Foraging behaviour at the fourth trophic level: a comparative study of host location in aphid hyperparasitoids

In studies of foraging behaviour in a multitrophic context, the fourth trophic level has generally been ignored. We used four aphid hyperparasitoid species: Dendrocerus carpenteri (Curtis) (Hymenoptera: Megaspilidae), Asaphes suspensus Walker (Hymenoptera: Pteromalidae), Alloxysta victrix (Westwood) (Hymenoptera: Alloxystidae) and Syrphophagus aphidivorus (Mayr) (Hymenoptera: Encyrtidae), to correlate their response to different cues with their ecological attributes such as host range and host stage. In addition, we compared our results with studies of primary parasitoids on the same plant–herbivore system. First, the olfactory response of females was tested in a Y‐tube olfactometer (single choice: plant, aphid, honeydew, parasitised aphid, aphid mummy, or virgin female parasitoid; dual choice: clean plant, plant with aphids, or plant–host complex). Second, their foraging behaviour was described on plants with different stimuli (honeydew, aphids, parasitised aphids, and aphid mummies). The results indicated that olfactory cues are probably not essential cues for hyperparasitoid females. In foraging behaviour on the plant, all species prolonged their total visit time and search time as compared to the control treatment (clean plant). Only A. victrix did not react to the honeydew. Oviposition in mummies prolonged the total visit time because of the long handling time, but the effect of this behaviour on search time could not be determined. No clear correlation between foraging behaviour and host stage or host range was found. In contrast to specialised primary aphid parasitoids that have strong fixed responses to specific kairomones and herbivore‐induced synomones, more generalist aphid hyperparasitoids seem to depend less on volatile olfactory stimuli, but show similarities with primary parasitoids in their use of contact cues while searching on a plant.     

Within-patch search flights by Pachyneuron aphidis (Hym., Pteromalidae): a potential strategy to compensate reduced foraging speed by foot

Search flights of the aphid hyperparasitoid Pachyneuron aphidis were observed under field conditions. The pattern and duration differed from those described for aphid primary parasitoids. Forty per cent targeted landings close to aphid colonies indicate that P. aphidis females locate potential host sites during slow and extraordinary long flights in the host plant Cirsium arvense . A comparison between different aphid hyperparasitoids showed that the walking speed of P. aphidis was significantly slower than that of other polyphagous species. This suggests that search flights were performed in order to compensate for a reduced efficiency of foraging on foot.     

Influence of adult experience on host microhabitat location by the generalist parasitoid,Campoletis sonorensis (Hymenoptera: Ichneumonidae)

The effect of adult experience on microhabitat location behavior of the generalist parasitoid Campoletis sonorensis (Cameron)was examined in a wind tunnel bioassay. Responses were tested to the odors of two host plants (cotton and sesame) of Heliothis virescens (F.) or a nonhost plant (potato), either damaged and infested with host larvae and host products (host/plant complex) or intact, clean and uninfested. Parasitoid females remained naive or were allowed one oviposition experience on either of the plants, 1 min, 2 h, or 24 h prior their introduction into the wind tunnel. In a no-choice test, parasitoids experienced 1 min prior to bioassay completed significantly more flights to sesame and potato host/plant complexes than did naive parasitoids. However, 24 h after experience, only females experienced on potato completed more flights to the host/ plant complex than did naive females. Parasitoids experienced 1 min prior to flight to undamaged plants showed a slight increase in flight response (significant only for potato) but, after 24 h, completed only as many flights as naive parasitoids. In a dual-choice situation, parasitoids did not show a preference for either of the two host plants but did prefer a host to a nonhost plant. This innate plant preference was not changed by a single oviposition experience. The potential significance of these results to the microhabitat location behavior of C. sonorensisin the field is discussed.     

Phenological matching rather than genetic variation in host preference underlies geographical variation in host plants used by orange tip butterflies

An insect species that shows variation in host species association across its geographical range may do so either because of local adaptation in host plant preference of the insect or through environmentally or genetically induced differences in the plants, causing variation in host plant suitability between regions. In the present study, we experimentally investigate the host plant preference of Anthocharis cardamines (orange tip butterfly) in two populations from the UK and two from Sweden. Previous reports indicate that A. cardamines larvae are found on different host plant species in different regions of the UK, and some variation has been reported in Sweden. Host plant choice trials showed that females prefer to oviposit on plants in an earlier phenological stage, as well as on larger plants. When controlling for plant phenological stage and size, the host species had no statistically significant effect on the choice of the females. Moreover, there were no differences in host plant species preference among the four butterfly populations. Based on our experiment, the oviposition choice by A. cardamines mainly depends on the phenological stage and the size of the host plant. This finding supports the idea that the geographical patterns of host–plant association of A. cardamines in the UK and Sweden are consequences of the phenology and availability of the local hosts, rather than regional genetic differences in the host species preference of the butterfly.     

