Alien herbivores and native parasitoids: rapid developments and structure of the parasitoid and inquiline complex in an invading gall wasp Andricus quercuscalicis (Hymenoptera: Cynipidae)

Abstract.

• 1 Rapid and substantial changes have occurred in the parasitoid and inquiline community associated with the agamic galls of Andricus quercuscalicis since it invaded Britain in the late 1950s. The number of parasitoid and inquiline species has risen from one to thirteen over a 15-year period. Although the number of species has been relatively consistent over the last 8 years, the species composition has changed considerably and in a highly characteristic way during this period.
• 2 The parasitoid complex can be divided into two broadly distinct sets of parasitoid species; one set attacks only the gall former whereas the other set concentrates on the inquilines living in the wall of the gall.
• 3 The most dramatic change, however, is in the abundance of inquilines which were reported to be virtually absent in earlier studies on this community in Britain. Over a period of only 5 years, between 1988 and 1993, inquiline attack rose from less than 0.01 to an average of 0.26 inquilines per gall. The intensity of inquiline attack is geographically heterogenous, with high inquiline numbers restricted to south-east England. Because of the relatively high specificity of the parasitoids, high inquiline abundance is positively correlated with parasitoid species richness in knopper galls.
• 4 Parasitism rates, particularly on the gall former, were generally low (<10%). Over the last 5 years, however, seven parasitoid species have been consistently recorded and the mortality caused by these species has increased continuously. The species composition of the community associated with this alien gall wasp in Britain has quickly converged to the community known from its native range in continental Europe. Parasitoid species known to attack the galls of A.quercuscalisis on the continent have been recorded from it in Britain for the first time mainly in areas where inquilines have recently become abundant.
• 5 Since rates of parasitism of the gall former are still low, parasitoids are unlikely to play a major role in the population dynamics of this invading gall wasp at present, but the rapidly increasing inquiline and parasitoid attack could be a source of increased mortality for native cynipid species which are the alternative hosts of those parasitoid species.

     

Environmental thermal levels affect the phenological relationships between the chestnut gall wasp and its parasitoids

Studies of thermal level‐related asynchrony in a host–parasitoid relationship are necessary to understand the effects of climate change on new host–parasitoid interactions. In the Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae) and its Chalcidoidea parasitoids, phenological synchrony is assumed to be weather‐dependent in a new area of expansion. To evaluate the effects of environmental thermal regimes on the host, a phenology model for different cynipid stages (larvae, pupae, adults, and adult emergence) and a host–parasitoid phenological estimator are developed in three chestnut fields during two successive growth seasons and subsequently validated in areas with chestnut fields at two different altitudes. Comparisons of the timings of the juvenile and adult stages with those of the parasitoid complex demonstrate that the shortest period of occurrence for cynipids within galls has negative effects on the host–parasitoid relationships at higher temperature levels, thereby increasing phenological asynchrony for some parasitoids species. Reducing the development time of pupae and adults decreases the likelihood of success for some parasitoid species at higher temperature levels. We also record the extension of the gall wasp development time (approximately 15 days) at higher altitudes (linked to a lower mean temperature of approximately 1.5 °C). These results highlight how parasitization on the new hosts is dependent on the host phenology and, in the present study, is limited by the short duration of the presence of the host in galls, which could explain the considerable differences in cynipid gall wasp parasitization recorded at different altimeters.     

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

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

Parasitoid pressure and the radiation of a gallforming group (Cecidomyiidae: Asphondylia spp.) on creosote bush (Larrea tridentata)

Summary We tested the Enemy Impact Hypothesis, which predicts that communities of one tropic level are organized by the tropic level above. In the case of gallforming insect communities, the hypothesis predicts that gall morphology will diverge, minimizing the number of parasitoids shared among species. We used the monophyletic group of gallforming cecidomyiids (Asphondylia spp.) on creosote bush (Larrea tridentata) to test this hypothesis, predicting that species with thicker gall walls should exclude species of parasitoids with shorter ovipositors and have lower levels of parasitism. Of 17 parasitoid species reared from Asphondylia galls on creosote bush, 9 accounted for over 98% of parasitism. Seven of these 9 species had ovipositors long enough to penetrate 10 of 13 gall morphs measured. There was no significant relationship between gall wall thickness and number of associated parasitoid species (r ²=0.01, P>0.05, n=13). There was no relationship between gall wall thickness and types of parasitoid species colonizing galls: parasitoids with the shortest ovipositors colonized all types of gall morphs and were dominant members of the parasitoid assemblages in galls with the thickest walls. Ultimately, there were no significant differences in percent parasitism among Asphondylia species, regardless of gall wall thickness. We found no difference in numbers of associated parasitoids or percent parasitism in galls with different textures (e.g. hairy versus smooth), different locations on the plant or different phenologies. Our results suggest that enemy impact has not influenced the diversity of this gall community. Gall wall thickness, phenology, location on the plant and surface structure do not appear to influence the distribution of parasitoid species. Other explanations are offered to account for diversity in gall morphology among these species.     

