Isolation and characterization of 14 microsatellite markers in <Emphasis Type="Italic">Macrodasyceras</Emphasis><Emphasis Type="Italic">hirsutum</Emphasis> (Hymenoptera: Torymidae)

Macrodasyceras hirsutum Kamijo is the seed parasitoid wasp of the bird-dispersed, dioecious tree, Ilex integra Thunb. The wasp reduces the level of dispersal mutualism between the Ilex tree and its frugivorous birds by manipulating the color of mature berries. The female trees do not blossom every year and sometimes change sex. Thus, the reproduction biology of I. integra affects the population size and structure of M. hirsutum in a forest and consequently influences the seed dispersal mutualism between the tree and birds, because of limited ability of adult locomotion. To investigate the wasp population structure with reference to the dispersal mutualism between trees and birds, we isolated 14 microsatellite loci of M. hirsutum wasps. Every locus was polymorphic among 20 females, with 3–13 alleles per locus, without linkage disequilibrium. The observed and expected heterozygosities ranged from 0.100 to 0.900 and 0.099 to 0.818, respectively, indicating their utility in molecular analyses of the wasp population.     

Fruit Colour Preferences of Redwings (Turdus iliacus): Experiments with Hand-Raised Juveniles and Wild-Caught Adults

Certain fruit colours and their contrast with the background coloration are suggested to attract frugivorous birds. To test the attractiveness of different colours, we performed three experiments in laboratory with controlled light conditions. In the first two experiments, we studied the fruit colour preferences of naive juvenile redwings. In the third experiment, we continued to investigate whether the contrast of the fruit colour with the background coloration affects the preference of both naive juveniles and experienced adult redwings. In the first experiment, juvenile birds preferred black, UV‐blue and red berries, to white ones. In pairwise trials, a new set of juveniles still preferred red berries to white ones. When testing the effect of contrasts on their choice, juveniles preferred UV‐blue berries to red ones on a UV‐blue background. However, no preference was found, when the background was either red or green. Adult redwings preferred UV‐blue berries to red ones on all backgrounds. According to these results, juveniles seem to have an innate avoidance of white berries. Furthermore, the foraging decisions of fruit‐eating birds are affected more by fruit colour than its contrast with background coloration, at least when contrasting displays are encountered from relatively short distances. Differences in preferences of adult and juvenile birds also indicate that learning seems to play a role in fruit choices.     

Influence of cabbage resistance and colour upon the diamondback moth and its parasitoid Oomyzus sokolowskii

Host plant resistance and biological control are vital integrated pest management tools against the diamondback moth (DBM), Plutella xylostella (L.) (Lepidoptera: Plutellidae), but to date no study has investigated this system including the DBM parasitoid Oomyzus sokolowskii (Kurdjumov) (Hymenoptera: Eulophidae). We examined oviposition and development of P. xylostella exposed to two commercial cabbage cultivars (green ‘Chato de quintal’ and red ‘Roxo’) and possible effects upon O. sokolowskii. Under free‐choice tests, DBM females laid significantly more eggs on plants of the green cabbage, even though several population growth parameters showed that DBM developed better on the red cabbage. Furthermore, a laboratory free‐choice test with artificially green‐ and red‐painted kale leaf discs demonstrated a similar oviposition preference pattern, with green colour being preferred over red colour. The preference was apparently visually mediated; olfactometer tests showed similar attraction of moths to both green and red cultivars in choice and non‐choice tests. Host plant cultivar had no statistically significant effect on female parasitoid behaviour towards DBM larvae, nor on parasitoid numbers or longevity. Moreover, wasps parasitizing DBM larvae reared on the green cultivar developed more quickly and in larger numbers per parasitized larva. Thus, feeding on green cabbage rather than red does not hinder, and potentially even enhances, control of DBM by O. sokolowskii. On a practical level, these results suggest that intercalating green cabbage cultivars as a trap crop might help protect more profitable red cultivars in growing fields.     

