Evolution of larval competitiveness and associated life‐history traits in response to host shifts in a seed beetle

Resource competition is frequently strong among parasites that feed within small discrete resource patches, such as seeds or fruits. The properties of a host can influence the behavioural, morphological and life‐history traits of associated parasites, including traits that mediate competition within the host. For seed parasites, host size may be an especially important determinant of competitive ability. Using the seed beetle, Callosobruchus maculatus, we performed replicated, reciprocal host shifts to examine the role of seed size in determining larval competitiveness and associated traits. Populations ancestrally associated with either a small host (mung bean) or a large one (cowpea) were switched to each other's host for 36 generations. Compared to control lines (those remaining on the ancestral host), lines switched from the small host to the large host evolved greater tolerance of co‐occurring larvae within seeds (indicated by an increase in the frequency of small seeds yielding two adults), smaller egg size and higher fecundity. Each change occurred in the direction predicted by the traits of populations already adapted to cowpea. However, we did not observe the expected decline in adult mass following the shift to the larger host. Moreover, lines switched from the large host (cowpea) to the small host (mung bean) did not evolve the predicted increase in larval competitiveness or egg size, but did exhibit the predicted increase in body mass. Our results thus provide mixed support for the hypothesis that host size determines the evolution of competition‐related traits of seed beetles. Evolutionary responses to the two host shifts were consistent among replicate lines, but the evolution of larval competition was asymmetric, with larval competitiveness evolving as predicted in one direction of host shift, but not the reverse. Nevertheless, our results indicate that switching hosts is sufficient to produce repeatable and rapid changes in the competition strategy and fitness‐related traits of insect populations.     

Adaptation to a novel host modifies host discrimination by the seed beetle Callosobruchus maculatus

Females of the seed beetle Callosobruchus maculatus avoid adding eggs to seeds that already bear eggs. Geographical variation in this behaviour has been thought to depend on differences in host size. In populations that attack small-seeded legumes, only one or two larvae can develop within a seed, and females are especially adept at detecting and rejecting occupied (egg-laden) seeds. We performed a mass-selection experiment in which replicate lines of a population associated with a small host (mung bean, Vigna radiata) were either maintained on this host or were transferred to a larger host (cowpea, Vigna unguiculata) that can support several larvae per seed. After more than 40 generations, we estimated the strength of host discrimination by presenting females a choice of egg-free and egg-laden seeds, and by quantifying how uniformly females spread their eggs among egg-free seeds. Compared to females maintained on mung bean, females from cowpea lines were more likely to accept occupied seeds in choice tests. They also distributed their eggs less uniformly, especially when cowpea served as the test host. Cowpea lines thus evolved to resemble populations that have long been associated with the larger host. A separate study showed that weaker host discrimination in the cowpea lines was accompanied by a decline in larval competitiveness, which may have further relaxed selection for avoidance of occupied hosts. Our results demonstrate that switching to a novel resource can produce rapid and predictable changes in a fitness-related insect behaviour. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

How labile are the egg‐laying preferences of seed beetles?

Abstract.  1. Previous studies have produced conflicting results with respect to the genetic lability of host preference in the seed beetle Callosobruchus maculatus .
2. In this study, replicate lines of an Asian population were kept on an ancestral host (mung bean) or switched to a novel host (cowpea). After 40+ generations, lines were assayed for host preference (in choice tests) and host acceptance (under no-choice conditions), and were compared to African lines chronically associated with cowpea.
3. Host preference diverged in the expected direction. When presented a mixture of cowpeas and mung beans, females from the cowpea lines laid a greater fraction of their eggs on cowpea than did females from the mung bean lines. Preference for cowpea was nearly as strong in the cowpea lines as it was in the cowpea-adapted African lines.
4. In contrast, the experimental host shift did not affect long-term host acceptance. African females laid more eggs if given cowpeas than if given mung beans, but realised fecundities in the cowpea and mung bean lines were similar on the two hosts. Females from all lines laid more eggs if they were reared on cowpea than on mung bean, but rearing host had no effect on either relative host acceptance or host preference.
5. Comparisons with earlier studies suggest that the lability of host preference varies among beetle populations, which precludes generalisation at the species level. Because lines were maintained under no-choice conditions, modification of host preference probably occurred via a lower acceptance threshold for the novel host, without a concomitant change in the long-term acceptance of the ancestral host.     

