Response of the egg parasitoids of the pine processionary moth to host density and forest cover at the southern edge of the range

1. Thaumetopoea pityocampa is the most important pine defoliator in the Mediterranean basin. Despite being attacked by a number of natural enemies, populations occur frequently at high density in several areas.
2. Egg parasitism was studied in 27 pine and cedar forests in Algeria, in relation to the host density (tents per tree) and the proportion of forest cover in the landscape.
3. Egg parasitism varied from 2% to 25%, accounted by two parasitoid species, the specialist Baryscapus servadeii and the generalist Ooencyrtus pityocampae.
4. Tent density was negatively correlated with parasitism by B. servadeii but not with that of O. pityocampae. Conversely, parasitism by O. pityocampae increased with the proportion of forest and agricultural cover, but not in the case of B. servadeii.
5. Maximum summer temperature showed no correlation with parasitism rates. Still, temperature frequently exceeded 40 °C during the period of adult parasitoid activity.
6. The low performance of the egg parasitoids at the southern edge of the host range could be explained by the reduced fecundity of the host, climate effects, and phenological mismatching between the parasitoids and the egg development. These and other factors potentially involved the need to be further explored with a long-term study of population dynamics.

     

The influence of aphid-produced honeydew on parasitoid fitness and nutritional state: A comparative study

Honeydew is a sugar-rich resource excreted by many hemipteran species and is a key food source for other insect species such as ants and parasitoid wasps. Here, we evaluated the nutritional value of 14 honeydews excreted by 13 aphid species for the generalist aphid parasitoid Lysiphlebus testaceipes to test a series of hypotheses concerning variation in the nutritional value of honeydew. There was a positive correlation between the body sugar content of honeydew-fed parasitoids and their longevity. This information is valuable for biological control researchers because it demonstrates that the nutritional state of honeydew-fed parasitoids in the wild can indicate their fitness, independently of the honeydew source they have fed on.Although the carbohydrate content and longevity of L. testaceipes differed greatly among the different honeydews, we did not find a significant effect of aphid or host plant phylogeny on these traits. This result suggests that honeydew is evolutionarily labile and may be particularly subject to ecological selection pressures. This becomes apparent when considering host aphid suitability: Schizaphis graminum, one of the most suitable and commonly used hosts of L. testaceipes, produced honeydew of the poorest quality for the parasitoid whereas Uroleucon sonchi, one of the few aphids tested that cannot be parasitized by L. testaceipes, excreted the honeydew with the highest nutritional value. These data are consistent with the hypothesis that hemipterans are subject to selection pressure to minimize honeydew quality for the parasitoids that attack them.     

Effects of honeydew sugar composition on the longevity of Aphidius ervi

Feeding on sugar‐rich foods such as nectar and honeydew is important for survival of many adult parasitoids. Especially in agricultural systems, honeydew is often the most prevalent carbohydrate source. However, relative to plant nectar, honeydew may be relatively unsuitable, as a result of an unfavourable sugar composition or the presence of secondary plant compounds. We studied survival of the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) on honeydew collected from various aphid species feeding on potato (Solanum tuberosum L., cv. Desiree) (Solanaceae), wheat (Triticum aestivum L., cv. Bobwhite) (Poaceae), or artificial diet, as well as the sugar composition of the different honeydews. Honeydews from the tested aphid species on potato, wheat, or artificial diet were found to be relatively suitable food sources for adult A. ervi, although not always as suitable as a 2 M sucrose solution. There were differences in honeydew sugar composition among the different aphid species on the various host plants. Multivariate statistics showed that the factor ‘aphid species’ had a significant influence on the sugar composition of the honeydew, explaining 27% of the variation in the potato system and 89% in the wheat system. When exploring the relationship between carbohydrate composition of the honeydews from aphids on potato and wheat plants, and their nutritional value for A. ervi, data revealed that differences in parasitoid longevity can to some extent be explained by carbohydrate composition. Furthermore, our results confirm that sucrose and its hexose components glucose and fructose are very suitable carbohydrate sources for hymenopteran parasitoids and show that parasitoid survival on an equimolar solution of the two monosaccharides glucose and fructose does not exceed performance on the disaccharide sucrose.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

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.      