Signals of profitability? Food colour preferences in migrating juvenile blackcaps differ for fruits and insects

Red is a common colour signal in both aposematic warning displays, and in fruit displays. One common feature is that red is conspicuous against the natural background of the prey and fruits. However, there is a potential conflict between fruits and aposematic prey in how a bird predator should react to red colours, where fruits aim to attract birds and aposematic insects aim to ward off, often the same bird individuals. Here we investigate possible differences in red/green colour preferences of frugivorous, wild-caught, young blackcaps (Sylvia atricapilla), when food is either a fruit or an insect. Birds in two groups were presented with a series of pairs of food items that had been artificially painted red and green, in the order of (I) fruits, crickets and maggots, or (II) crickets, fruits, and maggots. Birds first presented with crickets or fruits differed in first attacks directed at the two colours: They showed no colour preference between fruits, but showed a clear preference for green over red crickets. Also, birds in both experimental groups clearly preferred green to red maggots. These results provide evidence that wild, frugivorous birds are able to differentiate between prey types, and show different colour preferences depending on whether food is insect or fruit. We conclude that blackcaps show an attack bias against red insects, and that one important function of the signal in insects, is to inhibit attack after discovery. However, the lack of preference for red fruits suggests other functions to red fruit displays, such as facilitating discovery per se, rather than directly stimulating attack after discovery.     

The parasitoid wasp Eruga unilabiana Pádua & Sobczak,sp. nov. (Hymenoptera: Ichneumonidae) induces behavioral modification in its spider host

Some polysphinctine parasitoid wasps can alter the web building behavior of their host spiders. In this paper, we describe and illustrate a new species Eruga unilabiana sp. nov. and report for the first time, to the best of our knowledge, the interaction between this parasitic wasp and the linyphiid spider Dubiaranea sp. We investigated the wasp's host selection, development, and manipulation of host behavior. We found that most of the parasitized spiders were intermediate‐sized adult females that probably provide sufficient resources for parasitoid larvae and are less vulnerable for parasitoid females than larger host individuals at attack. The cocoon web of Dubiaranea sp. consists of a complex three‐dimensional tangle structure with several non‐stick radial lines that converge at the cocoon. In addition, E. unilabiana individuals construct their cocoons horizontally, which differ from cocoons of the majority of polysphinctine wasps. This study provides important information and discussion to further understand the evolution of parasitoid wasp–spider interactions.     

Frugivorous birds dispersing braconid parasitoids via endozoochory

Adult braconid wasps (Bracon sp.) emerged from the droppings of frugivorous birds (Turdus blackbirds and thrushes) collected in a rural environment in southern Europe. It was thus demonstrated for the first time that an insect parasitoid of a fruit‐infesting insect (lepidopteran tortricid) can survive bird ingestion and gut passage inside a seed (privet Ligustrum vulgare), constituting a case of an evolutionary tetrad.     

The influence of light environment on host colour preference in a parasitoid wasp

1. Visual chromatic cues and contrast effects are widely used by insects in behaviours involving host/prey/mate‐finding and recognition. However, naturally changing light conditions may challenge the visual perception of cues for these organisms. 2. We used the host/parasitoid system Acyrthosiphon pisum/Aphidius ervi to determine if apparent visual preference of the wasp for green over pink host aphids was a visually based choice or a post‐attack mechanism based on host susceptibility depending on anti‐parasitoid symbiotic bacteria. 3. The study tested the ability of the wasp to recognise and attack pea aphid clones expressing variation based on colour and/or symbionts under a broad range of LED‐controlled light environments mimicking natural variations. 4. Results showed that the amount of reflected light of pink morphs was about half that of the green morphs in the cyan‐green components. Both host colours were recognised and attacked under all tested light conditions, even red light (660 nm). The previously reported preference of A. ervi for green pea aphids, was clear only for naive females given a choice between two aphid colours under all light environments, but quickly disappeared. 5. Wasps showed no tendency of avoiding oviposition in clones with defensive symbionts. 6. These findings suggest that variable rates of pea aphid parasitism by A. ervi in fields do not depend on host colour discrimination, but rather on susceptibility variation among aphid clones in allowing larval development after egg‐laying. Further studies should consider deeper investigation of the impact of red lights used in modified light environments in greenhouses and the proportion of host colour morph available.     

