The spatial patterns of parasitism of eumenid wasps,Anterhynchium flavomarginatum andOrancistrocerus drewseni by the miltogrammine flyAmobia distorta

1. Spatial patterns of parasitism of eumenid wasps Anterhynchium flavomarginatum and Orancistrocerus drewseni by the miltogrammine fly Amobia distorta were studied in Kyoto, Japan during 1980–1984.
2. In generations of low (<5%) and medium (5–20%) parasitism, percent parasitism per shed (the habitat of the hosts) increased as a function of host density. Conversely, in generations of high (>20%) parasitism, percent parasitism was rather constant over different host densities.
3. The spatial distributions of adult miltogrammine flies among sheds were censused in generations of low and medium parasitism. The frequency of observations of adult miltogrammine flies was higher at sheds of higher host density (aggregative behavioral response), but on the other hand, the adult miltogrammine flies distributed in an underdispersed (or regular) manner in relation to other conspecifics.
4. The spatially density independent relationship between host density and percent parasitism in generations of high parasitism was explained in relation to parasitoid dispersal from patches of high parasitoid density.

     

Optimal use of patches by parasitoids with a limited fecundity

1. a mathematical model is presented which predicts the expected optimal-patch-use strategy for solitary parasitoids with a limited fecundity.
2. The model predicts that the quality of the patches is determined by the proportion of unparasitized hosts and not by the density of those hosts, and that throughout the searching period the parasitoids should maintain the level of parasitism equal in all the patches irrespective of the host density per patch.
3. The spatial pattern of parasitism among field patches by a parasitoid with a low fecundity, Praestochrysis shanghaiensis, was in agreement with the prediction of the model, i.e., a similar level of parasitism in different patches was observed when the ratio of female parasitoids to hosts in the whole study area exceeded 0.07. When the ratio was less than 0.05, however, the level of parasitism per patch showed an inverse relation to the host density, and was positively correlated with the female parasitoid-host ratio.
4. The model assumes that the parasitoids move between patches without cost and have perfect information about patch quality. Consideration of the cost of moving and sampling bridges the gap between the observed and predicted rates of parasitism found when the female parasitoid-host ratio in the whole study area was low

     

Spatio-temporal analysis of the population dynamics of the oriental moth,Monema flavescens (Lepidoptera: Limacodidae)

The comparative analysis of life tables of the oriental moth, Monema flavescens, obtained in 6 patches for 8 generations in 4 years revealed the following:

1. The ratio of maximum to minimum of cocoon density for each patch ranged from 5.34 to 22.50, each value being more than 3.20, the ratio for the whole study area.
2. The density change from adult to cocoon in the next generation caused most of the spatial variation in density change per patch. The rate of adult-to-cocoon population change showed spatial density dependence in some generations but not in others. When the change rate lacked spatial density dependence, it was the key-factor for spatial variation in adult density for the following few generations till the change rate recovered spatial density dependence. This was because of flooding, which killed the spatial density dependence existing potentially in the adult-to-cocoon change rate and damaged the same patches during the few successive generations.
3. The rate of population change from overwintered generation adults (summer ones) to first generation cocoons was not only the key-factor for the rate of throughout-the-year change but temporally density dependent in each patch. Therefore, the density for the whole study area is considered to fluctuate within a range. However, the strong equilibrium seen in the cocoon density for the whole study area was due to the floods that happened to occur when the density was near and at its maximum, and it is considered that such a strong equilibrium does not always occur.
4. In the population change from summer adults to first generation cocoons, temporal density dependence was found in all the patches, but it was found only in one patch in the population change from autumn adults to second generation cocoons. This was because the spatial density dependence seen in the former corresponded to the absolute density of adults, while that in the latter corresponded to the relative density.