Prey‐mediated effects of glucosinolates on aphid predators

1. Plant resistance against herbivores can act directly (e.g. by producing toxins) and indirectly (e.g. by attracting natural enemies of herbivores). If plant secondary metabolites that cause direct resistance against herbivores, such as glucosinolates, negatively influence natural enemies, this may result in a conflict between direct and indirect plant resistance. 2. Our objectives were (i) to test herbivore‐mediated effects of glucosinolates on the performance of two generalist predators, the marmalade hoverfly (Episyrphus balteatus) and the common green lacewing (Chrysoperla carnea) and (ii) to test whether intraspecific plant variation affects predator performance. 3. Predators were fed either Brevicoryne brassicae, a glucosinolate‐sequestering specialist aphid that contains aphid‐specific myrosinases, or Myzus persicae, a non‐sequestering generalist aphid that excretes glucosinolates in the honeydew, reared on four different white cabbage cultivars. Predator performance and glucosinolate concentrations and profiles in B. brassicae and host‐plant phloem were measured, a novel approach as previous studies often measured glucosinolate concentrations only in total leaf material. 4. Interestingly, the specialist aphid B. brassicae selectively sequestered glucosinolates from its host plant. The performance of predators fed this aphid species was lower than when fed M. persicae. When fed B. brassicae reared on different cultivars, differences in predator performance matched differences in glucosinolate profiles among the aphids. 5. We show that not only the prey species, but also the plant cultivar can have an effect on the performance of predators. Our results suggest that in the tritrophic system tested, there might be a conflict between direct and indirect plant resistance.     

Testing for host‐associated differentiation in two egg parasitoids of a forest herbivore

Several studies underline the importance of ecological barriers and differential selection in driving sympatric speciation. Host‐associated differentiation (HAD) has been proposed as one of the mechanisms leading to sympatric speciation. However, it is still unclear how common HAD is or which are the factors that could promote it. In particular, not much is known about HAD in predators and parasitoids of herbivorous insects. One of the characteristics postulated to pre‐dispose insects to HAD is parthenogenesis as it may favour adaptive responses to particular environments, amplifying selected gene complexes. In this study, we used amplified fragment length polymorphism (AFLP) markers to determine whether HAD is present in two parthenogenetic egg parasitoids attacking the same herbivore species – the pine processionary moth, Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera: Notodontidae) – on two host Pinus species. A total of 100 loci for 59 individuals sampled in four populations of Baryscapus servadeii (Domenichini) (Hymenoptera: Eulophidae), a specialist parasitoid, and 106 loci for 117 individuals sampled in six populations of Ooencyrtus pityocampae Mercet (Hymenoptera: Encyrtidae), a generalist parasitoid, were analysed. Levels of genetic differentiation were also assessed with an outlier analysis, checking for alleles associated to host plants. No evidence of HAD was detected in any of the two parasitoid species. We hypothesize that both the lack of strict parthenogenetic reproduction and the ectophagous nature of the insect host could explain the absence of HAD. The genetic variation observed in the generalist parasitoid responded to a pattern of local adaptation, whereas no relationship with either host or geography was found in the specialist parasitoid.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

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.     

Egg parasitoids of Thaumetopoea pityocampa (Den. & Schiff.) (Lep., Thaumetopoeidae) and their impact in SW Turkey

