associated with preferential access to food. Data from studies of wild habituated bonobos in the Lomako Forest, Democratic Republic of the Congo, are examined for evidence of both female feeding priority and female social dominance using similar criteria as used for prosimians. Bonobos showed evidence of female feeding priority in small, but not in large, food patches. Male-male competition for mating opportunities at the start of the food bout was related to some, but not all, differences in time spent feeding between the sexes. Female dominance similar to that seen in prosimians was not observed in these bonobos. Males were consistently dominant in dyadic interactions. Female feeding priority with male dyadic social dominance implies that male deference during feeding cannot be excluded as one explanation of interpretations of female dominance in bonobos. Additionally, dominance of male bonobos by females appears to require the presence of female coalition partners. As in other primates with female feeding priority, bonobo females express this trait where food is economically defendable. Unlike prosimians, however, bonobo female feeding priority may result from male deference and the importance of female coalitions in nondyadic interactions.     

The Function of Male Dominance in the Eusocial Wasp, Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Males of a Neotropical eusocial wasp, Mischocyttarus mastigophorus , are dominant over their female nest mates. Mischocyttarus mastigophorus males behave aggressively toward females, while females rarely bite or chase males. Aggressive interactions between the sexes are behaviorally indistinguishable from dominance interactions among females. Males are long-lived as adults, and can reside on nests for periods of at least one month. Furthermore, males comprise a large proportion of post-emergence colony populations throughout much of the colony cycle. Males on the nest perform maintenance tasks at low rates, but contribute little other labor to their colonies. Males do not forage, but consume a disproportionate amount of the food (nectar and insect prey) collected by workers. Males in some colonies direct disproportionate amounts of aggression toward their queens, which may further contribute to males' food procurement. These data suggest that adult males represent a considerable energetic and labor cost to their colonies. I hypothesize that the dominance structure of M. mastigophorus directs food resources to adult males, with the function of increasing their longevity. Increased male longevity may be selectively advantageous in tropical species such as M. mastigophorus that found new colonies throughout much or all of the year. When females initiate new nests over much of the year, individual males' mating opportunities may be temporally distributed, favoring longer adult lifespans. Male dominance is predicted to occur in other populations of independent-founding Neotropical Polistinae with asynchronous colony foundation.     

Swarming patterns of light trapped individuals of caddisfly species (Trichoptera) in Central Europe

This study is on the light trapping of caddisfly species (Insecta: Trichoptera) related to the proportion of males and females and the moon phases. The data collected includes 24 species in 9 light-trap stations, for 49 swarming events between the years 1980 and 2000. We found the massive emergence of adults happens fractionally in swarming intervals. This is connected with the phenology and life cycle of each species. The percentage of males and females of the same species during different swarming events cannot be considered equal. The proportion of males and females are different in the swarming of different species. We found that the number of male and female individuals is substantially synchronized with each other within the swarming, but it can be different in the case of each species. The duration of the swarming, even in the same species, are not always uniform. The effect of the Moon cannot be clearly identified in any species, even if data from several swarmings are available. The swarming peaks appear near different Moon quarters.     

褐带蚜小蜂成虫生物学特性

[目的]褐带蚜小蜂Aphelinus maculatus是近几年发现的棉蚜Aphis gossypii重要寄生蜂之一.目前关于该寄生蜂生物学特性包括行为习性方面的研究非常少.为有效挖掘利用褐带蚜小蜂,对其成虫生物学特性进行了研究.[方法]在室内采取体视显微镜下观察和记录褐带蚜小蜂的羽化节律、交配行为、交配各阶段持续时间...     

Flight of the Chinese white pine beetle (Coleoptera: Scolytidae) in relation to sex, body weight and energy reserve

The flight distance, flight time and individual flight activities of males and females of Dendroctonus armandi were recorded during 96-h flight trials using a flight mill system. The body weight, glucose, glycogen and lipid content of four treatments (naturally emerged, starved, phloem-fed and water-fed) were compared among pre-flight, post-flight and unflown controls. There was no significant difference between males and females in total flight distance and flight time in a given 24-h period. The flight distance and flight time of females showed a significant linear decline as the tethered flying continued, but the sustained flight ability of females was better than that of males. The females had higher glycogen and lipid content than the males; however, there was no significant difference between both sexes in glucose content. Water-feeding and phloem-feeding had significant effects on longevity, survival days and flight potential of D. armandi, which resulted in longer feeding days, poorer flight potential and lower energy substrate content. Our results demonstrate that flight distances in general do not differ between water-fed and starved individuals, whereas phloem-fed females and males fly better than water-fed and starved individuals.     