Multifaceted determinants of host specificity in an aphid parasitoid

The host specificity of insect parasitoids and herbivores is thought to be shaped by a suite of traits that mediate host acceptance and host suitability. We conducted laboratory experiments to identify mechanisms shaping the host specificity of the aphid parasitoid Binodoxys communis. Twenty species of aphids were exposed to B. communis females in microcosms, and detailed observations and rearing studies of 15 of these species were done to determine whether patterns of host use resulted from variation in factors such as host acceptance or variation in host suitability. Six species of aphids exposed to B. communis showed no signs of parasitism. Four of these species were not recognized as hosts and two effectively defended themselves from attack by B. communis. Other aphid species into which parasitoids laid eggs had low suitability as hosts. Parasitoid mortality occurred in the egg or early larval stages for some of these hosts but for others it occurred in late larval stages. Two hypotheses explaining low suitability were investigated in separate experiments: the presence of endosymbiotic bacteria conferring resistance to parasitoids, and aphids feeding on toxic plants. An association between resistance and endosymbiont infection was found in one species (Aphis craccivora), and evidence for the toxic plant hypothesis was found for the milkweed aphids Aphis asclepiadis and Aphis nerii. This research highlights the multifaceted nature of factors determining host specificity in parasitoids.     

Companion planting – behaviour of the cabbage root fly on host plants and non-host plants

Six‐hundred individual female cabbage root flies (Delia radicum L.) (Diptera: Anthomyiidae) were each observed for 20 min under laboratory conditions to record how they behaved after landing on a host or a non‐host plant. Fly movements were recorded on host plants [cabbage –Brassica oleracea var. capitata (Cruciferae)] and non‐host plants [clover –Trifolium subterraneum L. (Papilionaceae)] surrounded by bare soil and on cabbage surrounded by clover. The most frequently observed behaviours made by the flies were (1) hops/spiral flights and (2) walks/runs. In the bare soil situation, the 50 individual flies observed in each treatment made 66 hops/spiral flights on the cabbage and 94 on the clover. When the two plants were tested together the movements were not additive as, instead of the expected 160 hops/spiral flights in the mixed plant treatment, the flies made 210 hops/spiral flights when they landed initially on cabbage but only 130 when they landed initially on clover. Few of the flies that landed initially on clover moved onto the host plant, even though the host plant was only a few centimetres away. The duration of the individual walks and runs made by the cabbage root flies were similar on both the host and non‐host plants. The only differences were the numbers of walks/runs made and the time the flies remained inactive. On the host plants, the females made four walks/runs, each of about 12 s duration, interspersed by rest periods that totalled 1.5 min. In contrast, on the non‐host plants the females made 10 walks/runs, each of about 9 s duration, interspersed by rest periods that totalled 7 min. Therefore, after landing on a plant, the flies, on average, left the host plant after 2.25 min and the non‐host plant after 8.5 min. Our conclusion is that the protracted time spent on the non‐host plants is the mechanism that disrupts insects from finding host plants in diverse plantings. Hence, the flies were arrested by non‐host plants rather than being repelled or deterred as suggested in earlier studies.     

The role of ecological availability and host plant characteristics in determining host use by the bogus yucca moth Prodoxus decipiens

