Foraging behaviour and resource utilization of the aphid parasitoid Pauesia unilachni: adaptation to host distribution and mortality risks

Pauesia unilachni is a parasitoid of the grey pine aphid, Schizolachnus pineti which feeds on needles of Scots pine. P. unilachni females displayed a dual movement system when foraging within a pine. They left a colony usually by foot and searched nearby needles intensively, thereby ensuring a maximum probability to encounter neighbouring S. pineti colonies. If search by foot was not successful, parasitoids changed their behaviour and searched needles by flight to cover larger distances. This behaviour was consistent both on uninfested and S. pineti-infested trees in the laboratory and in the field. In total, females allocated a 15-fold amount of time into search by foot compared to search by flight, although the overall foraging success, i.e. the number of discovered host colonies, did not differ significantly between the two movement types. However, the reduction of searching time by flight involved also higher travel mortality risks. In the field, P. unilachni got frequently entangled in spider nets when searching by flight and were killed.     

Foraging behaviour at the fourth trophic level: a comparative study of host location in aphid hyperparasitoids

In studies of foraging behaviour in a multitrophic context, the fourth trophic level has generally been ignored. We used four aphid hyperparasitoid species: Dendrocerus carpenteri (Curtis) (Hymenoptera: Megaspilidae), Asaphes suspensus Walker (Hymenoptera: Pteromalidae), Alloxysta victrix (Westwood) (Hymenoptera: Alloxystidae) and Syrphophagus aphidivorus (Mayr) (Hymenoptera: Encyrtidae), to correlate their response to different cues with their ecological attributes such as host range and host stage. In addition, we compared our results with studies of primary parasitoids on the same plant–herbivore system. First, the olfactory response of females was tested in a Y‐tube olfactometer (single choice: plant, aphid, honeydew, parasitised aphid, aphid mummy, or virgin female parasitoid; dual choice: clean plant, plant with aphids, or plant–host complex). Second, their foraging behaviour was described on plants with different stimuli (honeydew, aphids, parasitised aphids, and aphid mummies). The results indicated that olfactory cues are probably not essential cues for hyperparasitoid females. In foraging behaviour on the plant, all species prolonged their total visit time and search time as compared to the control treatment (clean plant). Only A. victrix did not react to the honeydew. Oviposition in mummies prolonged the total visit time because of the long handling time, but the effect of this behaviour on search time could not be determined. No clear correlation between foraging behaviour and host stage or host range was found. In contrast to specialised primary aphid parasitoids that have strong fixed responses to specific kairomones and herbivore‐induced synomones, more generalist aphid hyperparasitoids seem to depend less on volatile olfactory stimuli, but show similarities with primary parasitoids in their use of contact cues while searching on a plant.     

Influence of host distribution on foraging behaviour in the hyperparasitoid wasp, Dendrocerus carpenteri

The foraging behaviour of Dendrocerus carpenteriCurtis (Hymenoptera: Megaspilidae), an ectophagous hyperparasitoid of aphidiine wasps inside mummified aphids, was examined in the laboratory with an experimental system consisting of broad bean, Vicia fabaL, the pea aphid, Acyrthosiphon pisumHarris, and a primary parasitoid, Ephedrus californicusBaker. Pea aphids parasitised by E. californicusoften disperse from their feeding sites (or off host plants) before dying and mummifying. Response of female hyperparasitoids to host distribution was evaluated at two spatial scales. At the first scale, behaviour of hyperparasitoids was examined on individual plants with different densities of hosts. At the second scale, habitat complexity and host location were manipulated in large foraging cages containing several plants. I show that patterns of density-dependent hyperparasitism can result from the foraging behaviour of D. carpenteri. However, dispersal of parasitised aphids may not reduce the incidence of hyperparasitism if hyperparasitoids systematically search the habitat.     

