Searching at different spatial scales: the foraging behaviour of the aphid parasitoid Aphidius rosae in rose bushes

The searching patterns of the aphid parasitoid Aphidius rosae were analysed at three different spatial scales: leaves, shoots and bushes. Parasitoid females searched aphid-infested leaves and shoots intensively and remained on average more than twice as long on infested than on uninfested shoots. Patch times and oviposition numbers per shoot were highly variable both between females and for different shoot visits within females. However, at the shoot and bush level low oviposition numbers were generally found. The time spent on different behavioural patterns (searching, resting, feeding, host handling) changed significantly during subsequent shoot visits of individual females but oviposition success was not influenced by this change. Parasitoids searched individual leaves and shoots mainly by walking, while moving between shoots occurred exclusively by flight. The travel time between shoots (i.e. flight time) accounted for less than 1% of the residence time in a bush. At the bush level foraging was characterized by a high ability to localize infested shoots and consequently little time was wasted in searching on uninfested shoots. The pattern of resource exploitation of individual females was consistent with the distribution of A. rosae larvae in field samples taken from individual rose bushes.     

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.     

Lysiphlebus testaceipes,an introduced aphid parasitoid in portugal [Hym.: Aphidiidae]

Lysiphlebus testaceipes (Cresson), an exotic parasitoid species introduced in 1973–74 into southern France and subsequently widespread over the whole mediterranean France, Italy and Spain has become established even in Portugal since 1985. It was found on the southern coast of Portugal parasitizingAphis nerii Boyer de Fonscolombe. In 1986 it spread towards the North, its host range expanded and now includesAphis chloris Koch,Aphis citricola van der Goot,Aphis fabae Scopoli,Aphis frangulae Kaltenbach,Aphis gossypii Glover,A. nerii, Aphis parietariae Theobald,Aphis sedi Kaltenbach,Toxoptera aurantii (Boyer de Fonscolombe). The proved complete development and a high parasitization ofL. testaceipes onA. citricola is the 1st case in the whole Mediterranean, from where only incomplete development and a negligible percentage of parasitization (Italy) have been reported. Both the host range composition and development ofL. testaceipes indicate an increase of genetic polymorphism in the established populations. Interspecific relations ofL. testaceipes and the indigenous parasitoid species indicate an increasing dominance of the exotic biocontrol agent.      

Winter ecology of the cabbage aphid Brevicoryne brassicae (L.) (Homo., Aphididae) and its parasitoid Diaeretiella rapae (McIntosh) (Hym., Braconidae: Aphidiidae)

Abstract:  Flower strips near crops may stimulate natural enemies by the provision of nectar and hibernation sites. However, these habitats may also be beneficial for potential pest species. We investigated the dynamics of the cabbage aphid Brevicoryne brassicae (L.) (Homo., Aphididae) and its primary parasitoid Diaeretiella rapae (McIntosh) (Hym., Braconidae) in brussels sprout fields and adjoining flower plots in winter. A wide variety of 14 plant species were included in the study, each established as monoculture plots. Brussels sprout fields and flower plots were established at two sites. One site was located in an open agricultural landscape, the other in a landscape dominated by mixed forest. Brevicoryne brassicae and D. rapae were found on brussels sprout plants but not in the flower plots. Brevicoryne brassicae was initially more abundant in the open landscape, but as their densities declined rapidly in time, no living aphids were recovered at both sites by February. The density of aphids parasitized by D. rapae showed a similar trend, but densities of eight mummies per brussels sprout plant were still present by the end of February. These findings suggest that (i) flower species under investigation do not function as sources of B. brassicae and (ii) brussels sprout plants that are not harvested may not only harbour D. rapae populations that may sustain biological control, but are also likely to act as sources of B. brassicae infestation.     