1. Specialisation in host plant use is strongly correlated with speciation in many plant‐feeding insects. Specialised taxa, however, could be restricted in host range due to limits in ecological availability of host plant species rather than trade‐offs in using alternate host species. 2. Moths in the genus Prodoxus are extreme specialists on Yucca and speciation is closely tied to host plant shifts. However, many Yucca ranges are allopatric. This study examined whether the bogus yucca moth Prodoxus decipiens is limited in host range because of biogeographic factors or due to differences in the characteristics of host plant species. 3. In a common garden, local P. decipiens moths that use Yucca filamentosa were exposed to individuals of five Yucca species, two that are known hosts of P. decipiens in other parts of its range and three that are used by its sister species, Prodoxus quinquepunctellus. 4. Local moths were attracted to flowers of all Yucca species and females attempted oviposition in the flowering stalks of all species. However, larvae successfully completed development to diapause in only one of the five host plant species. Larval development on non‐natal Yucca species was significantly reduced compared with the local host. 5. The results suggest that differences in host plant characteristics among Yucca species would result in strong natural selection during a host shift. Thus, specialisation in host plant use is probably due to trade‐offs involved with using novel host plant species as well as ecological availability.     

Identifying bird cherry-oat aphid Rhopalosiphum padi emigrants, alate exules and gynoparae: application of multivariate methods to morphometric and anatomical features

The host microhabitat location behavior of females of the generalist parasitoid Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae) was studied in a wind tunnel. Visual cues associated with the host plant cotton, Gossypium hirsutum L., were important and significantly more parasitoids completed flights to a damaged 4-leaf cotton plant bearing a Heliothis virescens (F.) (Lepidoptera: Noctuidae) larva and frass than to a similarly damaged single leaf with frass and a larva. This difference in completed flights was not due to differences in amounts of volatiles released by the two stimuli. Both naive and experienced parasitoids responded differently to an undamaged cotton leaf, a mechanially damaged leaf, a naturally damaged leaf with the host removed and a naturally damaged leaf with a host larva. Parasitoids completed significantly fewer flights to the undamaged sources of volatiles than to damaged sources of volatiles. Experienced females responded strongly to all types of damage. The number of flights completed by naive females to the three types of damage differed but not significantly and was less than the number completed by experienced females. Components of the preflight experience were varied to determine which factors were responsible for the higher response of experienced females to the host/plant complex. Oviposition was the most important component of this experience. Contact with host frass or plant damage followed by oviposition did not increase the response over that exhibited by females allowed oviposition only. When frass or damaged plant material were contacted without subsequent oviposition, females completed fewer flights than naive females.     

The role of plant chemical cues in determining host preference in the generalist aphid parasitoid Aphidius colemani

The host preference behaviour of the generalist aphid parasitoid Aphidius colemaniwas investigated using a Y-tube olfactometer. Female A. colemanishowed a preference for the host-plant complex on which they had been reared, even though the same aphid host was involved, demonstrating a host plant preference. This preference was not evident when the parasitoids were dissected from their mummies prior to adult emergence. Host plant preference exhibited during host selection appeared to be induced by chemical cues encountered on the mummy case at the time of emergence, but preferences could be changed by subsequent foraging experiences. It is concluded that plant chemical cues play a major role in determining initial preferences through a process of emergence conditioning but that learning processes, involving cues encountered during oviposition in or contact with the host, can modify these initial preferences.     

Population level correlation between pre-alighting and post-alighting host plant preference in the Glanville fritillary butterfly

1. In many populations of the Glanville fritillary butterfly Melitaea cinxia, ovipositing females exhibit a post‐alighting preference for one of the potential host plant species available. The work reported here aimed to establish whether females with different post‐alighting preferences can discriminate between their host plant species prior to alighting, and whether pre‐alighting and post‐alighting preferences are correlated at the population level. 2. Alighting and oviposition events were recorded for groups of females from six populations in greenhouse and field experiments. 3. Landing frequencies did not change with experience, indicating that M. cinxia females did not learn from previous encounters with host plants. 4. Females from populations exhibiting post‐alighting preference searched efficiently for their host plants in the sense that they landed mainly on the species on which they oviposited predominantly. Pre‐alighting and post‐alighting preferences were correlated at the population level. 5. The correlation between pre‐alighting and post‐alighting preferences helps to explain why in nature, where the host plants often occur in distinct patches, females are more likely to colonise habitat patches in which their preferred host plant is abundant.     

Winter Host Plant Specialization in a Host-Alternating Aphid

The host alternating aphid, Rhopalosiphum padi (L.), feeds in summer on several different species of grasses but is monophagous on its winter host, Prunus padus L. The monophagy on P. padus could be a result of the restricted host range of the several different generations colonizing, or feeding, on this host during autumn–winter–spring. This study shows that the winter host plant specificity of R. padi is controlled mainly by the preference of the females remigrating (gynoparae) to the winter host, P. padus, in autumn. The other generations living on the winter host, i.e., sexual females, males, and spring generations, all accept a broader range of winter hosts. One alternative host plant, Prunus spinosa L., could be utilized by all generations associated with the winter host, except for the females remigrating at autumn.     

