Comparative population studies of threePieris butterflies,P. rapae,P. melete andP. napi,living in the same area

Utilization of patchy habitats by adult populations of three Pieris butterflies, P. rapae, P. melete and P. napi was studied throughout the flight season in an area of their coexistence, about 3×1.5 km, in a farm village in the mountains in Inabu, Aichi Prefecture. Field study was by the mark-recapture method. Results were analyzed by dispersal distances and recapture duration decay curves for adults of different age-classes estimated on the basis of physical condition of their wings, together with supplementary information of daliy egg-laying rate of females, obtained in field cages. Sexually immature, mated femals of P. rapae after teneral stage showed a migratory flight. On the other hand, reproductive females and all males of P. rapae were strongly resident within suitable habitats, and reproductive females begun to lay eggs abundantly at sunny places of newly suitable areas within a short period. P. melete seemed to disperse gradually from emerged stites and females of this species continued to lay some constant numbers of eggs for more than ten days over a wider area. P. napi appeared more like P. melete than P. rapae. The habitats of the three species can be characterized as follows:P. rapae, temporary, continued for pre-reproductive females but localized for reproductive females and all males, and unstable;P. melete, permanent, widespread, and stable;P. napi, permanent, localized, and stable. The numbers of generations of P. rapae, P. melete and P. napi were estimated to be about six, three and three, respectively. Seasonal fluctuations in the number of adults were influenced by the stability of their habitats, i. e., the population size fluctuated sharply in P. rapae, but it was much more stable in P. melete and P. napi. In view of these results, it can be said that P. rapae fits the general characteristics of a r-strategist whereas P. melete and P. napi are more K-strategic than P. rapae.     

The role of parasitoids in evolution of habitat and larval food plant preference by three Pieris butterflies

This article attempts to explain that parasitoids provide the evolutionary pressure responsible for relationships between habitat use and larval food plant use in herbivorous insects. Three species of butterflies of the genus Pieris, P. rapae, P. melete, and P. napi use different sets of cruciferous plants. They prefer different habitats composed of similar sets of cruciferous plants. In our study, P. rapae used temporary habitats with ephemeral plants, P. melete used permanent habitat with persistent plants, although they also used temporary habitats, and P. napi used only permanent habitat. The choice experiment in the field cages indicated that each of the three butterfly species avoided oviposition on plants usually unused in its own habitat, but accepted the unused plants which grew outside its own habitat. Their habitat use and plant use were not explained by intrinsic plant quality examined in terms of larval performance. Pieris larvae collected from persistent plants or more long lasting habitats were more heavily parasitized by two specialist parasitoids, the braconid wasp Cotesia glomerata and the tachinid fly Epicampocera succincta. The results suggest that Pieris habitat and larval food plant use patterns can be explained by two principles. The evolution of habitat preference may have been driven by various factors including escape from parasitism. Once habitat preference has evolved, selection favors the evolution of larval food plant preferences by discriminating against unsuitable plants, including those which are associated with high parasitism pressures. Received: December 3, 1998 / Accepted: January 20, 1999     

Spatio-temporal distribution patterns of two tachinid flies,Epicampocera succincta andCompsilura concinnata parasitizingPieris

Larvae of genus Pieris in the northern part of Kyoto City are parasitized by two tachinid flies:Epicampocera succincta, a specialist on genus Pieris, and Compsilura concinnata, a generalist with very wide host-range. We surveyed the parasitism rates of Pieris by both flies for two years at six study areas. In these study areas, there lived three host species in the genus Pieris: P. rapae, P. melete, and P. napi, but neither tachinid parasitized P. napi to any significant extent. In the mountainous district, P. rapae and P. melete coexisted and their populations were relatively continuous, while in the lowland, only P. rapae larvae were abundant in spring and autumn, but even they disappeared in summer. Parasitisms by E. succincta occurred mainly in mountainous district and never in the lowland. C. concinnata parasitized Pieris in all the areas, but its parasitisms occurred mainly in autumn. We analyzed the factors affecting the spatial and temporal patterns of parasitism rates and presumed that the temporal discontinuity of host population restricted the distribution of the specialist parasitoid.     

