Survival time and resistance to desiccation of diapause and non-diapause eggs of temperate Aedes (Stegomyia) mosquitoes

Time-related changes in the patterns of host discrimination may have a complex form depending on whether or not hosts are marked following first parasitisation, the persistence of chemicals in these marks, and changes in the host as a result of being parasitised. Here we present an experimental design capable of detecting temporal patterns in host discrimination of solitary parasitoids. This method is tested against experimental data from Aphidius ervi attacking the aphid, Acyrthosiphon kondoi. The results demonstrate that there are statistically significant changes in the patterns of egg distribution in aphid hosts held for intervals between 1 and 48 h between parasitisation. These changes appear to be attributable to increasing oviposition restraint by the superparasitising female as the time interval between attacks is increased. The form of this relationship is not significantly different to a sigmoid function and the rate of change of restraint was greatest between 6–8 h after the first oviposition.     

Intra- and interspecific host discrimination in two closely related egg parasitoids

Intraspecific host discrimination is frequently found in solitary parasitoids, but interspecific host discrimination, where female parasitoids recognize hosts already parasitized by females of other species, is rare. This particular behaviour appears to be adaptive only under specific circumstances. In this paper, we quantified intraspecific host discrimination in Anaphes n. sp. (Hymenoptera: Mymaridae), an endoparasitoid of the eggs of Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) and interspecific host discrimination toward eggs parasitized by Anaphes sordidatus (Girault), a sympatric species competing for the same resource in similar habitats. To examine host discrimination, choice experiments were used where the females had to choose between different categories of eggs (unparasitized, parasitized by Anaphes n. sp. or A. sordidatus). Superparasitism and multiparasitism were avoided in experiments where the female had a choice between unparasitized hosts and hosts parasitized by the same female, by a conspecific or by a female A. sordidatus. When all hosts available were parasitized, conspecific superparasitism occurred more often than self-superparasitism or multiparasitism. These results indicated that females Anaphes n. sp. were capable of self-, conspecific and interspecific discrimination. Self-discrimination followed recognition of an external marking while interspecific discrimination occurred mostly after insertion of the ovipositor. Interspecific discrimination could result from the recent speciation of these species and could be associated with a genotypic discrimination. This behavior appears to be adaptive because of the competition for common hosts between the two parasitoid species.     

Blunt egg pole holds cues for alien egg discrimination: experimental evidence

Eggshell colour patterns play a crucial role in avian host–parasite coevolution. In contrast to many experiments investigating general host egg discrimination abilities, studies testing where specific recognition cues are located on the eggshells (on blunt, sharp or both egg poles) are lacking. Previous studies suggested that discrimination cues might be located at the blunt egg pole, where the shell patterning is typically concentrated. We tested this hypothesis experimentally in species subject to interspecific (great reed warblers Acrocephalus arundinaceus, reed warblers A. scirpaceus), and also intraspecific parasitism (song thrushes Turdus philomelos, blackbirds T. merula). We examined host responses towards two types of intraspecific eggs painted non‐mimetic immaculate blue either at blunt or sharp poles. All four species rejected eggs manipulated at the blunt pole at significantly higher rates, indicating that they perceive the critical recognition cues in the same egg part. Thus, the presence of egg recognition cues at the blunt egg pole may be a general phenomenon in birds parasitized by both intraspecific and interspecific parasites.     

Costs of reproduction and geographic variation in the reproductive tactics of the mosquito Aedes triseriatus

