Multiple mating by both sexes in an invasive insect species,Aethina tumida (Coleoptera: Nitidulidae)

Multiple mating by both sexes is common among sexually reproducing animals. Small hive beetles (SHB), Aethina tumida, are parasites of bee nests endemic to sub-Saharan Africa and have become a widespread invasive species. Despite the considerable economic damages they can cause, their basic biology remains poorly understood. Here we show that male and female small hive beetles can mate multiple times, suggesting that costs for mating are low in this species. In an invasive A. tumida population in the United States, a combination of laboratory experiments for males and paternity analysis with eight polymorphic DNA microsatellite markers for field-caught females were used to estimate the number of mating by both sexes. The data show that females and males can mate multiple times—females mated with up to eight males, whereas males mated with at least seven females. The results also showed that A. tumida displayed a skewed paternity, although this was not consistent among the tested females. Thus, first or last male advantage seem to be unlikely in A. tumida. Our observations that individuals of both sexes of A. tumida can mate multiple times opens new research avenues for examining drivers of multiple mating and determining the role it may play in promoting biological invasions.     

Male mating costs in a polygynous mosquito with ornaments expressed in both sexes

Male mate choice in species with conventional sex roles is difficult to explain and has, therefore, been the focus of many recent theoretical models. These models have focused on variance in female quality and, to a lesser extent, male investments/costs associated with mating. In this study, we investigate the costs of courtship and copulation in the polygynous mosquito Sabethes cyaneus. In this species, both males and females possess elaborate ornaments. Previous studies suggest that the most likely explanation for the presence of these ornaments is mutual mate choice. Thus, this system provides an excellent model for exploring the evolution of mutual mate choice in polygynous species. We disentangle the costs of courtship and copulation by monitoring male survival in three groups of males: housed alone (group 1); able only to court females (group 2); or able to court and copulate with females (group 3). We show that males incur a cost of courtship and copulation and that courtship intensity is negatively related to male longevity. Our results suggest that courtship and copulation carry additive costs to males. We discuss the implications of these results in the context of current mutual mate choice theory and suggest that courtship costs may be an unappreciated key factor in the evolution of male mate choice.     

Diapause affects cuticular hydrocarbon composition and mating behavior of both sexes in Drosophila montana

Envipnmental cues,mainly photoperiod and temperature,are known to control female adult reproductive diapause in several insect species.Diapause enhances female survival during adverse conditions and postpones progeny production to the favorable season.Male diapause(a reversible inability to inseminate receptive females)has been studied much less than female diapause.However,if the males maximized their chances to fertilize females while minimizing their energy expenditure,they would be expected to be in diapause at the same time as females.We investigated Drosophila montana male mating bchavior under short-day conditions that induce diapause in females and found the males to be reproductively inactive.We also found that males reared under long-day conditions(reproducing individuals)court reproducing postdiapause fermales,but not diapausing ones.The diapausing fies of both sexes had more long-chain and less short-chain hydrocarbons on their cuticle than the reproducing ones,which presumably increase their survival under stressful conditions,but at the same time decrease their attractiveness.Our study shows that the mating behavior of females and males is well coordinated during and afier overwintering and it also gives support to the dual role of insect cuticular hydrocarbons in adaptation and mate choice.     

Delayed mating in tortricid leafroller species: simultaneously aging both sexes prior to mating is more detrimental to female reproductive potential than aging either sex alone

