Preference and performance linkage in aPhyllocolpa sawfly on the willow,Salix miyabeana,on Hokkaido

A leaf-folding sawfly in the genusPhyllocolpa (Hymenoptera: Tenthredinidae) attackingSalix miyabeana (Salicaceae) was studied near Sapporo, Hokkaido, along the Ishikari River in 1993. Host plant individuals were young trees 4–7 years old which were growing rapidly, producing some long shoots with large leaves. On a gradient of shoot length classes from 0–5 cm long to over 80 cm long, shoots were much more abundant in the shorter shoot length classes. However, attacks by ovipositing females increased as shoot length increase from 0 attacks on the shortest shoots to 5.17 attacks per shoot on the longest shoots. The frequency of attack per leaf increased from 0 to 0.13 over the same range of shoot lengths. This pattern of attack resulted in a high frequency of larval establishment in feeding sites, between 0.96 and 1.00, in all attacked shoot length categories. However, probability of survival to a late instar larva increased with shoot length and corresponded to the attack pattern, indicating a preference-performance linkage between female ovipositional decisions and larval survival. The patterns found for thisPhyllocolpa species are similar for galling sawflies in North America and Europe, especially in the genusEuura, members of which make stem, bud and leaf midrib galls. Extending the pattern to aPhyllocolpa species broadens identification of pattern and ultimately the generality of the emerging theory on populations of galling sawflies.     

Growth of coyote willow and the attack and survival of a mid-rib galling sawfly,Euura sp.

We studied the relationship between variation in age and shoot characteristics of the host plant Salix exigua Nuttall (coyote or sandbar willow) and the attack and survival of Euura sp. (an unnamed leaf-midrib galling sawfly). Variation in shoot characteristics resulted from reduced growth as willow ramets aged. Mean shoot length per ramet and mean longest leaf length per shoot decreased by 95% and 50% respectively between 1- and 9-year-old willow ramets. All measured shoot characteristics-shoot length, longest leaf length, number of leaves per shoot, and mean internode length-were significantly negatively correlated with ramet age (r ² ranged from –0.23 to –0.41). Correlations between shoot characteristics were highly positive, indicating that plants also grew in a strongly integrated fashion (r ² ranged from 0.54 to 0.85). Four hypotheses were examined to explain sawfly attack patterns. The host-plant hypothesis was supported in explaining enhanced larval sawfly survival through reduced plant resistance. As willow ramets aged, the probability of Euura sp. attack decreased over 10-fold, from 0.315 on 1-year-old ramets to 0.024 on 2- to 9-year-old ramets. As shoot length increased, the probability of sawfly attack increased over 100-fold, from 0.007 on shoots <100 mm, to 0.800 on shoots in the 1001–1100 mm shoot length class. These attack patterns occurred even though 1-year-old ramets and shoots >500 mm each represented less than 2% of the total shoots available for oviposition. Host plant induced mortality of the egg/early instar stage decreased by 50% on longer leaves and was the most important factor determining survival differences between vigorous and non-vigorous hosts. Sawfly attack was not determined by the resource distribution hypothesis. Although shoots <200 mm contained 82% of the total leaves available, they contained only 43% of the galls initiated. The attack pattern also was not explained by the gall volume hypothesis. Although gall volume increased on longer shoots, there was no significant variation in mid or late instar mortality over shoot length, as would be expected if food resources within smaller galls were limited. The natural enemy attack hypothesis could not explain the pattern of oviposition since predation was greater on longer shoots and leaves. In addition, larval survival was related to oviposition behavior. Due to a 69% reduction in late instar death and an 83% reduction in parasitism, survival of progeny in galls initiated close to the petiole base was 2.8 times greater than in galls initiated near the leaf tip. A 75% reduction in gall volume over this range of gall positions may account for the observed increases in late instar mortality and parasitism.     