Intraspecific host discrimination is widespread in solitary parasitoids whose adult females forage for and evaluate host suitability, whereas interspecific discrimination is less common. In some parasitoid species, mostly Diptera and Coleoptera, the larva performs the last step of host searching. It has been suggested that host discrimination will rarely occur in such host-seeking larvae because their low mobility results in a low host encounter rate. We determined the extent to which the larvae of Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae), a solitary parasitoid of aggregated Diptera pupae: (1) discriminated between unparasitized hosts and hosts parasitized by conspecifics; (2) used semiochemical cues to discriminate; (3) were influenced by life expectancy, presence of conspecifics and host availability in their host acceptance decision; and the extent to which (4) A. bilineata and A. bipustulata L., a species exploiting the same hosts and occurring sympatrically, showed interspecific host discrimination. A. bilineata larvae were able to discriminate between unparasitized hosts and hosts parasitized by conspecifics in a choice experiment. Such behavior has never previously been described for a coleopteran parasitoid or for a parasitoid species whose larvae perform host searching. Host discrimination in this species was not based on the presence of visual or tactile cues (e.g., entrance holes) but rather on chemical cues. The life expectancy of A. bilineata larvae was significantly shorter in the presence than in absence of hosts, and older larvae had lower parasitism success than young larvae in a 24-h experiment. However, the host acceptance decision of A. bilineata larvae was not influenced by larval age or the presence of conspecifics when the ratio of hosts per larva was greater than or equal to 1. When hosts were scarce, the degree of superparasitism increased significantly with the number of foraging conspecifics and the age of the larvae. Both species of Aleochara showed intra- and interspecific host discrimination in a choice experiment. In contrast to A. bipustulata, A. bilineata larvae more frequently parasitized hosts parasitized by A. bipustulata than those parasitized by conspecifics. We suggest that host discrimination will be frequent in solitary parasitoids with host-seeking larvae when hosts are aggregated. Received: 4 June 1998 / Accepted: 1 September 1998     

Host discrimination by the solitary endoparasitoid Dolichogenidea tasmanica (Hymenopotera: Braconidae)

Successful parasitism of a host partly depends on a female's assessment of its quality, including whether the host has already been parasitised or not. We conducted experiments to elucidate host discrimination by Dolichogenidea tasmanica (Hymenoptera: Braconidae). It is the most commonly collected parasitoid of light brown apple moth, Epiphyas postvittana (Lepidoptera: Tortricidae). To assess the rate of superparasitism avoidance by D. tasmanica, female wasps were given choices between (1) unparasitised hosts versus freshly self-parasitised hosts, (2) unparasitised hosts versus hosts at 24 h post-self-parasitisation and (3) freshly self-parasitised hosts versus hosts freshly parasitised by a conspecific female. Results confirm that host discrimination occurs in D. tasmanica. Females avoid laying eggs in hosts that have been parasitised by themselves or conspecifics, even though the frequency of first encounter with either an unparasitised or a parasitised host was the same for all choices. Thus, it appears that females are not able to discriminate the host parasitisation status prior to contacting a host, but host acceptance is not random. Host discrimination is time-dependent, with greater avoidance of superparasitism after 24 h. The ability of female D. tasmanica to distinguish healthy from parasitised hosts suggests that it could be an effective biological control agent in regulation of host populations. It should also ensure production efficiency in parasitoid mass-rearing.     

Competition for hosts between two hyperparasitoids of aphids,Dendrocerus laticeps andDendrocerus carpenteri (Hymenoptera: Megaspilidae): The benefit of interspecific host discrimination

The two hyperparasitoids of aphids, Dendrocerus carpenteri(Curtis) and Dendrocerus laticeps(Hedicke), differ in their interspecific host discrimination behavior: D. carpenterireadily accepts hosts for oviposition previously parasitized by D. laticeps,whereas D. laticepsoften refrains from ovipositing on hosts previously parasitized by D. carpenteri.The two species also differ in their competitive abilities: D. carpenteriis competitively superior to D. laticepswhen the time between ovipositions is 1 h. D. carpenteri's superiority probably arises from (a) the action of the venom produced by the female to paralyze the host, as evidenced by the comparison of emergence patterns from naturally vs artificially parasitized hosts, and (b) its more rapid egg development. Because of its competitive abilities, D. carpenterigains only limited benefit from discriminating interspecifically, at least when encountering a host parasitized by a competitively inferior species, such as D. laticeps.In contrast, D. laticepsgains considerable benefit from recognizing hosts previously parasitized by D. carpenteri,as the probability to produce offspring from such hosts is greatly reduced.     

