Sex ratio regulation in Coccygomimus turionella Linnaeus (Hymenoptera: Ichneumonidae) and its ecological implications

Abstract. 1. Sex regulation and its relevance to the ecology of a k-selected pupal parasitoid was investigated in Coccygomimus turionellae.
2. The ratio varied with host size, females predominating in large hosts.
3. Egg transfer experiments and comparative mortality rates among the progeny of virgin and inseminated females demonstrated that the phenomenon was due to parental behaviour.
4. The study of the mechanism confirmed Aubert's hypothesis of host size estimation with the addition that the host had to be exposed from the poles and had to contain the proper host kairomones.
5. Host size was highly correlated to parasitoid fecundity via influences on size and longevity but not the number of ovarioles per female or daily egg production.
6. Host encounters are more limiting than egg production and host size had no effect on host acceptance.
7. Sex regulation allows a maximal host encounter rate at the same time that it maximizes the reproductive potential of female progeny.
8. Low host density increased the production of female progeny resulting in more offspring searching for the limited resource. Concealed hosts and a high host density resulted in a shift towards a Fisher 1:1 sex ratio.     

Experience-induced preference for oviposition repellents derived from a non-host plant by a specialist herbivore

Foraging adults of phytophagous insects are attracted by host‐plant volatiles and supposedly repelled by volatiles from non‐host plants. In behavioural control of pest insects, chemicals derived from non‐host plants applied to crops are expected to repel searching adults and thereby reduce egg laying. How experience by searching adults of non‐host volatiles affects their subsequent searching and oviposition behaviour has been rarely tested. In laboratory experiments, we examined the effect of experience of a non‐host‐plant extract on the oviposition behaviour of the diamondback moth (DBM), Plutella xylostella, a specialist herbivore of cruciferous plants. Naive ovipositing DBM females were repelled by an extract of dried leaves of Chrysanthemum morifolium, a non‐host plant of DBM, but experienced females were not repelled. Instead they were attracted by host plants treated with the non‐host‐plant extract and laid a higher proportion of eggs on treated than on untreated host plants. Such behavioural changes induced by experience could lead to host‐plant range expansion in phytophagous insects and play an important role in determining outcome for pest management of some behavioural manipulation methods.     

Foraging behaviour of a leafminer parasitoid in the field

Abstract. 1. The searching behaviour of the parasitoid Sympiesis sericeicornis Nees (Hymenoptera, Eulophidae) is analysed under field conditions.
2. Females were tracked when flying around the canopy of apple trees infested with the tentiform leafminer Phyllonorycter cydoniella (D. & S.)(Lepidoptera, Gracillariidae) and their behaviour studied at the level of a single mine and at the level of several mines on the same leaf (patch level).
3. The presence of mines is detected during flight. The foraging behaviour at the patch level is efficient: the female is able to distinguish the suitable from the unsuitable hosts and handle the suitable hosts first. Not all mines are visited before departure from the leaf and those left unvisited are exclusively mines without suitable hosts. The patch leaving rule is discussed.
4. An ethogram of the behaviour on the mine is constructed. Selected behavioural categories are analysed for different combinations of mine content and outcome of the visit to the mine. The assessment of the mine content is done in a differential way, quickly and without mistakes.
5. The results are discussed in the context of two environmental factors, wind speed and temperature, which reduce the available time for searching. They are also related to the growing percentage of unsuitable hosts during each host generation. Tentiform mines remain attractive for a long period of time, even after parasitism or emergence of the adult, and their examination reduces the rate of host discovery.     

Plant structure and the foraging success of Aphidius rhopalosiphi (Hymenoptera: Aphidiidae)

Abstract. 1. A study of searching behaviour on wheat and host preference of Aphidius rhopalosiphi (DeStefani-Perez) is described.
2. Parasitoids divided their time equally between the leaves but spent very little time on the ear.
3. After contact with honeydew or an aphid host, parasitoids were arrested in a particular area and increased the time spent searching.
4. Aphids feeding on the ear were parasitized less successfully, since their position between the grains protected them from parasitoid attacks.
5. A.rhopalosiphi exhibited no preference for Metopolophium dirhodum (Walker) or Sitobion avenae (Fabricius), although the handling time for the latter was significantly longer; this resulted in fewer S.avenae being parasitized when it was abundant. Parasitoids did not switch between hosts in these experiments.
6. As a result of its searching behaviour, A.rhopalosiphi will encounter and parasitize M.dirhodum feeding on the leaves more frequently than S.avenae, which feeds on the ear. This will limit the parasitoid's ability to regulate populations of the cereal aphid S. avenae.     

