Novel observations of midge and mosquito larval population dynamics in leaves of the northern pitcher plant,Sarracenia purpurea L.

The invertebrate communities in Sarracenia purpurea were characterized by surveying the contents of 474 leaves from six pitcher plant populations in the eastern United States. More than 8200 invertebrate specimens, both living and dead, were recovered. Midge and mosquito larvae comprised 7907 (96%) of the specimens, with a mean of 16.9 dipteran larvae per leaf. Although midge larvae were present at all collection sites, mosquito larvae were absent from two of the sites. In the latter sites, the number of midge larvae per leaf was greater. This report documents for the first time that there is no statistical difference between the average number of Diptera larvae per leaf for sites that contain both midge and mosquito larvae compared to sites that contain only midge larvae. It appears that pitcher plants maintain an average number of larvae per leaf based on a number of interacting factors which we do not fully understand. This investigation documents that in the absence of mosquito larvae, more midge larvae are found per leaf per leaf than would be predicted based on data from sites that contain both larval taxa.     

Oviposition Behavior of the Predatory Midge Aphidoletes aphidimyza in Response to Aphid Patch Quality

Female Aphidoletes aphidimyza confronts serious challenges from both aphid prey and conspecifics. These challenges constitute strong selective pressures on the predatory midge to have an adaptive oviposition strategy. We did laboratory experiments to investigate clutch size of A. aphidimyza in response to the aphid Rhopalosiphum padi (density and age) and the presence of conspecific eggs (density) and larvae (age) in aphid colonies. The results of our study show that A. aphidimyza female adjusts its foraging effort by assessing the quality of prey patches. The number of eggs laid increased in response to aphid density. However, patches consisting of older aphids received fewer eggs. The number of eggs laid decreased in response to the presence of conspecific eggs, and in response to the presence of 2-day old conspecific larvae. Our study reveals deterrent effects on A. aphidimyza oviposition decisions on clutch size in response to older aphids and the presence of more conspecific eggs and older larvae.     

Preference for plants damaged by conspecific larvae in ovipositing cabbage root flies: influence of stimuli from leaf surface and roots

In laboratory dual-choice assays females of the cabbage root fly, Delia radicum, prefer for oviposition plants with roots damaged by conspecific larvae to undamaged controls. Cauliflower and kale plants were inoculated with root fly eggs (25 per plant) and the hatching larvae were allowed to feed on the roots for various periods of time (1–17 days). After 4 (cauliflower) or 5 (kale) days of larval feeding the oviposition preference was most pronounced and flies laid between 64% and 68% of their eggs near plants with damaged roots. Later, with increasing damage but fewer surviving, and thus actively feeding, larvae, the magnitude of the preference declined. The preference for plants already damaged by conspecific larvae may contribute to the previously observed aggregated distribution of D. radicum eggs in Brassica crop fields.Further experiments revealed that the sensory cues inducing this oviposition preference originate from the complex consisting of the damaged roots, the surrounding substrate (soil) and associated microbes, rather than from the aerial plant parts. In choice assays using the root-substrate complex of damaged and control plants (aerial parts removed), the observed preference for damaged roots was similar to that found for the entire plant but was more pronounced. The damaged roots alone, compared to control roots, received up to 72% (cauliflower) and 75% (kale) of the eggs. By contrast, surrogate leaves sprayed with methanolic leaf surface extracts from the most preferred plants which had been damaged were not discriminated from surrogate leaved sprayed with extracts of the respective control plants. Analysis of glucosinolate levels in methanolic leaf surface extracts revealed that root damage resulted in enhanced concentrations of indole-glucosinolates on the leaf surface in kale but not in cauliflower. Although indole-glucosinolates are oviposition stimulants for the cabbage root fly, the induced changes were apparently too small to influence oviposition behaviour.     