Selection on fruit traits is mediated by the interplay between frugivorous birds,fruit flies,parasitoid wasps and seed‐dispersing ants

Every organism on Earth must cope with a multitude of species interactions both directly and indirectly throughout its life cycle. However, how selection from multiple species occupying different trophic levels affects diffuse mutualisms has received little attention. As a result, how a given species amalgamates the combined effects of selection from multiple mutualists and antagonists to enhance its own fitness remains little understood. We investigated how multispecies interactions (frugivorous birds, ants, fruit flies and parasitoid wasps) generate selection on fruit traits in a seed dispersal mutualism. We used structural equation models to assess whether seed dispersers (frugivorous birds and ants) exerted phenotypic selection on fruit and seed traits in the spiny hackberry (Celtis ehrenbergiana), a fleshy‐fruited tree, and how these selection regimes were influenced by fruit fly infestation and wasp parasitoidism levels. Birds exerted negative correlational selection on the combination of fruit crop size and mean seed weight, favouring either large crops with small seeds or small crops with large seeds. Parasitoids selected plants with higher fruit fly infestation levels, and fruit flies exerted positive directional selection on fruit size, which was positively correlated with seed weight. Therefore, higher parasitoidism indirectly correlated with higher plant fitness through increased bird fruit removal. In addition, ants exerted negative directional selection on mean seed weight. Our results show that strong selection on phenotypic traits may still arise in perceived diffuse species interactions. Overall, we emphasize the need to consider diverse direct and indirect partners to achieve a better understanding of the mechanisms driving phenotypic trait evolution in multispecies interactions.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

Starvation and herbivore-induced plant volatiles affect the color preferences of parasitic wasps

Using light-emitting diode spotlights, we examined the responses of Cotesia vestalis, a parasitoid of diamondback moth (DBM), Plutella xylostella larvae, with different hunger level to different chromatic cues. Naïve satiated female wasps showed no significant preference for either green, yellow, orange, or red spotlighted areas over a control area with background fluorescent light. When starved for 2 h, female wasps preferred yellow and green light over the control area, but not orange or red light. We also tested the effects of DBM-larvae-induced cabbage-plant volatiles, which attract female wasps, on wasp responses to green versus yellow light. In control experiments with no plant volatiles, starved wasps showed no color preference. However, when synthetic volatiles were present, the wasps preferred green over yellow light. We concluded that both hunger level and herbivore-induced plant volatiles were important factors affecting the response of parasitic wasps to light of different color.     

Field superparasitism by Phymastichus coffea, a parasitoid of adult coffee berry borer, Hypothenemus hampei

Superparasitism by Phymastichus coffea LaSalle (Hymenoptera: Eulophidae), a parasitoid of adults of the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Curculionidae: Scolytinae), was recorded under field conditions in a coffee plantation in Colombia. Parasitoid adults were released 1, 5, and 9 days after artificial infestations of 90‐, 150‐, and 210‐day‐old coffee berries with H. hampei females. The position of the beetle inside the berry and the number of P. coffea larvae per female host were assessed 10 days after each parasitoid release. Under laboratory conditions, P. coffea usually lays two eggs per host, one female and one male. In our studies, we often recorded more than six P. coffea larvae in an individual host and mean numbers of larvae per host ranged from two to 4.45. Superparasitism by P. coffea under field conditions was influenced by the age of the coffee berries, which is the most important factor determining the speed of penetration by H. hampei, and therefore the time the beetles are exposed to a P. coffea attack. The number of parasitoid larvae in each H. hampei female gradually decreased with the age of the berry, and also linearly decreased with the time of parasitoid release. Age‐dependent effects of coffee berries that alter the ratio of available hosts to searching parasitoids by providing refuges to the herbivore, largely determine the extent of superparasitism of H. hampei by P. coffea under fields conditions in Colombia.     