Secondary host choice by the autoparasitoid Encarsia pergandiella

Autoparasitoids are species of parasitic wasps in the family Aphelinidae which produce females as solitary primary endoparasitoids of homopterans such as whitefly and scale insects (primary hosts), and males as solitary hyperparasitoids. Males generally develop on immature conspecific females or on individuals of other primary parasitoid species (secondary hosts). Encarsia pergandiella is an autoparasitoid that has been introduced to Italy for control of greenhouse whitefly Trialeurodes vaporariorum, in greenhouses and field crops. In this study we examined the secondary host selection behaviour of this species with regard to conspecific females and females of two thelytokous species, E. formosa and E. meritoria. Encarsia formosa has been used successfully for greenhouse whitefly control in Northern Europe, but has not been effective in Southern Italy in winter crops in unheated greenhouses. E. meritoria has recently spread in Italy, and may have potential for biological control of whitefly in the greenhouse environment. In the first experiment, female E. pergandiella were exposed to one of three pair-wise combinations of the three species in petri dish arenas. Parasitism was determined by dissection of the hosts. The number of hosts parasitized by E. pergandiella females did not differ with host species. However, significantly greater numbers of eggs were laid in E. meritoria in both treatments in which it was present; these hosts were more likely to be superparasitized. In a second experiment, observations of females in arenas with equal numbers of all three host species indicated that females encountered and parasitized all host species with approximately equal frequency, although the length of time females spent in the oviposition posture differed with host species.     

Effect of mechanical damage on emission of volatile organic compounds from plant leaves and implications for evaluation of host plant specificity of prospective biological control agents of weeds

Assessment of host plant specificity is a critical step in the evaluation of classical biological control agents of weeds which is necessary for avoiding possible damage to non-target plants. Volatile organic compounds (VOCs) emitted by plants likely play an important role in determining which plants attract and are accepted by a prospective arthropod agent. However, current methods to evaluate host plant specificity usually rely on empirical choice and no-choice behavioural experiments, with little knowledge about what chemical or physical attributes are stimulating the insect. We conducted experiments to measure the quantitative and qualitative effects on emission of VOCs caused by simple mechanical damage to leaves of plants known to differ in suitability and attractiveness to a prospective agent. More VOCs were detected from damaged than from undamaged leaves for all three species tested. Discriminant analysis was able to correctly distinguish the taxonomic identity of all plants based on their VOC profiles; however, the VOCs that discriminated species among undamaged leaves were completely different from those that discriminated among damaged leaves. Thus, damaged and undamaged plants present different VOC profiles to insects, which should be considered when conducting host plant specificity experiments. An unacceptable non-target plant, Centaurea cineraria, emitted all except one of the VOCs that were emitted by its preferred host plant, Centaurea solstitialis, indicating the importance of compounds that are repellant in host plant specificity. Centaurea cyanus emitted fewer VOCs than C. solstitialis, which suggests that it lacked some VOCs important for host plant recognition.     

Male and female Trybliographa rapae (Hymenoptera: Figitidae) behavioural responses to food plant, infested host plant and combined volatiles

Many parasitoids use volatiles produced by plants as important cues during their food and host search process. We investigated the attraction of the parasitic wasp Trybliographa rapae Westwood (Hymenoptera: Figitidae) to volatiles emitted from plants infested by the cabbage root fly Delia radicum L. (Diptera: Anthomyiidae), as well as to volatiles from a nectar food plant. Behavioural choice tests showed that male parasitoids were not attracted to any volatiles from plants infested by D. radicum or from nectar plants, while females showed clear attraction to both volatile sources. Young females were more attracted to combined volatiles of host and food plants over those from only the host plant, whereas older females showed no differences in attraction to the two odour sources. This suggests that intercropping attractive flowers with host plants could potentially be used to recruit newly emerged parasitoids from surrounding fields while older parasitoids invest more energy in host location than in additional food search. Volatiles from a whole infested plant were chosen over those emitted from separated above- and below-ground parts from infested plants. It is important to consider the availability of both energy and host resources for parasitoids when designing an eco-compatible management of a vegetable crop system.     