     

Recruitment of native parasitoids by an exotic leaf miner, Cameraria ohridella: host-parasitoid synchronization and influence of the environment

Abstract

• 1 The horse‐chestnut leaf miner, Cameraria ohridella, is a moth of unknown origin that has recently invaded Europe and severely defoliates the European horse‐chestnut, an important ornamental tree.
• 2 Several indigenous parasitoids have colonized this new host, but parasitism remains low. One of the hypotheses suggested to explain the low parasitism is that candidate parasitoids emerge too early in spring to attack the first host generation and, thus, need early‐occurring leaf miners as alternate hosts. This hypothesis was tested by observing the synchronization between the phenology of the moth and that of its main parasitoids, and by comparing parasitism rates and parasitoid richness in different environments with various levels of biological diversity.
• 3 In spring, the bulk of the parasitoids emerge at least 5 weeks before the occurrence of the first suitable larvae of C. ohridella whereas most parasitoid adults reared outdoors die within 5 weeks after emergence.
• 4 Parasitism rates and parasitoid richness do not increase with biological diversity, suggesting that most parasitoids attacking the first generation of C. ohridella do not come from alternate hosts. Parasitism does not increase later in the year in the subsequent generations, when host‐parasitoid synchronization becomes less critical.
• 5 We conclude that, although the spring emergence of parasitoids is not synchronized with the phenology of C. ohridella, the parasitoids attacking the first generation are probably old or late‐emerging adults of the overwintering generation. The lack of synchronization is probably not the only reason for the poor recruitment of native parasitoids by C. ohridella.

     

Biology of Centistes delusorius, a parasitoid of adult apple blossom weevil

Abstract

• 1 Natural control of apple blossom weevil, Anthonomus pomorum (L.), deserves attention, as the pest is regaining importance with the declining use of non‐selective pesticides in apple and pear orchards. In this study the biology of Centistes delusorius (Förster), a specific parasitoid of adult apple blossom weevil, is investigated.
• 2 The parasitoid hibernates as young larva in an adult weevil, and juvenile development is resumed in early spring. The fully grown parasitoid larvae leave their hosts during full bloom at the end of April and early May, to pupate. The adults emerging in May oviposit into the newly emerged weevils, which initially feed on apple leaves.
• 3 Centistes delusorius was detected in six out of 15 host‐weevil infested orchards, but was only common in two with larger apple trees standing in grass. There, parasitism levels of around 30% were usual in hosts taken from treebands in winter.
• 4 The delicate larva is vulnerable, and the thin cocoon provides little protection against either desiccation or drowning on a weedless orchard floor. Observations indicate that successful pupation of C. delusorius demands stable humid conditions and some shelter, such as that found in grass or woodland soils.
• 5 Parasitoid females, provided with honey, lived for a mean of 6.3 ± 2.1 days under outdoor conditions in June. Their life span was similar whether they had access to and oviposited in hosts, or not. The species is pro‐ovigenic, and potential fecundity is about 40 eggs. Oviposition usually takes a few seconds. Parasitized female hosts do not reproduce.
• 6 Up to 95% of the parasitoid eggs laid in May develop into a second generation, the adults of which appear in July, when the host has entered aestivation. Older (British) records of C. delusorius outside orchards suggest that some parasitized hosts, like the healthy ones, leave the orchard prior to aestivo‐hibernation, so that the latter do not escape parasitoid attack in July.
• 7 A trapping sample in late June, when most non‐parasitized weevils have gone into aestivo‐hibernation, is probably the most efficient method to detect parasitized weevils.
• 8 The (near‐)absence of C. delusorius in many orchards is probably due not only to pesticide side‐effects, or scarcity of its host, but also to the absence of suitable pupation sites for the wasp.

     

The effect of host developmental stage at parasitism on sex‐related size differentiation in a larval endoparasitoid

