Larval and adult seed consumption affected by the degree of food specialization in Amara (Coleoptera: Carabidae)

Many carabid beetles (Coleoptera: Carabidae) are known to feed on plant seeds, but the level of specialization on this food differs. This is the first study in which seed consumption is assessed for all larval instars and adults of ground beetles. Three species of Amara with syntopic occurrence, Amara aenea (DeGeer), Amara familiaris (Duftschmid) and Amara similata (Gyllenhal), were examined. Larvae of all three instars and adults were fed seeds of Stellaria media (L.) Vill., Capsella bursa‐pastoris (L.) Med. and Taraxacum officinale Wick. ex Wigg. in a laboratory no‐choice experiment. In general, larvae, particularly the first instar, showed greater differences in seed consumption than the adults, although the latter showed similar but less marked pattern. Amara aenea consumed all offered seed diets in all life stages. All three larval instars of granivorous A. familiaris almost exclusively fed on seeds of S. media and the adults also ate significantly more of this than other seeds. Amara similata consumed mostly seeds of C. bursa‐pastoris in the first instar and adult stages, whereas the larvae of the later instars seemed to be unspecialized on particular seed diet. Differences in seed‐specific consumption between larval instars in granivorous carabids are reported for the first time. The results provide further support for the parallel evolution of various degrees of granivory in the genus Amara, which may ultimately facilitate species coexistence. The daily seed consumption by the larvae was comparable or (in case of the third instar) even higher than that by the adults. Hence, we suggest that larvae may be the important consumers of seed in the field and should not be forgotten when seed predation is assessed.     

Feeding ecology of granivorous carabid larvae: a stable isotope analysis

Although most carabids are primarily carnivorous, some carabid species are omnivorous, with mainly granivorous feeding habits during the larval and/or adult stages (granivorous carabids). This feeding habit has been established based on laboratory and field experiments; however, our knowledge of the feeding ecology of these beetles in the field is limited owing to the lack of an appropriate methodology. In this study, we tested the utility of stable isotope analysis in investigations of the feeding ecology of granivorous carabids in the field, using two closely related syntopic species, Amara chalcites and Amara congrua. We addressed two issues concerning the feeding ecology of granivorous carabids: food niche differentiation between related syntopic species during the larval stage and the effect on adult body size of supplementing seeds with an animal diet during the larval stage. To investigate larval feeding habits, we analysed newly emerged adults, most somatic tissues of which are considered of larval origin. In the two populations examined, both δ¹⁵N and δ¹³C were significantly higher in A. chalcites than A. congrua, suggesting that the two species differentiate food niches, with A. chalcites larvae being more carnivorous than A. congrua larvae. The two isotope ratios of A. chalcites samples from one locality were positively correlated with body size, suggesting that more carnivorous larvae become larger adults. However, this relationship was not detected in other species/locality groups. Thus, our results were inconclusive on the issue of diet supplementation. Nevertheless, overall, these results are comparable with those of previous laboratory‐rearing experiments and demonstrate the potential utility of stable isotope analysis in field studies on the feeding ecology of granivorous carabids.     

Carabid larvae as predators of weed seeds: granivory in larvae of Amara eurynota (Coleoptera: Carabidae)

Up to date we do not have much information about predation on seeds by larvae of ground beetles. One of the reasons why such knowledge is important is that granivorous larvae contribute to predation of weed seeds. In this study, the food requirements of larvae of autumn breeding carabid species Amara eurynota (Panzer) were investigated in the laboratory and a hypothesis, that they are granivorous was tested. Insect diet (Tenebrio molitor larvae), three seed diets (seeds of Artemisia vulgaris, Tripleurospermum inodorum or Urtica dioica or a mixed diet (T. molitor + A. uulgaris) were used as food. For larvae of A. eurynota, seeds are essential for successful completion of development, because all those fed pure insect diet died before pupation. However, differences in suitability were observed between pure seed diets. Larvae fed seeds of A. vulgaris had the lowest mortality and fastest development of the seed diets. Those fed seeds of T. inodorum had also low mortality, but the development was prolonged in the third instar. In contrast, development of larvae reared on seeds of U. dioica was slowest of the tested diets and could not be completed, as all individuals died before pupation. When insects were included to seed diet of A. vulgaris (mixed diet), the duration of development shortened, but mortality remained the same when compared to seed diet of A. vulgaris. According to the results it was concluded that larvae of A. eurynota are granivorous. A mixed diet and seed diets of A. uulgaris and T. inodorum were suitable and insect diet and seeds of U. dioica were unsuitable diets in this experiment.     

