Life history strategy and taxonomic position of gall midges (Diptera: Cecidomyiidae) inducing leaf galls on Styrax japonicus (Styracaceae)

Four gall midge species (Diptera: Cecidomyiidae) that induce leaf galls on Styrax japonicus (Styracaceae) were identified to generic level based on larval morphology. Three of these gall midges, which induce whitish hemiglobular galls, flattened subglobular galls, and purple globular galls, respectively, were identified as three genetically distinct species of Contarinia, and the remaining species, which induces globular galls with dense whitish hairs, was identified as a species of Dasineura. Field surveys in Fukuoka, Japan, revealed that adults of these gall midges emerged and oviposited in late March to mid‐April at Mount Tachibana (approximately 200 m a.s.l.) and in late April to early May at Mount Sefuri (about 1050 m a.s.l.), coinciding with the leaf‐opening season of S. japonicus. Larvae of these gall midges mostly developed into third instars by June and then left their galls and dropped to the ground. These species therefore have a life history strategy that differs from that of another S. japonicus‐associated gall midge, Oxycephalomyia styraci, which overwinters as the first instar in ovate swellings, matures rapidly in spring, and emerges directly from the galls.     

A new Asphondylia species (Diptera: Cecidomyiidae) and a eulophid wasp (Hymenoptera) inducing similar galls on leaf buds of Schoepfia jasminodora (Schoepfiaceae), with reference to their ecological traits and a description of the new gall midge

Different gall inducers belonging to distinct insect orders are rarely known to induce similarly shaped galls on the same host plant organs. We report that Asphondylia tojoi Elsayed & Tokuda sp. nov. (Diptera: Cecidomyiidae) and Ceratoneura sp. (Hymenoptera: Eulophidae) induce galls on leaf buds of Schoepfia jasminodora Sieb. et Zucc. (Schoepfiaceae). We describe the gall midge species as new to science and report a phylogenetic analysis for known Japanese Asphondylia species. We also describe life histories of the two species, based on monthly surveys during 2015–2017: although both species are multivoltine, A. tojoi overwinters as first instars in galls, whereas Ceratoneura sp. possibly does so as adults outside the galls. In addition, the internal structure of galls differed between the two species. Galls containing A. tojoi consist of a single chamber with inner walls clearly covered with whitish fungal mycelia after the gall midges develop into second instars. Those containing the Ceratoneura sp. have multiple chambers with hard black inner walls. Although some eulophids are known to be inquilines of galls induced by Asphondylia species, we consider that the Ceratoneura sp. is probably a true gall inducer because of the different gall structure and absence of fungal mycelia in their galls. This is the first report detailing the annual life history of a Ceratoneura species. Asphondylia tojoi represents the first example of monophagous Asphondylia species with a multivoltine life history on a deciduous tree.     

Dasineura ingeris sp.n. (Diptera: Cecidomyiidae) on Salix viminalis in Sweden, including comparisons with some other Dasineura species on Salix

Infestations of an undescribed gall midge species were discovered in Southern Sweden in biomass plantations and nurseries of Salix viminalis L. Terminal leaf buds are damaged and side shoots subsequently develop. This midge, Dasineura ingeris sp.n., closely resembles three other species occurring on Salix: D.terminalis (H. Loew) on S.alba L. and S.fragilis L.; D.iteobia (Kieffer) recorded from S.caprea L. and S.cinerea L.; and D.schreiteri (Stelter) (= comb.n. for Rabdophaga schreiteri Stelter) originally found as an inquiline in the galls of another gall midge, Dasineura rosaria (H. Loew), on S.repens L. No qualitative differences in morphology were observed between these four gall midge species, but results of morphometric analyses show significant differences between all of them. In oviposition preference trials, which included host plants of all four midge species, D.ingeris laid eggs mainly on S. viminalis, D.schreiteri preferably on S.repens , and D.iteobia exclusively on S.caprea. In larval performance trials D.ingeris produced many galls on S. viminalis , one gall also on S.caprea , but no galls on either S.alba or S.repens.     