Parasitoid and inquiline attack in the galls of four alien, cynipid gall wasps: host switches and the effect on parasitoid sex ratios

1. Four alien cynipid gall wasps of the genus Andricus are established and still spreading in the British Isles. The order, according to the northerliness of their distribution boundary, is: A. corruptrix→A. quercuscalicis→A. lignicola→A. kollari. All four aliens have a sexual generation in spring on Quercus cerris (introduced to Britain) and an agamic generation in autumn on native oak species. 2. For 2 years 1994 and 1995, galls of both generations of the four alien species were sampled at eight sites from the south of England to the north of Scotland to determine the parasitoid and inquiline species that attack the new galls. The spring generations of the invading species shared a parasitoid complex of four pteromalid species. Five species of inquilines and 11 species of parasitoids emerged from the autumn galls. 3. Two colonisation events were recorded for A. lignicola and A. corruptrix. On both occasions, the spring generations were found first at the new sites, indicating that the agamic generation provides the colonisers for these invading species. After colonisation, the galls of both species were attacked by parasitoids in their first season. 4. In spring, the invading species were among the most abundant cynipids at all eight sites. By sampling the whole local community of cynipid galls, it was found that the parasitoid species attacking the spring galls of the invaders seemed to have shifted their attack to the new hosts. 5. The secondary sex ratios of the parasitoid species emerging from the sexual galls of A. quercuscalicis (the smallest of the four) showed a strong and significant male bias at all sites and in both years. Parasitoid emergence from the galls of the sexual generations of the other three species (all about equal in size) was between 60 and 70% male, and variable among sites and between years.     

Organization of a parasitoid community associated with a complex of galls on Atriplex spp. in southern California

Abstract. 1. Atriplex canescens (Pursh) Nuttall and A.polycarpa (Torrey) Watson (Chenopodiaceae) support twelve morphologically distinct gall types in southern California. Thirty-seven common species of parasitoids, predators and inquilines are associated with these galls. 2. The galls incited by eight members of the Asphondylia atriplicis Cockerell (Diptera: Cecidomyiidae) species complex are linked into a single, interacting community through shared hymenopterous parasitoids and inquilines. 3. Cluster analysis (UPGMA) grouped the fifteen most common species of Chalcidoidea into three host guilds of five species each: (1) specialists in tumour stem and blister leaf galls on A.canescens, (2) specialists in woolly stem galls on A.poiycarpa, and (3) generalists that attack all galls. Guild 1 dominated the galls with which it was primarily associated, while guild 3 dominated the remainder. 4. The abundances of the parasitoids of the tumour stem and blister leaf galls were negatively correlated with the abundances of two organizer species, a gall-forming inquiline, Tetrastichus cecidobroter Gordh and Hawkins, and an internal, larval—pupal parasitoid, Tetrastichus sp. B. The abundances of nine of the twelve most common chalcidoids were not correlated with the abundances of all coaccurring species in six other galls. 5. Host seasonality partly determines parasitoid population dynamics and guild structure. Parasitoid dominance increased with gall duration, suggesting that parasitoid competition depends on resource stability. The two continuously available galls were dominated by their specialist guild, while all seasonal galls were dominated by generalists. The subdominant specialists of woolly stem galls may represent competitively inferior species that utilize those galls opportunistically, because of the gall's widespread distribution and 9–10 month yearly availability. 6. Sites in the Colorado Desert and chaparral that supported several gall types showed stable relative abundances of the major parasitoid species, whereas sites in the Mojave Desert that supported only woolly stem galls had unpredictable parasitoid species assemblages. 7. The competitive success of Atriplex gall parasitoids may depend primarily on voltinism (multivoltine species dominated univoltine species) and mode of feeding (phytophagous, mixed entomophagous—phytophagous and facultatively hyperparasitic species in general dominated strict primary parasitoids).     