The effect of delayed mating on reproductive potential, longevity and oviposition period of female redbanded leafroller, Argyrotaenia velutinana (Walker) and Pandemis leafroller, Pandemis pyrusana Kearfott, was investigated in the laboratory. Virgin female or male moths of each species were held for 1, 2, 4, 6 or 10 days prior to pairing with one-day-old virgin conspecifics of the opposite sex. In addition, reproductive potential was assessed when both sexes of each species were aged for those periods prior to pairing. The expected reproduction of female A. velutinana was reduced by 34, 53, 71 and 81% for 2, 4, 6 and 10-day delays in female mating, respectively. For P. pyrusana, expected reproduction was reduced by 47, 74, 85 and 93% for 2, 4, 6 and 10-day delays in female mating, respectively. Increasing male age at mating in both species had a lesser effect on female reproductive output compared with increasing female age at mating. As male A. velutinana age at mating increased, the expected reproduction of female A. velutinana was reduced by 15, 45, 54 and 70% for 2, 4, 6 and 10-day delays, respectively. Comparing male P. pyrusana of various ages at mating, expected reproduction was reduced by 14, 42, 64 and 79% for 2, 4, 6 and 10-day delays in mating, respectively. The decrease in female reproduction when both sexes were aged prior to mating was higher than when either sex alone was aged prior to pairing with a one-day-old virgin of the opposite sex. The expected reproduction of female A. velutinana was reduced by 60, 83, 96 and 98% for 2, 4, 6 and 10-day delays in mating of both sexes, respectively. Only 7.5% of female eggs hatched when both sexes of A. velutinana were aged ten days prior to mating. When simultaneously aging both sexes of P. pyrusana prior to mating, expected reproduction was reduced by 71, 93, 96 and 99% for 2, 4, 6 and 10-day delays in mating, respectively. No P. pyrusana eggs hatched after a ten-day delay of mating for both sexes. For both species, female longevity increased and duration of oviposition period decreased with increasing female age at mating. Our results demonstrate that delayed mating in both females and males negatively affects female reproductive output in both species and that simultaneous aging of both sexes prior to mating has a greater effect than aging either sex alone. Our results suggest that laboratory studies that have paired aged females or aged males with conspecifics of optimal reproductive maturity have likely underestimated the effects of delayed mating on reproductive output.     

A bioassay of the pheromone of Trogoderma granarium males as an attractant for both sexes of the species

The pheromone of Trogoderma granarium males was subjected to two bioassays intended to examine its effect on the two sexes of the species. The tests were made with concentrations ranging from 0.1 to 200 males per ml of solvent. The weakest reaction of males and females was observed in concentrations of 0.1 and 0.4/cc respectively. The peak reaction of the males was attained in concentrations of 12.5/cc and of the females in 50/cc. Both the males and the females show a marked reduction of response at higher concentrations. A significant difference in response was observed between treated and control specimens (p < 0.001) at concentrations corresponding to peak response; the difference was nonsignificant near the lowest concentrations of the two sexes used.The initial response was seen to rise steeply during the first 15 min.; this primary rise was followed by a more gradual secondary rise later on.The results of our study seem to suggest that the volatile substance secreted by T. granarium is more in the nature of an aggregation agent, rather than being a sex attractant.

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Zusammenfassung Das Pheromon von Trogoderma granarium-Männchen wurde zwei Biotesten unterworfen, um seine Wirkung auf die beiden Geschlechter dieser Art zu prüfen. Die Versuche wurden mit Konzentrationen von 0,1 bis 200 Männchen pro Milliliter Lösungsmittel vorgenommen. Die schwächste Reaktion der Männchen und Weibchen wurde bei Konzentrationen von 0,1 bzw. 0,4/cc beobachtet. Die Spitzenreaktion der wurde bei Konzentrationen von 12,5/cc, die der bei 50/cc erreicht. Sowohl wie zeigen einen merklichen Rückgang der Reaktion bei höheren Konzentrationen. Ein signifikanter Reaktionsunterschied (p < 0,001) zwischen behandelten und unbehandelten Individuen wurde bei Konzentrationen beobachtet, die der Gipfelreaktion entsprachen; der Unterschied war nicht signifikant in der Nähe der schwächsten Konzentration, die für die beiden Geschlechter benutzt wurden.Es wurde beobachtet, daß die Anfangsreaktion während der ersten 15 Minuten steil anstieg; auf diesen primären Anstieg folgte später eine allmähliche sekundäre Zunahme.Diese Ergebnisse der Untersuchung scheinen darauf hinzudeuten, daß die von T. granarium ausgeschiedene flüchtige Substanz ihrer Natur nach eher ein Aggregationsmittel als einen Geschlechtslockstoff darstellt.

     

Territorial behavior of both sexes in the water striderMetrocoris histrio (Hemiptera: Gerridae) during the mating season

Territorial behavior of overwintered individuals of Metrocoris histriowas observed in an upstream area. Adults of both sexes held territories, but male territories were larger than those of females. Severe competition occurred among males for territories which give them access to receptive females. The effects of male body length and midleg length on establishment of territories were not significant. The effect of female midleg length on activity of females entering preferred foraging sites was equally not significant. Instead, territorial behavior increased with male age and males stayed longer at prime sites. Females of intermediate age were likely to occupy prime sites. Females had longer territory residence time than males. The sexes were dimorphic with respect to midleg length, and dimorphism in M. histriomay be related to a difference in life history, in that sexual selection may be relaxed due to asynchronous adult emergence patterns.     