• 1 For their larval development, parasitoids depend on the quality and quantity of resources provided by a single host. Therefore, a close relationship is predicted between the size of the host at parasitism and the size of the emerging adult wasp. This relationship is less clear for koinobiont than for idiobiont parasitoids.
• 2 As size differentiation in host species exhibiting sexual size dimorphism (SSD) is likely to occur already during larval development, in koinobiont larval endoparasitoids the size of the emerging adult may also be constrained based on the sex of the host caterpillar.
• 3 Sex‐specific growth trajectories were compared in unparasitised Plutella xylostella caterpillars and in second and fourth instar hosts that were parasitised by the solitary larval koinobiont endoparasitoid Diadegma semiclausum. Both species exhibit SSD, where females are significantly larger than males.
• 4 Healthy female P. xylostella caterpillars developed significantly faster than their male conspecifics. Host regulation induced by D. semiclausum parasitism depended on the instar attacked. Parasitism in second‐instar caterpillars reduced growth compared to healthy unparasitised caterpillars, whereas parasitism in fourth‐instar caterpillars arrested development. The reduction in growth was most pronounced in hosts producing male D. semiclausum.
• 5 Parasitism itself had the largest impact on host growth. SSD in the parasitoid is mainly the result of differences in growth rate of the parasitoid–host complex producing male and female wasps and differences in exploitation of the host resources. Female wasps converted host biomass more efficiently into adult biomass than males.

     

Do distances among host patches and host density affect the distribution of a specialist parasitoid?

The effect of spatial habitat structure and patchiness may differ among species within a multi-trophic system. Theoretical models predict that species at higher trophic levels are more negatively affected by fragmentation than are their hosts or preys. The absence or presence of the higher trophic level, in turn, can affect the population dynamics of lower levels and even the stability of the trophic system as a whole. The present study examines different effects of spatial habitat structure with two field experiments, using as model system the parasitoid Cotesia popularis which is a specialist larval parasitoid of the herbivore Tyria jacobaeae. One experiment examines the colonisation rate of the parasitoid and the percentage parasitism at distances occurring on a natural scale; the other experiment examines the dispersal rate and the percentage parasitism in relation to the density of the herbivore and its host plant. C. popularis was able to reach artificial host populations at distances up to the largest distance created (at least 80 m from the nearest source population). Also, the percentage parasitism did not differ among the distances. The density experiment showed that the total number of herbivores parasitised was higher in patches with a high density of hosts, regardless of the density of the host plant. The percentage parasitism, however, was not related to the density of the host. The density of the host plant did have a (marginally) significant effect on the percentage parasitism, probably indicating that the parasitoid uses the host plant of the herbivore as a cue to find the herbivore itself. In conclusion, the parasitoid was not affected by the spatial habitat structure on spatial scales that are typical of local patches.     

The parasitoid complex affecting Choristoneura rosaceana and Pandemis limitata in organically managed apple orchards

Parasitism and density of obliquebanded leafroller,Choristoneura rosaceana (Harris), and thethree-lined leafroller, Pandemis limitata(Robinson) (Lepidoptera: Tortricidae), were studied in 1998–2001 in organically managed apple orchards in the southern interior of British Columbia, Canada. The density of the overwintered generation of leafrollers in spring was significantly higher than thedensity of the first generation in summer. There was a significant inverse correlation between the density of leafrollers and the percent parasitism within generation and therefore parasitoids may play a role in controlling leafroller density. Parasitism of the overwintered generation (means between 5.5 and 24.7%) was significantly lower than parasitism documented in the summer generation (means up to 67.9%). The parasitoid complex recorded as emerging from these leafrollerpopulations in 1998–1999 included 30 species, of which eight have not been previously recorded in the literature as parasitoids of either leafroller species. The most frequently occurring parasitoids included Apophuasimplicipes (Cresson) and two species of Glypta(Hymenoptera: Ichneumonidae), Macrocentrus linearis (Nees), Meteorus trachynotus Viereck, Apantelespolychrosidis Viereck, Apanteles atra (Ratzeburg) and Microgaster epagoges Gahan (Hymenoptera: Braconidae), Colpoclypeus florus (Walker) and one Sympiesisspecies (Hymenoptera: Eulophidae). All of these more common parasitoids, except Glypta sp. 1 and M. epagoges, were recorded parasitizing leafroller hosts in both leafroller generations. The number of days to adult parasitoid emergence, when field collected parasitized hosts were held at 20°C, was recorded for four of the parasitoid species. Meteorus trachynotus was found to emerge early enough in spring toparasitize the few remaining overwintered early instar leafrollers. Glypta sp. 1, A. simplicipes and one Apanteles species emerged to coincide with the first generationin the summer.     

