Egg-laying strategy under natural conditions of Leptopilina boulardi, a hymenopteran parasitoid of Drosophila spp

The hypothesis of optimal host species selection predicts that when a parasitoid has the choice between two host species, it will choose the species thay gives the best survival chances for its progeny. We confirmed this hypothesis by laboratory experiments with Leptopilina boulardi Barb. et al., a cynipid parasitoid which prefers Drosophila melanogaster Meigen (the host species most suitable for parasitoid survival) above D. simulans Sturt. As far as fitness parameters are concerned, the fertility of L. boulardi is higher with D. melanogaster; the egg laying can be spread out over a long period when this host is relatively scarce. This does not occur with D. simulans in which parasitic oviposition stops soon when this host is not abundant.Investigations of this foraging strategy were done under more complex natural conditions. We found that L. boulardi has a type III functional response with D. melanogaster only; furthermore, it seems that a switching effect may exist with this host. Parasitoid females appear to distribute their eggs more regularly on D. melanogaster, thus avoiding superparasitism. This seems to be independent of the relative frequency of this host. However, superparasitism of D. simulans did occur more frequently when this host was scarce.

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Stratégie de ponte de Leptopilina boulardi (hyménoptère parasite de drosophiles) dans les conditions naturelles

Résumé Le concept de réponse optimale d'un parasite vis-à-vis de l'hôte le plus favorable pour son développement demeure surtout théorique et n'a pu être vérifié que dans les conditions de laboratoire. Nous avons montré que Drosophila melanogaster s'avère être, par rapport à D. simulans, l'hôte le plus favorable pour le développement du cynipide parasite Leptopilina boulardi. Une étude sur le terrain a démontré que ce parasite présente une réponse fonctionnelle densité dépendante vis-à-vis de D. melanogaster et non vis-à-vis de D. simulans, avec un effet de bascule. D'autre part, il s'avère que ce parasite exploite beaucoup mieux son hôte, en évitant le superparasitisme, ceci étant démontré au laboratoire et dans la nature. Enfin, il apparaît qu'il est capable d'allonger sa période de ponte lorsque cet hôte est rare, ce qui ne se produit pas avec D. simulans.

     

Influence of galls of Phanacis taraxaci on carbon partitioning within common dandelion, Taraxacum officinale

We examined how leaf galls, induced by the cynipid wasp Phanacis taraxaci, influence the partitioning of photoassimilates within the host, the common dandelion, Taraxacum officinale. Galled and ungalled plants were exposed to ¹⁴CO₂ and the labelled photoassimilates accumulating within galls and other parts of the host were measured. During the growth phase of the gall they were physiological sinks for photoassimilates, accumulating 9% to 70% of total carbon produced by the host, depending upon the number of galls per plant. High levels of ¹⁴C assimilation in the leaves of galled plants compared to controls, suggest that galls actively redirect carbon resources from unattacked leaves of their host plant. This represents a significant drain on the carbon resources of the host, which increases with the number and size of galls per plant. Active assimilation of ¹⁴C by the gall is greatest in the growth phase and is several orders of magnitude lower in the maturation phase. This finding is consistent with physiological and anatomical changes that occur during the two phases of gall development and represents a key developmental strategy by cynipids to ensure adequate food resources before larval growth begins.     

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.     

Wolbachia prevalence patterns: horizontal transmission,recombination, and multiple infections in chestnut gall wasp-parasitoid communities

The Oriental chestnut gall wasp, Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), is a global invasive pest that causes serious damage to almost all chestnut species belonging to the Castanea genus (Fagaceae). Dryocosmus zhuili Liu et Zhu is a recently described sibling species of D. kuriphilus, which induces galls on Castanea henryi (Skan) Rehd. et Wils. There are many indigenous parasitoid species in China which play an important role in the natural regulation of their population dynamics. Wolbachia is a maternally inherited α-proteobacterium widely found in arthropods. This study screened for the presence of Wolbachia in the two chestnut gall wasps and in six parasitoid species from 12 populations, to investigate the prevalence patterns of Wolbachia in the chestnut gall wasp-parasitoid communities. We found that D. zhuili and four parasitoid species were infected with Wolbachia; among them, all individuals of the two populations of Megastigmus sp. had multiple Wolbachia infections. By using multilocus sequence types to characterize bacterial strains, three new sequence types were identified. The Wolbachia strains infecting D. zhuili (ST-507), Torymus sinensis Kamijo (ST-508), and Sycophila variegata (Curtis) (ST-508) belonged to supergroup A, whereas the Wolbachia strain infecting Megastigmus nipponicus Kamijo (ST-503) belonged to supergroup B. Our results also suggested that horizontal transmission of Wolbachia occurs between chestnut gall wasps and their parasitoids. Moreover, multiple Wolbachia infections of Megastigmus sp. may be due to gene recombination and horizontal transmission.     

New records of Cynipidae (Hymenoptera) from Italy

New records of Cynipid gall wasps and inquilines for the Italian peninsula and Sicily and their new host plants for the Palaearctic Region are listed and commented on. Among them we find: Cerroneuroterus cerrifloralis (Müllner 1901) as new for Italy and new for the Palaearctic region as host on Quercus suber; Andricus multiplicatus Giraud 1859 on Q. suber, as new host for the Palaearctic region; Aylax papaveris (Perris 1839), reported in Italy over a century ago, but later overlooked; Cerroneuroterus minutulus (Giraud 1859), also reported more than a century ago from Sicily, but later overlooked. Among the inquilines are here listed: Synergus variabilis Mayr 1872, emerged from Janetia cerris (Kollar 1850) galls (Diptera Cecidomyiidae), and found for the first time in the Palaearctic Region as host on Q. suber; Saphonecrus haimi (Mayr 1872) and Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey 1986, are new records for Italy.     