Abstract: Egg parasitoids (Hym., Chalcidoidea) and egg parasitism were studied in 132 egg batches of the pine processionary moth collected from stands of Pinus nigra Arnold and P. brutia Ten. in SW Turkey, at four places in heights of 800–1010 m a.s.l. in October 1998, beyond the caterpillars’ hatch. The egg batches deposited on P. nigra were high‐significantly smaller than those found on P. brutia. In this mountainous regions, Baryscapus servadeii (Dom.) and Ooencyrtus pityocampae (Mercet) were the most abundant egg parasitoids of Thaumetopoea pityocampa (Den. & Schiff.). Considering the different sampling places, Trichogramma sp. was found maximally in 3.1% of the parasitized eggs. Additionally, the hyper‐parasitoid Baryscapusransversalis Graham (mean infestation 1.8%) and Pediobius bruchicida (Rond.) (0.1%), both occurring in both sexes, and Anastatus bifasciatus (Fonsc.) (0.1%), only females, were established. The primary parasitoid O. pityocampae dominantly parasitized the eggs placed near the top of the batches, and B. servadeii was found mainly in eggs near the base. The rate of parasitism of the egg batches was very variable. The impact of egg parasitoids and predators varied from 24 to 35.9% depending on the sampling place. The hatching rate of the caterpillars was reduced to 50.7–65.6%. Under laboratory conditions, the emergence dynamics of the parasitoids showed a peak of B. tranversalis in December of the year, in which the host eggs were laid. Baryscapus servadeii emerged mainly in March–May of the following year, when the second part of O. pityocampae appeared. A large part of both parasitoids emerged before sampling of the batches. In O. pityocampae 1.65% and in B. servadeii 2.23% males developed under laboratory conditions. A rate of 15.7% of O. pityocampae and 7.9% of B. servadeii died in the eggs. The polyembryonic Trichogramma sp. (see also Tsankov et al., 1996a ) attacked 1.8% of the eggs. Up to 20 (mean 12) dead pupae were found in one egg. Minor parts of eggs were undeveloped (2.9%) and 0.6% totally empty.     

The role of host plant species in the phenotypic differentiation of sympatric populations of Aleiodes nolophanae and Cotesia marginiventris

The ecology of parasitoids is strongly influenced by their host plant species. Parasitoid fitness can be affected by a variety of plant traits that could promote phenotypic differentiation among populations of parasitoids. Generalist parasitoids are expected to be more affected by plant traits (e.g., plant defensive traits) than specialist parasitoids. Data are presented on phenotypic differences of two braconid parasitoid wasps ovipositing on the same insect host species on two different host plant species. Adult mass, adult longevity, and percent parasitism are compared for the generalist parasitoid Cotesia marginiventris Cresson and the specialist parasitoid Aleiodes nolophanae Ashmead (both Hymenoptera: Braconidae) emerging from green cloverworms, Hypena scabra Fabricius (Lepidoptera: Noctuidae), feeding on two host plant species, alfalfa (Medicago sativa L.) and soybean (Glycine max L. Merr.) (both Fabaceae), at three locations. Specialist wasps that parasitized the green cloverworm on alfalfa had a significantly larger mass than the ones that parasitized the green cloverworm on soybean at the three study sites. Generalist wasps that parasitized green cloverworms on alfalfa had a larger mass than wasps parasitizing green cloverworms on soybean only at one of the study sites (i.e., Prince George's County, MD, USA). Similarly, both specialist and generalist wasps lived longer when parasitizing green cloverworms on alfalfa than when parasitizing them on soybean at only one of the study sites (i.e., Prince George's County). In Prince George's County, percent parasitism on alfalfa by the specialist parasitoid was higher than on soybean for three consecutive years and percent parasitism by the generalist parasitoid was the same on alfalfa and soybean every year. Thus, phenotypic differences among populations associated with different host plant species vary geographically (i.e., parasitoid phenotype associated with different host plant species differ at some sites while it is the same at other sites). The implications of geographic variation for biological control are discussed.     