Biology and host selection behaviour of the aphid hyperparasitoid Alloxysta victrix in association with the primary parasitoid Aphidius colemani and the host aphid Myzus persicae

The biology of the aphid hyperparasitoid Alloxysta victrix was investigated with respect to its interaction with the primary parasitoid Aphidius colemani and the host aphid Myzus persicae. Laboratory investigations of host selection behaviour indicated that naive A. victrix females could rapidly distinguish between unparasitized aphids and those parasitized by A. colemani, but showed little tendency to discriminate between aphids parasitized by the primary parasitoid three, five, seven, or ten days earlier (even though the latter were already mummified). Host suitability studies, in contrast, indicated that aphids parasitized by A. colemani became progressively less suitable as the primary parasitoid developed. When aphids were exposed to A. victrix more than seven days after being parasitized by A. colemani, survivorship of both primary and secondary parasitoid species decreased, with a significantly higher proportion of mummies producing no viable adult parasitoid of either species. The developmental period (egg to adult eclosion) of surviving hyperparasitoids was also longer in older host stages, and females developed significantly more slowly than males (14.2 days versus 13.6 days, respectively). The sex ratio of hyperparasitoid broods emerging from all host stages was consistently male-biased (31–46% female), but adult longevity was higher for females than for males (37.1 days versus 23.6 days).     

A potent, morph-specific parturition stimulant in the overwintering host plant of the black bean aphid, Aphis fabae

Abstract. The black bean aphid, Aphis fabae Scopoli, has a host‐alternating life cycle. Winged female autumn migrants (gynoparae) develop on numerous summer host plants but as adults will only colonize the winter host (spindle, Euonymus europaeus L.). When stylet activities of gynoparae were electrically recorded during access to a spindle leaf, the insects spent the majority (75%) of the 6‐h experimental period penetrating the plant surface and a large proportion of it (44%) ingesting from either phloem or xylem vessels. Most (95%) gynoparae initiated reproduction on spindle, producing 4.15 ± 0.59 offspring per adult by the end of the experiment (mean ± SEM). By contrast, gynoparae placed on a seedling of their natal, summer host (broad bean, Vicia faba L.) penetrated the plant for only 39% of the available time, rarely ingested plant sap and never reproduced. The number of nymphs deposited on spindle leaves was not correlated with the occurrence or duration of ingestion from vascular tissues, suggesting that parturition stimulants are detected before feeding, probably during penetration of nonvascular cells. Presentation of an aqueous spindle extract to the aphids in artificial feeding chambers showed that water‐soluble spindle factors evoke stimulation of parturition by gynoparae in 72‐h bioassays. The stimulant was extremely potent, remaining active until the total extracted material was diluted to less than 10 p.p.m. Stylet activities and reproductive responses were also evaluated for summer winged females (alate virginoparae), which have a broad host range, and will colonize both bean and spindle under laboratory conditions. On both of these plant species, virginoparae often ingested plant sap and deposited nymphs during the 6‐h electrical recording experiment (producing 4.60 ± 0.48 offspring on bean; 2.70 ± 0.35 on spindle: mean ± SEM), but no significant correlations were found between reproduction and the occurrence or duration of particular stylet activities. Aqueous host‐plant extracts had no effect on the numbers of offspring deposited by virginoparae in artificial feeding chambers, showing that this form of the aphid is not responsive to the spindle‐derived parturition stimulant. The results highlight the need for more information on the factors determining host acceptance and parturition by polyphagous aphid phenotypes.     

Within-patch foraging behaviour of the aphid parasitoidAphidius funebris: plant architecture, host behaviour, and individual variation

The influence of plant architecture, host colony size, and host colony structure on the foraging behaviour of the aphid parasitoidAphidius funebris Mackauer (Hymenoptera: Aphidiidae) was investigated using a factorial experimental design. The factorial design involved releasing individual parasitoid females in aphid colonies consisting of either 10 or 20 individuals ofUroleucon jaceae L. (Homoptera: Aphididae) of either only larval instar L3 or a mixture of host instars, both on unmanipulated plants and on plants that had all leaves adjacent to the colony removed. Interactions between the parasitoid and its host were recorded until the parasitoid had left the plant. The time females spent on the host plant and the number of eggs laid varied greatly among females. Host colony size significantly affected patch residence time and the number of contacts between parasitoids and aphids. Plant architecture influenced the time-budget of the parasitoids which used leaves adjacent to the aphid colony for attacking aphids. Female oviposition rate was higher on unmanipulated plants than on manipulated plants. No further significant treatment effects on patch residence time, the number of contacts, attacks or ovipositions were found. Oviposition success ofA. funebris was influenced by instar-specific host behaviour. Several rules-of-thumb proposed by foraging theory did not account for parasitoid patch-leaving behaviour.     