Egg load influences search intensity,host selectivity,and clutch size inBattus philenor butterflies

Numerous studies have documented the influence of environmental factors such as host plant species and host quality on the oviposition behavior of female insects. This paper shows that an internal physiological factor, the number of mature eggs a female carries (egg load), correlates with host selectivity and clutch size in unmanipulated natural populations of the pipevine swallowtail butterfly, Battus philenor.In addition, search intensity and host selectivity differed among females whose egg loads were manipulated experimentally before they were released and followed in the field. Females with many eggs searched more intensely for hosts and were less selective when they encountered them.     

Oviposition in Yponomeuta cagnagellus: the importance of contact cues for host plant acceptance

Small ermine moths (Yponomeutidae: Lepidoptera) are specialist herbivores. Species within the genus Yponomeuta are each specialized on a limited number of plant species, mainly within genera belonging to the Celastraceae. European Yponomeuta species have developed new specialized host affiliations, mainly on rosaceous hosts. Since these host shifts are reputed to be of consequence for speciation, the role of the ovipositing female is of particular interest. Study of the pre-oviposition behaviour of gravid Y. cagnagellus (Hb.) moths on host (Euonymus europaeus), non-host (Crateagus monogyna) and artificial oviposition substrates, provided information on the nature of the cues used for host plant acceptance and the insect’s perception of these cues. Host selection by adult females occurs with contact chemoreceptors probably located on the antennae or tarsi. MeOH-soluble, non-volatile phytochemical compounds washed from the host plant’s surface and applied on an artificial twig are sufficient to stimulate a complete sequence of behavioural elements leading to oviposition. Volatiles do not have a large effect on the pre-oviposition behaviour.     

Thrips see red – flower colour and the host relationships of a polyphagous anthophilic thrips

Abstract 1. The common blossom thrips, Frankliniella schultzei, is a polyphagous anthophilic species that colonises a wide range of host‐plant species across different plant taxa. The environmental cues used by these polyphagous insects to recognise and locate host plants are not known. We therefore determined if colour is an important environmental signal used by F. schultzei to recognise flowers of eight of its more significant host‐plant species. 2. The effect of flower colour on the colonisation of different host plant species by F. schultzei was investigated by collecting and analysing the following: (a) numbers of thrips from different heights and aspects of the primary host plant Malvaviscus arboreus, (b) thrips distribution within flowers of Hibiscus rosasinensis, (c) colour reflectance from flowers of eight different host‐plant species, and (d) reflectance from different coloured sticky traps and the number of thrips trapped on them at different times of the day and on different dates. 3. The results indicate that: (a) the thrips (both sexes) concentrate towards the top of the primary host plant M. arboreus and are not distributed differentially according to sunny or shady aspect of the plant, (b) the number of female thrips on H. rosasinensis was higher in anthers compared to petals (corolla) and the basal parts of the flower, and males were as numerous on the petals as were females, and (c) there is a common floral reflectance pattern (but with different intensities) across the eight host plant species, mainly in the red part of the spectrum (600–700 nm). 4. Results of colour sticky trapping show that red attracts more female thrips compared to any other colour and that most were caught between 09.00 and 11.00 hours. By contrast, more male thrips were trapped between 07.00 and 09.00 hours. Males were more evenly distributed across the different colours but the highest numbers were associated with the yellow traps. 5. The higher densities of thrips at the top of their host plant may be related to the early morning (07.00–11.00 hours) activity of the thrips, when the top portions of the plant are more exposed to sunlight. The sex‐related distributions of F. schultzei thrips across time, coloured sticky traps, and various parts of the flowers seem to be related to mating swarm formation by the males, on the one hand, and the relative frequency and intensity of the use of M. arboreus by the females, on the other, as a feeding and oviposition site. Frankliniella schultzei females respond more strongly to red than to any other colours, so it is predicted that the spectral properties of colour recognition by this species will correlate with the predominant red reflectance of its primary host, M. arboreus, and that there may well be a sex‐related difference in colour recognition within this species.     