Predictable modification of body size and competitive ability following a host shift by a seed beetle

Interfertile populations of the seed beetle Callosobruchus maculatus differ genetically in several behavioral, morphological, and life-history traits, including traits that affect the intensity of larval competition within seeds. Previous studies have suggested that this variation depends on differences in host size. I performed a selection experiment in which replicate beetle lines were either maintained on a small, ancestral host (mung bean) or switched to a larger, novel host (cowpea). After 40 generations, I estimated survival, development time, and adult mass on each host, both in the presence and absence of larval competition. The shift to cowpea substantially reduced body size; irrespective of rearing host, adults from the cowpea lines were more than 10% lighter than those from the mung bean lines. Switching to cowpea also improved survival and reduced development time on this host, but without decreasing performance on the ancestral host. The most striking effect of the shift to a larger host was a reduction in larval competitiveness. When two even-aged larvae co-existed within a seed, the probability that both survived to adult emergence was > or = 65% if larvae were from the cowpea lines but < or = 12% if they were from the mung bean lines. The adverse effects of competition on development time and adult mass were also less severe in the cowpea lines than in the mung bean lines. By rapidly evolving smaller size and reduced competitiveness, the cowpea lines converged toward populations chronically associated with cowpea. These results suggest that evolutionary trajectories can be predictable, and that host-specific selection can play a major role in the diversification of insect life histories. Because host shifts by small, endophagous insects are comparable to the colonization of new habitats, adaptive responses may often include traits (such as larval competitiveness) that are not directly related to host use.     

Genetic architecture of population differences in oviposition behaviour of the seed beetle Callosobruchus maculatus

Few studies have examined the genetic architecture of population differences in behaviour and its implications for population differentiation and adaptation. Even fewer have examined whether differences in genetic architecture depend on the environment in which organisms are reared or tested. We examined the genetic basis of differences in oviposition preference and egg dispersion between Asian (SI) and African (BF) populations of the seed beetle, Callosobruchus maculatus. We reared and tested females on each of two host legumes (cowpea and mung bean). The two populations differed in mean oviposition preference (BF females preferred cowpea seeds more strongly than did SI females) and egg dispersion (SI females distributed eggs more uniformly among seeds than did BF females). Observations of hybrid and backcross individuals indicated that only the population difference in oviposition preference could be explained by complete additivity, whereas substantial dominance and epistasis contributed to the differences in egg dispersion. Both rearing host and test host affected the relative magnitude of population differences in egg dispersion and the composite genetic effects. Our results thus demonstrate that the relative influence of epistasis and dominance on the behaviour of hybrids depends on the behaviour measured and that different aspects of insect oviposition are under different genetic control. In addition, the observed effect of rearing host and oviposition host on the relative importance of dominance and epistasis indicates that the genetic basis of population differences depends on the environment in which genes are expressed.     

Rapid Evolution of Lifespan in a Novel Environment: Sex-Specific Responses and Underlying Genetic Architecture

Animal lifespans can vary substantially among closely related species and even among conspecific populations, but it is often difficult to identify environmental and genetic factors producing such variation. We used experimental evolution to examine how transfer to a novel environment affects adult lifespan and rates of senescence in a seed-feeding beetle. Three replicate lines of Callosobruchus maculatus (F.) were switched to a new host plant (cowpea), and each evolved shorter adult lifespans compared to a line maintained on the ancestral host (mung bean). However, the evolution of lifespan differed between the sexes; female lifespan was reduced by ~11% in all cowpea replicates, whereas male lifespan decreased by an average of only 5.6% and the magnitude of the reduction varied among replicates. Reduced lifespan in lines switched to cowpea mirrored the shorter lifespan observed in a separate population chronically associated with cowpea. We then performed crosses between the mung bean and cowpea lines to estimate the genetic architecture underlying the rapid evolution of a shorter lifespan on cowpea. Dominance (overdominance) contributed substantially to the difference between the cowpea and mung bean lines for female lifespan but not for male lifespan. However, details of the genetic architecture varied among the three replicate crosses, so that the convergent evolution of shorter female lifespan in the different cowpea lines did not arise from identical allelic substitutions. Our study demonstrates that insect lifespan can be predictably modified by a switch to a novel host plant, that both the magnitude of this response and its underlying genetic architecture can be sex-specific, and that convergent evolution of a complex trait such as lifespan can arise from different genetic mechanisms.     

