Impact of cocoon predation and parasitism on endemic populations of the common pine sawfly, Diprion pini (L.) (Hymenoptera, Diprionidae) in different forest types

1 Predation and parasitism on litter‐buried cocoons of the common pine sawfly Diprion pini (L.) were compared in different forest types with endemic sawfly populations by field exposure of laboratory‐reared cocoons during three consecutive years (1993–1995). 2 The impact of cocoon predation was dependent on season and forest type. The highest predation (up to 95%) was found during autumn in forest stands with a dense understory vegetation. 3 Cocoon parasitism varied between year, season and forest type. The highest parasitoid attack was observed in pure pine forests with more or less barren soils, but did not exceed 24% of exposed cocoons. 4 Cocoons were exposed in small patches. Predators tended to exploit all cocoons of a patch, whereas parasitoids only attacked a few cocoons of a patch. Predation was similar on cocoons placed in the litter and those buried more deeply in the soil, whereas parasitism of soil‐buried cocoons was rare. 5 These results indicate that predators can have a remarkable potential for limiting endemic sawfly densities, if habitat conditions in a forest maintain their population and support their foraging behaviour. A notable effect of parasitoids on sawfly cocoons deposited in the litter is obviously restricted to typical pure and barren pine forests, but may play there a similar role as predation.     

Response to host age of the egg parasitoid Anaphes n.sp. (Hymenoptera: Mymaridae)

The mymarid Anaphes n.sp. was observed to parasitize the eggs of the carrot weevil Listronotus oregonensis (LeConte) up to 138 h old at 23±1°C and 18L:6D. Adult emergence of the parasitoid was observed in 85%‐97.5% of the weevil eggs 42–138 h old and host embryogenesis was interrupted in nearly 100% of these eggs. In host eggs of 18 h old, parasitism was very high but not all parasitoids completed their development and their subsequent emergence was generally retarded. An.sp. could parasitize host eggs 162 h old, but such an attack rarely retarded the normal development of the host. The parasitoid was unable to complete its development in sterile carrot weevil eggs.     

Evidence that the availability of suitable pine limits non‐native Sirex noctilio in Ontario

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Food web structure of aphids and their parasitoids in Belgian fruit agroecosystems

Community structures of aphids and their parasitoids were studied in fruit crop habitats of eastern Belgium in 2014 and 2015. Quantitative food webs of these insects were constructed separately for each year, and divided into subwebs on three host‐plant categories, fruit crop plants, non‐crop woody and shrub plants and non‐crop herbaceous plants. The webs were analyzed using the standard food web statistics designed for binary data. During the whole study period, 78 plant species were recorded as host plants of 71 aphid species, from which 48 parasitoid species emerged. The community structure, aphid / parasitoid species‐richness ratio and trophic link number varied between the two years, whereas the realized connectance between parasitoids and aphids was relatively constant. A new plant–aphid–parasitoid association for Europe was recorded. Dominant parasitoid species in the study sites were Ephedrus persicae, Binodoxys angelicae and Praon volucre: the first species was frequently observed on non‐crop trees and shrubs, but the other two on non‐crop herbaceous plants. The potential influence, through indirect interactions, of parasitoids on aphid communities was assessed with quantitative parasitoid‐overlap diagrams. Symmetrical links were uncommon, and abundant aphid species seemed to have large indirect effects on less abundant species. These results show that trophic indirect interactions through parasitoids may govern aphid populations in fruit crop habitats with various non‐crop plants, implying the importance for landscape management and biological control of aphid pests in fruit agroecosystems.     