Potential novel hosts for the lily leaf beetle Lilioceris lilii Scopoli (Coleoptera: Chrysomelidae) in eastern North America

Abstract.  1. Introduced insects often incorporate native plants into their diets and might be expected to show a predilection for novel hosts that are phylogenetically related to their normal hosts. The lily leaf beetle, Lilioceris lilii (Coleoptera: Chrysomelidae), is an introduced pest of cultivated lilies. Oviposition behaviour, larval behaviour, and development of L. lilii was examined on a range of potential host plants, as well as on the normal host, Asiatic hybrid lilies Lilium sp.
2. Neonate larval feeding behaviour was quantified on 15 food plant species: 10 from the Liliales, three from the Asparagales and two eudicots. Larvae fed plants closely related to the genus Lilium were more likely to initiate feeding, less likely to abandon their food leaf, and consumed more leaf area.
3. In no-choice tests, females oviposited on the novel hosts Lilium philadelphicum , Medeola virginiana , Clintonia borealis , Streptopus amplexifolius , and Polygonatum biflorum ; however, all but L. philadelphicum received very few eggs. Non- Lilium novel hosts were not used for oviposition when presented along with Asiatic lilies in choice tests.
4. A single individual was reared to the adult stage on the novel host S. amplexifolius . Several larvae survived to the pupal stage on M. virginiana , although no adults emerged from those pupae. Larvae reared on the native wood lily L. philadelphicum performed equally well or better than on the Asiatic cultivar.
5. Our results indicate that the lily leaf beetle poses a threat to native Liliaceae. Several native Lilium species, including L. philadelphicum , are threatened or endangered in certain jurisdictions throughout their range; these species should be monitored closely for colonisation by the beetle.     

Plant cues influence searching behaviour and parasitism in the egg parasitoid Trichogramma nr. brassicae

Abstract. 1. The effect of experience of plant cues at emergence on searching behaviour and parasitism was investigated in two strains of Trichogramma nr. brassicae (Hymenoptera: Trichogrammatidae).
2. Wasps reared on Sitotroga cereallela were allowed to emerge on a tomato plant or in a vial, with or without food. For one of the strains, females emerging on tomato searched significantly longer on a tomato seedling than females emerging in a vial.
3. The experience effect lasted between 1 and 2 days. It involves associating plant stimuli with the remains of the host, because females emerging from their host on a tomato plant had an increased searching response to tomato, but females transferred to the plant immediately after emergence did not.
4. The effect of emergence environment on parasitism of the host Helicoverpa punctigera on tomato and lettuce seedlings was tested. Wasps were allowed to emerge on seedlings of tomato or lettuce, or in an empty vial. For one strain, females experienced with tomato at emergence parasitized significantly more hosts on tomato than did females emerging on lettuce or in a vial. Parasitism on lettuce was not influenced by the treatments.     

The ecological significance of differences in host detection behaviour in coexisting parasitoid species

1. Six species of drosophilid parasitoids of the genus Leptopilina (Hymenoptera: Eucoilidae) exist in Europe. Laboratory experiments reveal that the fundamental niches for most of these species overlap. Field collections indicate that overlap in their realised niches is much smaller.
2. Here data are presented for the little known species Leptopilina longipes. Field and laboratory observations indicate that its niche overlaps with both that of L. clavipes and L. heterotoma .
3. The host detection behaviour of L. longipes is examined and it is shown that L. longipes uses vibrotaxis to detect its hosts. This behaviour of L. longipes is compared with that of L. heterotoma and L. clavipes , which both use the ovipositor in host detection.
4. The ways in which differences in host searching behaviour can contribute to niche segregation and coexistence of these species are discussed.     