Females of the weevil Zabrotes subfasciatus manipulate the size and number of eggs according to the host seed availability

Abstract. In some insects, the finding of oviposition substrate triggers the uptake into oocytes of yolk proteins that are stored in the fat body during post‐embryonic development. The main host of the bean weevil Zabrotes subfasciatus (Coleoptera; Chrysomelidae; Bruchinae; Amblycerini), in which larval resources are the sole source for future egg maturation, is Phaseolus vulgaris. Despite not feeding as adults, females of this species are able to lay eggs after encountering host seeds but it is not known how females react to changes in the availability of bean seeds. In the present study, the behaviour of Z. subfasciatus facing two very different environments for oviposition is investigated, as well as how this influences offspring fitness. The results obtained show that females of Z. subfasciatus react to variations in the availability of seeds belonging to the same host species by adjusting egg size and number. Females on low bean seed density lay larger and fewer eggs than those on high bean seed density, demonstrating a trade‐off between these reproductive traits. Moreover, females can adjust egg size to changing levels of host availability during the first 4 days of their oviposition period. Although no difference in offspring weight is found, those from small eggs (low competition environment) result in larger adults. No response to selection on these traits after rearing beetles on the same host for 40 generations is observed. This unresponsiveness may indicate that beetle populations behave according to their reaction norm that already allows rapid adaptation to a varying amount of host‐seed availability and better exploitation of the environments of this widespread stored‐seed pest.     

Mung bean (<Emphasis Type="Italic">Vigna radiata</Emphasis>) cultivars mediated oviposition preference and development of <Emphasis Type="Italic">Callosobruchus chinensis</Emphasis> (Coleoptera: Chrysomelidae: Bruchinae)

The Azuki bean weevil, Callosobruchus chinensis (L.), is a destructive pest of stored mung bean [Vigna radiata (L.) Wilczek] as well as other leguminous seeds. The development of resistant seeds to manage this pest is of current great interest to plant breeders. In this study, we investigated the oviposition preference and development of C. chinensis on two susceptible mung bean cultivars (Seonhwa and Gyeongseon) and one previously reported resistant cultivar (Jangan), compared to the susceptible cowpea (Vigna unguiculata L.), cultivar (Yeonbun) using both multiple-choice and no-choice tests. In addition, the development of C. chinensis was also examined at four constant temperatures (20, 25, 30, and 35 °C). Both tests found cowpea to be the most suitable seed for oviposition. Total developmental time from oviposition to adult emergence ranged from 27.01 to 38.2 days, being shortest on cowpea and longest on the mung bean, cv. Jangan. However, no successful development of C. chinensis larvae on mung bean, cv. Jangan, occurred at any temperature. The highest rate of adult emergence and the longest adult longevity both occurred on cowpea and certain mung bean cultivars (Seonhwa and Gyeongseon), with the dramatic exception of cv. Jangan. These results suggest that the higher preference and performance of C. chinensis on cowpea (3.3 egg/seed) and least on mung bean, cv. Jangan (0.4 egg/seed). This information may facilitate the exploration of resistant genetic materials and chemicals associated with seeds for successful breeding. Further studies should examine the chemicals associated with mung bean cultivars and its resistant mechanism to develop a control method against bruchines.     

Parasitism of grain legumes by Alectra species (Scrophulariaceae)

In glasshouse pot experiments in the United Kingdom, the host preference of nine seed samples of Alectra vogelii from Eastern, Western and Southern Africa and of two samples of A. picta from Cameroon and Ethiopia, to cultivars of cowpea, groundnut, bambara and mung bean, was assessed. A susceptible cowpea cultivar, Blackeye, and four cultivars of groundnut were attacked by all samples of both parasitic species regardless of whether the host of origin was cowpea, groundnut or bambara. Five “strains” of A. vogelii were distinguished using two criteria: their ability to parasitise bambara and/or mung bean and their ability to parasitise cowpea B301 and bambara TVU 870. The latter proved in an associated experiment to be resistant to collections of the parasite from some locations. Strain 1, including populations from Mali, Nigeria and Cameroon, attacked all groundnuts, cowpea cultivar Blackeye, but not cowpea line B301, mung bean or bambara. Strain 2, from Botswana, differed in attacking B301 and mung bean. Three other strains were identified which attacked susceptible lines of all four legume species. Strain 3 from Kenya failed to attack either cowpea B301 or bambara TVU 870, strain 4 from Malawi attacked cowpea B301, but not bambara TVU 870, while strain 5 from Northern Transvaal, South Africa, attacked bambara TVU 870, but not cowpea B301. Cowpea B359 was resistant to A. vogelii samples from all locations and also to A. picta, which has a similar host preference to strain 1 populations of A. vogelii from West Africa. Two out of 13 groundnut lines tested showed low susceptibility to A. vogelii from Cameroon suggesting there is scope for selecting resistance in this crop also.     