Effect of host species on the distribution of mutational fitness effects for an RNA virus

Knowledge about the distribution of mutational fitness effects (DMFE) is essential for many evolutionary models. In recent years, the properties of the DMFE have been carefully described for some microorganisms. In most cases, however, this information has been obtained only for a single environment, and very few studies have explored the effect that environmental variation may have on the DMFE. Environmental effects are particularly relevant for the evolution of multi-host parasites and thus for the emergence of new pathogens. Here we characterize the DMFE for a collection of twenty single-nucleotide substitution mutants of Tobacco etch potyvirus (TEV) across a set of eight host environments. Five of these host species were naturally infected by TEV, all belonging to family Solanaceae, whereas the other three were partially susceptible hosts belonging to three other plant families. First, we found a significant virus genotype-by-host species interaction, which was sustained by differences in genetic variance for fitness and the pleiotropic effect of mutations among hosts. Second, we found that the DMFEs were markedly different between Solanaceae and non-Solanaceae hosts. Exposure of TEV genotypes to non-Solanaceae hosts led to a large reduction of mean viral fitness, while the variance remained constant and skewness increased towards the right tail. Within the Solanaceae hosts, the distribution contained an excess of deleterious mutations, whereas for the non-Solanaceae the fraction of beneficial mutations was significantly larger. All together, this result suggests that TEV may easily broaden its host range and improve fitness in new hosts, and that knowledge about the DMFE in the natural host does not allow for making predictions about its properties in an alternative host.     

Differential host relations of the sexes in parasitic Hymenoptera

The divergent ontogeny of the male, a unique phenomenon characterizing certain genera of the Aphelinidae, has its physiological bases in the environmental control of sex permitted by the haploid origin of the male and the diploid origin of the female and in the efficiency of the chalcidoid type of spermatheca which, during the process of oviposition, brings the environmentally activated sperm into contact with the egg. This divergence is correlated with the sexual dimorphism and the sexual ditrophism of the early developmental stages. It is psychologically manifested in a number of species by a marked difference in the host relations of the aphelinid female before and after mating. The climax of differential host relations within a species is the production of the female in a coccid host and the production of the male in a lepidopterous host. The aberrant development of the male tends to increase the capacity of the species to survive at low host densities.

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Zusammenfassung Nachdem 1935 die obligatorisch wirtsverschiedene Geschlechterentwicklung bei den parasitischen Hymenopteren entdeckt wurde, ist dieses Phänomen bei ca 80% der bekannten Apheliniden, die in Schildläusen und Mottenschildläusen schmarotzen, gefunden worden. Die Kenntnis dieses Phänomens ermöglichte die Züchtung und Ansiedlung von 2 Arten solcher Parasiten in Californien. Die wirtsverschiedene Geschlechterentwicklung besteht in der aberranten Entwicklungsgeschichte des Männchens sowie der geschlechtsgebundenen Ditrophie mit Dimorphismus der Larvenstadien. Bei einigen Arten ist sie gekoppelt mit psychologischen Unterschieden zwischen begatteten und unbegatteten Weibchen, die dann verschiedene Wirtspräferenzen zeigen. Diese bemerkenswerte Spaltung der Wirtspräferenzen steht wohl teilweise in Zusammenhang mit der Funktion der Spermatheca des Chalcidoiden-Typs, die äußerst empfindlich auf wirtseigene Stimuli anzusprechen pflegt. Es handelt sich um ein Wirts-Beziehungs-Phänomen, welches die Wirtsfindungsrate der Art zu erhöhen pflegt. Die ontogenetischen Abweichungen in der Entwicklung der Männchen dürften zur Phylogenie der Apheliniden in keiner Beziehung stehen.

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Paper No. 1117, University of California Citrus Experiment Station, Riverside, California.     

Effects of male age on the mating behavior of both sexes in the sorghum plant bug, Stenotus rubrovittatus (Hemiptera: Miridae)

In the sorghum plant bug Stenotus rubrovittatus (Matsumura) (Hemiptera: Miridae), mating behavior consists of male courtship and female mating receptivity. Previous studies have found that female age is correlated with ovary development and that mature females with developed ovaries are more receptive to male courtship. Thus, we examined whether male age affects the mating behavior of both sexes and male accessory gland development. Unmated males 0–9 days after emergence and 3-day-old virgin females receptive to male courtship were studied. Immediately after emergence, only 20 % of the males courted females (n = 25). At 3 days old, 68 % of the males courted females (n = 25), the most active age. In contrast, more than 75 % of the courted females were receptive to the male courtship regardless of male age. These results indicate that only male courtship behavior is affected by male age, although it is not enhanced in proportion to male age. Male accessory glands developed with male age. However, no clear relationship was detected between male courtship behavior and accessory gland development. In S. rubrovittatus, it is difficult to explain male courtship behavior solely from male age and accessory gland development.     