Egg and larval load assessment and its influence on oviposition behaviour of the leaf beetle Galerucella nymphaeae

The oviposition behaviour of the water-lily beetle Galerucella nymphaeae was examined. This species is a specialist herbivore on the floating leaves of nymphaeids Nymphaeaceae and especially on the yellow water-lily, Nuphar lutea. Females lay their eggs in clutches on the leaves, and after hatching, the larvae feed on the leaves. The quality of the leaves decreases quickly after the larvae hatch, and eventually the leaves will sink below the water surface, whereupon the eggs, 1st-instar larvae and pupae are killed by drowning. The influence of conspecific eggs, larvae and feeding tracks on the oviposition preferences of the beetles was tested. Females were allowed to choose between fresh leaves and leaves with conspecific eggs and larvae as well as between leaves with larvae and leaves with feeding tracks but no larvae. An attempt was also made to determine whether eggs and larvae affect the oviposition rate of females when they are not given the opportunity to oviposit on untouched leaves. The results indicate that females tended to avoid leaves with conspecific larvae or to exhibit a decreased oviposition rate on such leaves. Females also avoided conspecific eggs, although the oviposition rate was not influenced by the presence of conspecific eggs. When females were allowed to choose between leaves with larvae and leaves with feeding tracks, possible discrimination against leaves with larvae just fails to reach the 5% level.     

Feeding damage by larvae of the mustard leaf beetle deters conspecific females from oviposition and feeding

Herbivorous insects may be informed about the presence of competitors on the same host plant by a variety of cues. These cues can derive from either the competitor itself or the damaged plant. In the mustard leaf beetle Phaedon cochleariae (Coleoptera, Chrysomelidae), adults are known to be deterred from feeding and oviposition by the exocrine glandular secretion of conspecific co-occurring larvae. We hypothesised that the exocrine larval secretion released by feeding larvae may adsorb to the surface of Chinese cabbage leaves, and thus, convey the information about their former or actual presence. Further experiments tested the influence of leaves damaged by conspecific larvae, mechanically damaged leaves, larval frass and regurgitant on the oviposition and feeding behaviour of P. cochleariae. Finally, the effect of previous conspecific herbivory on larval development and larval host selection was assessed. Our results show that (epi)chrysomelidial, the major component of the exocrine secretion from P. cochleariae larvae, was detectable by GC-MS in surface extracts from leaves upon which larvae had fed. However, leaves exposed to volatiles of the larval secretion were not avoided by female P. cochleariae for feeding or oviposition. Thus, we conclude that secretion volatiles did not adsorb in sufficient amounts on the leaf surface to display deterrent activity towards adults. By contrast, gravid females avoided to feed and lay their eggs on leaves damaged by second-instar larvae for three days when compared to undamaged leaves. Mechanical damage of leaves and treatment of artificially damaged leaves with larval frass or regurgitant did not affect oviposition and feeding of P. cochleariae. Since no adverse effects of previous herbivory on larval development were detected, we suggest that female P. cochleariae avoid Chinese cabbage leaves damaged by feeding larvae for other reasons than escape from competition or avoidance of direct negative effects that result from consuming induced plant material.     

Diamondback moth performance and preference for leaves of Brassica oleracea of different ages and strata

The preference–performance hypothesis predicts that moth behaviour links plant variations with caterpillar attack and distribution, and the plant‐age hypothesis states that specialist herbivores are more successful in exploring younger plant tissue. We integrated these predictions to investigate underlying mechanisms by which moths and caterpillars of Plutella xylostella L. (Lepidoptera: Plutellidae) track and exploit within‐plant variability of leaf age and stratification. We measured leaf proteins, glucosinolates and fibre, as well as larval choice, developmental performance, and moth oviposition preference with regard to leaf age classes (young, mature and senescent) of three varieties (collard, cauliflower and cabbage) of the main host plant Brassica oleracea L. Larvae consistently fit the prediction that specialist herbivores prefer and perform better on young, upper leaves that have the highest protein level, despite the highest content of defence compounds. Conversely, moths laid more eggs on fibrous and less nutritious leaves from the lower and senescent stratum. We argue that the leaf stratification of host plants imposes conflicting selective pressures concerning offspring feeding and protection on adult females. If egg mortality is catastrophic on the upper nutritious leaves in a particular microclimatic context (e.g. sun, heat, winds, drought or rain‐washing), then oviposition preference will remain for the suboptimal lower and senescent leaves. The ability of larvae to spread upwards over the plant to access the more nutritious leaf stratum is critical when eggs are preferentially laid on the protective low‐quality leaves.     