VARIABLE SELECTION ON EUROSTA'S GALL SIZE. II. A PATH ANALYSIS OF THE ECOLOGICAL FACTORS BEHIND SELECTION

We examined phenotypic selection exerted by natural enemies on the gall-making fly Eurosta solidaginis in an extensive field study of 16 populations, spanning four generations. Gall-makers that induce small galls are vulnerable to the attack of Eurytoma gigantea. This imposes upward directional selection on gall size. Insectivorous birds, predominantly the downy woodpecker, are more likely to attack larvae that induce large galls than small ones, and this imposes downward directional selection. We used path analysis to explore the relative contributions of these natural enemies to the net directional selection on gall size. The path models further examined several ecological factors that influence selection intensity through their effects on parasitoid and bird attack rates. Net directional selection varied more strongly with E. gigantea attack than bird attack. Competitive interactions among birds and the three parasitoid species, including E. gigantea, were evidenced by low winter bird attack rates in fields where a high proportion of galls contained the overwintering parasitoids. Eurytoma gigantea attack was heavier in fields where mean gall size was small and bird attack heavier in fields where mean gall size was large. Neither birds nor E. gigantea showed simple density-dependent attack. Data suggested a form of frequency-dependent attack by birds but not by E. gigantea.     

Variation in selection pressures on the goldenrod gall fly and the competitive interactions of its natural enemies

Summary Larvae of the tephritid fly Eurosta solidaginis induce ball-shaped galls on the stem of tall goldenrod, Solidago altissima. Survival probability depends on gall size; in small galls the larva is vulnerable to parasitoid oviposition, whereas larvae in large galls are more frequently eaten by avian predators. Fly populations from 20 natural old fields in central Pennsylvania were monitored in 1983 and 1984 to examine the distribution of the selection intensity imposed by natural enemies, the parasitoids Eurytoma gigantea and E. obtusiventris, the inquiline Mordellistena unicolor, and the predatory birds Dendrocopus pubescens and Parus atricapillus. Mordellistena and E. obtusiventris are able to attack galls of all diameters while E. gigantea and the predatory birds preferentially assaulted small and large diameter galls, respectively. Eurosta in intermediate sized galls had the highest survivorship, hence selection had a stabilizing component. However, parasitoid attack was more frequent than bird attack, and the two did not exactly balance, thus there was also a directional component. The mean directional selection intensity on gall size was 0.21 standard deviations of the mean, indicating that larger gall size was favored. Interactions among the insect members of the Eurosta natural enemy guild are complex and frequent.     

Can there be an escalating arms race without coevolution? Implications from a host-parasitoid simulation

Summary Under a restricted set of conditions, predator-prey or parasite-host systems may exhibit an escalating arms race over several generations that is not coevolutionary. Preconditions for such a process include high correlation between prey/host quality and defensive capability, and phenotypic plasticity in predator/parasite-counter defenses that responds to quality. We present simulation models based on the parasitoid waspEurytoma gigantea, which lays its egg in the goldenrod gall induced by the flyEurosta solidaginis. For the parasitoid to successfully lay an egg, the gall walls must be thinner than the parasitoid's ovipositor is long. Wall thickness is highly correlated with gall size, so probability of successful attack declines with gall size. However, since the parasitoid eats the gall tissue, individuals developing in small galls have little food and mature with shorter ovipositors than those which develop in large galls. The simulation showed that the population mean parasitoid size is set by mean gall size. Since small galls are more frequently parasitized, there is a selection pressure on the gallmaker to induce larger galls. But, an additional simulation showed that since parasitoid ovipositor length depends on gall size, an evolutionary increase in gall size will also result in a non-evolutionary increase in parasitoid body size and ovipositor length over several generations.     

