Effects of parasitization or injection of parasitoid-derived factors from the endoparasitic wasp Glyptapanteles porthetriae (Hym., Braconidae) on the development of the larval host, Lymantria dispar (Lep., Lymantriidae)

Second instar larvae of Lymantria dispar were parasitized or injected with parasitoid-derived factors such as venom, calyx fluid or parasitoid eggs from Glyptapanteles porthetriae . Growth and development of the host larvae were affected in all different groups compared to control larvae of the same age, injected with Ringer solution. The greatest impact on host growth and on the duration of the 3rd instar was caused by injecting parasitoid eggs. Treated larvae showed melanized capsules or nodules in the hemocoel. While the wasp age had no effect on parasitization efficiency or on the percentage of melanized particles in the hemocoel, significantly more encapsulations were found in larvae parasitized by old wasps as opposed to young wasps. Superparasitization (double or quadruple oviposition) increased the parasitization efficiency markedly. While none of the control larvae showed melanized particles, in the groups of single and superparasitized (2× and 4×) hosts a high percentage of melanized particles (capsules and nodules) occurred.     

Mating of Xenos vesparum (Rossi) (Strepsiptera, Insecta) revisited

The controversial mating of the strepsipteran Xenos vesparum was studied to investigate the possible sperm routes for fertilization. The female, which is a neotenic permanent endoparasite of Polistes wasps, extrudes only its anterior region, the "cephalothorax," from the host abdomen. This region has an opening where both mating and larval escape occur. Observations with scanning and transmission electron microscopy revealed spermatozoa not only in the hemocoel, but also in the "ventral canal" (an extragenital duct peculiar to strepsipteran females) and in the "genital ducts" (ectodermal invaginations connecting the ventral canal to the hemocoel) of recently mated females. Xenos vesparum spermatozoa can reach the oocytes either through the hemocoel as a result of a hypodermic insemination, or by moving along the extragenital ducts, which are later used by first instar larvae to escape. The hypothesis of hypodermic insemination is reconsidered in the light of behavioral and ultrastructural evidence.     

The midgut ultrastructure of the endoparasite Xenos vesparum (Rossi) (Insecta, Strepsiptera) during post-embryonic development and stable carbon isotopic analyses of the nutrient uptake

Females of the endoparasite Xenos vesparum (Strepsiptera, Stylopidae) may survive for months inside the host Polistes dominulus (Hymenoptera, Vespidae). The midgut structure and function in larval instars and neotenic females has been studied by light and electron microscope and by stable carbon isotopic technique. The 1st instar larva utilizes the yolk material contained in the gut lumen, whereas the subsequent larval instars are actively involved in nutrient uptake from the wasp hemolymph and storage in the adipocytes. At the end of the 4th instar, the neotenic female extrudes with its anterior region from the host; the midgut progressively degenerates following an autophagic cell death program. First the midgut epithelial cells accumulate lamellar bodies and then expel their nuclei into the gut lumen; the remnant gut consists of a thin epithelium devoid of nuclei but still provided with intercellular junctions. We fed the parasitized wasps with sugar from different sources (beet or cane), characterized by their distinctive carbon isotope compositions, and measured the bulk (13)C/(12)C ratios of both wasps and parasites. Female parasites developing inside the wasp hemocoel are able to absorb nutrients from the host but, after their extrusion, they stop incorporating nutrients and survive thanks to the adipocytes content.     

Growth and development of the endoparasitic wasp Apanteles congregatus: dependence on host nutritional status and parasite load

ABSTRACT. Previously we have shown that the number of Apanteles congregatus Say (Hymenoptera, Braconidae) larvae developing in Manduca sexta (L.) (Lepidoptera, Sphingidae) larvae that are parasitized in the first instar determines the timing of emergence of the parasites from the host. Here we show that the first larval ecdysis of the wasps occurs after the host ecdyses to the terminal stage, regardless of whether that stage is the host's fourth, fifth or supernumerary sixth instar. Starvation of newly ecdysed terminal stage host larvae prevents emergence of the parasites. When starvation is begun at progressively later times, then an increasing proportion of the hosts have parasites that emerge, suggesting a period of indispensable host nutrition exists during which the host must feed to satisfy the developmental requirements of the parasites. In hosts fed ad libitum , the weight of the host plus its parasites at the time of emergence is positively correlated with the number of parasites developing in the host. When the weight of the parasites alone is subtracted from the weight of the host—parasite complex, the data show that heavily parasitized hosts have a larger host mass than lightly parasitized larvae. In contrast, the wasp larvae, and the adult males and females that develop from them, have lower individual weights after development in heavily parasitized hosts.     