Heterogeneity and density dependence in a field study of a tephritid-parasitoid interaction

Abstract.

• 1 The spatial distributions of two tephritid flies (Urophora stylata (Fabricius) and Terellia serratulae L.) attacking thistle flower heads and the levels of parasitism from their associated parasitoid guilds were studied over a 7-year period.
• 2 Using these data it is possible to seek both temporal, density dependent relationships between average levels of parasitism and host density per generation, and also any spatial patterns of parasitism contributing to stability that may be operating within the same field system.
• 3 Parasitism by the two most important generalist parasitoids of T.serratulae is a direct function of average T.serratulae density per year. There is little evidence of any stabilizing heterogeneity arising from the spatial distribution of parasitism within generations.
• 4 Temporal density dependence of Urophora stylata cannot be confirmed from the 7 years of study but there is evidence of spatial heterogeneity which may have an important effect on the dynamics of the host population.

     

Density dependence of Anaphes sordidatus (Hymenoptera: Mymaridae) parasitism on eggs of Listronotus oregonensis (Coleoptera: Curculionidae)

Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) oviposits in carrot leaves and the larvae feed in carrot roots. Its eggs are parasitized by Anaphes sordidatus (Girault) (Hymenoptera: Mymaridae) throughout its oviposition period. This parasitoid is the major biotic mortality factor for L. oregonensis. Parasitism by A. sordidatus was studied for three years in sequentially sown plots of carrots, Daucus carota var. sativa. Over the scason, significantly fewer L. oregonensis eggs were oviposited in later sown carrots than in earlier sown carrots because oviposition started later in late sown plots of carrots. A positive temporal density-dependent relationship was observed each year between parasitism rates and host densities. This positive density dependence occurred in early and mid-summer for earlysown carrots where host density reached 1–2 host eggs per plant but disappeared in late summer when host density decreased while parasitism remained high. Latesown carrots had low host egg density (0.2 host egg per plant) and contributed little to the total number of eggs. In these late sown plots, parasitism increased rapidly to over 80% but no density dependence was observed. Spatially, few statistically significant regressions were found but all indicated a positive spatial density-dependence. Most non-significant regressions occurred because the range of egg density was too small between plots for a given date.     

Aggregation in field parasitoid populations: foraging time allocation by a population of Diadegma (Hymenoptera, Ichneumonidae)

ABSTRACT.

• 1 A field study was made of foraging time allocation by a population of parasitic wasps, Diadegma spp. (Ichneumonidae), to plants containing different densities of their hosts, the caterpillars of Plutella xylostella (L.).
• 2 The parasitoid population exhibited a clear aggregative response, spending more total time on higher density patches, which probably resulted from wasps making more and longer visits to these densities.
• 3 Despite this aggregation, positive density dependent parasitism was not found. The functional response of the Diadegma population exhibited an upper asymptote at high host densities, probably due to an increase in the proportion of time spent handling hosts, which countered the effect of aggregation.
• 4 While Diadegma may select and forage preferentially on plants with higher host density, they do not exhibit the tendency, predicted by some optional foraging models, to exploit progressively less profitable plants during a foraging bout. Some factors affecting patterns of parasitoid foraging in the field are discussed.

     

The effects of host weight at parasitism on fitness correlates of the gregarious koinobiont parasitoid Microplitis tristis and consequences for food consumption by its host, Hadena bicruris