Life history strategy and adult and larval behavior of Macrodiplosis selenis (Diptera: Cecidomyiidae), a species that induces leaf‐margin fold galls on deciduous Quercus (Fagaceae)

Life historical, behavioral and ecological traits of Macrodiplosis selenis, which induces leaf‐margin fold galls on Quercus serrata, Q. mongolica and Q. dentata (Fagaceae) in Japan and South Korea, were studied. Daily activity and larval development indicate that M. selenis is a diurnal and univoltine gall midge. In April, females lay their eggs both on upper and under surfaces of fresh leaves. The duration of the egg stage varies from 5 to 9 days, depending on daily temperatures. Hatched larvae crawl to the upper surface of the leaf margin, where they start to induce galls. Larvae become full‐grown in October, drop to the ground in November and overwinter in cocoons on the ground, while larvae of congeners mature in May and drop to the ground in June. A relatively long period of the second larval stadium from July to October on the host trees seems to be effective for M. selenis in avoiding summer mortalities caused by predation and aridity on the ground and by ectoparasitoids that attack mature larvae or pupae on the host leaves. The spatial distribution pattern of M. selenis leaf galls is contagious and the mean gall density per leaf is significantly correlated with the mean crowding. This study adds new insights of life history strategy and adult and larval behavioral pattern to the ecological knowledge of gall midges, and these kinds of information are essential for further studies of M. selenis population dynamics and interactions with other Quercus‐associated herbivores.     

Preference–performance relationship in the gall midge Rabdophaga rosaria: insights from a common‐garden experiment with nine willow clones

1. Oviposition preferences of herbivorous insects are predicted to match offspring performance on different host taxa or on conspecific host genotypes. In gall‐inducing insects, host‐plant properties such as growth rate and gall size, which are determined by plant genotype and growing conditions, may have a significant impact on offspring performance and, hence, should influence oviposition site selection. 2. The present study investigated host preference of the European rosette willow gall midge Rabdophaga rosaria (Loew) in relation to offspring success on seven clones of Salix myrsinifolia Salisb. and two naturally hybridised S. myrsinifolia × phylicifolia L. clones growing in a replicated design in an experimental field under two fertilisation regimes. For each clone, the average growth rate, number of shoot tips, and leaf and gall size were determined, and their effects on midge preference and larval survival were examined. 3. Main shoot height, number of shoot tips, and gall size were significantly affected by clone. The midges clearly preferred certain clones over the others, but preferences were not related to willow growth traits or to gall size. Survival probability was higher in large than in small galls, but females did not prefer large‐leaved clones that produced the biggest rosette galls. Midge oviposition was also uncorrelated with prior rates of leaf‐rust infection and with feeding preferences of voles and folivorous insects. 4. The weak preference–performance relationship of R. rosaria within S. myrsinifolia is probably explained by evolutionary constraints that prevent generalist insects from achieving an ability to discriminate among conspecific hosts of variable quality.     

The problem of optimal clutch size in a tritrophic system: the oviposition strategy of the thistle gallfly Urophora cardui (Diptera,Tephritidae)

The problem of optimal clutch sizes is a central theme in life history theory. Optimal allocation of eggs is especially complicated for insects in tritrophic systems. In this study we analyze some of the processes determining clutch sizes of the thistle gallfly Urophora cardui, a monophagous tephritid fly associated with Cirsium arvense. U. cardui forms multilocular shoot galls, which vary broadly in their size and number of their gall cells. We investigate various fitness consequences of gall size. An analysis of the number of cells per gall (which is correlated with gall diameter and gall weight) showed that in U. cardui there is mutual facilitation rather than larval competition. Increasing numbers of larvae per gall led to a decreasing mortality and increasing larval weight. Larval weight in turn was positively correlated with the probability of survival to adulthood and with adult weight and fecundity. Thus, all fitness parameters measured favoured large galls. Clutch sizes in oviposition experiments were distinctly larger than the number of gall cells of field populations and in cage experiments, suggesting high mortality of eggs and/or early larval instars. There was a significant relationship between the internal structure (i.e., the size of the growing point) of the bud and clutch size, suggesting that U. cardui females are able to measure bud quality and adapt clutch sizes accordingly. Clutch size was positively correlated with the female's age at first oviposition and negatively with the number of previous ovipositions and previously laid eggs. Since the potential egg capacity per female is higher than the average number of larvae it is likely to produce during its short adult lifespan, U. cardui females tend to be time-limited rather than egglimited, which might favour large clutches once an appropriate oviposition site has been located. As the development of the gall and hence the fate of a clutch depends on a number of unpredictable factors, exclusive concentration of eggs in a few large clusters would involve risks which could be avoided by increasing the number of clutches. Therefore we interpret the high variation of clutch sizes in U. cardui as a mixed strategy of bet hedging and gambling.     

Oviposition choice and larval survival of Dasineura marginemtorquens (Diptera: Cecidomyiidae) on resistant and susceptible Salix viminalis

Abstract.