Alcohol consumption as self-medication against blood-borne parasites in the fruit fly

Plants and fungi often produce toxic secondary metabolites that limit their consumption, but herbivores and fungivores that evolve resistance gain access to these resources and can also gain protection against nonresistant predators and parasites. Given that Drosophila melanogaster fruit fly larvae consume yeasts growing on rotting fruit and have evolved resistance to fermentation products, we decided to test whether alcohol protects flies from one of their common natural parasites, endoparasitoid wasps. Here, we show that exposure to ethanol reduces wasp oviposition into fruit fly larvae. Furthermore, if infected, ethanol consumption by fruit fly larvae causes increased death of wasp larvae growing in the hemocoel and increased fly survival without need of the stereotypical antiwasp immune response. This multifaceted protection afforded to fly larvae by ethanol is significantly more effective against a generalist wasp than a wasp that specializes on D. melanogaster. Finally, fly larvae seek out ethanol-containing food when infected, indicating that they use alcohol as an antiwasp medicine. Although the high resistance of D. melanogaster may make it uniquely suited to exploit curative properties of alcohol, it is possible that alcohol consumption may have similar protective effects in other organisms.     

The chemical strategies used by Polistes nimphus social wasp usurpers (Hymenoptera Vespidae)

Polistes foundresses can behave as facultative social parasites when, instead of founding their own nest, they usurp colonies of the same or a different species and temporary use the host workforce to raise their own brood. Conspecific usurpation appears to be common among Polistes wasps, but nothing is known about the mechanisms that these facultative social parasites use to have themselves accepted within usurped colonies. Using behavioural tests, we studied the chemical strategies employed by females of Polistes nimphus when they behave as facultative social parasites in colonies of the same or of a different species. We hypothesized that usurpers would mark host nests with their own odours and/or acquire host nest odours in order to camouflage their real identity from host workers. Our results indicated that P. nimphus usurpers used different chemical strategies depending on host nest species: they acquired conspecific host odours but marked heterospecific host combs with their own odours.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 505–512.     

Effects of brood parasitism on host reproductive success: evidence from larval interactions among dung beetles

This paper investigates the effect of brood parasitism in a dung beetle assemblage in an arid region of Spain. The study was conducted during the spring season (March-May 1994-1998) using mesh cylinders buried into the ground, filled with sand and with sheep dung on top. We quantified the proportion of nests containing larvae of parasitic beetles and their effect on host larvae survival. Experiments on the effect of parasitic larvae on host-larvae survival were conducted by placing scarab brood masses (raised from captive scarabs in the laboratory) in containers with and without aphodiid larvae. During the spring, dung desiccation is rapid, preventing aphodiids nesting in the dung, and forcing these species to adopt brood parasitism as a nesting strategy. Parasitic aphodiids were found in 12-47% of scarab nests of three species. The incidence of brood parasitization was positively related with the number of brood masses contained in the nests, being also higher in the most abundant species. Field data and experiments showed that brood parasites significantly reduced host larvae survival from 74.8% in non-parasitized nests to 8.8% in parasitized nests. Because different rates of nest parasitization and mortality were caused by parasites, brood parasitism had a differential effect on different host species. Thus, brood parasitism constitutes an important mortality factor reducing the reproductive success of the host species and potentially affecting the beetle abundance in the area.     