Host preference variation cannot explain microhabitat differentiation among sympatric Pieris napi and Pieris rapae butterflies

1. Often, closely related insect species feed on different host plant species, and the tremendous diversity of phytophagous insects is therefore attributed to host plant‐driven speciation. However, for most taxa, host use information comes from field observations of egg‐laying females or feeding caterpillars, which means that the underlying reason for a particular host‐affiliation is not easily determined. 2. Therefore, it is often unclear whether an insect feeds on a certain host because it prefers that plant to alternative hosts, or because the host distribution overlaps with the habitat requirements of the insect. 3. We ask to what extent a divergent host use in the field mirrors the host plant preferences of two closely related butterflies, Pieris napi and Pieris rapae (Pieridae). In nature, P. napi typically occurs in moister habitats than P. rapae. 4. We scanned several microhabitats at a field site in Southern Sweden during multiple years, and collected Pieris eggs from three different plants, Cardamine pratensis (wet meadows), Barbarea vulgaris (drier micro‐habitats) and Alliaria petiolata (intermediate areas). 5. As predicted, P. rapae eggs were more common than P. napi eggs on B. vulgaris, whereas all of the 358 individuals collected from C. pratensis were P. napi, indicating a divergence in host use between the Pieris species. However, under controlled laboratory conditions, both species had virtually identical oviposition preferences, laying eggs on all three plants, notably P. rapae also laying eggs on C. pratensis, indicating that habitat use, not plant preference, drives host plant use in nature.     

Body temperatures and behavioural thermoregulation strategies of threePieris butterflies in relation to solar radiation

Behavioural thermoregulation of 3Pieris butterfly species,P. rapae, P. melete andP. napi, was examined in relation to the intensity of solar radiation. To evaluate solar radiation intensity, the temperature (T_wr) was measured with a mercury thermometer whose bulb was covered with white cloth and exposed to direct sunlight. On clear days, the diurnal air temperature was between 16 and 28°C. The T_wt varied between 18 and 45°C, while the temperature in the shade was under 25°C. When the T_wt was under 28°C, the body temperatures (T_h) of butterflies closely coincided with it. Butterflies with T_b's under 26°C were resting, while those with T_b's between 26 and 28°C were basking. When T_wr was between 28 and 40°C, the butterflies were active and their T_b's were always lower than T_wr, never exceeding 36°C, though body temperatures could be artificially elevated easily up to the level of T_wr. When T_wr exceeded 40°C, butterflies showed species-specific heat-avoiding behaviour.P. rapae, whose habitat resources exist in the sun, intercepted solar radiation by closing the wings over the body.P. melete andP. napi, however, whose main habitat resources exist in the shade, moved into the shade. Strictly speaking, it is concluded that both butterflies, in many cases, leave shaded habitats for sunny habitats to elevate their T_b rather than enter the shaded habitats for heat-avoiding.     

Differential selection of host plants by twoPieris species: the role of oviposition stimulants and deterrents

Oviposition responses ofPieris rapae L. andP. napi oleracea Harris to nine crucifers, one Capparidaceae and one Tropaeolaceae were directly compared under controlled conditions. Chemical fractions from these plants were also tested on both insects for the presence of oviposition stimulants or deterrents. The results showed that plant chemistry is a key factor in differential selection of potential hosts by thesePieris species. Some plant species were equally acceptable to bothPieris species. However,P. rapae preferred cabbage over most test plants whereasP. napi oleracea strongly preferred plant species that were avoided byP. rapae. The observed preferences were explained in most cases by the presence of stimulants and deterrents in extracts of the plants. The twoPieris species have apparently evolved differential sensitivities to the chemical stimuli that trigger or deter oviposition. The balance of positively and negatively interpreted sensory signals evoked by plant chemicals obviously plays an important role in acceptance or rejection of a plant by both species. The role of specific glucosinolates and differing structure-activity relationships is suggested.     

Mate recognition in two sympatric species of butterflies,Pieris rapae andP. melete

Female butterflies should distinguish conspecific males from males of different species. The aim of the present study was to evaluate the female role in avoiding allospecific mating between 2 sympatric pierids,Pieris rapae crucivora (P. rapae hereafter) andP. melete. Since it is relatively rate to observe responses of a wild female to a courting male of different species, we substituted the wings of a female with those of a female of the other species. Such a female attracted allospecific males. FemaleP. melete took “mate-refusal posture” to courting males, regardless whether the male belonged to different species or not. FemaleP. melete discontinued the posture only when the male was conspecific. Such a behavioural difference indicates that the female discriminated its conspecific male. On the other hand, when only the mate-refusal posture was taken into account, a femaleP. rapae did not differently respond to courlship of males of bothP. melete andP. rapae. Having once approached, the maleP. melete attempted more frequently to copulate with its conspecific females than with allospecific ones, while the maleP. rapae indiscriminately attempted to copulate with both allospecific and conspecific females. The maleP. melete tended to persist in its courtship to females of bothP. rapae andP. melete who took the mate-refusal posture. The mate recognition system by male and female in these 2 species is discussed in connexion with the difference in mating behaviour pattern of the species.     