Within-plant vertical distributions of the scale insect Nipponaclerda biwakoensis and its five parasitoids that exhibit frequent successful multiparasitism on the common reed

Within‐shoot vertical distributions of the scale insect Nipponaclerda biwakoensis, five species of gregarious parasitoid wasps attacking the scale, and successful multiparasitism (emergence of multiple parasitoid species from a single host) by the parasitoids on the common reed were investigated. Each reed shoot collected was longer than 2.0 m, and was divided into pieces of 0.5 m each from the base of the shoot. The mean number of adult female scales per 0.5 m of shoot increased with height, but the mean rate of overall parasitism of the female scales decreased with height. The five parasitoids showed species‐specific within‐shoot distribution patterns, with respect to the rate of parasitism: Astymachus japonicus, Boucekiella depressa and Encyrtidae sp. 2 showed the greatest rates of parasitism at heights of 0–0.5, 0.5–1.0 and 1.0–1.5 m, respectively. The parasitism rate by Aprostocetus sp. was higher at the lower half of the shoots, whereas that by Encyrtidae sp. 1 was higher at the central positions. The distribution of the five parasitoids largely overlapped with each other at the level of scale aggregations. Nevertheless, successful multiparasitism occurred frequently only in several types of species combinations that involved two parasitoid species showing similar within‐shoot distribution patterns. The rate of successful multiparasitism by B. depressa with A. japonicus or Aprostocetus sp., and that by Encyrtidae sp. 2 with Encyrtidae sp. 1 was constantly high at different vertical positions on the shoots. This may be because B. depressa and Encyrtidae sp. 2 preferentially oviposit into scales previously parasitized by these other species.     

Temporal and geographic variation in parasitoid attack with no evidence for ant protection of the Melissa blue butterfly,Lycaeides melissa

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Parasitism of a pollinator fig wasp: mortalities are higher in figs with more pollinators,but are not related to local densities of figs

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Success of aphid parasitoids and their hosts varies with ant attendance: A field study

Aphid–ant associations are often described as mutually beneficial interactions in which honeydew is traded for protection from predators and parasitoids. The aim of the present study was to determine parasitization avoidance in ant‐tended aphid colonies. Field experiments were carried out on two host plants: hoary cress Lepidium draba (Brassicaceae) and Canadian teasel Cirsium arvense (Asteraceae). Lepidium was host to Acyrthosiphon gossypii (Aphididae) tended by the ant Lasius turcicus (Formicidae) and attacked by two parasitoids, Trioxys asiaticus (Braconidae) and Lysiphlebus fabarum (Braconidae). Cirsium was host to Brachycaudus cardui tended by Crematogaster sordidula and attacked by L. fabarum and Aphidius colemani (Braconidae). The per capita population growth rate of A. gossypii was significantly higher in the presence of ants, while B. cardui was negatively affected, albeit non‐significantly. The parasitism rate of A. gossypii decreased significantly when tended by Lasius turcicus, but the presence of Crematogaster sordidula in colonies of B. cardui significantly increased parasitism. Our results indicate that the effects of ant attendance vary between different aphid–ant interactions. Moreover, parasitoids can benefit from the presence of ants under some conditions.     

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.     

Parasitoid complex of the pistachio twig borer moth,Kermania pistaciella,in Iran

The parasitoid complex of the pistachio twig borer moth, Kermania pistaciella Amsel (Lepidoptera: Tineidae), a native pest of pistachio trees, was investigated at 27 pistachio plantation sites in Kerman province, the major pistachio growing area of Iran. The present study was conducted to document the naturally established parasitoid complex and to assess the need for improving the biological control of this species. In total, 22,390 moth cocoons were collected from 186 samples collected from commercial orchards during 2006–2008 and kept singly in controlled conditions to rear immature insects. An average of 2.8% of moth cocoons had been attacked by predators at time of sampling. Of the collected cocoons, on average 46.7% completed development and emerged as adult moths, no insects emerged from 8%, suggesting that the moth or wasp died before maturing, and parasitoids emerged from the remaining 42.5%. The overall percentage of host cocoons from which wasps emerged ranged from 25.6 to 59%. Fifteen hymenopterous parasitoid species were recovered from cocoons, of which three species were primary parasitoids, two were obligatory hyperparasitoids and the remaining 10 species were facultative hyperparasitoids. The primary parasitoid, Chelonus kermakiae (Tobias) (Hymenoptera: Braconidae), was the most abundant comprising almost 85% of the total emerging parasitoids. In addition, a further four species of larval parasitoid developed within the PTBM's larval tunnels in pistachio fruit cluster-stem tissue. Conservation of these parasitoids in the pistachio growing areas is recommended since a high level of parasitized moths’ cocoons was found at the majority of experimental sites.     