Evolutionary conservatism of geographic variation in host preference in Callosobruchus maculatus

Abstract.  1. A study of host preference of four pairs of populations of the cowpea weevil Callosobruchus maculates was carried out. The pairs had different geographical origins.
2. One population of each pair had been maintained for about 110 generations on cowpea Vigna unguiculata , the other population had been maintained on mung bean V. radiata . Half of the tested females from each population were raised on cowpea and exposed to this host prior to the assay; the other half was raised on mung bean. This design permitted assessment of the relative contributions of geographical origin, recent host use in the laboratory, and individual experience, to variation in host preference.
3. Host preference was assayed by letting the females oviposit on an equal-weight mixture of cowpea and mung seeds; two experiments were performed six generations apart.
4. Both experiments revealed a strong effect of geographical origin: populations originating from Nigeria laid a much greater proportion (68–86%) of their eggs on cowpea than those originating from Uganda and Yemen (30–42%); those from Cameroon were intermediate (56–60%). These preferences were not affected consistently by about 110 generations of laboratory evolution on one or the other host, or by experience of individual females.
5. These results indicate considerable geographical variation in host preference, and suggest that host preference is behaviourally inflexible and evolutionarily conserved.     

Direct and plant trait‐mediated effects of the local environmental context on butterfly oviposition patterns

Variation in the intensity of plant–animal interactions over different spatial scales is widespread and might strongly influence fitness and trait selection in plants. Differences in traits among plant individuals have been shown to influence variation in interaction intensities within populations, while differences in environmental factors and community composition are shown to be important for variation over larger scales. However, little is still known about the relative importance of the local environmental context vs. plant traits for the outcome of interactions within plant populations. We investigated how oviposition by the seed‐predator butterfly Phengaris alcon on its host plant Gentiana pneumonanthe was related to host plant traits and to local environmental variation, as well as how oviposition patterns translated into effects on host plant fruit set. We considered the local environmental context in terms of height of the surrounding vegetation and abundance of the butterfly's second host, Myrmica ants. The probability of oviposition was higher in plants that were surrounded by lower vegetation, and both the probability of oviposition and the number of eggs increased in early‐flowering and tall plants with many flowers in the three study populations. Flowering phenology, shoot height and flower production were, in turn, related to higher surrounding vegetation. Myrmica abundance was correlated with vegetation height, but had no effect on oviposition patterns. Oviposition and subsequent seed predation by the caterpillars strongly reduced host plant fruit set. Our results show that plant–animal interactions are context‐dependent not only because the context influences the abundance or the behavior of the animal interactor, but also because it influences the expression of plant traits that affect the outcome of the interaction. The results also demonstrate that heterogeneity in environmental conditions at a very local scale can be important for the outcomes of interactions.     

Larval competition within seeds: From the behaviour process to the ecological outcome in the seed beetle Callosobruchus maculatus

Abstract Environmental conditions experienced by organisms during development can have profound impacts on adult fitness and behaviour. Internally feeding larvae unable to leave the seed selected by their mother face limitations of resource suitability and competition. The host seed may guide the larval behaviour within the seed leading to differential intensity of competition and determining its process and outcome, which varies in strains of the legume seed beetle Callosobruchus maculatus (Coleoptera: Bruchidae). However, the intensity, process and outcome of larval competition in different hosts have yet to be simultaneously considered, the objective of the present study. Here we assessed the intensity, process and outcome of intrastrain larval competition as related to host type, and how they are interrelated. Larval competition was faced with two distinct strategies – scramble and contest competition depending on the insect strain and host seed species. The intensity of competition did not show any straight link with the process and outcome of competition. Only a single strain showed a contest competition process with likely interference between larvae, while the four other strains studied showed the process of scramble competition. The process of scramble competition, however, led to variable outcomes in mung beans based on larval competition curves. Such differences were not apparent on cowpea seeds and either the plateau or the peak expected on the larval fitness curves were not reached preventing the distinction of the competition outcome, a likely consequence of the egg laying behaviour of these strains limiting the maximum number of eggs laid per seed. Seed host species rather than seed size are the likely cause of the differences observed from the initial expectation. The strain showing the process of contest competition increased larval fitness with density of larvae emerged per seed regardless of the host species, an unexpected outcome based on theoretical models. In this case the egg laying behaviour of the adult female is probably the main fitness determinant of its progeny.     