Preference and performance of a gall-inducing sawfly: a test of the plant vigor hypothesis

To test the Plant Vigor Hypothesis, we determined female oviposition preference of Phyllocolpa leavitii (Hymenoptera: Tenthredinidae) for shoot lengths on 20 clones of Salix discolor (Salicaceae) and examined larval performance by determining larval survival based on clone, shoot length, and leaf length for each Phyllocolpa gall observed. Sawfly galls were found on significantly longer shoots compared to shoots without galls, and shoots with 2, 3, 4, and 5 galls were successively longer than shoots with fewer galls. Furthermore, a much higher proportion of longer shoots had galls that did shorter shoots. These data demonstrate strong sawfly oviposition preference for long shoots. However, when shoot length was adjusted for numbers of available leaves per shoot, the number of galls per available leaf showed a weaker, but still positive relationship with shoot size. Using a logistic regression on survival of Phyllocolpa larvae, we detected highly significant effects of clone, shoot length, leaf length, and the square of leaf length on survival. Additionally, much higher sawfly survival on the long shoots of one clone caused a significant clone by shoot-length interaction. Survival was positively correlated with shoot length, indicating directional selection favoring oviposition on longer shoots; still, shoot length explained only 11.8% of the variation in survival. The directional component of selection favored oviposition on smaller leaves as survival was inversely related to leaf length; however, disruptive selection for leaf length oviposition preference was also detected, with survival of Phyllocolpa galls lowest on intermediate leaf lengths. This study provided evidence supporting the preference prediction of the Plant Vigor Hypothesis. However, much of the data contradicted the performance prediction of the Plant Vigor Hypothesis, with only a modest amount of evidence supporting the performance prediction.     

Patterns of interspecific associations of stem gallers on willows

Abstract.  The pattern of interspecific associations of three stem-galling sawfly species ( Euura atra , E. elaeagnos , E. purpureae ) and three stem-galling gallmidge species ( Rabdophaga sp. 3–5) was investigated on five willow taxa ( Salix alba , S. fragilis , S.  ×  rubens , S. elaeagnos , S. purpurea ) at five natural sites in Central Europe. The willow species harboured specific species associations of two stem gallers, each pair consisting of one Euura and one Rabdophaga species. The stem gallers were patchily distributed and their densities varied significantly among willow host plant species, host plant individuals, and host plant sexes. Four of the six species showed a significant increase in galling rate with shoot length. The other two species were the sawfly and cecidomyiid pair that induce galls on S. purpurea . The preference of stem gallers to longer shoots was generally not related to higher larval performance in terms of survival. Only one species, Rabdophaga sp. 5, was found to be more abundant on male plants. The correlation of densities of the species pairs of stem gallers was independent of willow sexes. Species pairs of stem gallers co-occurring on the same willow species tended to attack different shoots within the same host plant individual. When species pairs co-occurred on shoots they were usually found in similar densities as when occurring alone on shoots. The stem-galling sawflies usually formed galls at the basal part of a shoot, whereas the gallmidge R . sp. 5 ( R . sp. 3 and R . sp. 4 showed no clear tendency) preferred the middle or distal part of a shoot. This is interpreted with differences of their phenology and oviposition period.     

Mixed significance of plant vigor: two species of galling Pontania in a hybridizing willow complex

Distributions of leaf galls and offspring performance of two Pontania sawfly species were explored in individual willows of the subarctic Salix caprea – starkeana hybrid complex. The more common sawfly, an undescribed species near the dolichura group (P1), had the highest gall numbers in trees with long shoots both in S. caprea and hybrids. While numbers were high on vigorously growing hybrids, offspring of P1 were aborted significantly more often on hybrids than on pure hosts. Further, non-aborted galls were smaller on hybrids. Fast shoot growth may be important for P1 sawflies, because females oviposit early in summer and larvae develop rapidly compared with the other species, Pontania pedunculi (P2). Distributions of P2 galls were related to tree height and not to shoot length in both parental and hybrid groups of willows. Like P1, also P2 offspring were frequently aborted on hybrids, but not significantly more often than on pure hosts, and P2 galls were equally large among the host groups. Survival of both species was related to abortion rates, while larvae were parasitized equally in all host groups. This study demonstrates that the significance of plant vigor may vary even for closely related galling sawflies exploiting the same hosts, or for the same species on different host plants. Vigorous growth may mislead gallers to oviposit on suboptimal plants.     