Oviposition choice by the Onopordum capitulum weevil Larinus latus (Coleoptera: Curculionidae) and its effect on the survival of immature stages

Female Larinus latus lay eggs into the capitula of their Onopordum spp. host plants from the onset of capitulum development until the completion of flowering. This tight linkage between insect life-history and plant development optimised larval survival, apart from some initial asynchrony between the readiness of the females to oviposit and the suitability of the very small capitula to physically support eggs. Eggs were laid on bracts and stems of capitula or directly in the florets, when these became available. Both the location of the egg and time of oviposition influenced mortality factors such as egg and larval parasitism, egg desiccation and larval establishment. Overall, survival of eggs laid later in the season into florets was higher, although the adults that emerged were smaller. The change in choice of oviposition site that occurs at the onset of flowering, not only favours survival, but ensures more efficient resource use by the larvae. This occurs because sequential flowering of Onopordum spp. drives a sequential oviposition pattern and spreads the egg load more evenly over the available capitula, reversing a trend earlier in the season to clump eggs that were laid directly on the bracts. Other potential constraints, such as capitulum size, absolute egg density and previous egg-laying do not play a major role in determining the oviposition pattern of L. latus. Larval survival was positively correlated with capitulum size and not strongly influenced by egg density. Competition for resources only appeared to play a role in smaller capitula, and manifested itself in a reduction in the size of emerging adults rather than the death of immatures. In the absence of strong interspecific competition, the oviposition behavior of L. latus has become geared to maximising resource use for larvae (i.e. its own potential competitiveness) rather than minimising interactions with other members of the capitulum endophage guild.     

Temporal distribution of Smittium culisetae in a wild population of Wyeomyia smithii from pitcher plants

The fungus Smittium culisetae is a trichomycete that develops in the hindguts of larval aquatic Diptera. This is the first report of S. culisetae from the pitcher plant mosquito, Wyeomyia smithii. Larvae of the mosquito were collected from the purple pitcher plant, Sarracenia purpurea, from a bog in Jackson County, North Carolina. The lowest proportions of colonized larvae occurred in December, January and July. The greatest proportions of colonized larvae occurred in October and March. The distribution of colonized larvae among pitchers did not differ significantly from a random distribution.     

Oviposition Deterrence Is Likely an Effect,Not a Mechanism,in the Leaf Beetle <Emphasis Type="Italic">Chrysophtharta bimaculata</Emphasis> (Olivier) (Coleoptera: Chrysomelidae)

Oviposition deterrence is common in many insects as an evolutionary mechanism to reduce subsequent larval competition. We investigated a suspected case of oviposition deterrence by the paropsine chrysomelid, Chrysophtharta bimaculata. In paired choice tests, gravid females were found to prefer ovipositing on host leaves without conspecific eggs, confirming the presence of an apparent oviposition deterrence mechanism. Washing egg batches in water, hexane, or ethanol did not change this preference, suggesting that a soluble marking pheromone was not involved. Furthermore, it is unlikely that a plant-derived oviposition deterring substance is produced as beetles showed no significant oviposition preference between leaves which had been oviposited upon, but then had the eggs removed, and those that had never been oviposited upon. In trials using artificial leaves and mimic egg batches, “leaves” with “egg batches” placed near the tip of the leaf (the preferred site of oviposition in this species) were significantly less likely to be laid upon than artificial leaves where mimic eggs were placed away from the tip. In combination, the results strongly infer that oviposition deterrence in C. bimaculata is due to the mechanical blocking of the oviposition site by the first laid egg-batch, rather than a specific oviposition deterring cue. The apparent oviposition deterrence in this insect may well be an outcome or evolutionary effect of oviposition-site selection, rather than a clear adaptive mechanism to decrease larval competition.     