Gregarious koinobiont parasitoids attacking a range of host sizes have evolved several mechanisms to adapt to variable host resources, including the regulation of host growth, flexibility in larval development rate, and adjustment of clutch size. We investigated whether the first two mechanisms are involved in responses of the specialist gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) to differences in the larval weight and parasitoid load of its host Hadena bicruris Hufn. (Lepidoptera: Noctuidae). In addition, we examined the effects of parasitism on food consumption by the host. Parasitoids were offered caterpillars of different weight from all five instars, and parasitoid fitness correlates, including survival, development time, and cocoon weight, were recorded. Furthermore, several host growth parameters and food consumption of parasitized and unparasitized hosts were measured. Our results show that M. tristis responds to different host weights by regulating host growth and by adjusting larval development rate. In hosts with small weights, development time was increased, but the increase was insufficient to prevent a reduction in cocoon weight, and as a result parasitoids experienced a lower chance of successful eclosion. Cocoon weight was negatively affected by parasitoid load, even though host growth was positively affected by parasitoid load, especially in hosts with small weights. Later instars were more optimal for growth and development of M. tristis than early instars, which might reflect an adaptation to the life‐history of the host, whose early instars are usually concealed and inaccessible for parasitism on its food plant, Silene latifolia Krause (Caryophyllaceae). Parasitism by M. tristis greatly reduced total host food consumption for all instar stages. Whether plants can benefit directly from the attraction of gregarious koinobiont parasitoids of their herbivores is a subject of current debate. Our results indicate that, in this system, the attraction of a gregarious koinobiont parasitoid can directly benefit the plant by reducing the number of seeds destroyed by the herbivore.     

Structure,organization, and response of a species-rich parasitoid community to host leafminer population dynamics

The parasitoid community dynamics of an agromyzid honeysuckle leafminer, Chromatomyia suikazurae (Agromyzidae, Diptera) were studied between 1981 and 1990 in a natural forest in Kyoto, Japan. The parasitoid fauna composed three koinobionts (all larval-pupal solitary parasitoids) and 22 idiodiont species (11 larval solitary, nine pupal solitary and one pupal gregarious). The parasitoid community was dominated by early-attacking oligophagous braconid koinobionts at early periods, but was gradually displaced by late-attacking polyphagous eulophid idiobionts. Accordingly, the diversity index of the parasitoid community peaked at an intermediate point in the intra-generational succession. The succeeding attack-in-waves by the late-attacking idiobionts greatly reduced not only the survival rates of early-attacking parasitoid larvae but also the survival rates of hosts. The density-dependence observed in the host pupal mortality was thought to result from density-dependent host-switching by a keystone polyphagous pupal idiobiont parasitoid, Chrysocharis pubens, whereas high host pupal mortality was potentially attained by an early-attacking koinobiont braconid. Supposed aggregation of polyphagous parasitoids at high host density resulted in intense within-host competition and in an increase of host-feeding attack, both of which contributed to low emergence rates of parasitoids at high host densities. Parasitoid emergence rates were also reduced at low host densities, probably by inter- and intra-specific hyperparasitism among oligophagous parasitoids for limited hosts. The regulation effects of the species-rich parasitoid community upon the host population dynamics are thought to derive from succeeding attack-in-waves by polyphagous late-attacking idiobionts, especially by the keystone species.     

The developmental strategies and related profitability of an idiobiont ectoparasitoid Sclerodermus pupariae vary with host size

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Behavioral manipulation of host caterpillars by the primary parasitoid wasp Cotesia glomerata (L.) to construct defensive webs against hyperparasitism

Many parasitoids control the behavior of their hosts to achieve more preferable conditions. Decreasing predation pressure is a main aim of host manipulation. Some parasitoids control host behavior to escape from their enemies, whereas others manipulate hosts into constructing defensive structures as barriers against hyperparasitism. Larvae of the parasitoid wasp Cotesia glomerata form cocoon clusters after egression from the parasitized host caterpillar of the butterfly Pieris brassicae. After the egression of parasitoids, the perforated host caterpillar lives for a short period and constructs a silk web that covers the cocoon cluster. We examined whether these silk webs protect C. glomerata cocoons against the hyperparasitoid wasp Trichomalopsis apanteroctena. In cocoon clusters that were not covered by silk webs (bare clusters), only cocoons hidden beneath others avoided hyperparasitism. In covered cocoon clusters, both cocoons hidden beneath others and those with a space between them and the silk web avoided hyperparasitism, whereas cocoons that contacted the silk webs were parasitized. The frequency of cocoons that were hidden beneath others increased with the increasing number of cocoons in a cluster, but the defensive effect of cluster size was thought to be lower than that of silk webs. However, the rate of hyperparasitism did not differ between covered and bare clusters when we allowed the hyperparasitoids to attack the cocoon clusters in an experimental arena. This result was thought to have been caused by low oviposition frequency by these hyperparasitoids. As a result, silk webs did not guard the cocoons from hyperparasitoids in our experiments, but would protect cocoons under high hyperparasitism pressure by forming a space through which the ovipositors could not reach the cocoons.     