Weak interactions between avian and insect frugivores: the case of Pistacia terebinthus L. (Anacardiaceae)

Vertebrate frugivores often feed on fruits upon or within which insects also feed, yet little information exists on the potential magnitude of interactions between these consumers. The Mediterranean shrub Pistacia terebinthus, the birds that consume its fruits, and the wasps that feed upon its seeds are examined in this study. P. terebinthus produces a highly variable fraction of final-sized red fruits that never become mature (green-colored). Red fruits can be immature, parthenocarpic, aborted, or attacked by wasps, and their pulp is much less nutritious than that of mature fruits. A total of 20 bird species consumed the fruits in the study area. Legitimate dispersers accounted for 39% of the total fruit removal, while pulp eaters and seed predators accounted for the remainder. Birds strongly preferred the mature fruits (only 4% of the fruits consumed were red). The incidence of wasps in the seeds ranged from 0 to 42% of the crop in 1989 and from 0 to 24% in 1990. The influence of avian and insect frugivore guilds on each other appears to be quite low because of the narrow overlap in resource utilization by birds and wasps, and an overall low intensity of wasp seed predation. From an evolutionary perspective, the possible ability of wasps to preclude fruit maturation appears not to be attributable to the present interaction with avian frugivores.     

Measuring the discrepancy between fecundity and lifetime reproductive success in a pollinating fig wasp

Lifetime reproductive success in female insects is often egg‐ or time‐limited. For instance in pro‐ovigenic species, when oviposition sites are abundant, females may quickly become devoid of eggs. Conversely, in the absence of suitable oviposition sites, females may die before laying all of their eggs. In pollinating fig wasps (Hymenoptera: Agaonidae), each species has an obligate mutualism with its host fig tree species [Ficus spp. (Moraceae)]. These pro‐ovigenic wasps oviposit in individual ovaries within the inflorescences of monoecious Ficus (syconia, or ‘figs’), which contain many flowers. Each female flower can thus become a seed or be converted into a wasp gall. The mystery is that the wasps never oviposit in all fig ovaries, even when a fig contains enough wasp females with enough eggs to do so. The failure of all wasps to translate all of their eggs into offspring clearly contributes to mutualism persistence, but the underlying causal mechanisms are unclear. We found in an undescribed Brazilian Pegoscapus wasp population that the lifetime reproductive success of lone foundresses was relatively unaffected by constraints on oviposition. The number of offspring produced by lone foundresses experimentally introduced into receptive figs was generally lower than the numbers of eggs carried, despite the fact that the wasps were able to lay all or most of their eggs. Because we excluded any effects of intraspecific competitors and parasitic non‐pollinating wasps, our data suggest that some pollinators produce few offspring because some of their eggs or larvae are unviable or are victims of plant defences.     

Effect of parasitoids,seed‐predators and ant‐mutualists on fruiting success and germination of Acacia drepanolobium in Kenya

Acacia drepanolobium is an obligate ant‐plant that bears dehiscent pods exploited by predispersal seed‐predators and parasitoids. Fruit set and seed germination in relation to ant‐association, bruchid and parasitoid infestation were investigated in a large‐scale multi‐year study in Kenya. Ant‐association had a significant impact on the overall numbers of fruiting trees. 94.8% of Crematogaster mimosae‐occupied trees, 25.6% of C. nigriceps‐occupied trees, and 82.2% of Tetraponera penzigi‐occupied trees set fruit. Within each tree, ant‐association had no significant impact on the amount of seed produced: C. mimosae‐occupied trees produced 47.5 seeds per branch, C. nigriceps‐occupied trees 44.9 seeds per branch and T. penzigi‐occupied trees 38.3 seeds per branch. Ant‐association did not limit seed‐predators or vary significantly by ant. Seeds infested by bruchids germinated in significantly lower proportions (6.2%) compared to uninfested seed (78.6%). Bruchid‐infested seed is also exploited by parasitoid wasps. Parasitoids appear to have a moderate but significant ‘rescue’ effect on bruchid‐infested seed with 18.4% of parasitoid‐infested seed germinating. Stable isotopes (δ15N) revealed the trophic structure of the seed‐associated insects, showing clearly that bruchids are seed predators and the parasitoid wasp Dinarmus magnus exploits the dominant Bruchidius sp.     