• 1 Substantial intraspecific variation exists in Salix viminalis resistance to the gall midge Dasineura marginemtorquens. Earlier work has found this variation to have a large genetic component. Willow clones are stable in their resistances between midge generations and different nutrient levels in both field and laboratory culture.
• 2 This study reports the results of laboratory experiments on female oviposition choice and larval survival on potted plants from clones that are very different in resistance as determined in field studies.
• 3 In choice experiments using pairs of plants, the average female midge did not prefer susceptible willow clones over resistant ones for oviposition. In about one third of the replicates, midges actually laid more eggs on the resistant clone. Further work is necessary to examine the nature of variation among midges in discrimination of these plant types.
• 4 Resistance is manifested as great differences in larval survival. Six days after oviposition survival was 92% on susceptible plants but only 6% on resistant ones. Galls developed on all of the susceptible plants, while in 73% of the resistant plants galls were not even initiated.
• 5 The plant traits causing resistance are enigmatic. Larval behaviour suggests that resistant plants interfere with feeding behaviour. On resistant plants, most larvae wander for more than 24 h without initiating any galls before dying. On susceptible plants many first instar larvae begin feeding and initiate galls within this period.

     

Oviposition site of and gall formation by the fruit gall midge Asphondylia aucubae (Diptera: Cecidomyiidae) in relation to internal fruit structure

Most gall insects use young developing plant organs for gall formation; however, little information is available on the histological identification of such tissues or the changes in their availability with plant growth. We investigated the oviposition site of and the tissue used for gall formation by the midge Asphondylia aucubae Yukawa and Ohsaki, which is responsible for galls on the fruit of Aucuba japonica Thunb., by comparing the internal structures of young developing fruit, mature intact (uninfested) fruit, and galled fruit. The midge deposited eggs between the integument and the carpel of young fruit. Larval chambers were made of callus‐like tissue and were formed between the embryo sac and the carpel, where the integument was initially situated. The integument and part of the carpel were thus identified as critical plant tissues used by A. aucubae in forming galls. The integument degenerates in mature intact fruit; therefore, the season of emergence and oviposition by the midge may be determined by the timing of integument degeneration.     

Ecological divergence among morphologically and genetically related Asphondylia species (Diptera: Cecidomyiidae), with new life history data for three congeners including the Alpinia fruit gall midge

Ecological data is crucial for determining the degree of reproductive isolation among closely related species, and in identifying the factors that have produced this divergence. We studied life history traits for three Asphondylia (Diptera: Cecidomyiidae) species that induce fruit galls either on Alpinia, Ligustrum or Aucuba, and we compared the traits with those published for three other closely related Japanese Asphondylia species. We found that the six species were significantly differentiated in important life history traits, such as host range, voltinism, lower developmental threshold temperature, thermal constant and diapausing season. The data indicate that divergence in the assessed life history traits evolves before morphological divergence, and such ecological divergence could strengthen isolating barriers among the taxa. We present scenarios on how host range expansion, host plant shift and host organ shift for galling initiate the early stages of speciation. We also highlight the importance of ecological data in identifying cryptic species. Specifically, we confirm that Alpinia intermedia (Zingiberaceae) is not an autumn–spring host of the soybean pod gall midge Asphondylia yushimai based on many differences in the life history traits between the Alpinia fruit gall midge Asphondylia sp. and A. yushimai.     

Intra‐plant heterogeneity influences the preference and performance of juveniles and adults of a gall midge

Abstract 1. Field studies were conducted to evaluate the preference and performance of a gall‐inducing midge (Harmandia tremulae) within the crown of trembling aspen (Populus tremuloides). Females did not select oviposition sites preferentially within leaves, but did lay preferentially on young leaves. 2. Larvae were the only life stage involved in gall site selection within leaves and in gall initiation and development. Gall size, which was positively related to survival, was highest for galls on mid veins that were located close to the petiole. However, one‐third of galls were located on lateral veins and most galls were not adjacent to the petiole, indicating that many larvae choose sub‐optimal gall initiation sites. 3. Gall density was positively associated with leaf length, and leaf length, was positively associated with gall size. However, gall density per leaf was not related to larval survival in galls. This latter result may be a result of an observed inverse relationship between gall size and gall density for similar‐sized leaves. 4. The results partially support the plant vigour and optimal plant module size hypotheses, which predict that galler fitness in successfully induced galls should be highest on large, fast‐growing plant modules. The lack of a strong preference‐performance link supports the confusion hypothesis, which predicts that oviposition and gall site selection may often be suboptimal in systems where galler lifespan is short. This study suggests that small‐scale variations in plant quality within leaves, can render gall site selection by juveniles as important as that previously reported for adult females.     