Effects of learning experience on behaviour of the generalist parasitoid Sclerodermus pupariae to novel hosts

Massicus raddei Blessig (Coleoptera: Cerambycidae), also referred to as the oak long‐horned beetle (OLB), is a non‐natural host for the generalist parasitoid Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae). To determine whether this generalist parasitoid might be a suitable agent for the control of OLB, the adaptive learning experience of adult female parasitoids to OLB larvae was investigated in the laboratory. A Y‐tube olfactometer bioassay was used to examine the effects of adaptive learning experience on the foraging ability of parasitoids for OLB larvae. The results indicated that parasitoids were significantly attracted by the volatiles of ash bark, Fraxinus velutina, with emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae and larval frass, after exposure to ash bark mixed with EAB larval frass (learning condition A). In contrast, after exposure to oak bark, Quercus liaotungensis, mixed with OLB larval frass (learning condition C), parasitoids showed significant preference for the volatiles of oak bark with OLB larvae and larval frass. On the basis of the results of no‐choice tests, we found that parasitoids exposed to learning condition C had greater paralysis efficiency and higher OLB larvae parasitism rates than those exposed to learning condition A or no experience. Furthermore, parasitoids fed on OLB larvae in learning condition C had significantly greater paralysis efficiency and higher OLB larvae parasitism rates than other parasitoids tested. Parasitoids fed on EAB larvae in learning condition A had the lowest paralysis efficiency and OLB larvae parasitism rates among the parasitoids tested. These findings suggested that adaptive learning significantly enhanced the ability of a generalist parasitoid to utilize a novel host. This may provide a new approach to controlling non‐natural hosts using generalist parasitoids.     

Herbivore species identity rather than diversity of the non‐host community determines foraging behaviour of the parasitoid wasp Cotesia glomerata

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species‐rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore‐induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non‐suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non‐host herbivores influence the behaviour of parasitoids, we created environments with different levels of non‐host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non‐host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non‐host species diversity nor non‐host identity influenced the preference of the parasitoid for herbivore‐infested plants. However, after landing on the plant, non‐host species identity did affect parasitoid behaviour, whereas non‐host diversity did not. One of the non‐host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non‐host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non‐host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.     

Seasonal phenology and stage-specific parasitism of the apple ermine moth, Yponomeuta malinellus Zeller, in Korea

The apple ermine moth, Yponomeuta malinellus Zeller (Lepidoptera: Yponomeutidae), is a tent caterpillar that feeds on Malus spp. in Korea. Populations of the moth in native areas appeared to be regulated by the assemblage of parasitoids. Phenological associations between host stages and parasitoids, susceptible stage(s) of the host for each parasitoid, and stage‐specific parasitism were studied. The egg larval parasitoid Ageniaspis fuscicollis (Dalman) had highest parasitism of first instar larvae (24%), with 14% parasitism of other larval stages. Dolichogenidea delecta (Haliday) was recovered from all larval instars with the highest parasitism rate of second instar larvae (20.1%), followed by 19.9% parasitism of mid‐larval hosts. Herpestomus brunicornis Gravenhorst was reared from second instar larvae through to pupal collection, and had the highest parasitism rate (29.9%) at the pupal stage. The larval pupal parasitoid Zenillia dolosa (Meigen) was recovered from mid‐larval to pupal stages with the highest parasitism rate (5.5%) occurring in third to fourth instar larvae. The host stages for developing A. fuscicollis completely overlap with those of D. delecta, and with those of H. brunicornis to some degree. A statistically significant negative correlation exists between A. fuscicollis and these dominant parasitoids, indicating competitive interaction within the host.     