Testing a new hypothesis: plant vigor and phylloxera distribution on wild grape in Arizona

Summary Longer, meaning more vigorous, shoots of a wild grape clone (Vitis arizonica) were more susceptible to attack by second and third generations of leaf-galling grape phylloxera,Daktulopsphaira vitifoliae, as the growing season progressed. Although there was no significant difference in mean shoot length between attacked and unattacked shoots within a clone at the beginning of shoot elongation, attacked shoots were significantly longer than unattacked shoots when elongation had ceased (P<0.01). Also, long attacked shoots had a significantly greater population of phylloxera galls than short attacked shoots (P<0.01) as the season progressed. The phylloxera population on long shoots increased rapidly while the population on short shoots remained the same. Longer shoots also produced significantly more axillary shoots than shorter shoots as the season progressed (P<0.001), and the number of axillary shoots accounted for 66 percent of the variance in number of attacked leaves on a shoot. Experimental evidence showed that there was a significantly greater percentage of available leaves attacked on long shoots than on short shoots (P<0.05) and the leaves on long shoots generally had a greater number of galls per leaf. The relationship between shoot length and probability of attack was also tested by comparing shoots lengths of 10 attacked clones and 10 unattacked clones at a second location. Mean shoot lengths of attacked clones were significantly longer than mean shoot lengths of unattacked clones (P<0.05), and mean shoot lengths of attacked shoots within a clone were significantly longer than unattacked shoots (P<0.001). Longer shoot length accounted for 81 percent of the variance in probability of attack. The reason for this pattern of attack was that long shoots produced newly expanding leaves over a longer time during the growing season and multivoltine phylloxera require undifferentiated tissue to initiate gall formation. Patterns of attack within a shoot were characterized by an uneven distribution of galls among leaves. This was due to development time between generations and the current availability of undifferentiated tissue at times of colonization. This study supports the hypothesis that some herbivore species are favored more by vigorous plants than by stressed plants.     

Induced tree response to herbivory influences the foraging preference and performance of a conifer‐defoliating sawfly

Herbivory often alters the growth and development of woody plants and can thereby render hosts less susceptible to subsequent herbivores. We carried out field surveys and experiments to investigate how previous herbivory influences adult egg lay, larval feeding preference, and associated survival of the yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer) (Hymenoptera: Tenthredinidae), within crowns of black spruce, Picea mariana (Mill.) Britton et al. (Pinaceae). Pikonema alaskensis females laid nearly twice as many eggs, and late‐instar larvae consumed nearly twice as much foliar biomass, on undefoliated vs. defoliated branches. The major factor driving the lower incidence of egg lay on defoliated branches was a significant reduction in the availability of preferred size classes of shoots. In general, adult females preferred medium‐sized shoots, whereas late instars preferred large shoots; previous defoliation generally decreased the availability of medium and large shoots. In a field bioassay, late‐instar survival generally declined with increasing defoliation, which again corresponded to significant reductions in shoot length on defoliated branches. The tendency of P. alaskensis to avoid shoots on severely defoliated branches is likely to increase the time lag between herbivory and the feedback effects of induction and could thus contribute to fluctuations in sawfly population density during outbreak. Our results differ from numerous past studies of closely related diprionid sawflies in conifers, most of which have reported weak or positive effects of previous herbivory on host susceptibility; we attribute the observed responses of spruce to the unusual sawfly feeding preference of P. alaskensis for developing conifer foliage.     