Oviposition substrate selection by Florida mosquitoes in response to pathogen‐infected conspecific larvae

The impact of the presence of larval mosquito pathogens with potential for biological control on oviposition choice was evaluated for three mosquito species/pathogen pairs present in Florida. These included Aedes aegypti infected with Edhazardia aedis, Aedes albopictus infected with Vavraia culicis, and Culex quinquefasciatus infected with Culex nigripalpus nucleopolyhedrovirus (CuniNPV). Two‐choice oviposition bioassays were performed on each host and pathogen species with one oviposition cup containing infected larvae and the other cup containing uninfected larvae (control). Both uninfected and E. aedis‐infected female Ae. aegypti laid significantly fewer eggs in oviposition cups containing infected larvae. Uninfected gravid female Ae. albopictus and Cx. quinquefasciatus oviposited equally in cups containing uninfected larvae or containing larvae infected with V. culicis or CuniNPV, respectively. Gravid female Ae. albopictus infected with V. culicis did not display ovarian development and did not lay eggs. The decreased oviposition by gravid Ae. aegypti in containers containing E. aedis‐infected larvae may indicate that the infected larvae produce chemicals deterring oviposition.     

From broadscale patterns to fine-scale processes: habitat structure influences genetic differentiation in the pitcher plant midge across multiple spatial scales

The spatial scale at which samples are collected and analysed influences the inferences that can be drawn from landscape genetic studies. We examined genetic structure and its landscape correlates in the pitcher plant midge, Metriocnemus knabi, an inhabitant of the purple pitcher plant, Sarracenia purpurea, across several spatial scales that are naturally delimited by the midge’s habitat (leaf, plant, cluster of plants, bog and system of bogs). We analysed 11 microsatellite loci in 710 M. knabi larvae from two systems of bogs in Algonquin Provincial Park (Canada) and tested the hypotheses that variables related to habitat structure are associated with genetic differentiation in this midge. Up to 54% of variation in individual‐based genetic distances at several scales was explained by broadscale landscape variables of bog size, pitcher plant density within bogs and connectivity of pitcher plant clusters. Our results indicate that oviposition behaviour of females at fine scales, as inferred from the spatial locations of full‐sib larvae, and spatially limited gene flow at broad scales represent the important processes underlying observed genetic patterns in M. knabi. Broadscale landscape features (bog size and plant density) appear to influence oviposition behaviour of midges, which in turn influences the patterns of genetic differentiation observed at both fine and broad scales. Thus, we inferred linkages among genetic patterns, landscape patterns and ecological processes across spatial scales in M. knabi. Our results reinforce the value of exploring such links simultaneously across multiple spatial scales and landscapes when investigating genetic diversity within a species.     

Avoidance of occupied hosts by the Hessian fly: oviposition behaviour and consequences for larval survival