Phytohormones in Japanese Mugwort Gall Induction by a Gall-Inducing Gall Midge

A variety of insect species induce galls on host plants. Liquid chromatographic/tandem mass spectrometric analyses showed that a gall midge (Rhopalomyia yomogicola) that induces galls on Artemisia princeps contained high levels of indole-3-acetic acid and cytokinins. The gall midge larvae also synthesized indole-3-acetic acid from tryptophan. Close observation of gall tissue sections indicated that the larval chamber was surrounded by layers of cells having secondary cell walls with extensive lignin deposition, except for the part of the gall that constituted the feeding nutritive tissue which was composed of small cells negatively stained for lignin. The differences between these two types of tissue were confirmed by an expression analysis of the genes involved in the synthesis of the secondary cell wall. Phytohormones may have functioned in maintaining the feeding part of the gall as fresh nutritive tissue. Together with the results in our previous study, those presented here suggest the importance of phytohormones in gall induction.     

Host range expansion by Rhopalomyia yomogicola (Diptera: Cecidomyiidae) from a native to an alien species of Artemisia (Asteraceae) in Japan

In 2001, subconical galls were found on the leaves of an alien Artemisia species (Asteraceae) in Ibusuki City, Kagoshima Prefecture, Japan. These galls were quite similar to those induced by Rhopalomyia yomogicola (Diptera: Cecidomyiidae) on Artemisia princeps, Artemisia montana, and Artemisia japonica in Japan. The morphological features of the pupal head and molecular sequencing data indicated that the gall midge from the alien Artemisia was identical to R. yomogicola. Usually, galling insects do not expand readily their host range to alien plants, but R. yomogicola is considered to have expanded its host range to the alien Artemisia by its multivoltine life history trait and oligophagous habit across two different botanical sections of the genus Artemisia. Adult abdominal tergites and sternites and immature stages of R. yomogicola are described for the first time and detailed biological information is presented.     

Nine new species of Dasineura (Diptera: Cecidomyiidae) from flowers of Australian Acacia (Mimosaceae)

Abstract. Thirteen species of Australian acacias are invasive plants in agricultural and native vegetation areas of South Africa. Biological control programmes for Australian acacias in South Africa have been implemented and are aimed at suppressing reproductive vigour and, in some cases, vegetative growth of these weeds. Gall-forming midges are under consideration as potential biological control agents for invasive acacias in South Africa. Entomological surveys in southern Australia found a diverse cecidomyiid fauna associated with the buds, flowers and fruits of Acacia species. Nine new Dasineura species are described and two species, D. acaciaelongifoliae (Skuse) and D. dielsi Rübsaamen, are redescribed. The newly described taxa are D. fistulosa sp.n. , D. furcata sp.n. , D. glauca sp.n. , D. glomerata sp.n. , D. oldfieldii sp.n. , D. oshanesii sp.n. , D. pilifera sp.n. , D. rubiformis sp.n. and D. sulcata sp.n. All eleven species induce galls on ovaries and prevent the formation of fruit. Two general types of gall are caused. Type A comprises woody, tubular galls with larvae living inside ovaries (D. acaciaelongifoliae, D. dielsi, D. fistulosa, D. furcata, D. glauca, D. glomerata, D. oldfieldii). Type B includes soft-tissued, globose galls that belong to four subtypes: inflated, baglike, hairy galls with larvae living between ovaries (D. pilifera); pyriform, pubescent swellings with larvae living inside ovaries (D. rubiformis); globose, hairy, swellings with larvae living superficially on ovaries in ovoid chambers (D. oshanesii); and inconspicuous, glabrous swellings with larvae living superficially on ovaries in shallow groovelike chambers (D. sulcata). The gall types are associated with a particular pupation pattern. In type A galls, larvae pupate within larval chambers in galls, whereas in type B galls pupation takes place between ovaries in galls or in the soil beneath the host tree. Gall midges responsible for the same general gall type are morphologically related and differ from species causing the other gall type. Phylogenetic analysis of a 410 bp fragment of the mitochondrial cytochrome b gene supports the division of the gall midge species into two groups except for D. sulcata, which appears as a subgroup of the group causing type A galls. The interspecific divergence values in group A species were between 0.5 and 3.9% with intraspecific divergence estimates of 0–0.2%. Gall midges causing type B galls had interspecific divergence values of 4.6–7.3% and intraspecific divergence values of 0–3.7%. Closely related biology and morphology together with low cytochrome b divergence estimates suggest a more recent speciation in group A when compared with species of group B. Dasineura rubiformis and D. dielsi are proposed as potential biological control agents for Acacia mearnsii De Wild. and Acacia cyclops A. Cunn. ex G. Don, respectively, in South Africa due to their narrow host range and ability to form high population densities that reduce seed formation. Both species produce galls with low biomass, which makes them compatible with commercial exploitation of their host species in Africa.     