Learning pays off: influence of experience on host finding and parasitism in Lariophagus distinguendus

Abstract. 1. The influence of experience on egg maturation, parasitism rate, and behaviour during host searching was investigated for Lariophagus distinguendus (Först.) (Hymenoptera: Pteromalidae) parasitizing larvae of the granary weevil Sitophilus granarius (L.) in grains of wheat Triticum aestivum L.
2. Dissection of female parasitoids and parasitism bioassays at high host density revealed that experience with hosts (e.g. by oviposition or by host feeding) is not required either for triggering oogenesis or for oviposition.
3. In parasitism experiments at low host density, when single host-infested grains were offered within a bulk of healthy grains, host finding and parasitism rate were increased by experience.
4. Behavioural observations revealed that searching time required for finding an infested grain was shorter for experienced parasitoids than for naive parasitoids, because travel time from grain to grain is shorter for experienced parasitoids, and because experienced parasitoids spend less time than naive parasitoids on non-infested grains.
5. In conclusion, experience due to host exposure increases parasitism and thereby the fitness of the parasitoids. It is discussed that this increase is more likely due to learning than to different egg load dynamics of experienced parasitoids.     

The probing behaviour of nymphs of Vanduzeea arquata and Enchenopa binotata (Homoptera: Membracidae) on host and non-host plants

Abstract. 1. Nymphs of Vanduzeea arquata Say have been found to be more host-specific in nature and to show a higher degree of selectivity in host discrimination experiments than nymphs of Enchenopa binotata (Say), It was hypothesized that this differential selectivity would be reflected in the probing behaviour of individuals placed on twigs of host and non-host plants. Probing behaviour was examined by direct observation of nymphs and by sectioning and staining the probed plant tissues.
2. All nymphs probed readily and for extended periods on both host and non-host twigs. E.binotuta nymphs showed no consistent differences in probing behaviour on hosts versus non-hosts, but V.atquuta nymphs were more likely to withdraw their stylets within 60 s when on non-host twigs and produced honeydew only when on their host species. V.urquatu nymphs reached the phloem sieve elements only when on host twigs and broke many cells in peripheral plant tissue layers while probing. E.binotata nymphs broke few cells and often reached the phloem of non-host as well as host plants.
3. Nymphs of V.arquata always reject non-host plants, apparently in the course of probing and prior to encountering the phloem sap. Chemical compounds released from ruptured parenchyma cells may act as probing stimulants or inhibitors. E.binotura nymphs often feed on non-host plants in a non-choice situation; their preferential settling on host twigs in discrimination experiments may reflect a tendency to abandon non-host twigs more readily than host twigs.     

Oviposition behaviour and food plant discrimination in leafhoppers of the genus Oncopsis

Abstract. 1. Field host plants of the six British species of Oncopsis are described. O. avellanae is unusual in being associated with both Corylus and Alnus.
2. Laboratory experiments on nymphal feeding preferences, nymphal and adult survival, and adult oviposition preferences are described for all species, using Carpinus, Corylus, Alnus and Betula species as test plants.
3. All experiments demonstrated considerable specificity in feeding, and especially oviposition.
4. It is suggested that discriminative oviposition behaviour is the major determinant of observed field distribution patterns.     

The influence of host plant diversity and food quality on larval survival of plant feeding heteropteran bugs

Abstract. 1. In a laboratory experiment, the influence of host plant diversity and food quality, in terms of nitrogen content, on the larval survival of two oligophagous bug species (Heteroptera, Miridae: Leptopterna dolobrata L., Notostira erratica L.) was investigated. Both species are strictly phytophagous and capable of feeding on a wide range of grass species. Moreover, they typically change their host plants during ontogenesis; it has been suggested that this behaviour is a response to the changing protein content of the hosts.
2. To investigate the importance of host plant diversity for these insects, the development of insects reared on grass monocultures was compared with that on mixtures of four grass species. In addition, the host grasses were grown under two nitrogen regimes to test whether nitrogen content is the key factor determining host plant switching.
3. Both species had a significantly higher survival rate when feeding on several host plants but only L. dolobrata showed a significant response to food nitrogen content. Furthermore, there was no correlation between the nitrogen content of the host plants and the survival rate of N. erratica larvae.
4. The study suggests that at least some Stenodemini need a variety of host plants during larval development but that the level of host plant nitrogen is not the main factor responsible for the observed diversity effect.     

The biology and host searching behaviour of a Cynipoid parasite of aphidophagous syrphid larvae

Abstract. 1. Callaspidia defonscolombei was found to be a common, solitary, thelytokous parasitoid of aphidophagous syrphid larvae. In 1977 it was bivoltine with adults being most numerous in early August and late September. It is probably polyphagous.
2. An analysis of host searching behaviour suggests that females respond to an aphid odour bringing them to the region of an aphid colony.
3. Syrphid hosts are located within the aphid colony by response to a contact chemical. There is a host size preference, only larger second and third instar larvae being attacked.
4. Females inject a venom, which temporarily paralyses host larvae, and place the egg inside a cerebral ganglion.     

Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper

Abstract.  1. Both host plant nutrition and mortality from natural enemies have been predicted to significantly impact host plant selection and oviposition behaviour of phytophagous insects. It is unclear, however, if oviposition decisions maximise fitness.
2. This study examined whether the salt marsh planthopper Pissonotus quadripustulatus prefers higher quality host plants for oviposition, and if oviposition decisions are made so as to minimise mortality at the egg stage.
3. A controlled laboratory experiment and 4 years of field data were used to assess the rates of planthopper oviposition on higher quality 'green' and lower quality 'woody' stems of the host plant Borrichia frutescens . The numbers and percentages of healthy eggs and eggs that were killed by parasitoids or the host plant were recorded.
4. In all years, including the laboratory experiment, Pissonotus planthoppers laid more eggs on lower quality woody stems than on higher quality green stems. While host plant related egg mortality was higher in woody stems, the percentage of eggs parasitised was much greater in green stems. This resulted in a lower total mortality of eggs on woody stems.
5. The results of this study demonstrate that, although Pissonotus prefers lower quality host plants for oviposition, this actually increases fitness. These data seem to support the enemy free space hypothesis, and suggest that for phytophagous insects that experience the majority of mortality in the egg stage, oviposition choices may be made such that mortality is minimised.     

Cardiac glycosides and oviposition by Danaus plexippus on Asclepias fruticosa in south-east Queensland (Australia), with notes on the effect of plant nitrogen content

Abstract. 1. Previous studies which attempt to relate oviposition by female monarchs, Danaus plexippus (L.), to the cardiac glycosides of their milkweed host plants are reviewed and found to be defective.
2. Here we followed ovipositing females in the field and recorded their oviposition behaviour on individual Asclepias fruticosa (L.) plants. The cardenolide glycoside and per cent nitrogen of each of these plants and also of a control set of plants was recorded.
3. Milkweeds accepted by females, on which egg laying took place, had significantly lower mean cardenolide levels compared with rejected plants and a control sets of plants.
4. Most oviposition ( c. 70%) took place on plants with intermediate cardenolide (CG) levels (=200–300 μg/0.1 g DW).
5. The mean CG level in A.fruticosa was 345 μg/0.1 g DW (SD = 138, n =122, range 119–719).
6. There was no relationship between monarch oviposition behaviour and plant per cent nitrogen.     

The influence of food plant dispersion on caterpillar searching success

Abstract. 1. Collards (Brassica oleracea L.) grown in clumps were more difficult for unstarved, early instar cabbage white caterpillars (Pieris rapae L.) to find than collards spaced at regular intervals, although total plant densities were identical.
2. Early instar, unstarved larvae are less mobile than later instar larvae (our data) or starved larvae (Jones' 1977 data).
3. Starved larvae and late instar larvae tend to find plants more readily than do unstarved or younger larvae.
4. We contend that larval mobility governs the success of searching caterpillars and their sensitivity to variations in plant dispersion.
5. Our results show that the dispersion of targets can influence searching success even though target density is constant. This means that the degree of clumping in host plants is capable of influencing their availability to herbivores.     

Limits to the abundance of rare species: an experimental test with a tree aphid

Abstract. 1. The birch ( Betula )-feeding aphid, Monaphis antennata, is always found at low densities on individual hosts and has low local abundance, but another birch-feeding aphid, Euceraphis betulae , is often found at high densities on individual hosts and has high local abundance.
2. This work attempts to establish whether the interaction between M. antennata and its host or the behaviour of individuals limits its densities.
3. Both species were reared on saplings, and population sizes were monitored for 6 weeks. Two levels of host quality were used and feeding space was kept constant throughout the experiment. Adults were prevented from leaving the saplings by clipping their wings, and predators were excluded.
4. On plants of similar host quality, both species achieved similar population sizes.
5. It is concluded that resource availability or the interactions between individuals are unlikely to be important causes of rarity.     