Very long diapause and extreme resistance to population disturbance in a galling insect

1. Some insects have a prolonged diapause – a dormancy that extends over more than 1 year. In most species prolonged diapause involves one or a few extra years, but in extreme cases diapause may surpass 10 years. Few cases of very long diapause have been described, and very little is known about the population consequences of the temporal refuge formed by the diapausing individuals. 2. The gall midge Contarinia vincetoxici Kieffer galls the flowers of a long‐lived herb Vincetoxicum hirundinaria Med. After completing development, larvae leave the galls for the ground where they enter diapause. Extending an earlier published inoculation experiment, we show that the diapause may last up to at least 13 years, with a median duration of at least 6 years. 3. The gall midge is attacked by two parasitoid species. Dissections of gall midge larvae for presence of parasitoids revealed that Omphale salicis Haliday had a maximum 2 year diapause and Synopeas acuminatus Kieffer a maximum 4 years. The very long diapause of the gall midge may thus provide a temporal refuge from these enemies. 4. In a 15‐year field experiment all galls were removed every year from six isolated habitat patches. Density changes in experimental populations were not statistically different from control populations for over a decade. After 14–15 years a modest decline could be observed. This slow response illustrates that prolonged diapause in C. vincetoxici provides a very strong population buffer against mortality during the galling stage.     

Ant mutualists alter the composition and attack rate of the parasitoid community for the gall wasp Disholcaspis eldoradensis (Cynipidae)

Abstract.  1. The strength or density dependence of pairwise species interactions can depend on the presence or absence of other species, especially potential mutualists.
2. The gall wasp Disholcaspis eldoradensis induces plant galls that secrete a sweet honeydew from their top surfaces while the wasp larvae are active. These galls are actively tended by Argentine ants, which collect the honeydew and drive off parasitoids attempting to attack the gall wasp.
3. When ants were excluded, the total rate of parasitism by seven species of parasitoids increased by 36%, and the rate of gall-wasp emergence decreased by 54%.
4. The total percentage parasitism was affected by gall density when ants were excluded but not when ants were unmanipulated, suggesting a change in parasitoid functional responses due to ant tending.
5. In addition, excluding ants significantly altered the proportions of different parasitoid species that emerged from galls; one parasitoid species increased from 1% to 34%, and another decreased from 46% to 19%.
6. The invasive Argentine ants studied are capable of maintaining the mutualism with the gall wasps that evolved in the presence of different ant species and also act as a selective filter for the local community of generalist parasitoids trying to attack this gall species.     

Non-random distribution among a guild of parasitoids: implications for community structure and host survival

Abstract 1. Immature stages of the gall midge, Asphondylia borrichiae, are attacked by four species of parasitoids, which vary in size and relative abundance within patches of the gall midge’s primary host plant, sea oxeye daisy (Borrichia frutescens). 2. In the current study, a bagging experiment found that the smallest wasp, Galeopsomyia haemon, was most abundant in galls exposed to natural enemies early in the experiment, when gall diameter is smallest, while the wasp with the longest ovipositor, Torymus umbilicatus, dominated the parasitoid community in galls that were not exposed until the 5th and 6th weeks when gall diameter is maximal. 3. Moreover, the mean number of parasitoids captured using large artificial galls were 70% and 150% higher compared with medium and small galls respectively, while stem height of artificial galls significantly affected parasitoid distribution. Galls that were level with the top of the sea oxeye canopy captured 60% more parasitoids compared with those below the canopy and 50% more than galls higher than the plant canopy. 4. These non‐random patterns were driven primarily by the differential distribution of the largest parasitoid, T. umbilicatus, which was found significantly more often than expected on large galls and the smallest parasitoid of the guild, G. haemon, which tended to be more common on stems level with the top of the plant canopy. 5. Large Asphondylia galls, especially those located near the top of the Borrichia canopy, were more likely to be discovered by searching parasitoids. Results using artificial galls were consistent with rates of parasitism of Asphondylia galls in native patches of sea oxeye daisy. Gall diameter was 19% greater and the rate of parasitism was reduced by almost 50% on short stems; as a result, gall abundance was 24% higher on short stems compared with ones located near the top of the plant canopy. 6. These results suggest that parasitoid community composition within galls is regulated by both interspecific differences in ovipositor length and preferences for specific gall size and/or stem length classes.     