The interaction between host and host plant influences the oviposition and performance of a generalist ectoparasitoid

The successful development of parasitoids of herbivores depends on the quality of their host, which is often affected by the host plant. Therefore, a parasitoid’s oviposition decisions will directly depend on the host, but also on plant quality. Here, we investigated the direct effects of host species and the indirect effects of the host’s food plant on the oviposition decisions and performance of the gregarious ectoparasitoid Euplectrus platyhypenae Howard (Hymenoptera: Eulophidae). With a series of no‐choice experiments, we determined the oviposition and performance of the parasitoid on: (1) two caterpillar species, fall armyworm, Spodoptera frugiperda JE Smith (Lepidoptera: Noctuidae), and velvet armyworm, Spodoptera latifascia Walker, reared on maize (Zea mays L., Poaceae), (2) the same caterpillars reared on maize, bean (Phaseolus vulgaris L., Fabaceae), or squash (Cucurbita pepo L., Cucurbitaceae) leaves, and (3) S. latifascia caterpillars reared on leaves of wild and cultivated lima bean, Phaseolus lunatus L. All these insects and plants originate from Mesoamerica where they have coexisted for thousands of years in the traditional agricultural system known as Milpa in which maize, beans, and squash are planted together. We found that the preferred and best combination of host and host plant for parasitoid performance was S. frugiperda on maize. Parasitoids laid larger clutches, had higher survival, and more females and larger adults emerged from S. frugiperda reared on maize. However, when both caterpillar species were reared on squash, S. latifascia was the better host. Contrary to the literature, S. frugiperda was not able to develop on bean plants. Results from the lima bean experiment showed that parasitoid performance was best when S. latifascia was reared on leaves of cultivated compared to wild lima bean. These findings are discussed in the context of mixed cropping in which the ability of generalist parasitoids to switch among hosts and host plant species could be advantageous for pest management.     

Lack of evolution in a leaf beetle that lives on two contrasting host plants

The interactions between plant‐eating insects and their hosts have shaped both the insects and the plants, driving evolution of plant defenses and insect specialization. The leaf beetle Trirhabda eriodictyonis (Chrysomelidae) lives on two shrubs with differing defenses: Eriodictyon crassifolium has hairy leaves, whereas E. trichocalyx has resinous leaves. We tested whether these beetles have differentiated onto the two host plants, and if not, whether the beetles prefer the better host plant and prefer mates who are from that host plant. In feeding tests, adult beetles strongly preferred eating E. trichocalyx regardless of which host they came from. In addition, females laid more eggs if they ate E. trichocalyx than E. crassifolium. So, E. trichocalyx is generally the better host. However, beetle mate preference was not in line with food choice. Males did not prefer to mate with females from E. trichocalyx. Females from E. crassifolium did prefer males from E. trichocalyx over males from E. crassifolium, but did not lay more eggs as a result of these matings. We conclude that the beetle populations we studied have not differentiated based on their host plants and may not have even adapted to the better host. Although to humans these host plant defenses differ dramatically, signs that they have caused evolution in the beetles are lacking. The case of T. eriodictyonis stands counter to many other studies that have seen the differentiation of ecotypes and/or adaptive coordination of an herbivore's life cycle based on host plant differences.     

The influence of host plant variation and intraspecific competition on oviposition preference and offspring performance in the host races of Eurosta solidaginis

1. A series of experiments was conducted to measure the impact of plant genotype, plant growth rate, and intraspecific competition on the oviposition preference and offspring performance of the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and Solidago gigantea (Asteraceae). Previous research has shown that both host races prefer to oviposit on their own host plant where survival is much higher than on the alternate host plant. In this study, neither host race showed any relationship between oviposition preference and offspring performance in choosing among plants of their natal host species. 2. The larval survival of both host races differed among plant genotypes when each host race oviposited on its natal host species. In one experiment, altissima host race females showed a preference among plant genotypes that was not correlated with offspring performance on those genotypes. In all other experiments, neither the altissima nor gigantea host race demonstrated a preference for specific host plant genotypes. 3. Eurosta solidaginis had a preference for ovipositing on rapidly growing ramets in all experiments, however larval survival was not correlated with ramet growth rate at the time of oviposition. 4. Eurosta solidaginis suffered high mortality from intraspecific competition in the early larval stage. There was little evidence, however, that females avoided ovipositing on ramets that had been attacked previously. This led to an aggregated distribution of eggs among ramets and strong intraspecific competition. 5. There was no interaction among plant genotype, plant growth rate, or intraspecific competition in determining oviposition preference or offspring performance.