Distribution and oviposition preference of galling sawflies in arctic Canada

The distribution and ecology of insects in arctic regions are poorly known. The aim of this study was to determine the distribution of galling sawflies in the Canadian arctic and their oviposition preference. The Swedish Tundra Northwest 1999 expedition visited 17 sites in the Canadian arctic. We determined the occurrence of galling sawflies at all the sites and studied the oviposition preference of two leaf-galling sawflies, Eupontania arctica and Pontania nivalis, on Salix reticulata and S. glauca, respectively. Galling sawflies were abundant at only one site, the mainland site at Ivvavik National Park. Only a few galls in total were found at the remaining sites, suggesting that galling sawflies are rare in the higher arctic, and potential explanations for this pattern are discussed. Shoots with leaf galls were longer than shoots without galls on both S. reticulata and S. glauca. These differences could not be explained by a higher number of leaves on longer shoots. This suggests that long shoots are preferred by sawflies because of faster development and better survival of larvae on long shoots.     

Why does the bud-galling sawfly,Euura mucronata,attack long shoots?

Summary The bud-galling sawfly, Euura mucronata, attacked longer shoot length classes on its host, Salix cinerea, more frequently than shorter shoots. Shoot length accounted for 76 to 93 percent of the variance in number of galls per 100 shoots in three habitats: forest, watermeadow, and lakeside. The reasons for this pattern were addressed with studies on shoot length in relation to: 1. Number of resources (buds) per shoot; 2. Success in establishment of larvae in galls; 3. Gall size and resources per gall; and 4. Survival of larvae after establishment as influenced by plant resistance and natural enemy attack. The most important factors proved to be success in establishment of larvae, with percent of variance accounted for ranging from 57 to 77 percent in three of four sites where relationships were significant, and survival after establishment of larvae, with variance accounted for ranging from 40 to 54 percent in the same three sites. The pattern of survival was dictated by plant resistance and not by natural enemies. These two additive factors resulted in a general relationship across all sites of increasing emergence of fully developed larvae per cohort as shoot length increased, accounting for 78 percent of the variance. These adaptive advantages to attacking longer shoots are sufficient to account for the pattern of increased probability of shoots being attacked as they increase in length.     

Multiple effects of precipitation onSalix lasiolepis and populations of the stem-galling sawfly,Euura lasiolepis

Relatively low winter precipitation (e.g., 18–28 cm from October to May compared to 45 to 65 cm) caused reduced growth of the arroyo willow,Salix lasiolepis, with number of shoots per stem initiated and shoot length reduced. Resources were reduced for the stem-galling sawfly,Euura lasiolepis, which declined in numbers after the relatively dry winter of 1980–81. Sawfly phenology was advanced relative to willow phenology in the 1981 generation, causing an additional reduction in resource availability. These direct effects of precipitation on the sawfly were increased by indirect effects on survivorship of the 1981 generation. Egg death in galls increased on water-stressed willow plants, forming the major mortality in the generation. This resulted in very low survival in the generation and an even lower population in the 1982 generation, even though resources recovered after high precipitation during the 1981–82 winter (46.7 cm). An experiment using high, medium, and low water treatments on potted willow plants demonstrated that the effects on willows and sawflies could be reproduced using only water as a variable.     

Long term oviposition preference and larval performance of Schizomyia macrocapillata (Diptera: Cecidomyiidae) on larger shoots of its host plant Bauhinia brevipes (Fabaceae)

The Plant Vigor Hypothesis (PVH) proposes that natural selection on female oviposition choice results from higher fitness of larvae on more vigorous and larger plant modules. For six consecutive years we tested the PVH predictions by investigating the effect of shoot size of Bauhinia brevipes (Fabaceae) on the oviposition preference and offspring survival of the gall-midge Schizomyia macrocapillata (Diptera: Cecidomyiidae). Additionally, we analyzed the effects of bottom-up and top-down mortality forces on the system. The hypersensitive reaction (bottom-up effect) accounted for more than 90% larval mortality of S. macrocapillata, making available few galls to be found and killed by natural enemies (top-down effect). Smaller shoots were always more abundant while longer shoots were rare. Nevertheless, the percent number of galls induced by S. macrocapillata was up to 10-fold greater on the largest shoots, corroborating the preference prediction of the PVH. Schizomyia macrocapillata should use over-exploit larger shoots to maximize the preference for, and consequently increase the performance on these shoots. Our results partially support the performance prediction of the PVH: (1) the observed survival was higher than expected on longer shoots, and (2) the ratio of survival per shoot was positively related with shoot length only in 2 years. Thus, we found a link between female preference and larval performance on large-sized shoots, at least in some years. The gall-midge attack pattern in this study might be an evolved response to maximize the female preference and increased larval performance on longer shoots of the host plant.     