The parasitoid community of <Emphasis Type="Italic">Andricus quercuscalifornicus</Emphasis> and its association with gall size,phenology, and location

Plant galls are preyed upon by a diverse group of parasitoids and inquilines, which utilize the gall, often at the cost of the gall inducer. This community of insects has been poorly described for most cynipid-induced galls on oaks in North America, despite the diversity of these galls. This study describes the natural history of a common oak apple gall (Andricus quercuscalifornicus [Cynipidae]) and its parasitoid and inquiline community. We surveyed the abundance and phenology of members of the insect community emerging from 1234 oak apple galls collected in California’s Central Valley and found that composition of the insect community varied with galls of different size, phenology, and location. The gall maker, A. quercuscalifornicus, most often reached maturity in larger galls that developed later in the season. The parasitoid Torymus californicus [Torymidae] was associated with smaller galls, and galls that developed late in the summer. The most common parasitoid, Baryscapus gigas [Eulophidae], was more abundant in galls that developed late in the summer, though the percentage of galls attacked remained constant throughout the season. A lepidopteran inquiline of the gall (Cydia latiferreana [Tortricidae] and its hymenopteran parasitoid (Bassus nucicola [Braconidae]) were associated with galls that developed early in the summer. Parasitoids and inquilines, in general, had a longer emergence period and diapause within the gall than the gall-inducer. The association of different parasite species with galls of different size and phenology suggests that different parasite species utilize galls with slight differences in traits.     

Cecidophagy in adults of Demotina fasciculata (Coleoptera: Chrysomelidae) and its effect on the survival of Andricus moriokae (Hymenoptera: Cynipidae) inhabiting leaf galls on Quercus serrata (Fagaceae)

Females of Demotina fasciculata (Coleoptera: Chrysomelidae) were found to prefer to feed on galls induced by Andricus moriokae (Hymenoptera: Cynipidae) rather than on leaves of its host plant, Quercus serrata (Fagaceae). This is the first record of cecidophagy by adult chrysomelid beetles. Demotina fasciculata did not infest healthy galls induced by another unidentified cynipid species on the same host trees, but did feed on galls inhabited by an inquiline species Synergus quercicola (Hymenoptera: Cynipidae), presumably because such galls remained on the host trees longer than healthy galls. Galls of A. moriokae were infested more severely than cynipid galls inhabited by the inquiline. Therefore, higher density and thicker gall wall in A. moriokae galls seem to make them more suitable targets for D. fasciculata to attack. Larval chambers of A. moriokae galls were stripped by the infestation of gall walls and readily dropped to the ground, resulting in 100% death of cynipid larvae due to desiccation, while 62.5% of pupae survived when they had developed to the late stadium before the fall of larval chambers.     

The effect of hosptlant and parasitoids on the reproductive success of the parthenogenetic gall wasp Diplolepis rosae (Hymenoptera,Cynipidae)

Summary The univoltine cynipid gall wasp Diplolepis rosae reproduces by an obligate homozygosity promoting system known as gamete duplication. The wasp is confined to roses (Rosa spp) on which it induces large, complex and multichambered galls. In southern Sweden, D. rosae was found to parasitize Rosa canina, R. dumalis, R. rubiginosa, R. villosa, R. sherardi and R. rubrifolia, but not R. majalis and R. rugosa. The distribution of galls shows that there are differences in the relation between wasp and hosplant with respect both to species and individual plants. There is a positive correlation between wasp size and gall (clutch) size. Parasitoid pressure was found to be high, causing D. rosae an estimated average larval loss of approximately 75%, mainly due to the attack of the ichneumonid wasp Orthopelma mediator. The very common cynipid inquiline Periclistus brandtii does not seem to have any negative effects. Overall parasitism and probability of no hatched offspring per gall decrease with increasing gall (clutch) size. The probability of loosing all of a given number of offspring decreases with the number of galls produced. It is suggested that D. rosae, in order to escape parasitoids, needs high ability to establish new colonies. Hence the production of many comparatively small galls, which increases the chance of leaving any offspring, rather than the production of few large galls, maximizing the number of offspring, should be favoured by selection.     

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.     

Indirect competitive effects of stemborers on a gall community

Interspecific competition between phytophagous insects using the same host plant occurs frequently and can strongly affect population densities of competing species. Competition between gallmakers and stemborers could be especially intense because both types of herbivore are unable to avoid competition by relocation during their immature stages. For apical meristem gallmakers the main result of competition is likely to be the interruption of resources to the gall by the stemborers' devouring of stem contents. The proximate effect of such competition could be to reduce gall size, thereby increasing the number of chambers per gall unit volume, and reducing the size and potential reproductive output of the gallformer. In addition, smaller galls may be more susceptible to attack from size‐limited parasitoids, resulting in a second indirect effect of competition. Using a community of galling and stemboring insects on the saltmarsh shrub Iva frutescens L. (Asteraceae), we measured for indirect effects of competition. We examined the primary indirect effect of competition on gall midge crowding and the secondary effects on parasitism rates and parasitoid guild composition. Results indicated that galls co‐occurring with stemborers were smaller, crowding of gall inhabitants was 22% greater, and the composition of the parasitoid guild was altered relative to galls on unbored stems. The overall parasitism rate was not different between galls on bored vs. unbored stems. These results show that competition resulting from the presence of stemborers has the potential to affect the gall midge Asphondylia borrichiae Rossi & Strong (Diptera: Cecidomyiidae) and secondarily to affect its guild of hymenopteran parasitoids.     

