Determinants of oviposition in Acanthoscelides obtectus: a nonconformist bruchid

Abstract. Acanthoscelides obtectus is atypical of bruchids found in stored seeds because females do not attach eggs individually to host seeds but scatter them irregularly among potential hosts. In other respects, its life history resembles that of other species and comparison provides evidence for the adaptive nature of their behaviour. Individual females differ dramatically in the temporal pattern of their oviposition. The hypothesis is proposed that this is an adaptation to life in heterogeneous habitats. Oviposition is not regulated by host availability to the same extent as in other bruchids, providing evidence that regulation of the number of eggs laid by species which stick eggs to host seeds is an adaptation to the abundance of hosts. The continued presence of males and opportunity for multiple matings does not affect fecundity or egg viability. Females preferentially lay on clean seeds when provided with a choice between them and seeds previously exposed to other adult beetles. However, they do not appear to be deterred by, or cannot detect, the presence of larvae infesting seeds. Other bruchid species avoid such seeds that they can identify by the occurrence of egg cases and pheromones. The number of eggs laid is influenced by the host cultivar available for oviposition. Oviposition by A. obtectus appears to be an adaptation to both storage environments and the heterogeneous environment of a growing crop or uncultivated land. Its behaviour suggests an ancestral condition in bruchids and the changes which might have occurred in the ecology of other bruchid species as they have adapted to become pests of human seed stores.     

Plasticity in the egg-spacing behavior of a seed beetle: Effects of host deprivation and seed patchiness (Coleoptera: Bruchidae)

Egg-laying females of the seed beetle Callosobruchus maculatusdiscriminate between egg-free and egg-laden seeds and produce a nearly uniform distribution of eggs among seeds. We examined plasticity in this trait in response to both an internal factor (level of host deprivation) and an environmental one (the spatial configuration of available seeds). Responses to each factor were measured in genetically divergent strains that show a relatively strong (S strain) or weak (B strain) tendency to spread eggs evenly among seeds. Following a modest (10-h) period of host deprivation, S-strain females distributed their eggs less uniformly among seeds; the proportion of females committing at least one oviposition mistake increased from 20 to 50%. Similarly, S-strain females distributed their eggs less uniformly if seeds were presented in multiple, discrete patches instead of in a single, large patch. The higher frequency of oviposition mistakes in the multiple-patch arena was caused in part by females maintaining a uniform distribution of eggs within patches but not among patches. In contrast, females from the sloppier B strain were unaffected by either host deprivation or resource dispersion. Responses to seed patchiness are discussed in relation to the role of learning in the egg-spacing behavior of C. maculatus.     

Trophic-egg production in a subsocial bug: adaptive plasticity in response to resource conditions

Adomerus triguttulus (Heteroptera: Cydnidae) females provision host Lamium spp. seeds to their nymphs. Females also produce trophic eggs, which are inviable and usually function as a food supply for hatched nymphs. Here we report experimental evidence of the potentially adaptive maternal effects of this unusual resource investment. To investigate the effects of food-resource environments on trophic-egg production, we reared females under different resource conditions prior to oviposition and then compared the subsequent allocations of trophic eggs. Females that had been supplied with less-developed seeds produced fewer viable eggs than those supplied with well-developed seeds. However, there was no difference in the number of trophic eggs produced between the two treatments, and thus the trophic-egg ratio, i.e. the number of trophic eggs per viable egg, was higher in females supplied with less-developed seeds. Trophic-egg feeding by hatched nymphs enhances their growth or survival, and a higher trophic-egg ratio would be adaptive particularly under limited resource conditions. In A . triguttulus , adults and offspring depend on the same food resource, i.e. host seeds, and the resource conditions experienced by females prior to oviposition should be well correlated with those experienced later by the offspring. This may enable females to adjust the trophic-egg ratio in response to varying resource environments prior to oviposition.     