Mortality factors and resource use of the bud-galling sawfly, Euura mucronata (Hartig), on willows (Salix spp.) in arctic Eurasia

We studied egg and larval mortality factors in arctic populations of the bud-galling sawfly, Euura mucronata (Hartig) on three willow species (Salix glauca L., S. phylicifolia L., and S. lapponum L.) and the quality of resources (shoot length of willows) required for egg-laying. The survival was independent of latitude. There was a positive correlation in survival on different willow species among sampling sites, indicating that similar, locally operating factors affected survival. Host plant-based mortality factors were dominant and caused 17.9–48.0% mortality in eggs and 6.6–44.1% mortality in larvae. Parasitoids and inquilines caused relatively low and variable rates of mortality. Parasitoids were absent from several of the northernmost populations, but caused up to 11.0% mortality at southern sites. Mortality caused by inquilines was minor in other areas except in some sites in the Taymyr Peninsula, where it varied from 0 to 23.1%. E. mucronata laid eggs on the longest shoots available. Plant vigour as measured by shoot length decreased toward the north, and densities of galls were positively correlated with plant vigour. The difference in length between galled and ungalled shoots was 2.0- to 4.6-fold. Absence of E. mucronata in the most northern populations of willows was apparently caused by insufficient shoot growth. Short shoots failed to provide sufficient resources for successful development of galls. Our results suggest that the quality of host plants is the main factor determining abundance and distribution of E. mucronata in arctic areas. Received: 10 February 1997 / Accepted: 2 May 1997     

Hibernation adaptation and eclosion synchrony in leaf-galling sawflies in subarctic Norway

The winter dormancy adaptation in gall-inducing sawflies is poorly known. Diapause termination and the following post-diapause quiescence enhance synchronous eclosion in spring. This is probably the most critical part in the life history in gall-inducing sawflies, as there is only a short phenological window of opportunity for mating and oviposition. In a 2?years’ study, diapause duration, termination, survival and eclosion synchrony were experimentally investigated for three gall-inducing sawfly species (Symphyta: Tenthredinidae: Pontania nivalis, P. glabrifrons and P. arcticornis). Field-collected galls, sampled from willows (Salix spp.) in early autumn, were kept under natural temperatures outdoors until next spring. Subsamples were successively transferred to the laboratory at 14-day intervals from October to April in a test of development time to eclosion and survival ratio. The time to eclosion decreased throughout the experiment, whereas the proportion successfully eclosed and eclosion synchronicity increased, all indicative of prepupae entering a diapause in early autumn. The diapause terminates midwinter, and the prepupae enter a post-diapause quiescence until the temperature in spring allows a direct development and contributes to a nearly synchronous eclosion. In all three species, males eclosed 1–2?days prior to females (protandry). We hypothesize that synchronous eclosion as well as protandry enhance mating and oviposition success. Our finding indicates that gall-inducing sawflies are well adapted to its harsh subarctic and arctic environment.     