Comparing floral nectar and aphid honeydew diets on the longevity and nutrient levels of a parasitoid wasp

We compared the effects of floral nectar from buckwheat, Fagopyrum esculentum Moench, and honeydew produced by the soybean aphid, Aphis glycines Matsumura (Homoptera: Aphididae), on longevity, nutrient levels, and egg loads of the parasitoid Diadegma insulare Cresson (Hymenoptera: Ichneumonidae). Diadegma insulare lived for 2 days in control treatments of water or clean soybean leaves, for 6–7 days with honeydew, and in excess of 2 weeks with buckwheat nectar. Potential reasons for the superiority of buckwheat nectar over soybean aphid honeydew for extending the longevity of parasitoids include: (i) parasitoids ingest more sugars from floral sources, (ii) oligosaccharides in honeydew have a lower nutritional value than nectar sugars, and (iii) honeydew has antagonistic compounds. Overall sugar levels were lower in honeydew‐ vs. nectar‐fed female wasps, suggesting a lower feeding rate, but other explanations cannot be excluded. Diadegma insulare eclosed with high levels of lipids and glycogen, and low levels of gut and storage sugars. All carbohydrates increased over the life of both nectar‐ and honeydew‐fed wasps, but remained low or decreased in starved wasps. Lipid levels declined over the lifespan of female wasps, but females fed floral nectar showed the slowest rate of lipid decline. Diet did not affect egg load, probably because the females were not given hosts in the experiment.     

Comparing and contrasting life history variation in four aphid hyperparasitoids

1. In primary parasitoids, significant differences in life history and reproductive traits are observed among parasitoids attacking different stages of the same host species. Much less is known about hyperparasitoids, which attack different stages of primary parasitoids. 2. Parasitoids exploit hosts in two different ways. Koinobionts attack hosts that continue feeding and growing during parasitism, whereas idiobionts paralyse hosts before oviposition or attack non‐growing host stages, e.g. eggs or pupae. 3. Koino‐/idiobiosis in primary parasitoids are often associated with different expression of life history trade‐offs, e.g. endo‐ versus ectoparasitism, high versus low fecundity and short versus long life span. 4. In the present study, life history parameters of two koinobiont endoparasitic species (Alloxysta victrix; Syrphophagus aphidivorus), and two idiobiont ectoparasitic species (Asaphes suspensus; Dendrocerus carpenteri) of aphid hyperparasitoids were compared. These hyperparasitoids attack either the parasitoid larva in the aphid before it is killed and mummified by the primary parasitoid or the parasitoid prepupa or pupa in the dead aphid mummy. 5. There was considerable variation in reproductive success and longevity in the four species. The idiobiont A. suspensus produced the most progeny by far and had the longest lifespan. In contrast, the koinobiont A. victrix had the lowest fecundity. Other developments and life history parameters in the different species were variable. 6. The present results reveal that there was significant overlap in life history and reproductive traits among hyperparasitoid koinobionts and idiobionts, even when attacking the same host species, suggesting that selection for expression of these traits is largely association specific.     

Intrinsic prey suitability in specialist and generalist Harmonia ladybirds: a test of the trade‐off hypothesis for food specialization

The trade‐off hypothesis posits that increased performance on a given resource comes at the cost of decreased performance on other resources, and that this trade‐off is a driving force of food specialization in both predators and herbivores. In this study, we examined larval survival and performance in two sibling ladybird species, Harmonia yedoensis Takizawa and Harmonia axyridis Pallas (Coleoptera: Coccinellidae), fed on one of four prey species. Harmonia yedoensis is a specialist predator that preys mostly on pine aphids in the field, whereas H. axyridis is a generalist predator with a broad prey range. We experimentally showed in the laboratory that larval survival and performance were not higher when H. yedoensis was fed on pine aphids, compared with the other prey species. Rather, prey suitability was similar in both ladybird species, and H. yedoensis larvae developed as well or even better on prey species that they never utilize in nature. These results suggest that the host range in H. yedoensis may not be limited by the intrinsic suitability of the aphid species per se. Moreover, as shown by our previous study, the pine aphid is a highly elusive prey that is difficult for small ladybird hatchlings to capture, which means that the cost of utilizing this prey is high. Therefore, we conclude that some factor other than prey suitability is responsible for the observed food specialization in H. yedoensis.     