Host suitability, respiration rate and the outcome of larval competition in strains of the cowpea weevil, Callosobruchus maculatus

Abstract.  Variation between strains of Callosobruchus maculatus in several life-history traits is well known. Differences in functional anatomy of egg pores and larval respiration rates have also been reported in strains from Brazil and Yemen. The response of five strains of C. maculatus to seeds of two host species, cowpea ( Vigna unguiculata ) and mung bean ( Vigna radiata ), was measured along with the larval respiration rates of the same strains on both hosts. There was significant variability of response to the two hosts. Strains with higher larval respiration rate (µL O₂/insect/day) showed higher seed consumption, which significantly affected adult emergence per seed on both hosts. This finding provides support for the hypothesis that differential feeding rate is an important mechanistic component of the larval competition outcome observed in strains of C. maculatus .     

Evolution of host acceptance and its reversibility in a seed beetle

1. Adapting to a low‐quality plant may require modification of an insect's digestive physiology, oviposition behaviour, or other host‐use traits. If colonising a marginal host entails a cost, a decay in adaptation would be expected after selection is relaxed, i.e. if populations on a novel host are reverted to their high‐quality ancestral host. 2. Replicate lines of the seed beetle Callosobruchus maculatus (F.) rapidly adapted to lentil seeds; larval survival rose from approximately 1 to ≥ 90%, and oviposition on lentil increased more than two‐fold. This study compared egg‐laying behaviour in lines that either remained on lentil or were reverted to the ancestral host, mung bean, for 22–62 generations. 3. Consistent with the trade‐off hypothesis, females from two reverted sublines showed decreased oviposition on lentil (estimated as lifetime fecundity), but host acceptance in a third subline was unchanged. In a short‐term assay, acceptance of lentil by newly emerged females was lower in each reverted subline than in the corresponding non‐reverted one. Because effective population sizes (determined from genome resequencing) were large throughout the experiment, this decline in host acceptance is unlikely to be explained solely by genetic drift. 4. Variation among replicates in the magnitude of the reversion effect was also observed in a previous study of larval survival. However, the pattern of variation for survival was not congruent with the pattern of variation for host acceptance in this study. Thus, genes mediating improved performance on lentil appear to be largely independent of those responsible for increased oviposition.     

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.     

Quantifying the effects of host discrimination on egg‐laying decision of the cowpea seed beetle,Callosobruchus maculatus

Many parasitic and endophagous insect species are capable of discriminating among the quality of their hosts. However, there is no appropriate way to quantify their discrimination performance. In this study, we quantified how oviposition of the cowpea seed beetle, Callosobruchus maculatus (Fabricius) (Coleoptera: Bruchidae), was affected by the relative contributions of both egg number and host size discrimination. The effect of egg density and resource heterogeneity on these discrimination performances was also explored. Egg‐distribution predictions were made by combining time‐dependent available resource fitness (egg discrimination) and host weight factors (size discrimination). The χ² test was then used for goodness‐of‐fit testing. The effects of both egg and size discrimination on oviposition in environments with different levels of resource heterogeneity were compared. It was found that host size, rather than the number of eggs on the host, plays a larger role in the egg‐laying decision for most individual seed beetles, especially when egg density is high. Host size discrimination behavior was reinforced when the beetles experienced increasing resource heterogeneity, but the performance might reach a plateau. This is the first quantitative evaluation of the effect of host discrimination on egg‐laying decisions of seed beetles.     

Loss of adaptation following reversion suggests trade‐offs in host use by a seed beetle

