Host-feeding and oviposition strategies of parasitoids in relation to host stage

Until now, mathematical models of parasitoid-host interactions have not incorporated the tendency for destructively host-feeding parasitoids to partition their feeding and oviposition behaviour in relation to different host stages. A literature survey reveals a trend for female parasitoids to feed preferentially or exclusively on earlier host stages and to oviposit preferentially or exclusively in/or later ones. We explore the relative advantages to host-feeding parasitoids of a number of possible host stage selection strategies. We develop hypotheses, formalizing and testing them using modifications to our earlier simulation model of host-feeding strategies (Jervis and Kidd, 1986). We conclude from our modelling that the advantage to be gained from feeding on early host stages and ovipositing in late ones is likely to be associated with: 1) reduced handling times when feeding on early stage hosts; 2) reduced wastage of progeny from mortality factors other than host-feeding by the parent parasitoid, achieved by confining oviposition to late host stages; and 3) reduced probability of progeny mortality resulting from the parent's host-feeding activities.     

Host-feeding and acceptance by a parasitic wasp (Hymenoptera: Ichneumonidae) as influenced by egg load and experience in a patch

Optimal host selection models based on dynamic programming predict that the physiological state of a foraging insect, i.e. egg load, energy reserves etc., influences behavioral decisions. To test this prediction, the effect of physiological state on host acceptance of the ectoparasitic wasp Agrothereutes lanceolatus was investigated. Female wasps in plastic cups (regarded as patches) were presented with hosts, and their responses to the hosts were continuously observed. After observations, the wasps were dissected and the number of mature and immature eggs they carried were counted. The results showed that behavioral decisions by the female wasps were influenced by mature egg load, but not by wasp size or immature egg load. Hence the wasps with higher egg loads were more likely to oviposit. The number of hosts previously encountered in a patch (i.e. wasp experience) also had an independent effect on females' host acceptance, indicating that female informational state was updated during foraging in that patch. Female wasps host-fed only when mature egg load approached zero. Concurrent host-feeding was not observed. Parasitoid survival was almost zero when parasitoid eggs were transferred onto hosts that were fed upon, indicating that concurrent host-feeding could cause a high degree of offspring mortality. These three results supported the assumption and prediction of optimal host-feeding models. Parasitoid host selection and host-feeding are discussed in the context of recent models.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

Host-feeding and oviposition by parasitoids in relation to host stage: Consequences for parasitoid-host population dynamics

Among parasitoids which host-feed destructively, there is a tendency for females to partition their feeding and oviposition behaviour in relation to different host stages, feeding preferentially or exclusively on earlier host stages and ovipositing preferentially or exclusively in (or on) later ones. We explored the dynamic implications of this behaviour for parasitoid-host population dynamics, using modifications of the age-structured simulation models of Kidd and Jervis (1989, 1991). Using the new versions of the models, we compared the situation where parasitoids practice host stage discrimination with respect to feeding and oviposition, with the situation where they do not. Additionally, we examined the effects of host stage discrimination on populations by (a) having generations either discrete or overlapping, (b) varying initial age structure, (c) having varying degrees of density dependence acting on host adult mortality, and (d) varying parasitoid develoment times in relation to the length of host development. With either discrete or overlapping generations of the host population, a reduction in the parasitoid development time had a destabilizing influence on the parasitoid-host population interaction. With discrete generations stage discrimination had no effect on the risk of extinction, irrespective of either the degree of density dependence acting on the host population, or the initial age structure of the host population. When parasitoid search was uncoupled from the insect's adult energy requirements, the interaction was always unstable. With continuous generations, stage discrimination affected stability at certain parasitoid development times, but not at others. The relative lengths of parasitoid and host development times also influenced the tendency of the host population to show discrete or overlapping generations.     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non‐significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

Effect of host diet and adult parasitoid diet on egg load dynamics and egg size of braconid parasitoids attacking Anastrepha ludens