Genomic analysis reveals an exogenous viral symbiont with dual functionality in parasitoid wasps and their hosts

Insects are known to host a wide variety of beneficial microbes that are fundamental to many aspects of their biology and have substantially shaped their evolution. Notably, parasitoid wasps have repeatedly evolved beneficial associations with viruses that enable developing wasps to survive as parasites that feed from other insects. Ongoing genomic sequencing efforts have revealed that most of these virus-derived entities are fully integrated into the genomes of parasitoid wasp lineages, representing endogenous viral elements (EVEs) that retain the ability to produce virus or virus-like particles within wasp reproductive tissues. All documented parasitoid EVEs have undergone similar genomic rearrangements compared to their viral ancestors characterized by viral genes scattered across wasp genomes and specific viral gene losses. The recurrent presence of viral endogenization and genomic reorganization in beneficial virus systems identified to date suggest that these features are crucial to forming heritable alliances between parasitoid wasps and viruses. Here, our genomic characterization of a mutualistic poxvirus associated with the wasp Diachasmimorpha longicaudata, known as Diachasmimorpha longicaudata entomopoxvirus (DlEPV), has uncovered the first instance of beneficial virus evolution that does not conform to the genomic architecture shared by parasitoid EVEs with which it displays evolutionary convergence. Rather, DlEPV retains the exogenous viral genome of its poxvirus ancestor and the majority of conserved poxvirus core genes. Additional comparative analyses indicate that DlEPV is related to a fly pathogen and contains a novel gene expansion that may be adaptive to its symbiotic role. Finally, differential expression analysis during virus replication in wasps and fly hosts demonstrates a unique mechanism of functional partitioning that allows DlEPV to persist within and provide benefit to its parasitoid wasp host.     

Parasitoid attack of the seed-feeding beetle Bruchus loti enhances the germination success of Lathyrus japonicus seeds

Hard seeds of some legume species can germinate after seed-feeding insects bore through the seed coat and consequently break seed dormancy. Larvae of bruchine beetles are the main seed feeders attacking many legume species. Boring of the hard seed coat by bruchine beetle larvae enhances the germination percentage of legume species, but consuming too much of a single seed may reduce the chances the seed will survive. We hypothesise that the early mortality of bruchine larvae due to parasitism contributes positively to seed germination because larvae are killed before consuming too large a quantity of the seed. Here, we tested this hypothesis using Lathyrus japonicus seeds and Bruchus loti, the main seed feeder attacking this plant. B. loti larvae were mainly parasitised by two species of idiobiont parasitoids—Pteromalus sp. and Dinarmus sp. The seeds from which Pteromalus wasps emerged germinated more successfully than did the seeds from which B. loti adults emerged. B. loti larvae parasitised by the two wasp species consumed the seeds less intensively than did unparasitised larvae. Thus, the results of experiments supported our hypothesis. However, the germination success varied significantly between the seeds from which Pteromalus and Dinarmus wasps emerged. The difference in the size of seeds the two wasp species chose for parasitism may have influenced the germination percentage.     

Beeren und Farbenwahl durch V?gel

Summary Three chance observations on colour preference in birds feeding on berries are presented.Blackbirds (Turdus merula) in Bavaria fed in a cherry-tree irrespective of the colour of the cherries. They began feeding when the first cherries ripened and turned from greenish-yellow to light red. As the cherries were offered in different colours at the same time, and pell-mell in the same tree, the birds presumably quickly learned by trial and error to eat cherries of different colours.A little later Blackbirds, which must have been partly if not exclusively the same individuals, started to feed on red currants. Red and white currants were offered to them at the same time, and side by side but on different bushes. In this case there was a very clear spontaneous selection of red in preference to white. White berries were only eaten after the red bushes had been cleared up by the birds.Sardinian Warblers (Sylvia melanocephala) in Sardinia, feeding on berries ofRhamnus alaterna, which are coral red and hard when unripe and turn to blackish and soft getting ripe, prefered the black ones. It is presumed that they conditioned themselves to black and soft by trial and error learning. Generalisations as to birds preferring certain colours in berries they feed on can not be drawn from any special case.     