Abstract 1. Unoccupied wheat plants and wheat plants occupied by conspecific eggs or larvae were presented to ovipositing female Hessian flies in choice tests.
2. The presence of conspecific eggs on the leaf surfaces of wheat plants did not appear to have any effect on the responses of ovipositing Hessian fly females.
3. The presence of conspecific larvae at the base and nodes of wheat plants for 1, 6, or 11 days had significant effects on Hessian fly oviposition. Eggs oviposited on plants were inversely proportional to larval densities and days of larval occupation.
4. Feeding by Hessian fly larvae is associated with several changes in wheat plants. One of these changes, the growth arrestment of the plant, was measured by recording the heights of plants used in oviposition tests. Plant heights were inversely proportional to both larval densities and days of occupation. Plant heights were directly proportional to eggs oviposited on plants.
5. The consequences of adult female avoidance of plants occupied by conspecific larvae were investigated by allowing females to oviposit on unoccupied plants and 1-day, 6-day, and 11-day larval occupied plants, then scoring at the end of the first larval instar the survival of the offspring that resulted from this oviposition.
6. Survival during the first larval instar was 88% for the offspring of females that oviposited on unoccupied plants, decreasing to 82, 31, and 4% on the 1-day, 6-day, and 11-day occupied plant treatments. On these four plant treatments, a positive correlation was found between larval performance (i.e. survival) and the preferences of ovipositing females.
7. On the four plant treatments, relationships between first-instar larval density and first-instar larval survival varied significantly. On unoccupied plants, survival was inversely proportional to density. On plants oviposited on at 6 days of larval occupation, survival was directly proportional to density.     

Morphological correlates of necromass accumulation in the traps of an Eastern tropical pitcher plant, Nepenthes ampullaria Jack, and observations on the pitcher infauna and its reconstitution following experimental removal

I studied the trap morphology, necromass accumulation rates and pitcher infauna of an eastern tropical pitcher plant, Nepenthes ampullaria, that grew in `kerangas' heath forest in the Sungei Ingei Conservation Area, Brunei. I surveyed 164 pitchers distributed among 35 plants and extracted the necromass and larval infauna from the pitchers and then resampled the pitcher contents after 14 days. Plants varied significantly in the morphology of their pitchers, in their rate of necromass accumulation per pitcher and in the abundance and composition of the pitcher infaunas. On average, pitchers accumulated 11.5 mg dry weight over 14 days, but larger pitchers accumulated more necromass than smaller ones. Pitcher morphology explained 45% of the variation in necromass accumulation among plants. On average, pitchers initially contained 26.3 individual larval inquilines. Collectively, the larval infauna was composed of nine taxa of dipteran larvae and infrequent anuran tadpoles. These ten taxa were never found together in a single pitcher and the mean species richness per pitcher was 4.0. Of the six taxa that could be assessed, all except Toxorhynchites spp. had a contagious distribution among the pitchers. Pitcher morphology and necromass accumulation explained only 15% of the variation in inquiline abundance among plants. I found little evidence for the existence of density-dependent interactions between inquiline species: a partial correlation analysis detected only one statistically significant pairwise relationship between the abundances of inquiline taxa, which was a positive association. Fourteen days after being emptied, pitchers contained an average of 9.6 inquilines. There was no evidence that the species composition of the infauna recolonising each pitcher was related to that of its pre-removal infauna. Received: 2 June 1997 / Accepted: 9 September 1997     

Possible Age-Dependent Variation in Egg-Loaded Host Selectivity of the Pierid Butterfly, Anthocharis scolymus (Lepidoptera: Pieridae): A Field Observation

To understand better the effects of age on host selection through conspecific egg detection by Anthocharis scolymus females, field observations were performed at a graveyard where the insect population was isolated and the host plant was limited to the cruciferous plant, Turritis glabra. We chased females and recorded their oviposition behaviors and the conditions of plants which females approached. Older females tended to avoid ovipositing on egg-loaded host plants and selectively laid eggs on nonloaded host plants. This result was not confounded by other factors such as seasonality, air temperature, plant height, plant phenological stage, surrounding vegetation, host plant density, and extent of plant damage. We discuss the possibility that females make an oviposition decision on how to lay their limited number of eggs during their remaining lifetime.     

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.     