The role of host species, age and defensive behaviour on ovipositional decisions in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

The main objective of this study was to determine the extent to which host acceptance behaviour as related to host species, age, and defensive behaviour might explain the differences in host use that exist between two congeneric and sympatric species of parasitic wasps. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). Cotesia species differed in their responses to host species (P. brassicae (L.), P. napi (L.) and P. rapae) and developmental stage (early and late 1st, 2nd and 3rd instars). In no-choice tests, host acceptance by C. rubecula was higher for p. rapae and females did not distinguish among the 6 host ages. In contrast, when foraging for P. brassicae and P. napi, C. rubecula females more readily attacked early first instar. Cotesia glomerata showed a higher degree of behavioural plasticity towards acceptance of Pieris host species and host age than did C. rubecula. Cotesia glomerata females parasitized the three Pieris species and showed higher acceptance of first and second instars over third instar. Oviposition success was also influenced by host defensive behaviour. The frequency and the effectiveness of defensive behaviour rose with increasing age of the host, P. brassicae being the most aggressive Pieris species. Furthermore, the mean duration of C. glomerata oviposition was significantly reduced by the defensive reactions of P. brassicae, which would likely affect parasitoid fitness as oviposition time is positively correlated to clutch size in C. glomerata. Acceptance frequencies corresponded well to field reports of Pieris-Cotesia associations and to patterns of parasitoid larval performance, suggesting that the acceptance phase might be used as a reliable indicator of Cotesia host-specificity.     

Cue hierarchy for host plant selection in Pieris rapae

During oviposition, female insects utilize a wide variety of cues to find an optimal host for their offspring. These cues may be encountered simultaneously or sequentially, and females must rely on a hierarchical categorization to rank cues for optimal ovipositional choice. During alighting, cues are often encountered at different distances, which may influence hierarchical order. Cues that are observed at longer distances and are experienced first may be more influential on oviposition choice than those experienced at shorter distances. We tested the effects of two long‐distance cues, plant size and habitat structure, and two short‐distance cues, plant nitrogen level and predator presence, on Pieris rapae L. (Lepidoptera: Pieridae) oviposition choice through a series of two‐ and four‐choice tests in both greenhouse and field settings. We found that the long‐distance cues of plant size and habitat structure had the largest impact on P. rapae oviposition in laboratory and field settings, with females preferring large plants without habitat structure. Plant nitrogen level had a smaller impact on oviposition, and the presence of predators did not affect oviposition choice. However, plant size and plant nitrogen level had a synergistic relationship, with more eggs laid on large high‐nitrogen plants compared to large low‐nitrogen or small high‐nitrogen plants, suggesting that optimal cues throughout the entire searching process may be important for P. rapae in choosing a host plant.     

Plant-based host preference and larval competition of the tachinid parasitoid Epicampocera succincta

We investigated the causes of among-plant variation in the parasitism rate of Pieris melete larvae (Lepidoptera: Pieridae) by the parasitoid fly Epicampocera succincta (Diptera: Tachinidae). The rate of parasitism by E. succincta was much higher on the bitter cress Cardamine appendiculata than on any other cruciferous plants. Adult female flies were found to be more attracted to C. appendiculata than to other cruciferous plants for searching for hosts. The parasitoid appeared to be unable to distinguish P. melete from an alternative host, Pieris rapae larvae, that coexisted with P. melete on most crucifer plants. Similarly, E. succincta failed to avoid P. rapae parasitized by the braconid wasp Cotesia glomerata. C. glomerata is a superior competitor to E. succincta if occupying the same host, killing the host before the E. succincta larva can grow and depriving the larva of the chance to survive. E. succincta attacked P. rapae larvae on most cruciferous plants, many of which were already occupied by C. glomerata; only on Cardamine appendiculata was E. succincta free from interspecific competition, because only P. melete was found on this particular plant. However, the strong preference for the hosts on C. appendiculata incurred heavy intraspecific competition among larvae on this plant, killing as great a proportion of larvae as interspecific competition did on the other plants. The balance between the strength of intraspecific competition on the preferred plant and of interspecific competition on the other plants appeared to maintain plant preference by E. succincta. Received: November 13, 2000 / Accepted: April 27, 2001     