Phenology of native weevils (Coleoptera: Curculionidae) in New Zealand pastures and parasitism by the introduced braconid,Microctonus aethiopoides Loan (Hymenoptera: Braconidae)

Abstract

The phenology of native brachycerine weevil species at seven pasture sites in Otago, Canterbury and Waikato was studied by regular quantitative sampling of adults. Weevils were identified to species, and dissected to record reproductive status and parasitism by introduced braconid parasitoids in the genus Microctonus. Climatic data assisted in the interpretation of some population density patterns. Weevil population density was estimated for periods of two to five years at the selected sites. Species in the Entimini (species of Irenimus and Nicaeana) were generally univoltine, with adults emerging in winter‐spring. The main period of reproductive activity was spring, and parasitism by Microctonus aethiopoides reached its highest incidence in January. Low level parasitism of native weevil species by M. aethiopoides was detected at all sites, and by M. hyperodae at two sites. At one site in Otago, parasitism by M. aethiopoides was higher and could have affected the population density of Irenimus aemulator (Broun) and Nicaeana sp. Most parasitism occurred after the main reproductive period of weevils in spring, but a putative second generation in some species might be more affected by parasitoid attack. A native rhytirhinine species, Steriphus variabilis, differed from the entimines because adults emerged in autumn and spring, and may be bivoltine. Mechanisms of M. aethiopoides parasitism of non‐target species in the field are discussed.     

DEFENSE TRAITS OF LARVAL DROSOPHILA MELANOGASTER EXHIBIT GENETICALLY BASED TRADE‐OFFS AGAINST DIFFERENT SPECIES OF PARASITOIDS

Populations of Drosophila melanogaster face significant mortality risks from parasitoid wasps that use species‐specific strategies to locate and survive in hosts. We tested the hypothesis that parasitoids with different strategies select for alternative host defense characteristics and in doing so contribute to the maintenance of fitness variation and produce trade‐offs among traits. We characterized defense traits of Drosophila when exposed to parasitoids with different host searching behaviors (Aphaereta sp. and Leptopilina boulardi). We used host larvae with different natural alleles of the gene Dopa decarboxylase (Ddc), a gene controlling the production of dopamine and known to influence the immune response against parasitoids. Previous population genetic analyses indicate that our focal alleles are maintained by balancing selection. Genotypes exhibited a trade‐off between the immune response against Aphaereta sp. and the ability to avoid parasitism by L. boulardi. We also identified a trade‐off between the ability to avoid parasitism by L. boulardi and larval competitive ability as indicated by differences in foraging and feeding behavior. Genotypes differed in dopamine levels potentially explaining variation in these traits. Our results highlight the potential role of parasitoid biodiversity on host fitness variation and implicate Ddc as an antagonistic pleiotropic locus influencing larval fitness traits.     

Non-target parasitism of the endemic New Zealand red admiral butterfly (Bassaris gonerilla) by the introduced biological control agent Pteromalus puparum

The New Zealand red admiral butterfly, Bassaris gonerilla (F.) (Lepidoptera: Nymphalidae), has been known as a non-target host for the introduced biological control agent Pteromalus puparum (L.) (Hymenoptera: Pteromalidae) for at least 35 years, but the level of parasitism has never been quantified. Pre-imaginal mortality in B. gonerilla was assessed over the southern summer of 2000/01 at six field sites in the Christchurch area of the South Island, New Zealand. Individual eggs and larvae were identified by tagging the stem of the Urtica ferox Forst.f. plant on which they were found and the fate of these individuals was checked weekly. These data were used to construct a partial life table for B. gonerilla. Egg mortality was very high (95%), with parasitism by an unidentified Telenomus sp. Haliday (Hymenoptera: Scelionidae) causing 57% mortality. Mortality in the larval and pupal stages increased at a constant rate with age and the major mortality factor was disappearance, which was assumed to be a result of predation and dispersal of larvae. The introduced biological control agent P. puparum parasitized 14% of B. gonerilla pupae sampled. However, parasitism by another exotic parasitoid, the self-introduced Echthromorpha intricatoria (F.) (Hymenoptera: Ichneumonidae), was even higher at 26%. A survey of pupal parasitism in three regions of New Zealand (Wellington, Christchurch, and Dunedin) revealed overall parasitism levels of 67% by E. intricatoria and 8% by P. puparum, but due to the difference in emergence times of B. gonerilla and its parasitoids, these are likely to be overestimates of percent parasitism. It is concluded that P. puparum has permanently enhanced mortality in B. gonerilla, but the level of mortality is low relative to egg parasitism by Telenomus sp., larval disappearance mortality, and pupal mortality due to E. intricatoria parasitism. To determine if this level of pupal parasitism has had population effects will require more data and the development of a population model for B. gonerilla.     