Host discrimination and interspecific competition of Trissolcus nigripedius and Telenomus gifuensis (Hymenoptera: Scelionidae), sympatric parasitoids of Dolycoris baccarum (Heteroptera: Pentatomidae)

Trissolcus nigripedius Nakagawa and Telenomus gifuensis Ashmead (Hymenoptera: Scelionidae) are solitary egg parasitoids of Dolycoris baccarum L. (Heteroptera: Pentatomidae), a polyphagous seed-sucking bug of agricultural crops. Field examinations revealed that the two parasitoids are the most common species, but only a single species emerge from a host egg mass. To explain this observation we tested two hypotheses of interspecific host discrimination and asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis. Trissolcus nigripedius and Te. gifuensis could discriminate host egg pre-parasitized by either self or conspecific like other scelionid parasitoids that use external mark on host eggs. When provided with host eggs pre-parasitized by each other, both Tr. nigripedius and Te. gifuensis multiparasitized 94% and 100% of the host eggs without interspecific host discrimination, respectively. Interestingly, from the multiparasitized eggs irrespective of oviposition sequence, progeny of Te. gifuensis always survive better than Tr. nigripedius. Telenomus gifuensis is superior in immature competition probably due to shorter egg incubation period after oviposition, hence progeny of Te. gifuensis become first instar faster than that of Tr. nigripedius. However, adult Tr. nigripedius is always superior competitor in possessing and guarding the host eggs even after oviposition against Te. gifuensis. Therefore, the asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis may explain the emergence of a single species from a host egg mass in the field in spite of no interspecific host discrimination.     

Efficiency,intrinsic competition and interspecific host discrimination of Copidosoma desantisi and Trichogramma evanescens,two parasitoids of Phthorimaea operculella

Trichogramma evanescens West. (Hymenoptera: Trichogrammatidae) and Copidosoma desantisi Annecke & Mynhardt (Hymenoptera: Encyrtidae) are potential parasitoids of the potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae) in Egypt. Discrimination of a parasitized host from an unparasitized host would prevent wasting of time, eggs and reduce competition with conspecifics or heterospecifics. Therefore, we evaluated interspecific host discrimination, multiparasitism and intrinsic competition between the two wasp species. In a choice test, females of T. evanescens showed high interspecific host discrimination only when they were offered 2-day-old C. desantisi parasitized and unparasitized PTM eggs. In contrast, C. desantisi showed high host discrimination and preferred unparasitized eggs to PTM eggs harboring 2-h- or 2-day-old T. evanescens’ eggs. We also evaluated the effect of different introduction sequences on the efficacy of the two wasps. Dissection data indicated that the two parasitoids had a negative impact on each other. There was a significant reduction in the total number of deposited eggs as well as total number of parasitized hosts by each parasitoid. Regarding the rearing experiment, the total number of T. evanescens-induced black eggs or C. desantisi formed mummies in combined treatments was significantly lower than in single parasitoid treatments (control). Moreover, C. desantisi was inferior and did not develop from any multiparasitized host regardless of oviposition order. It was suggested that combined release of the two wasps would not elevate rate of parasitism over that of single parasitoid treatments and competition between them would reduce their efficacy.     

Host discrimination of a larval parasitoid: the quick movement of Microplitis demolitor

Many parasitoids discriminate previously parasitised hosts from unparasitised ones to avoid mortality of offspring. Parasitoids that parasitise aggressive hosts such as lepidopteran larvae are known to attack hosts very quickly to avoid being attacked. However, little is known about host discrimination of such quick attacking parasitoids. We investigated host discrimination of Microplitis demolitor (Wilkinson) (Hymenoptera: Braconidae) a quick attacking parasitoid of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). Results showed that ratios of female wasps that rejected the hosts after antennal examination did not differ between parasitised and unparasitised hosts, indicating that M. demolitor did not discriminate hosts by antennal examination. However, 95% of females that inserted ovipositor into unparasitised hosts actually laid eggs, whereas it was only 31% for parasitised hosts, indicating that females discriminated hosts by oviposition insertion. Analyzing video recordings revealed that the ovipositor exploration of the host took 0.3 s. Female wasps that had experienced high-host density of unparasitised hosts readily rejected parasitised hosts, while wasps with experience of low host availability of parasitised hosts tended to accept parasitised hosts. This suggests that host discrimination behaviour of M. demolitor is affected by previous experience of different host availability and host quality.