Fitness consequences of superparasitism and mechanism of host discrimination in the stemborer parasitoid Cotesia flavipes

The fitness consequences of superparasitism and the mechanism of host discrimination in Cotesia flavipes, a larval parasitoid of concealed stemborer larvae was investigated. Naive females readily superparasitized and treated the already parasitized host as an unparasitized host by allocating the same amount of eggs as in an unparasitized host. However, there was no significant increase in the number of emerging parasitoids from superparasitized hosts due to substantial mortality of parasitoid offspring in superparasitized hosts. Furthermore, the developmental time of the parasitoids in a superparasitized host was significantly longer than in a singly parasitized host and the emerging progeny were significantly smaller (body length and head width). Naive females entered a tunnel in which the host was parasitized 4 h previously and accepted it for oviposition. Experienced females (oviposition experience in unparasitized host) refused to enter a tunnel with a host parasitized by herself or by another female. In experiments where the tunnel and/or host was manipulated it was demonstrated that the female leaves a mark in the tunnel when she parasitizes a host. The role of patch marking in C. flavipes is discussed in relation to the ecology of the parasitoid.     

Alteration of hemolymph polypeptides in Manduca sexta larvae parasitized by Cotesia congregata: A two-dimensional electrophoretic analysis and comparison with major bacteria-induced proteins

Analyses using one-dimensional sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE) previously demonstrated that parasitization by the braconid wasp Cotesia congregata significantly alters the normal hemolymph polypeptide profile of host Manduca sexta larvae. In the present study two-dimensional gel analyses corroborated our earlier findings and provided additional evidence that multiple parasitism-specific polypeptides were induced, which varied according to the stage of development of the wasps. Parasitization additionally elicited changes in the total protein concentration detected in the blood. Initially an elevation was observed, with newly parasitized larvae exhibiting a twofold elevation in hemolymph protein concentration by 12–24 h postoviposition. In contrast, terminal-stage hosts with second instar parasites had significantly less protein in the hemolymph, likely due to reduced growth and inhibition of arylphorin synthesis by the fat body during the final stages of parasitism. Comparison of the array of hemolymph polypeptides produced in unparasitized larvae injected with 10⁶cells of the gram-negative bacterium Enterobacter cloacae with those proteins induced by parasitization indicated the two classes are different. Our findings confirm that the hostresponse to parasitism is a specific one, and not mimicked by bacterial challenge. Duringshort-term in vitro culture of wasp larvae dissected from the host hemocoel, several proteins were detected in the medium using SDS - PAGE, with their appearance in vitro suggestive of secretion by the wasps in vivo. Moreover, hemolymph from the parasites had significant amounts of putative host proteins, including an arylphorin - like polypeptide and a protein with a mobility similar to that of insecticyanin. Thus, a dynamic interchange of proteins may occur, with the parasites accumulating host proteins while simultaneously secreting a variety of factors into the host hemocoel.     

Biological notes on a fungus-growing ant, Trachymyrmex cf. zeteki (Hymenoptera, Formicidae, Attini) attacked by a diverse community of parasitoid wasps (Hymenoptera, Diapriidae)

A number of wasps in the family Diapriidae, subfamily Diapriinae (Proctotrupoidea), are parasitoids that specialize on ant larvae. These wasps are abundant and diverse in the Neotropics, but little is known about their biology. We studied parasitism rates by an array of diapriine wasps that attack the larvae of fungus-growing ants, Trachymyrmex cf. zeteki, in a single population (near Gamboa, Panamá). Relatively little is known about the biology and natural history of these ants, so we also present data on colony size and nest architecture. We excavated 136 colonies in central Panamá from June to September 2006, and 20 nests from July 2009. We reared six wasp morphotypes; two of them in the genus Mimopriella Masner and Garcia, one Oxypria Kieffer, two Szelenyiopria Fabritius and one Acanthopria Ashmead. The mean intensity of larval parasitism per ant colony was 33.9% (2006), and its prevalence across all ant populations was 27.2% (2006 and 2009). Parasitism rates were not positively correlated with host colony size. A single case of super-parasitism was documented in which two Oxypria males were reared from the same host larva.     

BIRD-INSECT NESTING ASSOCIATIONS IN AUSTRALIA.