Polyphagy and primary host plants: oviposition preference versus larval performance in the lepidopteran pest Helicoverpa armigera

Oviposition preference and several measures of offspring performance of Helicoverpa armigera (Hübner) were investigated on a subset of its host plants that were selected for their reputed importance in the field in Australia. They included cotton, pigeon pea, sweet corn, mungbean, bean and common sowthistle. Plants were at their flowering stage when presented to gravid female moths. Flowering pigeon pea evoked far more oviposition than did the other plant species and was the most preferred plant for neonate larval feeding. It also supported development of the most robust larvae and pupae, and these produced the most fecund moths. Common sowthistle and cotton were equally suitable to pigeon pea for larval development, but these two species received far fewer H. armigera eggs than did pigeon pea. Mungbean also received relatively few eggs, but it did support intermediate measures of larval growth and survival. Fewest eggs were laid on bean and it was also the least beneficial in terms of larval growth. Among the host plant species tested, only flowering pigeon pea supported a good relationship between oviposition preference of H. armigera and its subsequent offspring performance. Australian H. armigera moths are thus consistent with Indian H. armigera moths in their ovipositional behaviour and larval performance relative to pigeon pea. The results suggest that the host recognition and acceptance behaviour of this species is fixed across its geographical distribution and they support the theory that pigeon pea might be one of the primary host plants of this insect. These insights, together with published results on the sensory responses of the females to volatiles derived from the different host plant species tested here, help to explain why some plant species are primary targets for the ovipositing moths whereas others are only secondary targets of this polyphagous pest, which has a notoriously broad host range. Handling Editor: Joseph Dickens     

Assessment of patch quality by ladybirds: role of larval tracks

Gravid females of the two-spot ladybird, Adalia bipunctata (L.), were deterred from ovipositing when kept in petri dishes that had previously contained conspecific larvae but not conspecific adults, or the larvae of another two species of ladybird, Adalia decempunctata (L.) and Coccinella septempunctata L. The deterrent effect was density dependent and mediated via a chloroform-soluble contact pheromone present in the larval tracks. Similarly, gravid females of C. septempunctata were deterred from ovipositing by conspecific larval tracks and chloroform extracts of these tracks, but not by the tracks or extracts of tracks of A. bipunctata larvae. That is, in ladybirds the larvae produce a species-specific oviposition-deterring pheromone. In the field, the incidence of egg cannibalism in ladybirds increases very rapidly with the density of conspecific eggs or larvae per unit area. Thus, in responding to the species specific oviposition deterring pheromone female ladybirds reduce the risk of their eggs being eaten and spread their offspring more equally between patches. Received: 14 March 1997 / Accepted: 26 August 1997     

Protection of potato crop against Phthorimaea operculella (Zeller) infestation using frass extract of two noctuid insect pests under laboratory and storage simulation conditions

The oviposition deterrent effect of water extract of Spodoptera littoralis and Agrotis ipsilon larval frass on Phthorimaea operculella adult females was studied using two types of larval food “Natural host and Semi-artificial diet” under laboratory and storage simulation (semi-field) conditions. Extracted frass of fed larvae on semi-artificial diet showed complete oviposition deterrent effect at treatments with 4th, 5th and 6th instars of S. littoralis, also at treatments with 1st–3rd and 6th instars of A. ipsilon, while the same effect was observed when the larvae fed on castor oil leaves as a natural host only at treatment with frass extract of A. ipsilon 6th instar larvae. Presence of low amounts of phenols and flavonoids in water extract of A. ipsilon larval frass resulted in relatively more effect as oviposition deterrent to fertile adult females on treated oviposition sites, while the opposite effect was obtained in S. littoralis larval frass experiments. At semi-field experiments, the percentage reduction of laid eggs reached 100% after two?days at treatments with frass extracts of 4th and 5th S. littoralis larval instars and A. ipsilon 6th instar larvae fed on semi-artificial diet and/or castor oil leaves. Percentage reduction of laid eggs for untreated sacks reached 93.24 and 48.95% after 2 and 30?days, respectively, when placed between treated sacks, in comparison with the mean number of laid eggs for isolated control.     

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.