Interactions between the stem-mining weevils Ceutorhynchus napi Gyll. and Ceutorhynchus pallidactylus (Marsh.) (Coleoptera: Curculionidae) in oilseed rape

Abstract 1 The rape stem weevil Ceutorhynchus napi Gyll. and the cabbage stem weevil Ceutorhynchus pallidactylus (Marsh.) share the same habitat and food resource within the stems of oilseed rape, Brassica napus L. var. oleifera. Interactions occurring between these two sympatric species on this host were studied under both field and laboratory conditions. 2 The oviposition preference of C. pallidactylus and the within‐plant distribution of eggs and larvae were examined in field plots of oilseed rape. Female C. pallidactylus tended to lay their eggs in plants already infested by eggs and larvae of C. napi rather than in uninfested plants. The within‐plant distribution of the egg batches of C. pallidactylus did not differ significantly between uninfested plants and those preinfested by C. napi. Ovipositing females of C. napi and C. pallidactylus generally showed a significant preference for plants with larger stem diameter. 3 Laboratory choice tests provided further evidence for the oviposition preference of C. pallidactylus. Females laid significantly more eggs in leaves of plants that had been previously infested by C. napi than in leaves of previously uninfested plants. 4 Larvae of C. pallidactylus showed a significant shift of their feeding niche towards the stem base when feeding in individual plants attacked by both species. This possibly indicates ressource partitioning between C. pallidactylus and C. napi. The within‐plant distribution of C. napi larvae was not affected by the simultaneous attack of C. pallidactylus. 5 The size of the head capsule of full‐grown larvae of C. napi and C. pallidactylus was not significantly correlated with the diameter of the stem of their host plant or with the number of conspecific larvae within individual plants.     

Coexistence and niche segregation by field populations of the parasitoids Cotesia glomerata and C. rubecula in the Netherlands: predicting field performance from laboratory data

Field experiments with foraging parasitoids are essential to validate the conclusions from laboratory studies and to interpret differences in searching and host selection behaviour of parasitoid species. Furthermore, field experiments can indicate whether the parameters measured in the laboratory are relevant to elucidation of the ecological processes under study, such as adaptation or species interactions. In previous extensive laboratory studies we studied plant- and host-searching behaviour, host acceptance, host suitability; host plant preference, and learning of two congeneric parasitoids of Pieris caterpillars: the generalist Cotesia glomerata, which has been reported to attack several Pieridae species, and C. rubecula, a specialist of the small cabbage white Pieris rapae. In the present field study our aim was to verify the importance of these previous laboratory findings for explaining the performance of these two species in the field. We investigated experimentally whether parasitism on three Pieris species varied with parasitoid species and with food plant of the caterpillars. We exposed different types of host plants, infested with different Pieris species, to parasitism by natural populations of Cotesia species, by setting the experimental plants out in Brussels sprouts cabbage fields. Furthermore we made direct observations of parasitoid foraging in the field. In general, the field results confirmed our predictions on the range of host plant and host species used in the field. The two Cotesia species appear to coexist through niche segregation, since C. glomerata was mainly recovered from P. brassicae and C. rubecula from P. rapae. Although C. glomerata is a generalist at the species level, it can be a specialist at the population level under certain ecological circumstances. Our study shows the importance of variation in host plant attraction and host species acceptance in restricting host plant and host diet in the field. Furthermore the results suggest that, at least in the Netherlands, specialisation of C. glomerata on P. brassicae may occur as a result of C. rubecula outcompeting C. glomerata in P. rapae larvae. Received: 8 July 1999 / Accepted: 31 January 2000     

Antifeedant and oviposition-deterring activity of total ginsenosides against Pieris rapae

The purpose of this study was to evaluate the antifeedant and oviposition-deterring activity of total ginsenosides against P. rapae. Total ginsenosides exhibited increased antifeedant effects against P. rapae. The highest nonselective and selective antifeedant activity were observed at 2.0% concentration where ginsenosides caused antifeedant percentages of 86.09 and 88.90, respectively. The total ginsenosides showed significantly oviposition-deterring activity of 77.78% against oviposition of P. rapae at 1.0% concentration. Total ginsenosides had antifeeding activity against P. rapae and inhibitory effects on its oviposition. Ginsenosides could be used as an agent to prepare botanical new pesticidal formulations.     