Foraging behaviour and feeding success of the black-necked stork (Ephippiorhynchus asiaticus) in Dudwa National Park, Uttar Pradesh, India

Foraging behaviour of the black-necked stork (BNS) Ephippiorhynchus asiaticus was studied in 1996 and 1997 in Dudwa National Park (DNP), Uttar Pradesh, India. The storks were observed using tactile and visual techniques to catch fish. Of the 929 fish seen caught in 2 years, 894 (96%) were caught by a tactile mode of feeding and the remaining 35 (4%) by a visual mode of feeding. The rate of foraging attempts by BNS fluctuates with that of season in DNP and coincides with prey abundance. Immediately after the monsoon when the water level was high, BNS had to search for prey more often, as the prey became widespread. Whereas in summer when the water level decreased, the concentration of the fish was higher, which helps BNS to catch fish in quick succession. Prey behaviour and the condition of the wetland determined the selection of the tactile foraging technique among the BNS in DNP. The black-necked storks were more successful in the early hours of the day (06:00–10:00) and they were generally more successful or preferred to feed on medium-sized fish (i.e. 5–10 cm) in DNP. Prey profitability was highest for larger size fish and decreased as the prey size decreased. Principal component analysis showed that prey size, handling time of each prey, the month and water depth determined foraging success in 1996 and 1997.     

Effects of host age and cocoon position on attack rate by the hyperparasitoid,Eurytoma sp. [Hym.: Eurytomidae], on cocoons of the parasitoid,Cotesia (=Apanteles) glomerata [Hym.: Braconidae]

Host age preference of the hyperparasitoid,Eurytoma sp., attacking the cocoons of the gregarious parasitoid wasp,Cotesia (=Apanteles) glomerata L., was investigated in the field and laboratory. Under laboratory conditions of 20°C and L16D8 photoperiod.Eurytoma sp. parasitized cocoons of all ages, laying 7 to 10 eggs per cluster during a 24 h period. Field-collected cocoons also indicated that the host was parasitized regardless of its developmental stage. However, the mortality ofEurytoma sp. laid in cocoons on the day before host emergence was as high as 60%. Furthermore, progeny sex ratio (proportion males) reached 0.708 in eggs laid in the oldest cocoon clusters, whereas that for younger cocoons was strongly female-biased. Together, these facts suggest that older hosts are less suitable forEurytoma sp. than are younger ones, even though there was no significant decreasing tendency in the number of parasitized cocoons per cluster. In addition, the effect of cocoon position within a cluster was apparent, outer cocoons being more easily parasitized than inner ones. TheEurytoma sp. female oviposited at random on the free surface ofC. glomerata cocoons.     

The role of olfaction in aphid host location

Aphids are major economic pests of many of the worlds' crops, causing damage directly by feeding and by acting as vectors for plant viruses. By understanding how aphids locate their host plants, it may become possible to develop new means of controlling populations by taking advantage of these natural host location/nonhost avoidance behaviours. Aphids have also become important model organisms in the study of insect–plant interactions and an improved understanding of host location in aphids could yield insights into the behaviour and ecology of other insect orders. The use of olfaction by host‐seeking aphids is well documented and, in recent years, considerable information has been gained on how volatiles can encode host identity and suitability, as well as the specific behaviours they elicit from aphids. The purpose of this review is to highlight the major findings on how aphids respond behaviourally to volatile compounds and how they can use them to locate their host plants and avoid unsuitable hosts.     

Nocturnal activity and resource utilization in the aphid hyperparasitoid, Dendrocerus carpenteri

Abstract.

• 1 Females of the aphid hyperparasitoid Dendrocerus carpenteri (Curtis) search successfully for hosts during both day and night. Oviposition numbers per host patch did not differ significantly between day and night.
• 2 D.carpenteri females also displayed a nocturnal flight activity, showing that they are not only capable of searching on a given host plant but also of dispersing between host plants.
• 3 Nocturnal oviposition activity was mainly influenced by egg load. Females with a high egg load laid more eggs at night than females with a comparatively low egg load. Thus, D.carpenteri females may use nocturnal foraging to compensate for the lack of oviposition opportunities during day.
• 4 D.carpenteri females which foraged continuously for hosts both day and night (= for 24 h per day) benefitted from an 1.4-fold increase in lifetime reproductive success when compared to females which foraged only by day (= for 16 h per day).
• 5 The benefit of night foraging for this species is a significantly increased reproductive success.