Predicting the Impact of the Biocontrol Agent Aulacidea subterminalis (Cynipidae) on Growth of Hieracium pilosella (Asteraceae) under Differing Environmental Conditions in New Zealand

A gall wasp, Aulacidea subterminalis, was released into the field in New Zealand in 1999 for biocontrol of the grassland weed Hieracium pilosella. To predict its likely impact in the field, the effect of the gall wasp on plant growth was measured under stress-free conditions in a shade house trial with potted plants. Plants with galls showed a reduction in stolon length of 75%. They had slightly lower total dry matter and reduced root weight. In a glasshouse experiment to measure the impact of water stress, nutrient stress, and plant competition on growth of H. pilosella and performance of the gall wasp, the number and mean diameter of gall clusters were not significantly different between treatment and control plants. However, galled plants produced more, but shorter, stolons in all stress treatments, and stolons that were more branched in nutrient- and water-stressed plants, than ungalled plants. Under the plant competition treatment, total stolon length relative to biomass was greater with, than without, wasps, indicating that stolons were thinner. Galling by A. subterminalis is likely to reduce vegetative reproduction of H. pilosella whether or not the plants are stressed, indicating that the wasp may be a successful biocontrol agent.     

Support for the microenvironment hypothesis for adaptive value of gall induction in the California gall wasp, Andricus quercuscalifornicus

Three major hypotheses have been advanced for the adaptive nature of plant galls: nutrition, enemy-avoidance, and microenvironment. Of these, the microenvironment hypothesis has been frequently invoked, but rarely tested directly. We tested this hypothesis in a population of Andricus quercuscalifornicus (Bassett) (Hymenoptera: Cynipidae) wasps inducing galls on Quercus lobata Née (Fagaceae) trees in Northern California, USA. Relative humidity and temperature data gathered from both immature and mature galls in the field indicated that A. quercuscalifornicus larvae modify their microenvironments significantly by raising and stabilizing relative humidity levels inside galls to near saturation. In addition, excised larvae maintained under experimental conditions survived significantly longer under levels of high relative humidity. These data support the hypothesis that through gall induction, A. quercuscalifornicus manipulates its environment adaptively.     

Isozyme differences between the developmental stages of the parthenogenetic gall wasp Diplolepis rosae

The value of electrophoretic analysis of enzymes as an aid in connecting a morphologically deviating form to one of several possible species is dependent on the proportion of shared enzyme loci between the forms. In order to determine this proportion for the different instars of the parthenogenetic gall wasp Diplolepis rosae, 16 different enzyme systems were analysed. Out of the 37 loci detected, 25 (68%) were active in all instars. The larvae, pupae, and imagines had two, one and five unique loci, respectively. Larvae and pupae shared four loci not expressed in the imagines. In the ten enzymes analysed for comparison of males and females a total of 22 loci were detected, 21 found in both sexes and one unique to the females. The difference in isozyme pattern was found to be much more pronounced between pupa and imago than between larva and pupa. The chance to find a locus in an instar when already detected in another was calculated to 87%. If this estimate is valid for other forms within species, differential activity should not present a problem when electrophoretic analysis is used in order to connect such forms.     

Gall-inducing insects provide insights into plant systematic relationships

Field surveys of cynipid gall-inducer occurrences on Quercus species were conducted in Florida, North Carolina, and Pennsylvania, USA. All cynipids demonstrated strong host species and organ fidelity. One result of this specialization is effective niche partitioning among cynipids. The host-association patterns of these specialist herbivores should reflect similarities among oaks, thus we clustered oak species according to their cynipid distributions. Cynipids distinguished small differences among their hosts. A dendrogram of oak species based on cynipid distributions was largely congruent with botanical arrangernents. Cynipid occurrences offer information helpful to resolving some aspects of oak systematics. Collaborative efforts between taxonomic botanists and entomologists will be useful in resolving a variety of plant and insect systematic problems.     

A new subfamily of Figitidae (Hymenoptera, Cynipoidea)

Larvae of the parasitic wasp family Figitidae develop as internal parasitoids of other endopterygote insect larvae. The hosts are typically dipteran larvae living in other microhabitats but the earliest figitids probably attacked gall-inhabiting hymenopteran larvae. Here, we formally describe a new genus (Parnips) and subfamily (Parnipinae) for a species that is likely to be a surviving representative of these early gall-associated figitids. The species, P. nigripes , has been reared repeatedly from galls inside the seed capsules of annual poppies (Papaver dubium and P. rhoeas) in the Mediterranean region together with the gall inducer Barbotinia oraniensis belonging to the Cynipidae, the sister group of Figitidae. Parnips nigripes is strikingly cynipid-like and was first assumed to be a cynipid gall inducer of the genus Aulacidea. Phylogenetic analyses have since indicated that the similarity with the Cynipidae is symplesiomorphic and that P. nigripes belongs to the Figitidae, where it forms the sister group of all other extant figitids. Recently, it has also been shown that P. nigripes is a parasitoid of the gall-inducing Barbotinia oraniensis , consistent with its proposed phylogenetic position. Parnips nigripes shares several unusual morphological traits with its host. We speculate that many of these similarities are homologous even though the lineages separated at least 83 million years ago.