Role of granivory and insectivory in the life cycle of the carabid beetle Amara similata

1. The cereal field carabid beetle Amara similata was selected to elucidate the role of seeds and insects as food sources. Three experiments were performed to rank different weed seeds and insects in terms of food value: (i) fecundity in relation to adult diet, (ii) larval survival in relation to diet and (iii) larval survival in relation to parental diet.
2. Seeds were found to be of high value and insects of low value both for adults and larvae. Adding insects to a seed diet gave no significant improvement. The value of single species of seeds varied within adults and larvae, but also between adults and larvae. This is the first report that Amara larvae are granivorous.
3. The value of different insects varied by species. Cereal aphid species were of the lowest value both for adult and larval beetles. On a diet of mixed insects, the adults were able to produce a low number of eggs but the larvae all died before pupation.
4. There was no simple relation between survival of the larvae and the quality of the parents' diet.     

Food preferences and the value of animal food for the carabid beetle Amara similata (Gyll.) (Col., Carabidae)

Abstract:  Several studies have shown that the mainly granivorous carabid beetles, e.g. Amara spp., include animal food in their diet to a considerable extent. We therefore hypothesized that the performance of these beetles would be enhanced by dietary mixing including both seeds and animal food. In order to test this, we conducted laboratory feeding experiments with adults and larvae of Amara similata . Both adults and larvae were subjected to different diet treatments including: seeds, houseflies, grasshoppers, earthworms, slugs and snails in pure and mixed diets. Larval survival, development time, pupal and teneral weights were used as indicators of food quality for the larvae. For the adult beetles, mass change was used as an indicator of food quality. We found seeds to be high-quality food, while all pure animal diets were of low quality for both adults and larvae. Animal foods added to the seed diet had both positive and negative effects. A mixed diet of all foods enhanced the mass gain of adults compared with the seed diet, but reduced larval performance dramatically. Earthworms and grasshoppers added to seeds increased the pupal and teneral weights, while reduced larval survival. Thus, A. similata is omnivorous with a mainly granivorous feeding habit. It may gain benefits on some fitness parameters and incur costs on others from preying or scavenging on animal food. Therefore, the overall fitness consequences of a mixed seed-animal diet are uncertain.     

Effects of maternal diet on fecundity and larval development in the 'primitive' granivorous carabid Amara (Curtonotus) macronota

Granivory is one of the most specialized food habits in the Carabidae and has been reported for species from the tribes Zabrini and Harpalini. Most studies of carabid granivory have been conducted using specialized granivorous species, and few have examined primitive ones. This study examined effects of maternal diet on fecundity and larval development in Amara ( Curtonotus ) macronota (Solsky) (Coleoptera: Carabidae), a member of the most basal clade of the tribe Zabrini; a previous study indicated that larvae of this species are omnivores with a tendency toward carnivory. Three diet types, Tenebrio molitor L. larvae, mixed seeds [ Bidens frondosa L. (Asteraceae), Setaria spp., and Digitaria ciliaris (Retz.) Koeler (both Poaceae)], and T. molitor larvae + mixed seeds, were used as maternal diets, and larvae were reared on T. molitor larvae + mixed seeds (optimal diet) or T. molitor larvae (suboptimal diet). Fecundity differed significantly among treatments, with individuals fed the T. molitor larvae + mixed seeds diet having the highest fecundities and the mixed seeds diet producing the lowest values. Larval development [survival and duration of development through pre-overwintering stages (first and second instars)] was not significantly affected by either maternal or larval diets, but the addition of seeds to the maternal diet had a weak negative effect on larval survival. These results are in contrast with findings from specialized granivorous carabids, in which both adults and larvae performed best with pure-seed diets. Recent molecular phylogenies indicate that these specialized granivorous carabids belong to derived lineages, while A. macronota is the most basal clade of each tribe. These results are compatible with the hypothesis that carabid granivory has evolved gradually from ancestral carnivory, with omnivorous habits occurring as a transient state.     