     

Host selection and sex ratio in a heteronomous hyperparasitoid

Abstract.

• 1 Encarsia tricolor Förster is a heteronomous hyperparasitoid: females develop as primary endoparasitoids of Homoptera, whereas males develop hyperparasitically in primary endoparasitoids, including conspecific females.
• 2 When offered pupae of Encarsia inaron (Walker) or conspecific pupae, E.tricolor showed a distinct preference to exploit E.inaron for male production.
• 3 Parasitoids were given the opportunity to gain different types of ovipositional experience on male and female hosts and were then offered patches containing mixtures of male and female hosts in various ratios. The mean sex ratio which they laid was not dependent on the relative availability of each type of host, whereas prior ovipositional experience had a significant effect.
• 4 Adaptive reasons for the observed oviposition strategies are suggested.

     

Factors determining the host plant range of the phytophagous mite, Tetranychus urticae (Acari: Tetranychidae): a method for quantifying host plant acceptance

The host plant acceptance of the phytophagous mite Tetranychus urticae was experimentally quantified. Host plant acceptance is described as the proportion of adult females settling on the test plant on which they have been placed. On the other hand, the host plant suitability of T. urticae on different plant species is expressed as the mean number of eggs produced by the females within 5 days (hereafter 'fecundity'). An inbred T. urticae line was tested with regard to host plant acceptance and fecundity on 11 potential host plants. These two variables were positively correlated across host plants; host plant species on which the fecundity was low were also those on which females settled less readily compared to host plants with high fecundity. The characteristics of host plant acceptance of the T. urticae are discussed in light of their potential food resource under natural conditions.     

设施农业景观下破碎化麦田麦蚜及寄生蜂种群的最小适生面积

最小适生面积(MASH)指在一定的时空范围内物种能稳定存在的最小生境面积,它是种群生存力分析(PVA)的重要方法之一.本文采用基于种群数量-面积关系原理的MASH模型模拟了银川平原设施农业景观下破碎化麦田麦蚜、初寄生蜂与重寄生蜂种群发生的MASH.研究表明：密度 面积、增长速度-面积关系模型间存在反比例函数关系,不同物种存在的函数关系明显不同,尤其在不同营养级别的物种间,其函数关系差异更为明显.根据密度-面积关系,利用多项式回归模型计算了麦二叉蚜、麦长管蚜、燕麦蚜茧蜂、烟蚜茧蜂与蚜虫宽缘金小蜂的MASH,其营养级间的MASH差异显著.不同物种的MASH与营养级高低、体型大小、生境质量等有关.初寄生蜂最高的寄生率出现在800～1000 m²,可作为利用初寄生蜂自然控制麦蚜的依据,而不同营养级物种MASH差异可用于害虫的种群控制.     

Foraging behaviour and prey discrimination in the bluespotted maskray Dasyatis kuhlii

A study observing the foraging behaviours and prey discrimination of a common demersal stingray, the bluespotted maskray Dasyatis kuhlii was performed under controlled laboratory conditions. A selection of prey species and masses were offered at depths of 10 and 50 mm in sand. Foraging efficiency and prey selection at both burial depths were compared. Dasyatis kuhlii selected the ghost shrimps, Trypaea australiensis and T. australiensis >2·5 g, range ± 0·2 g though foraging errors represented by prey being excavated and not consumed suggested a limited discriminatory ability at the point of detection. Burial depth did not influence prey species, mass selection or discriminatory ability.     