Semiochemicals associated to spacing behaviour of the bird cherry-oat aphid Rhopalosiphum padi L. (Hem., Aphididae) do not affect the olfactometric behaviour of the cereal aphid parasitoid Aphidius rhopalosiphi De Stephani-Pérez (Hym., Braconidae)

Volatiles produced by the interaction of Rhopalosiphum padi and wheat, and semiochemicals which elicit the spacing behaviour of R. padi on wheat and oat, were evaluated in an olfactometer against the cereal aphid parasitoid Aphidius rhopalosiphi . The parasitoid was attracted by volatiles produced by the R. padi -wheat interaction at high density of aphids, but not by the R. padi -wheat interaction at low density. Compounds produced by the interaction of wheat or oat that was infested with a high density of aphids (approximately 9 aphids/cm²), 6-methyl-5-hepten-2-one (MHO), 6-methyl-5-hepten-2-ol (MHOH), 2-tridecanone (2-T), and methyl salicylate (MS), neither attracted nor repelled A. rhopalosiphi as pure compounds or in the naturally occurring mixture. The results are discussed in terms of the use of semiochemicals and parasitoids in the integrated management of aphid pests.     

Effects of wheat foliar fungicides on the aphid endoparasitoid Aphidius rhopalosiphi DeStefani-Perez (Hym., Aphidiidae) on glass plates and on plants

Abstract: The side-effects of several fungicides used in wheat to control disease at heading growth stage were assessed on the aphid parasitoid Aphidius rhopalosiphi by tests conducted in the laboratory on glass plates and in the greenhouse on young wheat plants. Very few formulations containing only one active ingredient (carbendazim, cyproconazole or epoxyconazole) or combinations of two (carbendazim + cyproconazole, carbendazim + hexaconazole) were harmless to A. rhopalosiphi in the glass-plate tests. There was no apparent synergism between fungicides tested in combinations. The parasitoid mortalities in tests carried out on plants were less and chlorothalonil, epoxyconazole, fenpropidin, fenpropimorph, flusilazole, flutriafol, prochloraz, tebuconazole, tridemorph and a number of combinations (carbendazim + flutriafol, chlorothalonil + cyproconazole, epoxyconazole + tridemorph, chlorothalonil + hexaconazole, chlorothalonil + flutriafol, cyproconazole + prochloraz, epoxyconazole + fenpropimorph, fenpropimorph + propiconazole, propiconazole + tridemorph, triadimenol + tridemorph) were harmless or only slightly harmful to the aphid parasitoid. Several combinations (carbendazim + epoxyconazole, carbendazim + fenpropimorph, carbendazim + flusilazole, carbendazim + tebuconazole, chlorothalonil + fenpropimorph, chlorothalonil + flusilazole, fenpropimorph + fenpropidin, fenpropimorph + prochloraz, fenpropidin + propiconazole, fenpropidin + tebuco nazole, tebuconazole + triadimenol) were toxic for wasps on plants. The parasitoid mortalities were less on plants than on glass plates but the wasps spent less time on treated leaves and in some cases parasitism of aphids was reduced to a large extent. These results suggest that in addition to study of the direct effects of pesticides on beneficial insects (mortalities, reduction of fertility) their effects on the behaviour of the insects should also be studied. Products that induced a repellent effect need further testing in field or semi-field conditions. However, many fungicide combinations that have little or no effect on A. rhopalosiphi can protect wheat against a wide range of diseases and the results obtained in this study indicate that an appropriate and effective protection of wheat at earing growth stage can be achieved with products that have no effects on aphid parasitoids.     

Apple sawfly Hoplocampa testudinea (Hym., Tenthredinidae) and its parasitoid Lathrolestes ensator in Dutch apple orchards (Hym., Ichneumonidae, Ctenopelmatinae)