Influence of synchronization between adult emergence and host plant phenology on the population density of Pseudasphondylia neolitseae (Diptera: Cecidomyiidae) inducing leaf galls on Neolitsea sericea (Lauraceae)

Synchronization between the appearance of herbivorous insects and their host-plant phenology is a critical event, especially for short-lived insects such as gall midges. We studied a natural population of Pseudasphondylia neolitseae (Diptera: Cecidomyiidae) that induces leaf galls on Neolitsea sericea (Lauraceae) to evaluate the effect of synchronization on gall density in the subsequent generation. To do so, we combined quantitative data on host resources with time lag between emergence and host-available seasons. The gamma distribution model was applied to the emergence curve of P. neolitseae and the normal distribution model to the daily changes in the number of host buds suitable for oviposition; the latter model was transformed into an available-resource curve based on the mean number of host buds required for a single female to realize her eggs. By superimposing the emergence curve on the available-resource curve and calculating overlapped area, the degree of synchronization was evaluated more accurately than previous studies, which had treated only the time lag. The number of females that synchronized with host buds affected gall density in the next generation.     

Biology and field host range of Ceutorhynchus cardariae,a potential biological control agent for Lepidium draba

Lepidium draba (Brassicaceae) is a clonal herb, originating from Eurasia, which is invasive in North America. A classical biological control project was initiated in 2001, and the gall‐forming weevil Ceutorhynchus cardariae was prioritized as a candidate agent. We studied its biology and field host range between 2003 and 2014 in the laboratory and a common garden in Switzerland and in the field in Romania. Ceutorhynchus cardariae is a univoltine to bivoltine species. In Switzerland, oviposition usually started at the beginning of March and can occur at temperatures as low as 2.5°C. Galls are formed on stems, leaf stalks and midribs of L. draba rosettes and bolting plants. Gall size increased with an increasing number of larvae per gall. The three larval instars feed inside the galls and leave the plant to pupate in the soil once mature. In Switzerland, development from egg to adult took about 12 weeks in spring. Adults emerged from May to July. After a brief feeding period, adults aestivate. From late summer, feeding recommenced and females may oviposit, forming a partial second generation. Eggs and all larval instars can be found in galls throughout winter. The rate of larval ectoparasitism reached 78%, while endoparasitism was low with a maximum of 2.3%. Lepidium draba populations differed in their suitability for development (number of C. cardariae produced), indicating that effectiveness of C. cardariae – in case released – may be variable. In the field, we observed that gall formation by C. cardariae can severely stunt or even kill shoots. Investigations on the field host range of C. cardariae indicated that only the closely related Lepidium campestre may act as an alternative host for the weevil in Europe. Host‐specificity tests are underway to determine its environmental safety before field release in North America is being considered.     

The role of olfactory cues in the sequential radiation of a gall‐boring beetle,Mordellistena convicta

1. Herbivorous insects often have close associations with specific host plants, and their preferences for mating and ovipositing on a specific host‐plant species can reproductively isolate populations, facilitating ecological speciation. Volatile emissions from host plants can play a major role in assisting herbivores to locate their natal host plants and thus facilitate assortative mating and host‐specific oviposition. 2. The present study investigated the role of host‐plant volatiles in host fidelity and oviposition preference of the gall‐boring, inquiline beetle, Mordellistena convicta LeConte (Coleoptera: Mordellidae), using Y‐tube olfactometers. Previous studies suggest that the gall‐boring beetle is undergoing sequential host‐associated divergence by utilising the resources that are created by the diverging populations of the gall fly, Eurosta solidaginis Fitch (Diptera: Tephritidae), which induces galls on the stems of goldenrods including Solidago altissima L. (Asteraceae) and Solidago gigantea Ait. 3. Our results show that M. convicta adults are attracted to galls on their natal host plant, avoid the alternate host galls, and do not respond to volatile emissions from their host‐plant stems. 4. These findings suggest that the gall‐boring beetles can orient to the volatile chemicals from host galls, and that beetles can use them to identify suitable sites for mating and/or oviposition. Host‐associated mating and oviposition likely play a role in the sequential radiation of the gall‐boring beetle.     