The quantity and quality of host nutrients can affect fitness‐related traits in hymenopteran parasitoids, including oogenesis. The present study tested the prediction that a high host quality will influence oogenesis‐related traits positively in synovigenic parasitoids, and that a high‐quality adult parasitoid diet can positively affect the same parameters, potentially compensating for development on low‐quality hosts. Four braconid parasitoid species with contrasting life histories are reared on a low‐quality diet [Anastrepha ludens Loew (Diptera: Tephritidae) larvae reared on mango] or a high‐quality (artificial) diet. Adult parasitoids are provided with a high‐quality (honey ad libitum), moderate‐quality (honey every other day) or low‐quality (guava pulp) diet. Generalist species that encounter high variation in host quality naturally are predicted to be more flexible in dealing with nutrient shortfalls than specialist species. By contrast to the predictions, low‐quality hosts yield parasitoids with higher egg loads in two species: Opius hirtus Fisher and Diachasmimorpha longicaudata Ashmead. However, as predicted, a high‐quality adult diet exerts a positive effect on egg load (Utetes anastrephae Viereck), egg size (Doryctobracon crawfordi Viereck) and egg maturation rate (D. longicaudata, O. hirtus and U. anastrephae). The generalist D. longicaudata varies in egg load and maturation rate depending on host quality and adult diet, respectively. Evidence of the combined effect of both factors on parasitoid fertility is presented for the specialist O. hirtus. The theoretical and practical implications of these findings are discussed.     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non-significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

卵表化合物对桔小实蝇产卵行为的影响

多种实蝇科害虫产卵后可释放寄主标记信息素来表明寄主果实已经被占据,从而调节自身或同种个体间的对寄主果实的产卵选择行为。为探明桔小实蝇卵表化合物的寄主标记作用,本文测定了桔小实蝇卵表甲醇提取物对桔小实蝇的产卵驱避活性,并观察了甲醇提取物处理对桔小实蝇产卵行为过程的影响。结果表明：桔小实蝇卵表甲醇提取物对桔小实蝇有明显的产卵驱避活性,且活性随浓度的增加而增强,当提取物浓度为100 mg/mL（虫卵与溶剂质量体积比,m/v）时,在选择性条件下的产卵驱避率为55-45%;非选择性条件下的产卵驱避率达到64%-79%。经卵表提取物处理后,桔小实蝇在寄主果实上的访问次数、试探产卵时间、试探产卵次数、实际产卵次数和产卵量均显著少于对照。证实了桔小实蝇卵表化合物具有明显的产卵寄主标记作用。     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

Influence of egg load and host size on host-feeding behaviour of the parasitoid Aphytis lingnanensis

Abstract.

• 1 Direct behavioural assays were used to investigate the influences of host size and parasitoid egg load on the decision to host feed versus oviposit made by the parasitoid Aphytis lingnanensis Compere. Egg load was manipulated without concurrent influences on the history of host contact by exploiting size-related variation in fecundity and by holding parasitoids at different temperatures to vary the rate of oocyte maturation.
• 2 Host feeding comprised a series of feeding bouts, separated by renewed probing of the scale insect body. Successive feeding bouts were progressively shorter, suggesting that hosts represent ‘patches’ yielding resources at a decelerating rate.
• 3 Parasitoids were significantly more likely to host feed on smaller hosts and oviposit on larger hosts.
• 4 Neither egg load nor the treatment variables (parasitoid size and holding temperature) exerted significant influences on the decision to host feed versus oviposit on second instar (low quality) hosts.
• 5 The failure to observe an effect of egg load on host-feeding decisions was not simply a reflection of the parasitoids being entirely insensitive to egg load; significant effects of egg load on parasitoid search intensity and clutch size decisions were observed.
• 6 Parasitoids developing on second instar (low quality) hosts experienced high levels of mortality late during development and yielded very small adults.
• 7 The discord between these experimental results and predictions regarding the importance of egg load underscores the need for additional work on the proximate basis for host-feeding decisions and the nutritional ecology of insect parasitoids.

     

Field oviposition rates and egg load dynamics of pollen beetles (Meligethes aeneus Fab.) (Colepotera: Nitidulidae)

Abstract 1 Egg loads from field collected pollen beetles (Meligethes aeneus Fab., Coleoptera: Nitidulidae) were determined by dissecting beetles caught at the beginning and end of the putative daily oviposition period. Field collected beetles were offered Brassica napus (L.) plants in cages for 8 (morning and early afternoon), 16 (overnight), and 24 h to ascertain the number of eggs laid during these time periods. 2 Most eggs were laid in the morning and early afternoon. The proportion of gravid females was higher at the beginning of the oviposition period than at the end. Most females in the morning carried two eggs, whereas one egg was more common in the afternoon. 3 We hypothesized that the number of eggs laid during the oviposition period would be equivalent to the difference between egg loads at the beginning and end of oviposition. This was not the case; differences in egg loads were significantly lower than number of eggs laid. However, the number of eggs laid was equivalent to the egg load at the beginning of the oviposition period, suggesting that eggs available in the morning are laid during the following day. 4 Population estimates of daily oviposition rates, approximately 0.7 eggs per beetle and day, were close to estimates from laboratory studies when the proportion of gravid females was taken into consideration.     