Australian records, beginning in 1866, of birds associating with aggressive insects for nesting purposes are assembled. Five species of the genus Gerygone are known to associate with wasps, more or less consistently, in tropical areas, and another species is reported as associating with wasps in the tropics and ants in temperate areas. Such birds deliberately seek the society of the insects, by which they are recognized and tolerated.
If protection is the motive, it would seem to be aimed against reptiles rather than against other birds or mammals. Other possible reasons for the practice ate suggested.
Australian records of insects frequenting birds' nests also are summarized; these include parasites such as the blood-sucking larvae of a fly, together with moth, beetle and fly larvae that serve a beneficial purpose as scavengers, and, in addition, wasps and beetles parasitic on the other insects. On the question of how the insects find the nests at appropriate times, it is suggested that some are guided by odour and others by some undetermined influence.     

Differential responses to related hosts by nesting and non-nesting parasites in a brood-parasitic duck

Host-parasite relatedness may facilitate the evolution of conspecific brood parasitism, but empirical support for this contention remains inconclusive. One reason for this disparity may relate to the diversity of parasitic tactics, a key distinguishing feature being whether the parasite has a nest of her own. Previous work suggests that parasites without nests of their own may be of inferior phenotypic quality, but because of difficulties in identifying these parasitic individuals, little is known about their host selection criteria. We used high-resolution molecular maternity tests to assign parasitic offspring to known parasites with and without their own nests in a population of Barrow's goldeneyes (Bucephala islandica). We determined whether parasite nesting status, host-parasite relatedness and distance between host and parasite nests affected the probability of parasitizing a host and the number of eggs laid per host. We also investigated whether nesting parasites, conventionally nesting females and non-nesting parasites differed regarding their age, structural size, body condition, nesting phenology or total brood size. The probability of engaging in parasitism increased with host-parasite relatedness and spatial proximity to host nests for nesting and non-nesting females alike. However, nesting parasites increased the number of eggs donated with relatedness to the host, while non-nesting parasites did not do so. Non-nesting parasites laid fewer eggs in total, but did not differ by any of the other quality measures from conventional nesters or nesting parasites. Our study provides the first demonstration that nesting and non-nesting parasites from the same population may use different host selection criteria.     

Neotrombicula autumnalis (Acari, Trombiculidae) as a vector for Borrelia burgdorferi sensu lato?

Larvae of the trombiculid mite Neotrombicula autumnalis were collected at 18 sites in and around Bonn, Germany, to be screened for infection with Borrelia burgdorferi s.l. by means of PCR. Questing larvae numbering 1380 were derived from the vegetation and 634 feeding ones were removed from 100 trapped micromammals including voles, mice, shrews and hedgehogs. In a laboratory infection experiment, a further 305 host-seeking larvae from the field were transferred onto Borrelia-positive mice and gerbils, and examined for spirochete infection at various intervals after repletion. In three cases borrelial DNA could be amplified from the mites: (1) from a larva feeding on a wild-caught greater white-toothed shrew (Crocidura russula), (2) from a pool of four larvae feeding on a B. garinii-positive laboratory mouse, and (3) from a nymph that had fed on a B. afzelii-positive laboratory gerbil as a larva. In the first case, borrelial species determination by DNA hybridization of the PCR product was only possible with a B. burgdorferi complex-specific probe but not with a species-specific one. In the second case, probing showed the same borrelial genospecies (B. garinii) as the laboratory host had been infected with. In the latter case, however, DNA hybridization demonstrated B. valaisiana while the laboratory host had been infected with B. afzelii. Subsequent DNA sequencing confirmed much higher similarity of the PCR product to B. valaisiana than to B. afzelii indicating an infection of the mite prior to feeding on the laboratory host. The negligible percentage of positive mites found in this study suggests that either the uptake of borrelial cells by feeding trombiculids is an extremely rare event or that ingested spirochetes are rapidly digested. On the other hand, the results imply a possible transstadial and transovarial transmission of borreliae once they are established in their trombiculid host. However, unless the transmission of borreliae to a given host is demonstrated, a final statement on the vector competence of trombiculid mites is not possible.     