Development, fecundity and egg dispersion of Zabrotes subfasciatus

Zabrotes subfasciatus Boh. (Coleoptera: Bruchidae) is the smallest of the bruchids commonly infesting stored legume seeds, yet its wild and cultivated hosts, Phaseolus lunatus and P. vulgaris, have large seeds. It is demonstrated that the maximum fecundity of females is around 55 eggs which are aggregated onto some of the available hosts. About 80% of the eggs normally hatch and development at 27°C and 70% relative humidity takes around 34 days. The sex ratio of emerging adults is slightly biased towards males. About 75% of the larvae in a seed produce adults at low and moderate initial densities and up to 20 adults can emerge from a single seed. Adult weight is not influenced by the initial larval density in the seed but there is a strong correlation between the weight of females at emergence and their fecundity. These results are considered in the light of existing knowledge of Z. subfasciatus, much of which is apparently contradictory or inconsistent. Many of these difficulties are resolved and it is demonstrated that the behaviour and bionomics are well adapted to the normal situation in which the beetle is found and that the differences between this species and other bruchids are explicable in this context.     

Role of kin recognition in oviposition preference and cannibalism by the predatory mite <Emphasis Type="Italic">Gynaeseius liturivorus</Emphasis>

Animals often select oviposition sites to minimize the predation risk for eggs and juveniles, which are more vulnerable to predation than adults. When females produce eggs in clusters, the eggs and juveniles are likely to suffer from cannibalism. Although cannibalism among siblings is known to be lower than among non-siblings, there have been few investigations into the possibility that females select oviposition sites that reduce the risk of cannibalism for the offspring. To test this possibility, we examined oviposition preference by adult females of the predatory mite Gynaeseius liturivorus in response to the presence of her own eggs and to eggs of other females, offering plastic discs as oviposition substrates. Although females did not clearly show a preference for plastic discs on which they had oviposited, they avoided plastic discs on which other females had oviposited. When eggs of other females were artificially placed on clean plastic discs, adult female mites avoided these discs, suggesting that the eggs were used as cues for oviposition preference. Cannibalism among juvenile siblings was lower than among non-siblings. These observations show that adult females and juveniles of G. liturivorus discriminate kin relationships among conspecific individuals. Therefore, oviposition preference by adult female G. liturivorus may lead to the reduced risk of cannibalism among offspring.     

Ability of ovipositing seed beetles to discriminate between seeds with differing egg loads

Abstract. 1. Previous work has shown that ovipositing females of Callosobruchus maculatus (F.) avoid seeds already bearing eggs, and thereby reduce competition among their larval progeny within seeds.
2. This study demonstrates that females also detect small differences in egg density, and prefer to oviposit on seeds with a lower-than-average number of eggs. A nearly uniform dispersion of eggs is thus maintained even after all seeds bear several eggs.
3. In addition, variation in egg load influences oviposition rate. Transfer of females from seeds with few eggs to seeds with many eggs inhibits oviposition; the reverse transfer stimulates it.
4. The upper surface of the egg chorion or egg 'cover' remains intact on the seed surface after the larva has entered the seed and continues to deter egg-laying for at least as long as the period required for larval development. 14-day-old egg covers provide as much deterrence as freshly laid eggs.     

昆虫寄主标记信息素

综述了昆虫寄主标记信息素的研究进展。昆虫寄主标记信息素是指由昆虫产生的用来标记寄主上有同种个体存在的化学物质。昆虫寄主标记信息素的主要生态学功能是调节昆虫的产卵行为,通过阻止自身或同种其它个体对已标记寄主的产卵选择,或减少产卵量来减少后代之间对寄主资源的竞争。寄主标记信息素也会给释放着带来不利的影响,如信息盗用和盗寄生现象等。昆虫寄主标记信息素也调节昆虫近缘种之间对共同寄主资源的竞争。近缘种昆虫对相互寄主标记信息素识别能力的差异反映了不同昆虫对同一寄主资源竞争能力的强弱。寄主标记信息素产生和贮存的部位一般与外分泌腺、消化系统或生殖系统相联系,杜氏腺、毒腺、前胸腺、腹腺、下唇腺、后产卵管、卵巢、中肠和后肠等是产生或贮存寄主标记信息素的常见部位。产生的寄主标记信息素一般在成虫产卵时由产卵器、口器或排泄口释放到寄主体内或体表。卵寄生蜂的寄主标记信息素一般标记在寄主的体表,雌成蜂用触角检测;其它寄生蜂的寄主标记信息素常产在寄主体内,用产卵器检测;植食性昆虫的寄主标记信息素只产在寄主表面,用触角或产卵器检测。昆虫的产卵器、口器、触角或跗节上着生有感受寄主标记信息素的化感器,可以检测到标记在寄主体内或体表的寄主标记信息素。昆虫寄主标记信息素的完全定性涉及活性化合物的分离、鉴定、合成以及行为测定等,已有几种昆虫的寄主标记信息素成分得到了分离鉴定。     