Spatial Distribution of Galls Caused by <Emphasis Type="Italic">Aculus tetanothrix</Emphasis> (Acari: Eriophyoidea) on Arctic Willows

The distribution of galls caused by Aculus tetanothrix (Acari: Eriophyoidea) on three Salix species was studied. The factors influencing this distribution were analysed, i.e. willow species, study area and shoot length. Spatial pattern of gall distribution within the shoot was also examined. The study was conducted in Russia, Kola Peninsula. Densities of galls caused by A. tetanothrix differed significantly among willow species. Considerably higher gall density was recorded in the White Sea coast than in the Khibiny Mountains. This may be explained by the influence of a milder maritime climate that favors mite occurrence compared to a harsh and variable mountain climate that limits mite abundance. There was no relationship between the gall density and the shoot length. The highest density of galls was recorded on the inner offshoots; within the offshoot, there was a maximum density on the fifth leaf. This pattern was repeatable for all shoots studied, independent of the study area, willow species and length of shoots, suggesting the optimal conditions for A. tetanothrix exist on leaves in the middle part of a shoot. This distribution pattern may be an effect of the trade-off between the costs and benefits resulting from leaf quality and mite movement along the shoot. This hypothesis, however, needs to be tested experimentally.     

Are galling insects better protected against parasitoids than exposed feeders?: a test using tenthredinid sawflies

ABSTRACT.

• 1 Data mostly from the published literature were used to assess the effect of galling on the number of parasitoid species per host species in the phylogeny of nematine sawflies from free external feeders (colonial and solitary) to leaf gallers and shoot gallers.
• 2 The strongest effects of galling were the total elimination of the species-rich cocoon-attacking guild of parasitoids, and eonymphal parasitoids, from the parasitoid community on shoot gallers, all of which are in the genus Euura.
• 3 All tachinid larval parasitoids were also eliminated by the galling habit.
• 4 The cumulative effects of these exclusions resulted in a decline in mean number of parasitoid species per host species from almost sixteen species on external colonial feeders to 4.0 species on shoot gallers.
• 5 General patterns in per cent parasitism by non-tachinid and tachinid larval parasitoids, eonymphal and cocoon parasitoids, on exposed feeders to shoot gallers, showed declines in non-tachinid attack and elimination of tachinid, eonymphal and cocoon parasitoids. But leaf gallers tended to be attacked more than exposed feeders by non-tachinid larval parasitoids.
• 6 The galling habit had a long-term impact by reducing the number of parasitoid species attacking nematine sawfly gallers and per cent mortality inflicted, so that natural enemies may have been important as a selective factor in the evolution of galling nematine sawflies.

     

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.     

Scale-dependent mechanisms in the population dynamics of an insect herbivore

A multiscale approach has lead to significant advances in the understanding of species population dynamics. The scale-dependent nature of population processes has been particularly clearly illustrated for insect herbivores. However, one of the most well-studied insect herbivores, the galling sawfly Euura lasiolepis, has to date been examined almost exclusively at fine spatial scales. The preference-performance, plant vigour and larval survival hypotheses are well supported by this species. Here, we test these hypotheses at a spatial scale larger than that previously considered, i.e. across a landscape in northern Arizona represented by an altitudinal gradient encompassing a series of drainages. We also develop a qualitative model for understanding the population dynamics of E. lasiolepis based on patterns of survival and mortality found in this study and previous ones. Gall density was highly variable across the altitudinal gradient, not explained by host plant variables, and thus a poor surrogate fot population abundance. These findings for the first time fail to support the plant vigour and preference hierarchy hypotheses for E. lasiolepis. Dispersal limitation most likely explains the lack of support for these hypotheses at this scale. By contrast, sawfly survival, gall abortion, parasitism and larval mortality were well explained by host plant quality variables and altitude. The larval survival hypothesis was well supported and is thus comparatively scale-invariant. A qualitative model developed here highlighted the importance of both willow water status and disturbance in determining host plant quality, as well as an apparent trade off between shoot length and plant moisture status in determining vital rates across the altitudinal gradient. This study thus demonstrated for the first time the scale-dependent nature of mechanisms underlying the population dynamics E. lasiolepis, and identified the interaction between parasitism and altitude as a novel mechanism underlying spatial patterns in the survival and mortality patterns of this species.     