Biology of Rhoophilus loewi (Hymenoptera: Cynipoidea: Cynipidae), with implications for the evolution of inquilinism in gall wasps

The sole described indigenous afrotropical cynipid, the South African endemic Rhoophilus loewi , was originally described as a gall inducer. As predicted by phylogenetic relationships, we confirm that R. loewi is instead an inquiline wasp. Although all other cynipid inquilines develop in galls induced by cynipid wasps, or very rarely cecidomyiid midges, R. loewi is the only cynipid inquiline of a lepidopteran gall, inducing secondary inquiline cells in galls induced by a cecidosid moth genus Scyrotis on Rhus species (Anacardiaceae). Rhoophilus is a lethal inquiline; its larval cells expand into the hollow interior of the host gall resulting in death of the gall inducer. Rhoophilus loewi is redescribed and the final-instar larva described for the first time. We describe host-plant associations, the morphology and phenology of gall formation, and the suite of parasitoid Hymenoptera associated with these galls. The community centred on Scyrotis galls is compared with those observed in other hosts of inquiline cynipids. Elucidation of the life history of R. loewi has fundamental implications for understanding the evolution of cynipid wasps.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 153–172.     

Determinants of assemblage size for the parasitoids of Cecidomyiidae (Diptera)

Summary The parasitoids known to attack 191 phytophagous species of gall midges (Cecidomyiidae) were used to examine factors influencing parasitoid assemblage size. The number of parasitoid species a midge species supports was tested against nine variables describing geographical, biological and ecological attributes of hosts. The apparency of midge larvae was found to have the greatest influence on parasitoid assemblage size; highly visible species support more parasitoids than less visible ones. Pupation site and midge voltinism also significantly affect associated parasitoids, at least for highly apparent hosts. Biogeographic region, host-plant architecture and the plant parts infested were found to be of secondary importance. The surface texture of infested plant parts, the number of midge larvae occupying galls and the diversity of plant tissues infested have minimal apparent effects on parasitoid richness. Parasitoid assemblage size and total parasitism rates were also found to be positively correlated for 73 galling and nongalling midge species, and gallers typically suffer higher levels of parasitism than non-gallers. Using these data to test the enemy hypothesis, which proposes that the galling habit has evolved to escape attack from parasitoids, we conclude that parasitoid pressure cannot account for the presence of galls in the Cecidomyiidae.     

VARIABLE SELECTION ON EUROSTA'S GALL SIZE,I: THE EXTENT AND NATURE OF VARIATION IN PHENOTYPIC SELECTION

Natural fluctuations in environmental conditions are likely to induce variation in the intensity or direction of natural selection. A long-term study of the insect, Eurosta solidaginins Fitch (Diptera; Tephritidae), which induces stem galls on the perennial herb Solidago altissima (Asteraceae) was performed to explore the patterns of variation in phenotypic selection. The intensity of selection imposed by parasitoids and predators on gallmaking larvae, for gall size, was measured across 16 populations over the course of 4 generations, for a total of 64 population-generations. Directional selection was quantified by i, the selection intensity, and variance selection by j‘, a measure of the intensity of selection on phenotypic variance. Size-dependent attack by parasitoids caused upward directional selection (mean i_p = 0.42; SE = 0.023), while size-dependent bird attack favored larvae that induced smaller galls (mean i_b = -0.07; SE = 0.013. The mean net directional selection intensity was 0.35 (SE = 0.030), which indicates that insects inducing larger galls are generally favored by selection. The opposing patterns of size-dependent attack resulted in stabilizing selection in half the population generations, with an overall average. j‘ of -0.11 (SE = 0.078). The magnitude of directional selection was strongly influenced by the population mean gall size and weakly by the optimal gall size. The intensity of variance selection was strongly influenced by the shape of the fitness function, with sigmoidal and Gaussian-like shapes causing greater depletion of phenotypic variance.