Variance in number of eggs per patch: oviposition behaviour and population dispersion in a seed parasitic moth

ABSTRACT. 1. The relationship between oviposition behaviour and population dispersion was analysed for Grey a subalba Braun (Lep., Incurvariidae), a species in which females usually, but not invariably. lay eggs in only a few schizocarps (seed-pairs) on each umbellet they visit on Lomatium dissectum (Umbelliferae). During each of four years females exhibited a constant probability of leaving an umbellet after each egg was laid. Hence, some umbellets received more eggs than others and the behaviour matched a geometric distribution of eggs among umbellets.
2. The variance in number of schizocarps attacked per umbellet by individual females did not result from females choosing schizocarps of particular sizes or avoiding schizocarps that already contained eggs or larvae. A potential, but speculative, explanation of the variance is that females distribute their eggs among umbellets in a way that maximizes unpredictability on larval dispersion to a searching parasitoid, The pattern of oviposition by Greya females fits the truncated geometric distribution that is predicted if information, as indexed by the Shannon-Wiener entropy measure, is minimized. None the less, other measures of unpredictability or minimum information are possible, and the results highlight the problem of how to identify patterns of movement that may result from selection for unpredictability to enemies.
3. In systematic surveys of the L.dissectum population during three consecutive years, virtually all plants had between 10% and 65% of their seeds attacked. Hence, the separation of oviposition bouts between umbellets and the constant probability of leaving an umbellet after each egg was laid resulted in a broad distribution of Greya attack among L.dissectum plants.     

Post-dispersal predation of Acacia farnesiana seeds by Stator vachelliae (Bruchidae) in Central America

Summary Post-dispersal seed predation by the bruchid beetle Stator vachelliae was investigated in Santa Rosa National Park, Costa Rica. This insect finds the seeds of the leguminous Acacia farnesiana in the feces of horses, deer, and ctenosaur lizards, the current major dispersers. Patterns of oviposition and pre-adult survival of beetles in the seeds were investigated in a series of experiments using fresh horse dung. S. vachelliae never minded into the dung balls, attacking only those seeds located on the surface. Fresh horse dung did not attract insects more readily than dry dung. The proportion of seeds attacked was not related to their density in a defecation, and was similar in three areas with different densities of the host plant. In a fourth area with no fruiting A. farnesiana shrubs all seeds survived insect predation. Bruchids attacked a greater proportion of seeds at 1 m than at 5 m from the edge of the shrub's crown. Seeds were mainly removed from horse dung by rodents with similar intensity in all areas and at both distances; this seed removal interfered with bruchid oviposition and probably with bruchid survival. S. vachelliae oviposited less frequently on seeds in dung fully exposed to sun. When oviposition on a dung pile was high, the distribution of eggs on the seeds was clumped, suggesting that some seeds were preferred to others. By the end of the dry season, bruchids stopped attacking the seeds. The results show that the fate of both seeds and bruchids is greatly influenced by the location and time of defecation.     

Smells like aphids: orchid flowers mimic aphid alarm pheromones to attract hoverflies for pollination

Most insects are dependent on chemical communication for activities such as mate finding or host location. Several plants, and especially orchids, mimic insect semiochemicals to attract insects for unrewarded pollination. Here, we present a new case of pheromone mimicry found in the terrestrial orchid Epipactis veratrifolia. Flowers are visited and pollinated by several species of aphidophagous hoverflies, the females of which also often lay eggs in the flowers. The oviposition behaviour of these hoverflies is mainly guided by aphid-derived kairomones. We show that the flowers produce α- and β-pinene, β-myrcene and β-phellandrene, and that these compounds attract and induce oviposition behaviour in female hoverflies. This floral odour profile is remarkably similar to the alarm pheromone released by several aphid species, such as Megoura viciae. We therefore suggest that E. veratrifolia mimics aphid alarm pheromones to attract hoverflies for pollination; this is the first time, to our knowledge, that such a case of mimicry has been demonstrated.     