     

Plant wound compounds from oviposition scars used in host discrimination by a stem-galling sawfly

The stem-galling sawfly Euura lasiolepisuses one or more plant wound compounds resulting from oviposition scars as cues in host discrimination (avoiding sites occupied by conspecifics). Four experiments were conducted to test hypotheses about how Euurapartitions resources. Experiment 1 demonstrated that Euuraavoids ovipositing on nodes with scars from previous ovipositions. Experiment 2 showed no evidence that the sawfly uses oviposition-deterring pheromones and indicated there is a time lag following oviposition before the oviposition scar becomes a deterrent. Experiment 3 showed that sawflies avoid artificially formed scars, demonstrating that a plant cue alone can lead to host discrimination. Experiment 4 showed that visual or tactile cues are not necessary for host discrimination and indicated that a plant wound compound functions as an oviposition deterrent. Both experimental results and field surveys showed that Euuraoviposition scars were more uniformly distributed than expected if sawflies were ignoring previous ovipositions.     

Can imperfect host discrimination explain partial patch exploitation in parasitoids?

1. Host discrimination by Aphidius rhopalosiphi (De Stefani Perez) (Hymenoptera: Braconidae) was first studied on the grain aphid Sitobion avenae (Fabricius) (Homoptera: Aphididae). Females tended to avoid oviposition in hosts parasitised 3 h earlier. No evidence of host discrimination ability on freshly parasitised hosts was suggested, however, and ovipositional experience had no effect on host discrimination. 2. The effects of host discrimination ability on the exploitation strategy of patches containing different proportions of unparasitised hosts and hosts parasitised for 3 h were studied. Females spent less time on patches with a higher proportion of parasitised hosts, reflecting the females' ability to perceive the potential profitability of the patch. This ability may be based on the nature of the hosts encountered (unparasitised or parasitised). 3. Incomplete exploitation of unparasitised hosts was also observed. It seems that this partial exploitation is related to the inability of A. rhopalosiphi to recognise freshly parasitised hosts. As a female may experience a risk of self‐superparasitism during patch depletion, this could promote early departure from incompletely exploited patches. 4. The effect of previous experience on the patch exploitation strategy was also assessed. Females were tested twice on two patches of the same quality. Results suggested that the experience acquired during a previous visit led the females to leave the patch sooner and to lay fewer eggs in parasitised hosts. 5. Patch exploitation strategy may therefore be the result of different factors such as host discrimination and experience. The evolutionary consequences of the results are discussed.     

The effect of egg load and host deprivation on oviposition behaviour in aphidophagous hoverflies

1. Two species of aphidophagous hoverfly, Episyrphus balteatus and Syrphus ribesii, were tested for the effects of egg load and host deprivation on oviposition choices. 2. Egg load affected the total number of eggs laid in E. balteatus but not in S. ribesii, however it did not affect the proportion laid on any one aphid in E. balteatus but did affect the proportion laid on any one aphid in S. ribesii. The rank order of preferences remained unchanged by age or host deprivation. 3. The dominant effects on host choices were aphid species (in both syrphids) and presentation order (in E. balteatus). 4. Being deprived of hosts increased egg load substantially in E. balteatus, and increasing time of deprivation also had an effect on discrimination; there was no effect of host deprivation in S. ribesii. 5. Reasons for these patterns are discussed.     

Superparasitism and host discrimination in the solitary ectoparasitoid Dinarmus basalis

In the solitary ectoparasitoid, Dinarmus basalis (Hymenoptera: Pteromalidae), the occurrence of superparasitism according to the unparasitised host density, and the nature of the host(s) provided was investigated in laboratory studies. In this species superparasitism was observed whatever the experimental conditions used, but the degree of superparasitism depended on the density of its host, Bruchidius atrolineatus (Coleoptera: Bruchidae). Superparasitism was due to successive egg-laying phases on the same host. However, females were able to discriminate between unparasitised hosts and hosts parasitised from 8 h to 72 h beforehand by themselves or by conspecifics. There was no conclusive evidence that superparasitism in the presence of a host parasitised 30 min before was linked to an absence of host discrimination. Host discrimination in this species is achieved by host-quality markers. These are individual-specific markers since conspecific superparasitism rates were often higher than self superparasitism rates. One deterrent substance is emitted by the females during oviposition onto the egg or released by the 16 to 24 h-old egg itself. Another host-quality marker is associated with the presence of a larva on its host. On the other hand, host discrimination ability did not always imply avoidance of superparasitism. In D. basalis there exists a positive relationship between the survival probability of the second egg and the tendency to superparasitise, and superparasitism could therefore result in a significant fitness gain. Under our experimental conditions, D. basalis females exhibited a wide range of oviposition behavioural plasticity in relation to the parasitoid developmental stage, the type of superparasitism, and the encounter rate with unparasitised hosts.     