Sexual attractiveness shared by both sexes mediates same‐sex sexual behaviour in the parasitoid wasp Telenomus triptus

The mating behaviour of a quasi‐gregarious egg parasitoid Telenomus triptus Nixon (Hymenoptera: Platygastridae), which exploits egg masses of a stink bug Piezodorus hybneri (Heteroptera: Pentatomidae), is examined in the laboratory. In this parasitoid wasp, male adults that emerge earlier stay at the natal egg mass and mate with subsequently emerging females. In the present study, a male adult that encounters the emergence of another male always waits for it to egress, and then mounts the newly emerging male. To examine why males of T. triptus show same‐sex sexual behaviour, male adults are presented with a parasitized host egg mass or a freshly killed wasp. Male adults are observed to remain at host egg masses from which only male wasp(s) had emerged. In addition, male adults attempt to copulate with freshly killed young male wasps. It is suggested that newly emerging male wasps are targets of same‐sex sexual behaviour because they possess cues for male sexual behaviour similar to the cues of females. Both the sex and age of freshly killed wasps affect the frequency of the sexual behaviour of male adults: females are more attractive than males, although their attractiveness declines with age. When the mating opportunity is restricted to the natal egg mass, the costs of failing to notice newly emerging female adults should be extremely high. Therefore, males are forced not to discriminate the sex, resulting in same‐sex sexual behaviour.     

Discrimination against previously searched, host-free patches by a parasitoid foraging for concealed hosts

1. The foraging behaviour of the parasitoid wasp Halticoptera laevigata Thomson (Hymenoptera: Pteromalidae) was studied. This wasp attacks the first‐instar larvae of the fruit fly Myoleja lucida Fallén (Diptera: Tephritidae) within honeysuckle Lonicera xylosteum L. (Caprifoliaceae) fruits. It was hypothesised that, to forage efficiently, wasps should be able to maintain a systematic or at least non‐random search pattern on the level of the microhabitat, i.e. branches with fruits, and/or be able to recognise previously searched fruits. 2. Comparisons of observed patch visitation patterns on branches bearing uninfested fruits with expected values for systematic and random visitation patterns indicated that H. laevigata wasps visit patches randomly and do not avoid visiting host‐free patches that they have visited previously. 3. While wasps did not alter their search effort on uninfested fruits searched previously by conspecifics, they searched for much shorter times and probed less frequently for hosts on uninfested fruits that they had searched previously. 4. The results indicate that H. laevigata wasps engage in partially systematic search and increase their search efficiency through the recognition of self‐searched fruits. The reasons why partial rather than fully systematic search should be adaptive in this parasitoid are discussed.     

SEX-RATIO MANIPULATION IN RESPONSE TO HOST SIZE BY THE PARASITOID WASP SPALANGIA CAMERONI: A LABORATORY STUDY

The prediction of Charnov et al.'s (1981) host-size model that there should be a negative relationship between host size and wasp sex ratio (proportion sons) was supported for Spalangia cameroni, a solitary parasitoid wasp. The relationship was shown to be a result of offspring sex manipulation by females in response to host size rather than a result of differential mortality of the sexes. A major assumption of the host-size model is that host size has a greater effect on the ultimate reproductive success of emerging female wasps than of males. This assumption was not supported. Host size had a positive effect on the size of both male and female S. cameroni. However, neither host size nor wasp size affected longevity, production of offspring by females, or ability of males to compete for mates. Host size may differentially affect the reproductive success of female and male wasps through effects on other aspects of reproductive success. Tests of the assumptions of offspring sex-ratio manipulation hypotheses are scarce but critical, not only for parasitoid wasps, but also for other organisms.