Flexible oviposition behavior in the golden egg bug (Phyllomorpha laciniata) and its implications for offspring survival

Female golden egg bugs show unique flexibility in their oviposition behavior. Females can lay eggs on plants, where they are leftunattended, or on the back of conspecifics, where they remainuntil hatching. In this article we show that eggs have greatersurvival rates when carried by an adult than when laid on plants.The main causes of egg mortality are predators and a parasitoidwasp. Our results suggest that, although predation pressureis similar, fewer eggs are attacked by parasitoid wasps whencarried by an adult than when laid on plants. In addition,we show that, when given a choice, females prefer to lay eggson the backs of conspecifics than on plants. Thus, female ovipositionchoice is adaptive and minimizes individual offspring mortality.The factors that may maintain such behavioral variation in current populations are discussed.     

Effects of egg load on finding and acceptance of host fruit in Ceratitis capitata flies

Abstract. In studies conducted on potted host trees in field cages and in the laboratory, we examined the influence of egg load on the finding and acceptance of high-ranking (kumquat) and lower-ranking (grapefruit) hosts for oviposition by wild-origin Mediterranean fruit fly females, Ceratitis capitata (Wiedemann). By prescribing the periods during which females had access to protein prior to testing, we generated four classes of females having progressively increasing egg loads but not differing in population origin, age, degree of protein hunger at testing, or amount of prior experience with host fruit (none). Egg load had no discernible effect on behaviour associated with finding either type of fruit but did have a significant effect on several behaviours associated with oviposition after alighting on fruit. Increasing egg load led to increasing propensity to engage in ovipositional-type behaviour on both kumquats and grapefruits. There was no evidence, however, to support a hypothesis that medflies would become less discriminating against grapefruits relative to kumquats as egg load increased. Relative to kumquats, grapefruits were accepted for oviposition by intermediate and high egg load females to a substantially greater degree in laboratory cages than on trees, suggesting that results of laboratory cage experiments on host discrimination by tephritid flies may poorly reflect differences in behavioural responses expressed under less constrained conditions.     

Influence of host deprivation on egg load and oviposition behaviour of Brachymeria intermedia, a parasitoid of gypsy moth

Abstract. Mated female Brachymeria intermedia (Hymenoptera: Chalcididae) deprived, since emergence, from pupae of their host Lymantria dispar (Lepidoptera: Lymantriidae), accumulated eggs but had a very low rate of hostacceptance. Parasitoids that were host-deprived after encountering pupae early in life also accumulated eggs, but maintained a high acceptance rate. Thus early exposure to hosts promoted active reproductive behaviour. Total egg production depended on the total number of pupae encountered, indicating that B.intermedia adjust their egg production to host availability. Hence, in B. intermedia both the physiological state of the parasitoid (age and egg load) and the informational state (in this case host-availability and experience) interact to shape oviposition behaviour.     

寄生蜂取食寄主特性及其在害虫生物防治中的作用

许多寄生性天敌昆虫的雌虫不仅寄生寄主, 而且还能取食寄主。在卵育型（synovigenic）寄生蜂类群中, 取食寄主行为是较为普遍的现象。本文综合近20年相关研究进展, 从寄生蜂类群、取食类型、生态学意义及影响因子等方面对寄生蜂的取食寄主行为进行了归纳总结。寄生蜂通过取食不仅可以杀死寄主, 直接起到控制害虫种群数量的作用, 还能通过取食策略为卵的成熟和再生提供营养来源, 对延长雌虫的寿命也有一定的帮助。对取食寄主行为的了解可为筛选优势寄生性天敌种类、评估寄生蜂在害虫生物防治中的作用提供重要信息。     

Trichostrongylus colubriformis: effect of anthelmintics on ingestion and oviposition

The effects of the anthelmintics ivermectin, levaminsole, oxfendazole, piperazine citrate, pyrantel pamoate, tetramisole, and thiabendazole on ingestion and oviposition by Trichostrongylus colubriformis were determined. Six of the compounds reduced in vitro feeding at the tested doses while all drugs reduced in vivo feeding after treatment of the host. Additionally, in vitro or in vivo exposure to most anthelmintics decreased oviposition during subsequent in vitro assay. Invermectin had the most pronounced effect on in vivo and in vitro feeding and egg release. The neuromuscular activities of pharyngeal pumping and egg ejection may be suitable systems for rapid determinations of anthelmintic effects.     