Abstract: Changes in population densities of the apple sawfly Hoplocampa testudinea (Klug) and its parasitoid Lathrolestes ensator (Brauns) were monitored in 15 apple orchards for a period of up to 4 years. The parasitoid species was found in all orchards except one, and was more numerous in plantings on sandier soils. Post‐bloom insecticide applications against other pests, and carbaryl for fruit thinning, often decimated both host and parasitoid in integrated pest management orchards. In organic orchards, where synthetic pesticides are banned, and Quassia is the only remedy against sawfly, the pest is more problematic. The low levels of parasitism in all organic orchards, except one, were possibly due to the application of wettable sulphur during the parasitoid flight period. The sawfly usually recovers more quickly than its parasitoid when chemical control is discontinued, because propagation of L. ensator is limited in various ways. The parasitoid is time limited, because suitable second instar host larvae are rarely available for more than a week on a single apple variety. Moreover, parasitism levels stay moderate because the parasitoid females do not avoid superparasitism. Finally, relatively more sawflies than parasitoids emerge after 10 months underground, because the incidence of prolonged diapause is more elevated in L. ensator than in the sawfly. It is suggested that both the high incidence of prolonged diapause and the inability to avoid superparasitism are useful in reducing the risk of local extinction. Elevated sawfly attack in a single early apple variety would reduce exploitation of suitable host larvae in other nearby varieties, in as far as the parasitoid is not able to distinguish fruitlets with accessible second instar host larvae from those with inaccessible older larvae. Although the former are available for a limited time, the latter may keep the parasitoids from moving to the less abundant second instar larvae in late(r) varieties. Reduction of elevated host density in such an early variety by a properly timed application of a short‐lived pesticide, such as Quassia, increases parasitism levels proportionally, and is expected to promote parasitoid movement to host larvae in other varieties nearby.     

Environmental and physiological factors for diapause induction and termination in the aphid parasitoid,Aphidius ervi (Hymenoptera: Aphidiidae)

In the aphid parasitoid Aphidius ervi Haliday, the incidence of diapause around the critical photoperiod was dependent on temperature. At a photoperiod of 12 h light:12 h dark, a significantly higher proportion of diapausing individuals was found at 12 degrees C than at 15 degrees C. The second larval stage of the parasitoid was most sensitive to photoperiod and localised-illumination experiments showed that the parasitoid responded independently of the aphid host. The sex ratio of the population which emerged after rearing in short-days and/or after diapause was male biased. Topical application of the juvenile hormone-mimic, pyriproxyfen, before or during diapause delayed emergence of Aphidius ervi but did not induce diapause or influence its termination. The ecdysteroid mimic RH 5849 induced diapause in a few treated individuals but did not affect diapause termination.     

Impact of the parasitoid Lathrolestes ensator (Hym., Ichneumonidae, Ctenopelmatinae) as antagonist of apple sawfly Hoplocampa testudinea (Hym., Tenthredinidae)

Abstract: Lathrolestes ensator (Brauns) was the only parasitoid species reared from larvae of apple sawfly Hoplocampa testudinea (Klug) collected in Dutch orchards. The life history parameters of apple sawfly and its parasitoid L. ensator were investigated in field and semi‐field experiments. Usually, the adult parasitoids emerge in synchrony with the preferred host stage, the second instar sawfly larvae. Parasitoid females carried 120–175 eggs, but never laid more than half this number. The mortality of sawfly eggs and larvae varied from 75 and 99%. The eggs and mining larvae are most vulnerable, as the older larvae survive for 90% on average. Three factors at least determine how many of the descending larvae become adult sawfly or parasitoid next spring. The failure of descending larvae to construct a cocoon varied from 7 to 31% and was highest in clay soil. Probably a similar mechanical hindrance prohibited more emerging adults, of both sawfly and L. ensator, from attaining the surface in heavier soils next spring. Relatively more sawfly prepupae than parasitoids died in the cocoon stage, from fungus disease or other causes, but more parasitoids than hosts stayed in prolonged diapause. Mainly due to this last factor, the overall result was a decrease of the parasitoid : host ratio during the first season underground. A life table based on survival rates during various life stages yields a net reproductive rate of up to 2.4 daughters per female sawfly. It indicates that 60% of the sawfly larvae need to be parasitized to stop population growth, or correspondingly less when more than 75% of the host eggs and young larvae are killed by predators or other causes.     