Experimental evolution has provided little support for the hypothesis that the narrow diets of herbivorous insects reflect trade‐offs in performance across hosts; selection lines can sometimes adapt to an inferior novel host without a decline in performance on the ancestral host. An alternative approach for detecting trade‐offs would be to measure adaptation decay after selection is relaxed, that is, when populations newly adapted to a novel host are reverted to the ancestral one. Lines of the seed beetle Callosobruchus maculatus rapidly adapted to a poor host (lentil); survival in lentil seeds increased from 2% to > 90% in < 30 generations. After the lines had reached a plateau with respect to survival in lentil, sublines were reverted to the ancestral host, mung bean. Twelve generations of reversion had little effect on performance in lentil, but after 25–35 generations, the reverted lines exhibited lower survival, slower development and smaller size. The most divergent pair of lines was then assayed on both lentil and mung bean. Performance on lentil was again much poorer in the reverted line than in the nonreverted one, but the lines performed equally well on mung bean. Moreover, the performance of the nonreverted line on mung bean remained comparable to that of the original mung‐bean population. Our results thus present a paradox: loss of adaptation to lentil following reversion implies a trade‐off, but the continued strong performance of lentil‐adapted lines on mung bean does not. Genomic comparisons of the reverted, nonreverted and ancestral lines may resolve this paradox and determine the importance of selection vs. drift in causing a loss of adaptation following reversion.     

Effect of experience on host preference inCallosobruchus maculatus (Coleoptera: Bruchidae): Variability among populations

Experiments using naive females established differences in host specificity among geographic strains of the seed beetleCallosobruchus maculatus; some females strongly preferred azuki bean over mung bean, and others failed to discriminate between the two hosts. We examined whether such congenital differences affect the degree to which host preference can be modified by experience. In choice tests, previous exposure to azuki bean increased the proportion of eggs laid on that host, but only in strains with a relatively low host specificity. Under more realistic, no-choice conditions, egg-laying experience affected oviposition rates mostly in strains with a high host specificity, but these experiments could not distinguish between the effects of a female's experience per se and her physiological state (i.e., egg load). Our results indicate that the likelihood of detecting an effect of experience on host choice depends on both the experimental protocol and the source of the test population. In natural populations ofC. maculatus, recent egg-laying experience probably plays little or no role in discrimination between host species but may influence discrimination between conspecific seeds that differ in quality.     

Oviposition deterrents of Callosobruchus maculatus (Coleoptera: Bruchidae)

Abstract Many bruchids distribute their eggs uniformly among the host seeds available to them and tend to avoid adding eggs to seeds bearing any eggs if pristine seeds remain. Several authors have postulated that the adult cowpea seed beetles, Callosobruchus maculatus (F.) (Coleoptera: Bruchidae), produce pheromones inhibiting oviposition by females exposed to them. It is shown that both sexes produce compounds which can be washed from glass surfaces in a variety of organic solvents which do not deter oviposition as such, but seeds coated with such washings are avoided by ovipositing females when they are selecting seeds for egg-laying and uncoated seeds are available. Some components, at least, of the products persist for at least 30 days, longer than the normal development time of insects in the seeds. The compounds affect the distribution of eggs among the available seeds. At least three different biotypes of the species produce, recognize, and respond in the same way to each other's products. It is emphasized that the eggs themselves may not be the site of the product produced by females but that the physical presence of eggs cannot be overlooked in any discussion of the possible role of pheromones in oviposition and egg distribution by bruchids.     

Host preference of the bean weevil Zabrotes subfasciatus

It is largely known that the range of an insect diet is mostly determined by oviposition behavior, mainly in species with endophytic larvae such as Zabrotes subfasciatus. However, the proximate factors determining host choice and the subsequent steps leading to the expansion or reduction of the host number and occasional host shifts are largely unknown. We analyzed various factors determining host preference of Z. subfasciatus through the evaluation of: (i) oviposition preference of a wild population of Z. subfasciatus on the usual host (bean) and unusual hosts (lentil, chickpea and soy), and the performance of the offspring; (ii) artificial selection for increasing preference for hosts initially less frequently chosen; (iii) comparison of oviposition behavior between two different populations (reared for ∼30 generations in beans or chickpeas, respectively); (iv) oviposition timing on usual and unusual hosts; and (v) identification of preference hierarchies. We found that when using unusual hosts, there is no correlation between performance and preference and that the preference hierarchy changes only slightly when the population passes through several generations on the less frequently accepted host. We also found a positive response to artificial selection for increasing oviposition on the less preferred host; however, when the host-choice experiment involved two varieties of the usual host, the response was faster than when the choice involved usual and unusual hosts. Finally, beetles reared on an unusual host (chickpea) for 26 generations showed similar good fitness on both usual and unusual hosts, indicating that the use of a new host does not necessarily result in the loss of performance on the original host. Nevertheless, this population showed lower fitness on the usual host than that of the original population, suggesting an underlying partial trade-off phenomenon which may contribute to a broadening of diet of this insect species.