Seasonal patterns of egg parasitism and natural biological control of rice brown planthopper in Indonesia

1 Egg mortality is known to be an important factor in brown planthopper (BPH) (Nilaparvata lugens) population dynamics in tropical Asia, but few quantitative data are available on the role of egg parasitoids. 2 Field studies are reported at two sites in Java, Indonesia, over a period of 4 years in both wet and dry seasons, where no chemical pesticides were used. Rice plants previously infested in the laboratory with BPH eggs were exposed to natural parasitism and predation in experimental fields for periods of 5 days. 3 Egg batches were dissected from both experimental and control plants after field exposure at seven fortnightly intervals through the rice growing season and BPH nymphs and adult parasitoids were allowed to emerge. 4 Parasitism by species of Oligosita (Hymenopera, Trichogrammatidae) and Anagrus (Hym., Mymaridae) varied between 18% and 61% in dry, and from 1% to 65% in wet, seasons. There was generally a trend from low to higher rates through wet seasons, contrasting with more uniform higher levels through dry seasons. 5 Reduction in BPH survival as a result of egg parasitism varied between 29% and 91%. Generally parasitism was density independent. 6 It is suggested that lower initial levels of parasitism in dry seasons may be due to the longer fallow period between dry and wet, than between wet and dry, resulting in lower local populations of appropriate natural enemies. 7 Unlike previous studies in Asia, little evidence was found for egg predation as an important mortality factor. 8 The conservation of egg parasitoids should form part of any rice pest management programmes in tropical Asia.     

Diamondback moth females oviposit more on plants infested by non‐parasitised than by parasitised conspecifics

Abstract 1. When offered a choice, female diamondback moths (Plutella xylostella) oviposited more eggs on plants with non‐parasitised conspecific larvae than on plants with parasitised larvae. 2. The leaf area consumed by parasitised larvae was significantly lower than that by non‐parasitised larvae. However, this quantitative difference in larval damage did not explain the female’s ability to discriminate between plants with parasitised and non‐parasitised larvae, as females showed an equal oviposition preference for plants infested by higher or lower densities of non‐parasitised larvae. 3. Pupal weight and duration of the larval stage of P. xylostella were independent of whether larvae were reared on plants that were previously infested by either non‐parasitised or parasitised larvae. 4. The larval parasitoid Cotesia vestalis did not distinguish between plants infested by non‐parasitised larvae and plants infested by larvae that had already been parasitised by conspecific wasps. 5. Based on these data, it can be concluded that the moth oviposition preference for plants infested by non‐parasitised conspecifics relative to plants infested by parasitised conspecifics was not explained by plant quality or by the attractiveness of plants towards wasps. It is hypothesised that one of the reasons for this preference is avoidance of plants where a relatively high risk of parasitism is expected due to the emergence of parasitoids from the parasitised host larvae.     

Seasonal changes in seaweed deposition,seaweed fly abundance,and parasitism at the pupal stage along sandy beaches in central Japan

Seasonal relationships among stranded wrack quantity, seaweed fly abundances, and parasitism at the pupal stage were studied along three sandy beaches in central Japan. The seasonal occurrence patterns of puparia of seaweed flies Coelopa frigida and Fucellia spp. generally corresponded to seaweed deposition, which peaked in May–July and October–December. Parasitoids use fly puparia in these seasons. However, the occurrence of seaweed flies and their parasitoids varied among the three sandy beaches and did not correspond to the wrack amounts. These findings suggest that populations of seaweed flies and their parasitoids are seasonally, but not spatially, regulated by bottom‐up processes. The parasitoid assemblage of fly puparia was composed of two Aleochara (Coleoptera: Staphylinidae), two Trichopria (Hymenoptera: Diapriidae), and five pteromalid species (Hymenoptera), but the rate of parasitism was less than 20% and might have had little effect on fly populations.