Low field performance of an aphid parasitoid,Aphelinus abdominalis,efficient in the laboratory [Hym., Aphelinidae]

Life history traits of two inbred lines of the aphid parasitoidAphelinus abdominalis (Dalman) were evaluated in the laboratory. Both lines exhibited high fecundities, produced a high proportion of female offspring and showed to be long-lived. However, release experiments in winter wheat fields failed to achieve establishment of the parasitoids. In addition, the wasps parasitized considerably fewer aphids in two types of field cages than they did in the laboratory. AsA. abdominalis eggs and/or larvae arrested in development were not found in dissected aphids from field cages, we conclude that the wasps were unable or unwilling to parasitize high numbers of hosts in the cages. The reason for this odd behaviour is unknown.     

Sucrose octanoate toxicity to brown citrus aphid (Homoptera: Aphididae) and the parasitoid Lysiphlebus testaceipes (Hymenoptera: Aphidiidae)

We report the toxicological effects of a new biorational, synthetic sucrose octanoate (AVA Chemical Ventures L.L.C., Portsmouth, NH), on brown citrus aphid, Toxoptera citricida (Kirkaldy), nymphs and adults and to its native parasitoid Lysiphlebus testaceipes (Cresson). Sucrose octanoate topically applied was equally toxic to brown citrus aphid adults and nymphs with LC50 and LC90 values ranging from 356 to 514 and 1029 to 1420 ppm, respectively. Mortalities of both stages did not differ significantly over time during the 3-24-h sampling period. Dry residues of sucrose octanoate exhibited similar levels of toxicity to both nymphs and adults. Mortality ranged from 60 to 70% at 6,000 ppm 4 h after exposure. L. testaceipes was not harmed by treatments as high as 4,000 ppm of sucrose octanoate as long as the parasitoid had mummified before treatment. Based on these results, sucrose octanoate would be a useful biorational in citrus integrated pest management programs.     

Studies on the host-finding ability of the aphid parasitoid,Trioxys complanatus (Hym.: Braconidae) in lucerne and clover

The spotted alfalfa aphid (SAA),Therioaphis trifolii (Monell)f. maculata, was first recorded in Australia in 1977. It is a major pest of lucerne and other pasture medics but not of clover (Trifolium spp.). The parasitoidTrioxys complanatus Quilis was introduced into Australia in late 1977 as part of a biological control program for SAA and made a substantial contribution to the management of the pest. Since 1989, aphids which are individually indistinguishable from SAA have been causing substantial yield loss to clover pastures in western and south-eastern Australia. This aphid, now known in Australia as the spotted clover aphid (SCA), is genetically distinct from SAA and has a much wider host range. Populations of SCA in clover are rarely parasitised byT. complanatus. A series of experiments was undertaken to determine whether the greater susceptibility of SAA in lucerne than of SCA in clover to parasitisation byT. complanatus is due to the attraction of the parasitoid preferentially to lucerne or to the differential attractiveness of the aphids themselves. SAA and SCA were offered to the parasitoids in arenas of increasing complexity from single trifoliate leaves of lucerne and clover on agar through single potted plants in small cages to groups of potted plants in a large cage. Results showed that the parasitoids exhibited a preference for lucerne though, in the cage situation, they did find and parasitise SCA on clover. They did not differentiate between SAA and SCA on lucerne.     

Aspects of host—parasitoid interactions of two aphid parasitoids, Aphidius urticae (Haliday) and Aphidius uzbeckistanicus (Luzhetski) (Hymenoptera, Aphidiidae)

Abstract. 1. Various aspects were studied of two host-parasitoid interactions: Aphidius urticae parasitizing the cocksfoot aphid Hyalopteroides humilis and Aphidius uzbeckistanicus parasitizing the cereal aphid Metopolophmm dirhodum. The work formed part of a study on the population dynamics of grassland and cereal aphids in Berkshire, England.
2. A.uzbeckistanicus had a higher instantaneous attack rate but a slightly longer handling time than A.urticae. In both cases the functional response was curvilinear.
3. A.urticae was specific to H.humilis and failed to develop in M.dirhodum. A.uzbeckistanicus could develop successfully in H.humilis but the functional response was sigmoid and the progeny were less fecund than those reared on the natural host. This species showed a clear preference for M.dirhodum when supplied with mixtures of M.dirhodum and H.humilis.
4. A.uzbeckistanicus had a faster rate of development and a lower developmental threshold than A.urticae , a situation paralleled by their natural hosts.
5. The results are discussed in relation to field observations. A.uzbeckistanicus may have considerable potential for reducing cereal aphid numbers. However, its preference for M.dirhodum mitigates against a reservoir being maintained in the economically unimportant grassland aphid, H.humilis.     