A preliminary investigation of pre-dispersal seed predation by Acanthoscelides schrankiae Horn (Coleoptera: Bruchidae) in Mimosa bimucronata (DC.) Kuntze trees

This is the first record of Acanthoscelides schrankiae Horn. feeding in seeds of Mimosa bimucronata (DC.) Kuntze. We investigated the pattern of oviposition and seed exploitation by A. schrankiae, and the distribution of mature fruits and seed predation in the inflorescences. We also compared the percentage of predated seeds, the total dry weight of fruits and non-predated seeds, the percentage of aborted seeds, and the percentage of non-emergent insects, among different quadrants of the M. bimucronata canopy. To determine the occurring species, the emergence of bruchids and parasitoids was observed in the laboratory, resulting altogether, only in individuals of A. schrankiae and Horismenus sp. (Hymenoptera: Eulophidae) species, respectively. Mean number of fruits produced in the median region of inflorescence was significantly higher than in the inferior and superior regions, and the frequencies (observed and expected) of predated and non-predated seeds differed among the different regions of inflorescence. Females of A. schrankiae laid their eggs on fruits, and larvae, after emergence, perforated the exocarp to reach the seeds. Most fruits presented one to three eggs and only one bruchid larva was observed in each seed. The highest value of the rate "number of eggs/fruit" and the highest percentage of predated seeds were recorded in April. Dry weight of fruits (total) and seeds (non-predated), proportions of predated seeds, seed abortions, and non-emergent seed predators, were evenly distributed in the canopy.     

Hydrologic and behavioral constraints on oviposition of stream insects: implications for adult dispersal

The supply of recruits plays an important role in plant and animal population dynamics, and may be governed by environmental and behavioral constraints on animals. Mated females of the mayfly genus Baetis alight on rocks protruding from streams, crawl under water and deposit a single egg mass under a rock. We surveyed oviposition and emergence of a bivoltine population of B. bicaudatus in multiple stream reaches in one high-altitude watershed in western Colorado over 3 years to establish qualitative patterns at a regional scale (entire watershed), and quantitative patterns over six generations at a local scale (one stream reach). We also measured characteristics of preferred oviposition substrates, performed experiments to test hypotheses about cues used by females to select oviposition sites, and measured mortality of egg masses in the field. Our goals were to determine whether: (1) hydrologic variation necessitated dispersal of females to find suitable oviposition sites; (2) the local supply of females could provide the supply of local recruits; and (3) local recruitment determined the local production of adults. The onset of oviposition corresponded with the decline of spring run-off, which differed dramatically among years and among sites within years. However, eggs appeared before any adults had emerged in 8 of 22 site-years, and adults emerged 2-3 weeks before any eggs were oviposited in 3 site-years. Furthermore, the size distribution of egg masses differed from that predicted by the size distribution of females that emerged from seven of nine stream reaches. Protruding rocks and eggs appeared earlier each summer in smaller tributaries than in larger mainstream reaches, suggesting that hydrologic and behavioral constraints on oviposition may force females to disperse away from their natal reach to oviposit, and possibly explain the predominantly upstream flight of Baetis females reported in other studies. Local oviposition rates in one third-order stream-reach increased rapidly as soon as substrates protruded from the water surface, and females preferred large rocks that became available early in the flight season. However, females oviposited on <10% of all available rocks, and <65% of preferred rocks as determined by an empirical model. These data indicated that the timing of appearance of suitable oviposition sites determined the phenology of local recruitment, but that preferred oviposition sites were not saturated. Thus, the magnitude of local recruitment was not limited by the absolute abundance of preferred oviposition sites. Only 22% of egg masses observed in the field suffered mortality during their embryonic development, and per capita Baetis egg mass mortality was significantly lower on rocks with higher densities of egg masses. Thus, we suspect that specialized oviposition behavior may reduce the probability of egg mortality, potentially compensating for the costs of dispersal necessary to locate suitable oviposition sites. Finally, the number of adults that emerged at one stream reach was independent of the number of egg masses oviposited over six generations of Baetis; and local recruitment was not a function of the number of adults of the previous generation that emerged locally. The patterns of oviposition and emergence of Baetis found in this study are consistent with the following hypotheses. Recruitment of eggs in a stream reach is not limited by the local supply of adults, but is a function of the regional supply of dispersing adults, which are constrained by the spatial and temporal distribution of preferred oviposition habitat. Furthermore, subsequent local production of adults is not a function of the supply of recruits, arguing for post-recruitment control of local populations by processes operating in the larval stage (e.g., predation, competition, dispersal, disturbance). Processes affecting larval and adult stages of Baetis act independently and at different scales, thereby decoupling local population dynamics of successive generations.     