Quantifying the effects of host discrimination on egg‐laying decision of the cowpea seed beetle,Callosobruchus maculatus

Many parasitic and endophagous insect species are capable of discriminating among the quality of their hosts. However, there is no appropriate way to quantify their discrimination performance. In this study, we quantified how oviposition of the cowpea seed beetle, Callosobruchus maculatus (Fabricius) (Coleoptera: Bruchidae), was affected by the relative contributions of both egg number and host size discrimination. The effect of egg density and resource heterogeneity on these discrimination performances was also explored. Egg‐distribution predictions were made by combining time‐dependent available resource fitness (egg discrimination) and host weight factors (size discrimination). The χ² test was then used for goodness‐of‐fit testing. The effects of both egg and size discrimination on oviposition in environments with different levels of resource heterogeneity were compared. It was found that host size, rather than the number of eggs on the host, plays a larger role in the egg‐laying decision for most individual seed beetles, especially when egg density is high. Host size discrimination behavior was reinforced when the beetles experienced increasing resource heterogeneity, but the performance might reach a plateau. This is the first quantitative evaluation of the effect of host discrimination on egg‐laying decisions of seed beetles.     

Landscape and oceanic barriers shape dispersal and population structure in the island nematode Pristionchus pacificus

In this study evolutionary host plant patterns at ranks from order to species were analysed using spatial evolutionary and ecological vicariance analysis (SEEVA), based on a multigene phylogeny of 45 ascomycete fungal species. The objective was to understand speciation events and host associations in Ophiognomonia (Gnomoniaceae). Species of this genus are perithecial fungi that occur as endophytes, pathogens, and latent saprobes on plants in the families of Betulaceae, Fagaceae, Juglandaceae, Lauraceae, Malvaceae, Platanaceae, Rosaceae, Salicaceae, and Sapindaceae. A second objective was to determine whether speciation events are influenced by host conservatism, host specialization, or host switching at different taxonomic host ranks. Host differences between sister clades were interpreted using the divergence index (D) from the SEEVA analysis, ranging from 0 for no divergence to 1 for maximum possible divergence. Several fungal subclades showed clear patterns of host order/family conservatism (D = 1.00) for hosts in Betulaceae, Fagaceae, Juglandaceae, and Rosaceae. Clear trends of host specialization at host genus and species ranks (D = 1.00) were suggested within these host families. Independent host jumps were observed for two species at the family rank and three at the order rank. As a result of this study, host specificity and specialization is hypothesized as a mechanism that can strongly contribute to speciation patterns in fungal pathogens. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 1–16.     

Identification of the Cues Used in the Host Finding Behavior During the Phoretic Association Ceroptera rufitarsis (Diptera: Sphaeroceridae) and Dung Beetles (Coleoptera: Scarabaeidae)

Ceroptera rufitarsis (Meigen) (Diptera: Sphaeroceridae) is a tiny fly phoretic on several roller dung beetles of the genus Scarabaeus L. (Coleoptera: Scarabaeidae). We investigated the attractiveness of Scarabaeus sacer L. and S. cicatricosus Lucas both under field and laboratory conditions. Several cues potentially responsible for the host finding behavior were tested in order to determine the basis of attraction of Ceroptera rufitarsis to Scarabaeus beetles. Visual cues, especially host movements, constituted the main factor responsible for host finding behavior and host choice while chemical cues did not play any role in host discrimination.     