Parasitism byAphidius ervi (Hym.: Aphidiidae): Preference for pea aphid and blue alfalfa aphid (Hom.: Aphididae) and competition withA. Smithi

The pea aphidAcyrthosiphon pisum (Harris) and the blue alfalfa aphidA. kondoi Shinji are pests of alfalfaMedicago sativa L. worldwide. The parasitoidAphidium ervi Haliday attacks both species andA. smithi is host specific to pea aphid Experiments were conducted to determine the preference ofA. ervi forA. pisum andA. kondoi at three densities of hosts using the percentages of parasitism and superparasitism, and the distribution of eggs among hosts as criteria. Also examined was the outcome of competition between the two parasitoids when exposed to the same hosts. A. ervi attacks moreA. pisum when provided alone (no choice test) and when provided together in equal numbers withA. kondoi (choice test). There was no significant difference in the number of progeny produced by either parasitoid when a female of each species was introduced simultaneously into the same test unit containingA. pisum hosts. The effects of the pathogenPandora neoaphidis on this competition is reviewed.     

Evidence for the importance of odour-perception in the parasitoid Rhopalicus tutela (Walker) (Hym., Pteromalidae)

Possible host location mechanisms in the chalcid wasp Rhopalicus tutela (Walker) (Hym., Pteromalidae), a parasitoid of the eight-spined spruce bark beetle, Ips typographus (L.) (Col., Scolytidae), were examined. This was carried out in order to repeat and complement former studies on parallel parasitoid–scolytid systems that had contradictory results. Morphological examinations of the parasitoid antennae were made using both scanning and transmission electron microscopy. Possible functions of the sensilla placodea (multiporous plate sensillum), and other sensilla present on the antennae, have been indicated. For the first time, the placoid sensilla in a pteromalid parasitoid have shown porous walls and numerous innervations, which are typical characteristics for chemoreceptors. Previously the placoid sensilla have been suggested to be an infrared receptor. In order to test the chemoreceptive ability of R. tutela females and males, a synthetic reference blend was analysed by combined gas chromatography and electroantennographic detector (GC-EAD). Their sensitivity to host-related volatiles (such as certain pheromone components and oxygenated monoterpenes) was significantly greater than that for host-tree-related compounds (monoterpene hydrocarbons). Employing an infrared thermo-scanner, the current study failed to detect 'hot spots' associated with susceptible hosts beneath the bark. Results from electrophysiology and electron microscopy revealed clear odour-perceptive functions of the parasitoid antennae. These results strongly support the major importance of volatiles in host location by the bark beetle parasitoid R. tutela .     

Biology and distribution of microbe-associated thelytokous populations of aphid parasitoids (Hym., Braconidae, Aphidiinae)

Abstract: Some phenomena of the biology and ecology of the microbe ( Wolbachia )-associated thelytokous populations of aphid parasitoids are presented for the first time. Thelytokous virgin females were found to refuse the mating attempts by males of biparental populations, although males occurred quite rarely among the offspring. The host range was found to be about identical in conspecific biparental and thelytokous populations, whereas the association of solely biparental populations with a certain host group might be considered as a marker of the presence of sibling species in the area. The relative frequency of conspecific biparental and thelytokous populations varied, depending upon the area and host species. The microbe-induced thelytokous populations were confirmed in three species of the genus Lysiphlebus Foerst. ( Lysiphlebus cardui Marsh., Lysiphlebus confusus Tremblay and Eady, Lysiphlebus fabarum Marsh.) and probably some other, as yet undescribed, species only in the West Palearctic subregion, whereas the other known congeneric species in the subregion as well as in the Nearctic and the Neotropical regions were determined as strictly biparental species. Microbe-induced thelytokous populations did not change due to host species alternation and/or distribution area (purposeful introductions as biocontrol agents).