Toxicity of secondary compounds to the seed-eating larvae of the bruchid beetle Callosobruchus maculatus

By incorporating various secondary compounds in the normal diet of larval Callosobruchus maculatus bruchids, we show that the effects of any particular compound are dosage-dependent. Alkaloids are generally the most toxic of the compounds tested. Non-protein amino acids are more toxic than protein amino acids but the latter can be toxic at 1 and 5% incorporation in the diet. The non-protein amino acid homoarginine has a salutary effect on larval survival at low concentrations. A variety of other secondary compounds found in seeds are toxic at various levels representative of those levels found in seeds in nature, and for all secondary compounds tested a 0.1–5% incorporation in the diet often has a detrimental effect on production of adult beetles. We conclude that many of the secondary compounds found in seeds are likely to be toxic to at least some animal, and thus are likely to be responsible at least in part for the extreme host-specifity shown by seed-eating insects.     

Egg performance on an egg-carrying bug. Experiments in the field

Selection of oviposition sites has direct influence on female fitness. Differences in offspring survival among sites should favour females to select oviposition sites with the highest survival. Golden egg bug females ( Phyllomorpha laciniata; Coreidae, Heteroptera) use conspecifics as oviposition substrates. Most eggs are laid on the back of the bug but they can be found on all body parts. Females never carry their own eggs, and males commonly carry eggs received without copulation with the donor. We examined differences in egg survival relative to paternity to the egg, host sex and body size and attachment of an egg on a host's body in the field. We also studied which bugs received eggs. Egg loss was surprisingly common: 30–80% of recaptured individuals lost eggs during a six-day period. Unexpectedly, host's paternity to the eggs did not affect egg survival. Also, egg loss did not differ among hosts in respect to other parameters studied. Males received more eggs than females, but survival of the eggs was not better on a male's back than on a female's back. Since egg loss is common in the golden egg bug, we suggest that female oviposition strategy to lay eggs on conspecifics is not perfect. This is due to active behaviour of potential hosts (for example, rejected oviposition attempts) or lack of suitable oviposition substrates (conspecifics) in the field. Since eggs do not survive unless carried, females may be acting opportunistically, doing the best job possible by laying eggs on available conspecifics.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

The role of chemical cues in host finding and acceptance by Callosobruchus chinensis

We studied the response of female Callosobruchus chinensis to chemical cues emitted by cowpea seeds at different stages of bruchid infestation (uninfested, egg carrying, L1-, and L4-infested). Olfactory attractiveness was determined in Y-tube olfactometer assays by testing individual seed categories against either clean air or uninfested seeds. Oviposition preferences between uninfested and infested seeds were determined in petri-dish choice-experiments. The olfactometer assays revealed that weevils discriminate between seeds containing different stages of developing bruchids on the basis of olfactory cues. While odors from uninfested and egg-carrying seeds acted as attractants, odors from L1- and L4-infested seeds failed to induce a positive response by the bruchids. When given a choice between uninfested and infested seeds in the olfactometer, weevils preferred uninfested seeds over L1- and L4-infested seeds, but failed to distinguish between uninfested and egg-carrying seeds. In the oviposition experiment as well, bruchids showed a distinct preference for uninfested seeds when offered in combination with L1- and L4-infested seeds. This experiment further showed a reduced acceptance of egg carrying seeds. Our results indicate that C. chinensis females use chemical information during both host searching and host acceptance. Volatiles from uninfested or egg carrying seeds act as attractants, while deterrence increases as development of bruchid immature stages progresses.     