Hyperparasitism behaviour of the autoparasitoid <Emphasis Type="Italic">Encarsia tricolor</Emphasis> on two secondary host species

Hyperparasitism by virgin female Encarsia tricolor was studied by direct observation of its behaviour when contacting two secondary host species (Encarsia formosa and E. tricolor) at different host stages (first and second larval stage, third larval stage, and pupal stage). The searching and hyperparasitism behavioural sequence of E. tricolor was independent of the host stage of the whitefly (Aleyrodes proletella), and was similar to several related primary parasitoid species. In experiments with equal numbers of secondary hosts, encounter frequencies were equal for both secondary host species in all developmental stages observed. However, rates of hyperparastism were different according to host stage and host species. Hosts in the late larval stages were most preferred for hyperparasitization and the heterospecific E. formosa was more preferred as a secondary host than the conspecific, E. tricolor, in particular from the prepupal stage onwards. The window of vulnerability, i.e., the duration of the period in which a secondary host is susceptible to hyperparasitism, was largely determined by the occurrence and rate of melanization after the onset of pupation. The duration of a successful hyperparasitization event was longer than one that failed. Superparasitism occurred only once in all cases. The potential effect of autoparasitoids on biological control programs and the consequences for selection and release of an effective, yet ecologically safe agent are discussed. Handling editor: Torsten Meiners.     

Effects of age at mating of both sexes on female longevity and fecundity performance in Ophraella communa LeSage (Coleoptera: Chrysomelidae)

Life-history strategies of animals are governed by fitness-related trait trade-offs. In particular, the age at which either one of the two sexes copulate has been shown to affect insect reproductive output and longevity. This suggests that trade-offs between longevity and reproduction might drive the choice of a mating partner based on their age and ultimately such choices might select for different life-histories. Although several studies indicate that female age at mating dictates subsequent insect longevity, fecundity, and egg hatch rate, it is unclear how male age at copulation affects these life-history traits. In this study, we simultaneously investigated the effects of female and male age on female fecundity, eggs hatch rate, and adult longevity in Ophraella communa LeSage (Coleoptera: Chrysomellidae), the primary biological control agent of the invasive common ragweed, Ambrosia artemisiifolia L. We found that young mature females lived significantly longer when they were mated with older males. Maximum female fecundity and subsequent egg hatch rate occurred when young females mated with 3-day-old males. On the other hand, females lived longer when mating with an older male. These findings are in accordance with the cost-of-reproduction concept.     

Sex allocation in a field population of an autoparasitoid

Autoparasitoid wasps lay fertilized eggs in homopteran nymphs, and these eggs develop into female primary parasitoids. Unfertilized, male-producing eggs are laid in immatures of the wasps' own or another primary parasitoid species; males then develop as secondary or hyperparasitoids. In the population of Encarsia pergandiella studied in Ithaca, NY, fertilized eggs were laid in the nymphs of the whitefly Trialeurodes packardi (primary hosts) and unfertilized eggs were laid almost exclusively in pupal females of their own species (secondary hosts). In the two years the population was studied, secondary hosts were always much less abundant than primary hosts at both sites. However, secondary hosts were parasitized at a significantly greater rate than primary hosts. In a laboratory experiment, the encounter rate of females with primary and secondary hosts was not significantly different. Moreover, there was no evidence from the field that wasps found leaves bearing secondary hosts more frequently than leaves without secondary hosts. Dissections of field-collected females showed them to be mated, and thus capable of laying both unfertilized and fertilized eggs. These results suggest that wasps did not encounter secondary hosts at a greater rate, nor were they constrained to lay unfertilized eggs, but rather secondary hosts were preferred. The oviposition sex ratios were influenced by the proportion of secondary hosts, but were less female-biased than would be predicted from the proportion of secondary hosts alone. The results do not support the predictions of Godray and Waage (1990) for either strictly host-limited autoparasitoids (sex ratio should reflect the proportion of secondary hosts) or for egg-limited autoparasitoids (sex ratio should be equal, and independent of the proportion of secondary hosts).     