Aphid Honeydew Quality as a Food Source for Parasitoids Is Maintained in Bt Cotton

Bt-transgenic cotton has proven to be highly efficient in controlling key lepidopteran pests. One concern with the deployment of Bt cotton varieties is the potential proliferation of non-target pests. We previously showed that Bt cotton contained lower concentrations of insecticidal terpenoids as a result of reduced caterpillar damage, which benefited the aphid Aphis gossypii. It is thus important that non-target herbivores are under biological control in Bt cotton fields. The induction or lack of induction of terpenoids could also influence the quality of aphid honeydew, an important food source for beneficial insects. We therefore screened A. gossypii honeydew for cotton terpenoids, that are induced by caterpillars but not the aphids. We then tested the influence of induced insect-resistance of cotton on honeydew nutritional quality for the aphid parasitoid Lysiphlebus testaceipes and the whitefly parasitoid Eretmocerus eremicus. We detected the cotton terpenoids gossypol and hemigossypolone in A. gossypii honeydew. Although a feeding assay demonstrated that gossypol reduced the longevity of both parasitoid species in a non-linear, dose-dependent manner, the honeydew was capable of sustaining parasitoid longevity and reproduction. The level of caterpillar damage to Bt and non-Bt cotton had no impact on the quality of honeydew for the parasitoids.These results indicate that the nutritional quality of honeydew is maintained in Bt cotton and is not influenced by induced insect resistance.     

Geographic and between-generation variation in the parasitoid communities associated with an invading gallwasp,Andricus quercuscalicis (Hymenoptera: Cynipidae)

The knopper gallwasp Andricus quercuscalicis Burgsdorf 1783 (Hymenoptera: Cynipidae) has invaded western and northern Europe from southern and eastern Europe over the last 400 years. A. quercuscalicis has two alternating generations, which differ in phenology, structure, and host oak species. This study describes geographic variation in the community in the tiny catkin galls of the sexual generation on Turkey oak, Quercus cerris, and compares the patterns obtained with those in the community attacking the alternate agamic generation. As predicted from considerations of parasitoid recruitment to the communities of invading phytophagous insects (Cornell and Hawkins 1993), in its native range the sexual generation shows (1) higher parasitoid community species richness, (2) higher total mortality due to parasitoid attack and (3) a higher ratio of specialist to generalist parasitoid species than is evident in the invaded range. Counter to predictions, there is no indication that parasitoid community richness in the invaded range has increased with time since the arrival of the new host. Higher host mortality in the native range is due principally to a single specialist, Aulogymnus obscuripes Mayr 1877 (Hymenoptera: Eulophidae), and is not distributed evenly among parasitoid species which attack the gall-former only in this area. This contrasts with the community in Britain, where three principal generalist parasitoids cause approximately equal mortalities. The agamic gall contains a taxonomically and structurally diverse guild of parasitoid and inquiline species, associated with the changing resource provided by a large, long-lived, complex gall. In contrast, the sexual community includes a taxonomically and structurally narrow guild, associated with a resource which is structurally simple, small in size and short-lived. No parasitoid species attacks the gall-former in both generations. Surprisingly, in spite of these differences in the nature of the gall resource in the two generations, over their entire range (native and invaded) the parasitoid guilds of the two are equally species rich.     

The impact of an ant–aphid mutualism on the functional composition of the secondary parasitoid community