Intraspecific competition in Zabrotes subfasciatus: Physiological and behavioral adaptations to different amounts of host

Abstract The effects of competition on populations of the bean weevil Zabrotes subfasciatus were analyzed during 41 generations under different competition levels. Three competition environments were established by maintaining the number of couples (6) and varying the amount of available host seeds: HC, high (limited availability of host: 1.35 g); IC, intermediate (intermediate availability of host: 6 g); and LC, low competition (abundance of host: 36 g). It was found that the distribution of the eggs laid on grains was different among treatments: in LC, for example, although females showed high fecundity (35.4 ± 5.6 eggs/female) the number of eggs laid on each grain was small (1.2 ± 0.4 eggs on each seed), thus avoiding larval competition of their offspring; whereas in HC treatment, females showed low fecundity (27.04 ± 4.5 eggs/female) but laid many eggs on each grain (15.03 ± 4.3 eggs). There were no changes in the ability to respond to different amounts of host via oviposition behavior (egg distribution) during 41 generations. However, HC females had more offspring than LC females under HC conditions. This suggests that HC insects evolved toward higher fitness in crowded conditions. In addition, after inverting the competition level, insects behaved independently of the treatment conditions they experienced through generations, thus showing that oviposition behavior is flexible. Taken together, our results show that Z. subfasciatus presents a broad range of behavioral and physiological responses which allows for quick and reversible adjustments to sudden changes in the amount of resources.     

Factors affecting egg maturation in the bean weevil Callosobruchus maculatus

Abstract. This paper concerns the effects of mate and seed availability on the rate of egg maturation in the bean weevil Callosobruchus maculatus (F.). Egg maturation starts before emergence from the seed and, provided that both oviposition sites and mates are available, eggs are laid at a rate determined by the number of oviposition sites, and mature at a similar rate. If seeds or mates are absent then a small number of eggs are laid, but oocytes continue to mature until the oviduct-capacity is approached. The number of eggs that a female can store is dependent on her body weight and does not correlate with the number of ovarioles. If, after a period in which oocyte development has been halted, conditions for egg-laying become suitable, then egg maturation can be re-started, but only after the oviducts have been emptied of eggs. The rate of egg maturation is then similar to that for females of the same age which have been maturing eggs since emergence.     

Identification of Communal Oviposition Pheromones from the Black Fly Simulium vittatum

The suite of pheromones that promote communal oviposition by Simulium vittatum, a North American black fly species, was identified and characterized using gas chromatography-mass spectrometry, electrophysiological, and behavioral bioassays. Behavioral assays demonstrated that communal oviposition was induced by egg-derived compounds that were active at short range and whose effect was enhanced through direct contact. Three compounds (cis-9-tetradecen-1-ol, 1-pentadecene, and 1-tridecene) were identified in a non-polar solvent extract of freshly deposited S. vittatum eggs that were capable of inducing the oviposition response. Electroantennography demonstrated that two of these three compounds (1-pentadecene and 1-tridecene) actively stimulated antennal neurons. Identification of the oviposition pheromones of this family may be helpful in developing control measures for nuisance black flies and for medically-important species such as Simulium damnosum sensu lato.     

Behavioral cues associated with oviposition by Trioza eugeniae

Trioza eugeniae oviposited significantly more eggs on Syzygium paniculatum leaves free of eggs compared to leaves with pre-existing eggs in a choice experiment, suggesting that females modify oviposition based on cues associated with the presence of eggs. In separate experiments, females avoided ovipositing on parts of leaves where eggs were present, but readily oviposited on areas of the same leaf without eggs. Epideictic pheromones are apparently not used by this species because females readily oviposited on areas of a leaf from which eggs had previously been removed. Females laid 57% of all their eggs along the margins of normal leaves, but also oviposited on artificial margins produced by cutting the leaves. The preference for natural margins over artificial margins was eliminated when the entire margin was coated with an antitranspirant compound to mask the damaged edge. These studies show that ovipositing psyllids respond both to the presence of eggs and to the suitability of the substrate.