The nutritional value of aphid honeydew for non-aphid parasitoids

Intake of sugar-rich foods by adult parasitoids is crucial for their reproductive success. Hence, the availability of suitable foods should enhance the efficacy of parasitoids as biological control agents. In situations where nectar is not readily available, homopteran honeydew can be a key alternative food source. We studied the impact of honeydew feeding on the longevity of the larval endoparasitoids Cotesia marginiventris, Campoletis sonorensis and Microplitis rufiventris, all natural enemies of important lepidopteran pests. Females of these wasps lived longer when feeding on honeydew produced by the aphid Rhopalosiphum maidis on barley compared to control females provided with water only. However, they lived shorter than females fed with a sucrose solution. Further investigations with C. marginiventris showed that access to honeydew also increases the number of offspring produced, but less so than access to a sucrose solution. Moreover, it was found that females of this species need to feed several times throughout their life in order to reach optimal longevity and reproductive output. Analyses of the sugars in the honeydew produced by R. maidis on barley revealed that it contains mainly plant-derived sugars, but also several aphid-synthesized sugars. The sugar composition of the honeydew changed as a function of aphid colony size and time a colony had been feeding on a plant. In general, the higher the aphid infestation, the smaller the percentage of aphid-synthesized sugars in the honeydew. Experiments with honeydew sugar mimics allowed us to reject the hypothesis that the relatively poor performance of the parasitoid on a honeydew diet was due to the sugar composition. Instead, the results from additional feeding experiments with diluted honeydew showed that the nutritional value of pure honeydew is primarily restricted by its high viscosity. The possible consequences of these findings for biological pest control are discussed.     

Age-associated male mating success in three swarming caddis fly species (Trichoptera: Leptoceridae)

Abstract. 1. In the three caddis fly species, Athripsodes albifrons (L.), A. cinereus (Curtis) and Mystacides azurea (L.) (Leptoceridae; Trichoptera), males swarm above the water surface of lakes or rivers. Receptive females fly to swarms and are chased and/or courted by males. After one of the swarming males has grasped an approaching female, the pair flies in tandem to the shore where they copulate.
2. In males, wing wear indices were negatively correlated with the ratio of fat/dry weight. In the only species in which comparisons were possible between newly emerged and swarming males (M. azurea), the former had significantly lower indices. Unmated females on average had lower wing wear indices than spent females. These facts suggest that wing wear reflects relative age.
3. The tandem males had significantly less wing wear than those in swarms, and are probably therefore younger. Age is therefore likely to be significant in relation to mating success.
4. Among males of the same relative age, tandem males had higher fat ratio than swarming ones, indicating that male mating success was also influenced by traits other than age. It is suggested that the shortest possible duration of the period of adult prematurity is adaptive, especially in insects with marginal adult food intake.     

Searching behaviour and mate recognition by males of the two-spot ladybird beetle, Adalia bipunctata

Abstract. 1. Adult males of the two-spot ladybird beetle, Adalia bipunctata , did not show a functional response to increase in aphid abundance and consumed markedly fewer aphids than do the females.
2. At high densities of prey, females spent more time in area-restricted search than when prey was scarce. Males were always less active than females and they did not respond to an increase in prey abundance by a change in searching behaviour.
3. After a brief encounter with a female, a male showed area-restricted searching behaviour. This behaviour occurred in response to encountering a female's elytra and in particular to a chloroform-soluble component (sex pheromone) present on or in the elytra.
4. Males needed to encounter a female in order to respond to her presence, which indicated the pheromone is a contact pheromone.
5. The searching behaviour of males appeared to be mainly directed towards locating females; that of females towards locating aphids. This difference between the sexes should be taken into account when quantifying the predatory response of ladybirds to aphid abundance in the field.     

Feeding-site utilization in three sympatric species of Petrotilapia (Pisces, Cichlidae) from Lake Malawi

Three sympatric sibling species of Petrotilapia , with similar dietary and macrohabitat requirements, occur near Monkey Bay, Lake Malawi. Males and females of all three species fed selectively on the rocky shores. Males utilized less than 6% of the space they defended as a breeding territory for feeding and although interspecific overlap of territories was considerable, only 2.5% of the feeding space used by territorial males was shared. Males primarily utilized rich food patches which occurred within the territories of highly aggressive Pseudotropheus species, whereas females tended to feed in undefended areas. Although Petrotilapia species are not highly aggressive fishes, there was a well-defined social dominance hierarchy amongst them and it is suggested that feeding-site utilization is dependent on social rank. Inter-and intraspecific differences in feeding-site utilization probably facilitate species coexistence and thus contribute to the maintenance of the high species diversity which is a characteristic of the ichthyofauna of Lake Malawi.