The effects of host age and superparasitism by the parasitoid, Microplitis rufiventris on the cellular and humoral immune response of Spodoptera littoralis larvae

To study the dynamics of stage-dependent immune responses in Spodoptera littoralis (Boisd.) larvae (Lepidoptera: Noctuidae), single and superparasitism experiments were carried out using the parasitoid Microplitis rufiventris Kok. (Braconidae: Hymenoptera). Compared to younger (preferred) host larvae, the older (non-preferred) host larvae displayed a vigorous humoral response that often damaged and destroyed the single wasp egg or larva. Superparasitism and host age altered both the cellular and humoral immune responses. Younger host larvae showed a stronger encapsulation response compared to older host larvae. Moreover encapsulation rates in younger hosts (e.g., second instar) decreased with increasing numbers of parasitoid eggs deposited/larvae. In older larvae, the encapsulation rate was low in fourth, less in fifth and absent in sixth instar hosts. Conversely, the order and magnitude of the cellular immune response in S. littoralis hosts were highest in second instar larvae with the first instar larvae being a little lower. The immune response steadily decreased from the third through to the fifth instar and was least obvious in the sixth instar. In contrast, the general humoral immune response was most pronounced in sixth instar larvae and diminished towards younger stages. The results suggest that both cellular and humoral responses are stage-dependent. Wasp offspring in younger superparasitized host larvae fought for host supremacy with only one wasp surviving, while supernumerary wasp larvae generally survived in older superparasitized larvae, but were unable to complete development. Older instars seem to have a method for immobilizing/killing wasp larvae that is not operating in the younger instars.     

Parasitism rates in relation to nest site in bees and wasps (Hymenoptera: Apoidea)

To account for differences in occurrence of social behavior in different lineages of bees, Michener (1985) hypothesized that ground nests are more easily located by parasites than are twig nests. In the former case parasites search in two dimensions, while in the latter they search in three-dimensional space. One prediction derived from this hypothesis is that ground nests will have higher rates of parasitism than twig nests. A survey of published reports on rates of cell parasitism for 92 species of nesting bees and wasps (Apoidea) shows no significant differences in mean parasitism rates between these two classes of nests. The analyses were repeated at the generic level (N=44), yielding the same pattern. These data may be biased due to phylogenetic effects. Paired comparisons (n=11 pairs) of related taxa that differ in nest site show that ground-nesting taxa more often have higher rates of parasitism than twig-nesters. The use of artificial trap-nests to study twig-nesters significantly enhances the success rate of parasites. This bias, as well as several other limitations, suggests that experimental studies of the host-searching capabilities of parasites and predators may be more efficacious than such comparative tests.     

槲栎超寄生橡子内象甲幼虫的适应对策

栎属植物的橡子常常受到象甲的侵害, 对橡子存活产生影响, 但有关橡子对象甲幼虫适合度影响的研究尚未见报道。本研究旨在通过对槲栎Quercus aliena种子雨进程以及象甲幼虫逃逸过程的调查, 研究二者之间的进化适应关系。结果表明： 当年槲栎的种子产量为51.92±29.26粒/m², 虫蛀率达到42.4%。虫蛀的橡子中65%以上被象甲超寄生, 且显著大于完好的橡子。在种子下落过程中, 早期从橡子中逃逸的象甲幼虫要比后期逃逸的大。对某一超寄生的橡子而言, 首先逃逸出的象甲幼虫要比随后逃逸的幼虫大。另外, 从早期脱落的橡子中逃逸的幼虫明显小于后期脱落的橡子。象甲幼虫的干重随着橡子内象甲幼虫数量的增加而显著下降。象甲幼虫寄生数量与橡子的长径和短径呈现明显的正相关关系。尽管超寄生降低了象甲虫的适合度, 但雌性成体象甲产卵过程中偏好选择大橡子在一定程度上可以减少这种不利影响。因此, 小橡子可能更容易逃脱象甲的超寄生而获得较大的存活机会。     

Laboratory Culture and Life History of Heleidomermis magnapapula in Its Host, Culicoides variipennis (Diptera: Ceratopogonidae)