Predation,thermoregulation, and wing color in pierid butterflies

Summary This paper explores two hypotheses about the relationships among predation, thermoregulation, and wing color in butterflies: First, that butterflies are susceptible to predation during thermally marginal periods (e.g., cool weather) when effective thermoregulation and flight are not possible; second, that Pieris butterflies are relatively unpalatable to visual predators, supporting the idea that the white wing pigment of Pieris represents aposematic coloration. Field experiments with Pieris and Colias in 1984 and 1985 demonstrate that substantial predation may occur during the morning period before butterflies are able to actively fly. Circumstantial evidence is presented to suggest that at least some of the predation is by small, cursorial mammals. Feeding experiments in the field using Grey Jays as predators indicate that Pieris napi and P. occidentalis are less palatable than other sympatric butterflies, including confamial Colias alexandra. These and previous results suggest that Pieris are edible but less preferred as prey by birds, and that the degree of palatibility may vary among Pieris species. The relatively low palatability of these Pieris is consistent with the hypothesis that their white pigmentation represents aposematic coloration; however, the cues by which potential bird predators might discriminate against Pieris have not been established.     

Constituents of host- and non-host plants deterring oviposition by the cabbage butterfly, Pieris rapae

Oviposition by Pieris rapae L. on cabbage was deterred by homogenized cabbage tissue sprayed onto intact plants. Ether extracts of cabbage also were deterrent, but water extracts were not. Hexane extracts of other host plants deterred oviposition and water extracts had little or no effect. Polar as well as non-polar extracts of non-host plants inhibited oviposition. Polar deterrents in the non-host crucifers, Erysimum cheiranthoides and Capsella bursa-pastoris may explain the avoidance of these plants by P. rapae. Chemical deterrents, as well as stimulants, apparently play a major role in the acceptance or rejection of plants as hosts by ovipositing female butterflies.

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Zusammenfassung Die Eiablage von Pieris rapae auf Kohl wurde gehemmt von einem Homogenisat aus Kohlblättern, das auf intakte Pflanzen gesprüht wurde. Ether-Extrakte der Kohlblätter wirkten im Gegensatz zu Wasserextrakten abschreckend. Ebenso zeigten Hexanextrakte anderer Wirtspflanzen eine hemmende Wirkung auf die Eiablage, während Wasserextrakte anderer Wirtspflanzen keine Wirkung aufwiesen. Dagegen aber verhinderten sowohl polare als auch unpolare Extrakte von Nicht-Wirtspflanzen die Eiablagen. Polare, abschreckende Substanzen in den Nicht-Wirtspflanzen Erysimum cheiranthoides und Capsella bursa-pastoris (Cruciferae) könnten Ursache dafür sein, dass P. rapae diese Pflanzen meidet. Chemische Substanzen mit sowohl abschreckender als auch stimulierender Wirkung spielen offensichtlich eine bedeutende Rolle für legebereite Schmetterlingsweibchen bei der Auswahl von Pflanzen als geeigneten Eiablageplatz.

     

A comparative study of the egg-laying behaviour and larval development of Pieris rapae L. and P. brassicae L. on the same host plants

Summary Pieris rapae and P. brassicae feed on the same host plants and have synchronized seasons. P. brassicae, whose larvae are twice the size of P. rapae, lays eggs in clusters of 40–100 eggs whereas P. rapae lays single eggs. In this paper we examine how egg clustering may be advantageous for P. brassicae. The larval development of each species was studied, and found not to differ significantly. P. brassicae larvae were observed to migrate from their host plant after defoliating it. A comparison of the efficiency of host plant utilization by the two pierid species was undertaken by measuring the effect of larval feeding on the growth of their host plants (kale and brussel sprouts). The results show that egg clustering is advantageous for larval fitness in terms of host resource exploitation, and we suggest that P. brassicae is adapted for ovipositing on clumped vegetation, while P. rapae is selected for exploiting isolated plants.     