Temperature influences host instar selection in an aphid parasitoid: support for the relative fitness rule

The relative fitness rule states that parasitoid females should adopt risk‐prone reproductive behaviours when expecting low reproductive success. Temperature influences the reproductive success of insects by affecting their basal metabolic rate during development, their egg load at emergence, and their life expectancy as adults. Using an aphid–parasitoid model system, we investigated the influence of developmental and adult temperature on the risk‐sensitive decision‐making of females. We considered the use of a low‐quality host nymphal instar by the aphid parasitoid Aphidius ervi to be a risk‐prone behaviour. Immature females were reared at 12, 20 or 28 °C and had access to the four nymphal instars of the potato aphid Macrosiphum euphorbiae for oviposition at one of these temperatures. Host selection behaviour was continuously recorded during exploitation of an aphid patch. We observed that warm‐developed females and parasitoids foraging at high temperature attacked low‐quality hosts more frequently than females from other treatments. These results support the hypothesis that a decrease in expected parasitoid reproductive success resulted in risk‐prone behaviours. To our knowledge, this is the first experimental evidence suggesting that temperature influences host stage selection and risk‐sensitive making decision in parasitoids, and the present study is the first to support the relative fitness rule. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.     

Foraging behaviour of the parasitoid Lysiphlebus testaceipes (Hymenoptera: Braconidae) in response to plant volatiles,with reference to biocontrol of aphids in peri-urban vegetable production systems

Foraging behaviour of Lysiphlebus testaceipes was studied in a Y-tube olfactometer. Naïve females responded to the Aphis gossypii/eggplant complex on which they had been reared, but did not show a preference for this aphid/plant complex over an uninfested eggplant, an eggplant that had been damaged by the aphid or an A. gossypii/okra complex. Females that had been allowed to forage on the A. gossypii/eggplant complex before being tested in the olfactometer showed a strong preference for an uninfested eggplant over an uninfested okra plant and for the A. gossypii/eggplant complex over an A. gossypii/okra complex, indicating that they had learned to associate eggplant-derived volatiles with host availability. When females that had been reared on the A. gossypii/eggplant complex were tested after foraging experience on the A. gossypii/okra complex, they did not show a preference for either complex, suggesting that they still retained a memory of the cues associated with the complex on which they had been reared. The efficiency of L. testaceipes as a biological control agent in peri-urban vegetable production, such as the Cuban organoponics system, potentially could be improved by producing and releasing females behaviourally primed to prefer specific host/plant complexes.     

Plant water stress intensity mediates aphid host choice and feeding behaviour

1. The effects of drought-induced changes in plant quality on aphid performance and population growth is well-studied. The response of aphid behaviour to plant water limitation has received less attention. Water limitation may affect host-plant colonization by altering the attractiveness of plants. Additionally, plant water limitation may inhibit feeding site establishment and phloem ingestion.
2. Our goal was to examine bird cherry-oat aphid (Rhopalosiphum padi L.) host selection and feeding behaviour under water limitation. We assessed aphid response to well-watered, mildly-stressed, and highly-stressed wheat (Triticum aestivum L.) by evaluating (i) host-plant selection through two-choice assays, (ii) feeding behaviour using the electrical penetration graph technique, and (iii) phloem ingestion by quantifying honeydew production.
3. Aphids were less likely to select highly stressed plants than a mildly stressed or well-watered alternative. Aphids did not distinguish between mildly stressed and well-watered plants. Aphid feeding behaviours, including duration of phloem ingestion, were not affected by water availability. However, honeydew production was reduced under both levels of water limitation. These results suggest that the volume of phloem ingested by aphids per unit time declined on stressed plants. The combination of lower colonization and diminished access to food on stressed plants may lead to a reduction in aphid abundance, independent of the direct effects of nutrition on individual aphid performance.
4. This study highlights the potential contribution of herbivore behaviour to documented changes in aphid abundance on stressed plants and underscores the important role of plant water stress intensity in mediating plant-herbivore interactions.

     

栗喉蜂虎捕食差异

2010年4～5月和2011年4～6月,用焦点动物取样法对云南省保山市潞江镇怒江河谷栗喉蜂虎捕食差异进行了观察。对栗喉蜂虎雌性成年鸟、雄性成年鸟、青年鸟捕食种类、数量、成功率、食物组成、食物资源量、处理食物成功率及处理时间进行了统计。结果表明,栗喉蜂虎捕食种类包括7类昆虫,以膜翅目Hymenoptera、蜻蜓目Odonate、鳞翅目Lepidoptera昆虫为主;成年雄鸟捕食频次高于成年雌鸟和青年鸟;成年鸟与青年鸟处理食物成功率无显著差异,但二者捕食成功率和处理食物时间差异显著。