The mermithid parasite Heleidomermis magnapapula was maintained in larvae of the midge Culicoides variipennis for 20 months in enamel pans containing nutrient-rich water and polyester pads as a substrate. Inseminated female mermithids were introduced to the pad surface when the host was in the late second or early third-instar. Host larvae were harvested from the pans 9 days after exposure and held in tap water for nematode emergence. Preparasite yield was positively correlated with female nematode size and averaged 1,267 preparasites/female. Male and female nematodes emerged an average of 12.2 and 13.4 days after host exposure, respectively. Supplemental host food (Panagrellus) during the final days of parasitism did not alter time of emergence. Parasites emerging singly were 64% females, whereas superparasitized hosts yielded males (up to nine/host). Nematode carryover into the adult midge normally occurred at a level of 0.5-2.5%. Parasite load (nematodes/ parasitized individual) in midge adults was lower than that of larvae from the same cohort, and adult midges were more likely to harbor female parasites. Exposure of fourth-instar host larvae resulted in higher levels of adult parasitism (up to 17%).     

Performance of Myrmica ant colonies is correlated with the presence of social parasites

1. The performance of ant colonies depends on different factors such as nest site, colony structure or the presence of pathogens and social parasites. Myrmica ants host various types of social parasites, including the larvae of Maculinea butterflies and Microdonmyrmicae (Schönrogge) hoverfly. How these social parasites affect host colony performance is still unexplored. 2. It was examined how the presence of Maculinea teleius Bergsträsser, Maculinea alcon (Denis & Schiffermüller), and M. myrmicae larvae, representing different feeding and growth strategies inside host colonies, is associated with worker survival, the number of foragers, and colony productivity parameters such as growth and reproduction. 3. It was found that the presence of social parasites is negatively associated with total colony production and the production of ant larvae and gynes. Male production was lower only in nests infested by M. teleius, whereas the number of worker pupae was significantly higher in all types of infested colonies than in uninfested colonies. Laboratory observations indicated that nests infested by Maculinea larvae are characterised by a higher number of foragers compared to uninfested nests but we did not find differences in worker survival among nest types. 4. The observed pattern of social parasite influence on colony productivity can be explained by the feeding strategies of parasitic larvae. The most negative effect was found for M. teleius, which feeds on the largest host brood and eliminates a high number of sexual forms. The strong, adverse influence of all studied parasite species on gyne production may result in low queen production in Myrmica populations exposed to these social parasites.     

The geographic structure of selection on a coevolving interaction between social parasitic wasps and their hosts hampers social evolution

Social parasites exploit societies, rather than organisms, and rear their brood in social insect colonies at the expense of their hosts, triggering a coevolutionary process that may affect host social structure. The resulting coevolutionary trajectories may be further altered by selection imposed by predators, which exploit the abundant resources concentrated in these nests. Here, we show that geographic differences in selection imposed by predators affects the structure of selection on coevolving hosts and their social parasites. In a multiyear study, we monitored the fate of the annual breeding attempts of the solitary nesting foundresses of Polistes biglumis wasps in four geographically distinct populations that varied in levels of attack by the congeneric social parasite, P. atrimandibularis. Foundress fitness depended mostly on whether, during the long founding phase, a colony was invaded by social parasites or attacked by predators. Foundresses from each population differed in morphological traits and reproductive tactics that were consistent with selection imposed by their natural enemies and in ways that may affect host sociality. In turn, parasite traits were consistent with selection imposed locally by hosts, implying a geographic mosaic of coevolution in this brood parasitic interaction.     

Parasitism affects worker size in the Neotropical swarm-founding social wasp, <Emphasis Type="Italic">Polybia paulista</Emphasis> (Hymenoptera,Vespidae)

Summary We discovered two kinds of parasites, i.e., a strepsipteran, possibly Xenos myrapetrus (Trois) and an undescribed gregarine in the Neotropical swarm-founding paper wasp, Polybia paulista (Ihering). Although proportions of workers that were infected by these parasites varied greatly among colonies analyzed, prevalence of infected workers was recognized. Five external characters were measured and compared among uninfected workers, stylopized (i.e., infected by Strepsiptera) workers and workers that were infected by gregarines. Uninfected workers were significantly larger than stylopized workers, while smaller than workers that were infected by gregarines. Nutrients of stylopized workers may be plundered during their growth period, and consequently their body size may be reduced. However, the gregarines may manipulate host larvae to solicit more food from adults or increase development time of larvae longer, and therefore produce more parasites from a larger host.Received 1 July 2003; revised 1 December 2003; accepted 4 December 2003.