豚草生育期和被利用状态对广聚萤叶甲产卵选择的影响

【背景】植食性昆虫对产卵场所的选择行为受多种因素的影响,其中寄主植物的发育阶段和被同种其他个体利用程度是影响其产卵选择的重要因素之一。【方法】采用野外罩笼的选择性试验,观察了广聚萤叶甲成虫对不同生长发育阶段、不同被取食程度、不同落卵量的豚草植株的产卵选择行为,分析产卵选择频次,用Cox比例风险模型拟合识别期以估计产卵选择偏好性。【结果】开花期豚草被选择产卵的风险(瞬间概率)仅为生长期的16.5%,苗期豚草被选择产卵的风险为生长期的34.1%;成虫对3种被食程度豚草的产卵选择频次之间存在显著差异(n=20,P<0.01),在3级被食程度豚草上产卵次数最多(12次),而在5级被食豚草上产卵次数最少(2次),表明成虫明显偏好将卵产于已被中等程度取食的植株上;成虫对不同落卵量植株(0、4、8个.株-1)未表现出显著的产卵偏好,对不同密度(高、低)斑块的豚草的产卵选择亦无显著差异。【结论与意义】豚草生育期和被取食程度影响广聚萤叶甲雌虫的产卵选择,而植株上的落卵量和豚草密度对产卵选择没有影响。     

The effect of egg load and host deprivation on oviposition behaviour in aphidophagous hoverflies

1. Two species of aphidophagous hoverfly, Episyrphus balteatus and Syrphus ribesii, were tested for the effects of egg load and host deprivation on oviposition choices. 2. Egg load affected the total number of eggs laid in E. balteatus but not in S. ribesii, however it did not affect the proportion laid on any one aphid in E. balteatus but did affect the proportion laid on any one aphid in S. ribesii. The rank order of preferences remained unchanged by age or host deprivation. 3. The dominant effects on host choices were aphid species (in both syrphids) and presentation order (in E. balteatus). 4. Being deprived of hosts increased egg load substantially in E. balteatus, and increasing time of deprivation also had an effect on discrimination; there was no effect of host deprivation in S. ribesii. 5. Reasons for these patterns are discussed.     

Changes in egg colour, egg weight and oviposition rate with the number of eggs laid by wild females of the small heath butterfly, Coenonympha pamphilus

ABSTRACT. 1. The colour of eggs laid by individual Coenonympha pamphilus (L.) (Lep., Satyridae) females changed over their lifespan. The first laid eggs were green, but after having laid about 100 eggs, females laid only yellow eggs.
2. By following females in the field, or by capturing wild females, and noting the colour of their eggs, it was established that younger females showed higher oviposition rates and laid heavier eggs than older females.
3. Hence, wild C.pamphilus females are unable to sustain a constant egg production by extracting amino acids or any other nutrients from nectar. We hypothesize that this is a general phenomenon in nectar feeding butterflies.
4. The egg colour did not seem to match the colour of the substrate on which the female chose to deposit the egg.     

Field studies on the behaviour of two egg parasitoids of the pine processionary mothThaumetopoea pityocampa

An investigation on the behaviour ofTetrastichus servadeii Domenichini (Hym.: Eulophidae) andOoencyrtus pityocampae Mercet (Hym.: Encyrtidae) was carried out in 2 stands ofPinus nigra Arnold in the Venitian Prealps (NE Italy) by quantitative observations in the field and by trapping with different baits. The results indicate that: 1)O.pityocampae is caught by traps baited with pityolure, the synthetic sex pheromone of the host, whileT. servadeii was never caught. 2) The host selection process ofT. servadeii seems to be associated with the host plant ofThaumetopoea pityocampa (Denis & Schiffermüller) and with the aggregative behaviour of the parasitoid females. 3) The frequency distribution ofT. servadeii on the egg masses follows the negative binomial distribution; the degree of aggregation decreases with the egg mass age. These results may be taken into account in the utilization of egg parasitoids for the biological control ofTh. pityocampa.