Phenotypic plasticity and similarity among gall morphotypes on a superhost,Baccharis reticularia (Asteraceae)

Understanding factors that modulate plant development is still a challenging task in plant biology. Although research has highlighted the role of abiotic and biotic factors in determining final plant structure, we know little of how these factors combine to produce specific developmental patterns. Here, we studied patterns of cell and tissue organisation in galled and non‐galled organs of Baccharis reticularia, a Neotropical shrub that hosts over ten species of galling insects. We employed qualitative and quantitative approaches to understand patterns of growth and differentiation in its four most abundant gall morphotypes. We compared two leaf galls induced by sap‐sucking Hemiptera and stem galls induced by a Lepidopteran and a Dipteran, Cecidomyiidae. The hypotheses tested were: (i) the more complex the galls, the more distinct they are from their non‐galled host; (ii) galls induced on less plastic host organs, e.g. stems, develop under more morphogenetic constraints and, therefore, should be more similar among themselves than galls induced on more plastic organs. We also evaluated the plant sex preference of gall‐inducing insects for oviposition. Simple galls were qualitative and quantitatively more similar to non‐galled organs than complex galls, thereby supporting the first hypothesis. Unexpectedly, stem galls had more similarities between them than to their host organ, hence only partially supporting the second hypothesis. Similarity among stem galls may be caused by the restrictive pattern of host stems. The opposite trend was observed for host leaves, which generate either similar or distinct gall morphotypes due to their higher phenotypic plasticity. The Relative Distance of Plasticity Index for non‐galled stems and stem galls ranged from 0.02 to 0.42. Our results strongly suggest that both tissue plasticity and gall inducer identity interact to determine plant developmental patterns, and therefore, final gall structure.     

Adult behavior of an ambrosia gall midge Illiciomyia yukawai (Diptera: Cecidomyiidae) and synchronization between its emergence and host plant phenology

The adult behavior of an ambrosia gall midge Illiciomyia yukawai (Diptera: Cecidomyiidae) that induces leaf galls on Illicium anisatum (Illiciaceae) was studied at the population level from 1977 to 1995 in Kagoshima, Japan. Most males emerged between 0:00 and 08:00 h and females between 05:00 and 11:00 h. Males swarmed around the host trees between 05:00 and 11:00 h. Mating occurred on the host leaves mainly between 06:00 and 08:00 h. Females then left the host trees for somewhere else, possibly to collect symbiont fungal conidia. From 08:00 to 16:00 h, females were observed ovipositing into the host shoots. The low development threshold temperature for overwintered larvae was 14°C, while the thermal constant for emergence differed with individuals. Thermal totals above 14°C up to the 50% emergence date varied yearly from 33.1 to 68.7 degree‐days. The 50% emergence date varied from 9 to 18 May. The thermal totals significantly correlated with the 50% emergence date but did not correlate with the date when 50% of shoots became suitable for oviposition. Thus, the host‐plant responded to thermal effects differently from the gall midge. Illiciomyia yukawai has been synchronizing well with the host‐plant phenology but will suffer from asynchrony when global warming becomes more conspicuous.     

Larval distributions of the ectoparasitoid wasp Eurytoma robusta relative to the host tephritid gall fly Urophora cardui

Parasitism may explain the patchy distributions of host populations. The present paper is a study of larval distributions of the parasitoid Eurytoma robusta in galls of the tephritid gall fly Urophora cardui. It focuses on E. robusta's choice of U. cardui gall and whether this changes relative to the rate of parasitism. Oviposition patterns were inferred by direct counts of larvae in galls and genetically, for both species, using indirect relatedness estimates between gall‐members. Furthermore, rates of parasitism in four populations were monitored for 4 years. The modal distribution of E. robusta larvae per gall was one and independent of the level of parasitism. The mean number of E. robusta per gall did not differ from Poisson distributions at different parasitism rates. We were not able to demonstrate a parasitoid preference for gall size. In contrast, parasitoids may have a negative effect on gall growth. Relatedness estimates showed that E. robusta gall members were often unrelated, whereas U. cardui were siblings. Thus, larval distributions of E. robusta suggest that oviposition behaviour is generally constrained and density independent. In four populations monitored over 4 years, parasitism was initially high (up to 70%), but suddenly declined with no apparent effect on fly (gall) abundance.