Do caterpillars disperse their damage?: larval foraging behaviour of two specialist herbivores,Euphydryas phaeton (Nymphalidae) and Pieris rapae (Pieridae)

Abstract. 1. To examine ecological and evolutionary aspects of caterpillar foraging behaviour, this study focused on observation of the individual foraging behaviour of two lepidopteran species, Pieris rapae L. and Euphydryas phaeton (Drury), on their respective host plants.
2. Periodic observations over the course of a day showed that the larvae move considerable distances, forage on the upper surfaces of leaves, and often immediately leave areas from which they have fed, leaving a pattern of dispersed herbivory.
3. Differences in foraging behaviour were not found between the two species, even though one species is aposematic and the other is cryptically coloured, but there were significant differences in the foraging patterns of P.rapae on the two host plants, broccoli and radish.     

Patterns of parasitism by insect parasitoids in patchy environments

Abstract. 1. This paper shows how the different spatial patterns of per cent parasitism in patches of different host density can be explained within a single model framework that takes into account the parasitoid's aggregative response, and the factors limiting the degree of host exploitation within patches.
2. Two contrasting laboratory examples are presented in which the distribution of searching parasitoids and the resulting levels of parasitism in different patches are both known for a range of parasitoid densities.
3. A model is described predicting the number of hosts parasitized per patch, in which the number of parasitoids searching is determined from a simple expression allowing different degrees of aggregation.
4. The model generates patterns of parasitism encompassing the two laboratory examples and a wide range of examples from the field.
5. The importance of density dependent spatial distributions of parasitism to population stability is briefly discussed.     

Time allocation of a parasitoid foraging in heterogeneous vegetation: implications for host-parasitoid interactions

1. Changing plant composition in a community can have profound consequences for herbivore and parasitoid population dynamics. To understand such effects, studies are needed that unravel the underlying behavioural decisions determining the responses of parasitoids to complex habitats. 2. The searching behaviour of the parasitoid Diadegma semiclausum was followed in environments with different plant species composition. In the middle of these environments, two Brassica oleracea plants infested by the host Plutella xylostella were placed. The control set-up contained B. oleracea plants only. In the more complex set-ups, B. oleracea plants were interspersed by either Sinapis alba or Hordeum vulgare. 3. Parasitoids did not find the first host-infested plant with the same speed in the different environments. Sinapis alba plants were preferentially searched by parasitoids, resulting in fewer initial host encounters, possibly creating a dynamic enemy-free space for the host on adjacent B. oleracea plants. In set-ups with H. vulgare, also, fewer initial host encounters were found, but in this case plant structure was more likely than infochemicals to interfere with the searching behaviour of parasitoids. 4. On discovering a host-infested plant, parasitoids located the second host-infested plant with equal speed, demonstrating the effect of experience on time allocation. Further encounters with host-infested plants that had already been visited decreased residence times and increased the tendency to leave the environment. 5. Due to the intensive search of S. alba plants, hosts were encountered at lower rates here than in the other set-ups. However, because parasitoids left the set-up with S. alba last, the same number of hosts were encountered as in the other treatments. 6. Plant composition of a community influences the distribution of parasitoid attacks via its effects on arrival and leaving tendencies. Foraging experiences can reduce or increase the importance of enemy-free space for hosts on less attractive plants.     

Host-plant selection by insects – a theory based on `appropriate/inappropriate landings'' by pest insects of cruciferous plants

Seven hypotheses, including the `Resource Concentration Hypothesis' and the `Enemies Hypothesis', have been put forward to explain why fewer specialist insects are found on host plants growing in diverse backgrounds than on similar plants growing in bare soil. All seven hypotheses are discussed and discounted, primarily because no one has used any of them to produce a general theory of host plant selection, they still remain as hypotheses. However, we have developed a general theory based on detailed observations of insect behaviour. Our theory is based on the fact that during host plant finding the searching insects land indiscriminately on green objects such as the leaves of host plants (appropriate landings) and non-host plants (inappropriate landings), but avoid landing on brown surfaces, such as soil. The complete system of host plant selection involves a three-link chain of events in which the first link is governed by cues from volatile plant chemicals, the central link by visual stimuli, and the final link by cues from non-volatile plant chemicals. The previously `missing' central link, which is based on what we have described as `appropriate/inappropriate landings', is governed by visual stimuli. Our theory explains why attempts to show that olfaction is the crucial component in the central link of host plant selection proved intractable. The `appropriate/inappropriate landings' theory is discussed to indicate the type of work needed in future studies to improve our understanding of how intercropping, undersowing and companion planting can be used to optimum effect in crop protection. The new theory is used also to suggest how insect biotypes could develop and to describe why pest insects do not decimate wild host plants growing in `natural' situations.