Role of chemical cues and natal rearing effect on host recognition by the parasitic wasp Melittobia digitata

Parasitoids are expected to have the ability to find, recognize, and perhaps to discern potential hosts that can best support the development of their progeny. Melittobia Westwood (Hymenoptera: Eulophidae) are gregarious ectoparasitoids, which primarily attack mud daubers (Hymenoptera: Sphecidae). How Melittobia females locate their host is not well known, but the process may involve host‐related chemical signals. In this study, we investigated the roles of host chemical cues and natal rearing effect in host recognition by Mel. digitata Dahms. In an olfactometer that contained prepupae of Trypoxylon politum Say (Hymenoptera: Sphecidae), Megachile rotundata (Fabricius) (Hymenoptera: Megachilidae), puparia of Neobellieria bullata (Parker) (Diptera: Sarcophagidae), empty or intact host cocoons, or nest mud, Mel. digitata females spent significantly more time in air fields that contained T. politum (prepupae + cocoon) and Meg. rotundata (prepupae + cocoon) than in N. bullata and control fields. Nest mud and natal host had no attraction for parasitoid host choice. Most first and last choices of Mel. digitata females in the olfactometer were not consistent, suggesting an initial random dispersion, although they responded positively towards hosts in cocoons.     

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.     

Performance of Microplitis tuberculifer (Hymenoptera: Braconidae) parasitizing Mythimna separata (Lepidoptera: Noctuidae) in different larval instars

The solitary parasitoid Microplitis tuberculifer (Wesmael) is an important biological control agent of various lepidopteran pests in Asia. We examined the preference of M. tuberculifer for different instars of its common host, Mythimna separata (Walker), host instar effects on parasitoid development, and the consequences of parasitism in different stages for growth and consumption of host larvae. The wasp successfully parasitized the first four larval instars of M. separata, but not the fifth, which appeared to be behaviorally resistant. First and second instars were parasitized at higher rates compared to thirds and fourths in no-choice situations, ostensibly due to longer handling times for the latter, but second instars were most preferred in a choice test that presented all stages simultaneously. Although later instar hosts yielded heavier cocoons, the fastest parasitoid development was obtained in second instars. Lower sex ratios were obtained from first instars as females appeared to lay a smaller proportion of fertilized eggs in small hosts. Both weight gain and food consumption of parasitized larvae were reduced significantly within 24 h of parasitism, regardless of the stage parasitized, and final body weights were less than 10% those of unparasitized larvae. Thus, M. tuberculifer has good potential as a biological control agent of M. separata, successfully parasitizing the first four larval instars and dramatically reducing plant consumption by the host in all cases.     

Patterns of parasitism by Trybliographa rapae, a cynipid parasitoid of the cabbage root fly, under laboratory and field conditions

ABSTRACT.

• 1 The spatial patterns of parasitism of the cabbage root fly caused by the cynipid parasitoid Trybliographa rapae (Westw.) have been studied in a laboratory system, within field cages and in a natural situation.
• 2 Continuous observations during the laboratory experiments showed the parasitoids to spend proportionately more time on the patches of high host density. This resulted in the per cent parasitism per patch being directly density dependent.
• 3 Similar patterns of parasitism were found from the field cage system, and also from experiments using the natural parasitoid population and either manipulated or natural host densities.
• 4 While mutual interference was marked in the laboratory experiments, there was little or no sign of it within the larger field cages.

     

The effect of multi‐species host density on superparasitism and sex ratio in a gregarious parasitoid

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