Willow leaf traits affecting host use by the leaf-gall-forming sawfly

Gall-forming insects usually have very restricted host ranges, but plant traits affecting patterns of host use have rarely been examined. The sawfly Phyllocolpa sp. (Hymenoptera: Tenthredinidae) forms leaf-roll galls on three of seven Salix (Salicaceae) species that occur together on riverbanks in central Japan. We have attempted to explain this host-use pattern by invoking three plant traits: synchrony of leaf flush with the oviposition period of the sawfly, intrinsic leaf quality as a potential larval food, and leaf morphology. Two Salix species frequently used by the sawfly, Salix eriocarpa and Salix pierotii, had similar leaf traits suitable for larval survival. The third species, Salix serissaefolia, was used relatively less often and the sawfly frequently stopped laying eggs on the plant during oviposition, suggesting ovipositional selection. S. serissaefolia had the smallest leaves, and survival of sawfly larvae was lower on S. serissaefolia than on S. eriocarpa and S. pierotii, because of gall destruction, by other herbivorous insects, and leaf-size restrictions. Among the four unused species, Salix chaenomeloides had a late leaf-flush phenology, Salix gracilistyla had inferior leaf quality, and Salix gilgiana had linear leaves; these seemed to be critical factors for non-use. Salix subfragilis was also unused, but the reason for this could not be explained by the three leaf traits studied.     

Plant stress and larval performance of a dipterous gall former

According to the plant vigour hypothesis, galling insects should respond positively and perform better on vigorous plants or plant parts, the opposite of the predictions of the plant stress hypothesis. I carried out field experiments to analyse the effects of sustained abiotic stress on the interactions between the common reed (Phragmites australis) and a gall-forming fly (Lipara lucens). The reed shoot diameter (a measure of plant vigour) is strongly affected by environmental conditions, where dry and/or nutrient-poor habitats produce thinner (stressed) shoots. L. lucens gall density is negatively correlated with shoot diameter. In a survival experiment with a wide range of shoot diameters, larval mortality was also highly correlated with shoot quality. Gall formation was higher on thinner, stressed shoots. An analysis of the gall tissues revealed that galls induced by L. lucens contain a high amount of a nutrient-rich feeding tissue. The impact of L. lucens is higher on thinner shoots. The results of this study showed that L. lucens performs better on stressed hosts, which contradicts the plant vigour hypothesis for galling insects. The low nutrient availability in the stressed shoots can be compensated by the production of galls with a nutrient-rich feeding tissue.     

Resource partitioning and exploitative competition in stem-galling midges mediated by the host plant

Izeniola obesula Dorchin and Stefaniola defoliata Dorchin (Diptera, Cecidomyiidae) are two cecidomyiid species inducing multichambered galls in young shoots of the salt marsh plant, Suaeda monoica Gmelin (Chenopodiaceae). The purpose of this work was to study the relationship between the two species, which co‐occur in time and space, and may therefore potentially compete for plant resources. Characterization of the resources available to ovipositing females revealed that shoots in apical positions are more vigorous than shoots in basal positions, but that the two species of gall midges react to this variability of resources in different ways, as reflected by the patterns of gall distribution. Izeniola obesula galls were primarily induced in the younger, more rapidly growing shoots, whereas those of S. defoliata were predominantly induced in older, less vigorous shoots. Nevertheless, despite the general partitioning of resources, a certain amount of niche overlap was found, resulting in the occasional induction of an S. defoliata gall proximal to that of I. obesula in the same shoot. Co‐occurrence of the two types of galls in time and space did not have apparent effect on S. defoliata galls, but did have a strong adverse impact on young I. obesula galls, 60% of which were killed without producing adult midges, and the remaining 40% produced a much smaller number of adults. These effects are attributed to asymmetric interspecific competition that is mediated by the plant, resulting from the difference in stem tissues utilized by the larvae of the two species. It is concluded that preponderance of vacant niches, low densities of galls, and small niche overlap between the species do not preclude the occurrence of interspecific exploitative competition in this system, although its role in shaping the populations of the species is limited.