Evidence for a marking pheromone in host discrimination byCephalonomia Stephanoderis (Hym.: Bethylidae)

The bethylidCephalonomia stephanoderis Betrem is an ectoparasitoid that prefers to oviposit on the prepupae and pupae of the coffe berry borerHypothenemus hampei (Ferrari) (Coleoptera: Scolytidae). It has the ability to distinguish unparasitized from parasitized hosts and rarely lays more than one egg per host. The mechanism of this host discrimination byC. stephanoderis was investigated under laboratory conditions. For this, parasitoid eggs that had been deposited on host pupae were removed and pupae were then offered (individually and collectively) to individual female wasps. A total of 92% of individually offered hosts and 93% of collectively offered hosts were not parasitized. It is concluded thatC. stephanoderis recognizes a marking pheromone deposited into or onto the host, preceding, during, or after oviposition which enables female parasitoids to avoid self and conspecific superparasitism.     

Recognition of heterospecific parasitism: Competition between Aphidiid (Aphidius ervi) and Aphelinid (Aphelinus asychis) parasitoids of Aphids (Hymenoptera: Aphidiidae; Aphelinidae)

The solitary parasitoids Aphidius erviHaliday (Hymenoptera: Aphidiidae) and Aphelinus asychisWalker (Hymenoptera: Aphelinidae) attacked but generally did not oviposit in pea aphids parasitized by the other species. Wasps selectively oviposited in unparasitized hosts when given a choice. Host discrimination depended on the recognition of internal cues. Females of A. asychiseither could not recognize or ignored A. ervi'sexternal host marking pheromone. Under most conditions, A. ervisurvived in superparasitized hosts, killing competing A. asychislarvae by physical attack and possibly physiological suppression. The outcome of larval competition was not affected by oviposition sequence or age difference between larvae; A. asychissurvived only when it had substantially completed larval development before the host was superparasitized by A. ervi.It is suggested that competition for host resources incurs a cost, for the winner in terms of reduced size or increased development time and for the loser in terms of lost progeny and searching time. Consequently, heterospecific host discrimination can be functional. Internal, and probably general, cues enable wasps to recognize and avoid oviposition in hosts already parasitized by an unrelated species.     

Evolutionary history of <Emphasis Type="Italic">Wolbachia</Emphasis> infections in the fire ant <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

Background  

Wolbachia are endosymbiotic bacteria that commonly infect numerous arthropods. Despite their broad taxonomic distribution, the transmission patterns of these bacteria within and among host species are not well understood. We sequenced a portion of the wsp gene from the Wolbachia genome infecting 138 individuals from eleven geographically distributed native populations of the fire ant Solenopsis invicta. We then compared these wsp sequence data to patterns of mitochondrial DNA (mtDNA) variation of both infected and uninfected host individuals to infer the transmission patterns of Wolbachia in S. invicta.     

Host-discrimination by <Emphasis Type="Italic">Phymastichus coffea</Emphasis>, a parasitoid of the coffee berry borer

Phymastichus coffea (LaSalle) (Hymenoptera: Eulophidae) is an African endoparasitoid of the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Scolytidae) that has been introduced to several countries to control this important pest. In the present study we performed a series of laboratory experiments in order to determine if there was evidence of host discrimination and superparasitism in P. coffea. Our choice experiments demonstrate that P. coffea females showed significant preference to attack unparasitized hosts, rather than those parasitized conspecifically. No significant preferences were detected in self-specific attacks between parasitized hosts and the healthy ones. A further dissection of hosts sequentially attacked either self or conspecifically, revealed that there were no more than two eggs per host. As P. coffea is a species that normally allocates two eggs per host in a single attack, we assumed that females were able to attack already parasitized hosts, but they did not lay eggs in them. Based on this fact, we conclude that there is a host discrimination ability in P. coffea females. With respect to the superparasitism by P. coffea using non-choice experiments, there was no significant difference between self-specific or conspecific attacks with respect to the control after one or two successive attacks. Conspecific attacks yielded the largest numbers of eggs after 3rd, 4th and 5th attacks and significant differences were found between this treatment and the control. The maximum number of eggs found in a single host was six individuals (conspecific treatment). These results confirmed that P. coffea usually laid two eggs per host; however, when there are no hosts available, conspecific attacks can result in the superparasitism in this species.