Hyperparasitism by an exotic autoparasitoid: secondary host selection and the window of vulnerability of conspecific and native heterospecific hosts

Encarsia transvena is an 'autoparasitoid' in the hymenopteran family Aphelinidae. In this species, female eggs are laid in whitefly nymphs. Male eggs are laid externally on immature parasitoids enclosed within the whitefly integument, either their own species, or other primary parasitoids. We explored parasitism by E. transvena of conspecific female immatures and those of a native primary parasitoid, Eretmocerus eremicus, in laboratory experiments. In the first experiment, female E. transvena were offered different combinations of two stages of E. transvena (late larvae – prepupae (ET2), and early pupae (ET3)), and one stage of E. eremicus (prepupae – early pupae (EE2)) in paired choice tests. The results indicated very little parasitism of ET3 relative to the host it was paired with, either EE2 or ET2. However, when EE2 was offered with ET2, there was no statistically significant difference in parasitism. In a no-choice experiment in which oviposition patterns and male progeny development were examined in four stages of both species of wasp, clear differences were observed between the host species. Only one stage of E. transvena (ET2) was parasitized and supported development of male E. transvena to any significant degree. In contrast, in E. eremicus, EE2, EE3 (red-eyed pupae), and EE4 (late pupae) were all parasitized, and male E. transvena emerged from all three stages, although fewer males emerged from EE4. In both species, wasp larvae that were still enclosed within the wet whitefly remains (ET1 and EE1) were parasitized at a very low rate. Lastly, an experiment that determined the length of the later developmental stages of E. transvena and E. eremicus suggested that the duration of the period in which E. transvena is susceptible to parasitism by conspecific females is less than half the period of susceptibility of E. eremicus. These results taken together suggest the potential for interference of E. eremicus by E. transvena, but other factors not examined here may also influence the outcome of interactions in the field.     

Effects of different sex pheromone compositions and host plants on the mating behavior of two Grapholita species

The two congener species Grapholita molesta and Grapholita dimorpha share two major sex pheromone components: cis-8-dodecenyl acetate (Z8-12Ac) and trans-8-dodecenyl acetate (E8-12Ac). In fact, commercial sex pheromone lures composed of only these two major components attract the males of both species. In this study, we aimed to determine the reproductive isolation components of these two species by analyzing the effects of the minor sex pheromone components and host plants. First, different ratios of the two major sex pheromone components were greatly favored by either male species. Sex pheromone gland extracts of G. dimorpha contained a lesser proportion of Z8-12Ac than that of G. molesta. In the three (apple, pear, and peach) orchards investigated in this study, a larger number of G. molesta males were attracted to the 95:5 pheromone mixture (Z8-12Ac and E8-12Ac, respectively), while a larger number of G. dimorpha males were attracted to the 85:15 mixture. Second, there was a significant variation in male attractions in different host plants. G. molesta males were more attracted to the sex pheromone lure in the apple orchards than that in the pear and peach orchards. In contrast, G. dimorpha males were more attracted to the lures in the pear and peach orchards than that in the apple orchard. Third, the minor sex pheromone components were important for reproductive isolation. Among the four minor components tested, addition of (Z)-8-dodecenol (Z8-12OH) to the major sex pheromone components significantly suppressed male attraction in G. dimorpha and slightly elevated male attraction in G. molesta. The discriminating effect of Z8-12OH was further validated using male electroantennogram analysis. These results suggest that reproductive isolation between two congeners can be achieved by variations in the minor sex pheromone components and in the host plants, as well as by changes in the ratio of the two major components.     

Localization of mating behavior of releasedBactrocera dorsalis flies on host fruit in an orchard

Oriental fruit flies,Bactrocera dorsalis, cultured in the laboratory for six generations, were released when 12–14 days old in an orchard of nonfruiting host trees that were furnished with either food and water, nonpunctured host fruit, punctured host fruit, or no resources. Nearly all flies of both sexes, all sexual behavior of males, and all mating pairs were observed on trees with fruit. Moreover, on trees with fruit, nearly all flies of both sexes, nearly all sexual behavior of males, and nearly all mating pairs occurred on the fruit itself rather than on the foliage or branches. In a subsequent test, both sexes were found to be strongly attracted to the odor of host fruit. For a polyphagous species of a tropical tephritid, these findings are the first to show a high level of male aggregation and a high amount of male sexual behavior on the fruit of host trees. Findings are discussed in relation to current knowledge of tephritid mating behavior. An additional quality-control test for laboratory cultured males used in the sterile insect technique of fly management is recommended.