1. Mutualistic and antagonistic interactions, although often studied independently, may affect each other, and food web dynamics are likely to be determined by the two processes working in concert. 2. The structure, and hence dynamics, of food webs depends on the relative abundances of generalist and specialist feeding guilds. Secondary parasitoids of aphids can be divided into two feeding guilds: (i) the more specialised endoparasitoids, which attack the primary parasitoid larvae in the still living aphid, and (ii) the generalist ectoparasitoids, which attack the pre‐pupa of the primary or secondary parasitoid in the mummified aphid. 3. We studied the effect of an ant–aphid mutualism on the relative abundance of these two functional groups of secondary parasitoids. We hypothesised that generalists will be negatively affected by the presence of ants, thus leading to a greater dominance of specialists. 4. We manipulated the access of ants (Lasius niger) to aphid colonies in which we placed parasitised aphids. Aphid mummies were collected and reared to determine the levels of endo‐ and ecto‐secondary parasitism. 5. When aphids were attended by L. niger the proportion of secondary parasitism by ectoparasitoids dropped from 26 to 8% of the total number of parasitised aphids, with Pachyneuron aphidis most strongly affected, while endoparasitoids as a group did not respond. However, among these Syrphophagus mamitus profited from ant attendance becoming the dominant secondary parasitoid, while parasitisation rates of Alloxysta and Phaenoglyphis declined. 6. The shift to S. mamitus as dominant secondary parasitoid in ant‐attended aphid colonies is likely due to the behavioural plasticity of this species in response to ant aggression, and a release from tertiary parasitism by generalist ectoparasitoids. 7. The reduction of secondary parasitism by generalist ectoparasitoids reduces the potential for apparent competition among primary parasitoids with consequences for the dynamics of the wider food web.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Optimal biological parameters for rearing Ooencyrtus pityocampae on the new laboratory host Philosamia ricini

Biological control programmes involving Ooencyrtus pityocampae Mercet (Hymenoptera: Encyrtidae) have proved effective at reducing the damage caused by the pine processionary Thaumetopoea pityocampa Denis & Schiffermüller (Lepidoptera: Thaumatopoeidae). In this study, the biological variables that influence the parasitism of O. pityocampae on the new laboratory host Philosamia ricini Danovan (Lepidoptera: Saturniidae) have been investigated. Laboratory experiments were conducted under the conditions of 25 ± 1°C, 65 ± 5% R.H (relative humidity) and a photoperiod of 16 : 8 h (L : D = light : dark). The host egg age and parasitoid age are often regarded as being key factors influencing the emergence rate of O. pityocampae. The optimal age of host eggs for parasitization was 1–2 days, and the emergence rate was highest with 5‐day‐old female parasitoids. Thus, our results define the optimal conditions for the effective and economic rearing of parasitoids as follows: one 5‐day‐old female parasitoid per 50 (1–2)‐day‐old host eggs. The development time of O. pityocampae ranged between 19.5 and 22.6 days. Parasitoids that were exposed to bio‐honey survived 10.5 times longer than those that did not receive supplemental food. O. pityocampae was reared for more than nine generations on the eggs of P. ricini. Consequently, P. ricini has been found as a suitable new laboratory host for the mass rearing of O. pityocampae for the use of biological control programmes against T. pityocampa in future.     

Metapopulation dynamics in an aphid-parasitoid system

Metapopulation theory makes a number of predictions concerning the effects of dispersal on the persistence of predator-prey or host-parasitoid systems. While the stabilising effects of dispersal have been shown in a number of laboratory studies, evidence from field studies remains scarce due to a lack of suitable model systems. I describe a host-parasitoid system that shows a classical metapopulation structure with frequent extinctions and colonisations consisting of the aphidiid Lysiphlebus hirticornisand the aphid Metopeurum fuscoviride. Both the parasitoid and the aphid are specialists on their respective hosts. I followed the dynamics of host and parasitoid on individually marked tansy (Tanacetum vulgare) plants, the host of M. fuscoviride. Dynamics of host and parasitoid populations were characterized by frequent extinctions and colonisations. Mean longevity of aphid colonies was only 3.1 weeks. Parasitism by L. hirticorniswas a main cause of extinction for the aphid as rates of parasitism often reached 100%, in particular towards the end of the field season. Patchiness in this system occurs at two spatial scales. Aphid colonies form on single tansy ramets = shoots but movements of aphid individuals among ramets within a particular tansy genet are frequent. Because aphids can persist on a genet for a large numer of generations, it is argued that local populations form on genets rather than ramets. The number of host and parasitoid extinctions described in this study exceeds the number of extinctions usually observed in field studies of host-parasitoid metapopulations. It is suggested that aphid-parasitoid systems such as the one studied in this paper may be good models to test the predictions of metapopulation theory.