Defense against Pieris rapae in cabbage plants induced by Bemisia tabaci biotype B

Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) and Pieris rapae L. (Lepidoptera: Pieridae) are serious pests of vegetables, that can occur simultaneously on a single cabbage plant [Brassica oleracea var. capitata L. (Brassicaceae)]. We determined whether pre‐feeding or infestation by B. tabaci on cabbage could induce physiological and biochemical responses of the plant against P. rapae. Developmental time, length, and weight of immature P. rapae, and defense‐related plant compounds (SOD, superoxide dismutase; POD, peroxidase; CAT, catalase; APX, ascorbate peroxidase) were measured. Development of P. rapae larvae was 21% slower on B. tabaci‐pre‐infested plants than on plants without B. tabaci infestation. When feeding on B. tabaci‐pre‐infested plants, 22% of P. rapae larvae pupated as compared with 83% on B. tabaci‐free plants. Weights of P. rapae from first to fourth instars that fed on B. tabaci‐pre‐infested plants were also reduced, whereas those of fifth instars and pupae were not. Similarly, body length of P. rapae from first to fourth instars was affected by B. tabaci pre‐infestation, whereas that of the fifth instars was not. Peroxidase and APX activities of the B. tabaci‐pre‐infested plants increased more than SOD and CAT. Peroxidase and SOD activities of B. tabaci and P. rapae co‐infested plants increased as compared with those of P. rapae‐infested plants; however, CAT and APX activities were not different between B. tabaci‐ and P. rapae‐infested plants. These results showed that B. tabaci infestation had a negative effect on P. rapae when they occurred simultaneously on the same host plant. The implications of the induced plant changes on the herbivore are discussed.     

DISSECTING CORRELATED CHARACTERS: ADAPTIVE ASPECTS OF PHENOTYPIC COVARIATION IN MELANIZATION PATTERN OF PIERIS BUTTERFLIES

In this study we address the question of how much of the covariation among phenotypic characters observed in natural populations is adaptive. We examine covariation among a set of phenotypic characters that describe the wing-melanization pattern of Pieris butterflies. Previous functional analyses of thermoregulatory performance allow us to predict a priori whether and how different wing melanic characters should be correlated. We quantify and analyze the variation in the wing-melanization pattern within species for a series of Pieris populations from relatively cool environments in North America and compare these results with the predictions based on our adaptive hypothesis. We consider adaptive covariation both for biogeographic variation among populations and for seasonal polyphenism (phenotypic plasticity) within populations. Our hypothesis correctly predicts many of the qualitative features of covariation in melanization among major regions of the wings, at the level of biogeographic variation among populations, for both males and females of Pieris occidentalis. When within-population variation is considered, agreement with the adaptive predictions varies considerably in different populations for both P. occidentalis and P. napi males and females. Agreement for P. napi, particularly the females, is generally poorer than for P. occidentalis. In both species, there is a consistent difference in melanization pattern between alpine and arctic sites; this difference is discussed in relation to the differences in the radiative environment between these two types of “cold” habitats. Our results suggest that some important aspects of phenotypic correlation among wing melanic characters in Pieris are adaptive. We emphasize the important distinction between covariation and co-occurrence of characters, and we discuss these results in relation to the extensive biogeographic variation and phenotypic plasticity (seasonal polyphenism) in Pieris wing-melanization patterns.     

Support for mending the matrix: resource seeking by butterflies in apparent non-resource zones

In conserving organisms, a bipolar view has generally been adopted of landscapes, in which resources are allocated to patches of habitats and the matrix ignored. Allocating additional resources to the matrix would depend on two conditions: first, that organisms search for resources in landscapes regardless of differences in vegetation types and resource availability; second, that when resources occur in the matrix they are used by species. Behavioural data linked to biotope and substrate types, on three pierid species (Pieris brassicae, P. rapae, P. napi), have been examined to explore the relationship between flight modes and resource availability. Search flight and resource use, taken as measures of resource quests, occur extensively in all biotopes and over all substrates in addition to direct linear flight, more typical of butterflies when migrating between habitat units. Resource seeking and using even exceed direct flight in some biotopes that have been regarded generally as lacking in resources for these butterflies. For example, such is the case for P. brassicae in woodland and for P. rapae in scrub. This finding supports the view for repairing the matrix and enhancing the general countryside outside habitat patches with resources increasingly being made for organisms. The proviso is that species’ requirements are adequately researched and resourced so as not to become sinks.     

Hypersensitivity reaction of Brassica nigra L. (Cruciferae) kills eggs of Pieris butterflies (Lepidoptera: Pieridae)

Summary Some individual plants of the mustard Brassica nigra in lowland California kill eggs of the Crucifer-specialist herbivores Pieris rapae and P. napi by producing a necrotic zone at the base of the egg, thereby apparently desicating it. This is a typical hypersensitivity reaction, but to an atypical stimulus. The eggs can be rescued by maintaining them in a saturated atmosphere. Attempts to demonstrate a bacterial or fungal agent associated with the reaction were unsuccessful.