Laboratory studies on the larval food habits of the ground beetle Amara (Curtonotus) gigantea (Motschulsky) (Coleoptera: Carabidae: Zabrini)

Ground beetles (Coleoptera: Carabidae) are important components of soil ecosystems in temperate zones, but our knowledge regarding the ecology of many species is limited. This study examined larval food habits in the ground beetle Amara (Curtonotus) gigantea (Motschulsky) using laboratory‐rearing experiments. Because this beetle is a member of the tribe Zabrini, which includes several species that exhibit granivory during larval and/or adult stages, three diet types were tested, with starvation as a control: Tenebrio larvae, mixed seeds (Bidens pilosa (Asteraceae), Setaria spp., Digitaria ciliaris (both Poaceae), and Humulus scandens (Moraceae)), and Tenebrio larvae + mixed seeds. Because of high larval mortality during overwintering under laboratory‐rearing conditions, survival and the duration of development through pre‐overwintering stages (first and second instars) were compared. Larvae fed the diet of Tenebrio larvae + mixed seeds showed the highest survival (89%), followed by the diet of Tenebrio larvae (83%), but the difference between the two treatments was not significant; similarly, developmental durations did not differ between these two diets. Larvae fed the mixed‐seeds diet showed markedly lower survival, and no larvae reached the second instar. Therefore, for A. gigantea larvae, animal food results in high larval performance, whereas seeds provide a low value diet; animal food is essential for larval development, but seeds provide no benefit, even as a dietary supplement. Overall, the present results suggest that A. gigantea larvae are omnivorous but with a mainly carnivorous food habit.     

Laboratory studies on larval food habits of two syntopic,related, granivorous ground beetles <Emphasis Type="Italic">Amara chalcites</Emphasis> and <Emphasis Type="Italic">A. congrua</Emphasis> (Coleoptera: Carabidae): a comparison with stable isotope analysis

This study examined larval food habits of two syntopic, related, carabids Amara chalcites Dejean and A. congrua Morawitz using a laboratory-rearing experiment, and the results were compared with those of a previous stable-isotope analyses of the same species. Larvae were reared on six different diets (seeds of Capsella bursa-pastoris (Linnaeus) Medikus, Stellaria media (Linnaeus) Villars, Taraxacum officinale Weber ex Wiggers; mixed seeds; Tenebrio molitor Linnaeus larvae; and Tenebrio larvae + mixed seeds). For both species, beetles fed seed diets exhibited moderate to high survival rates, with the exception of those fed Stellaria seeds, on which A. congrua showed low survival rates. The pure-animal diet resulted in high survival rates of A. chalcites but low survival rates of A. congrua. These results were consistent with those of the isotope analysis, in which A. chalcites larvae were more carnivorous than were A. congrua larvae, and the larvae of the two species appeared to prefer different types of seeds. In contrast, the effect of diet on adult weight differed between the two methods. In the laboratory experiments, supplementing seeds with animal food during the larval stage positively affected adult weight for both species, whereas in the isotope analysis, the positive effect only occurred in A. chalcites. Possible explanations for the discrepancy between the two methods and suggestions for future research are provided.     

Diet affects male gonad maturation,female fecundity,and larval development in the granivorous ground beetle Anisodactylus punctatipennis

Abstract 1. Granivory is a specialised food habit in the Carabidae and is considered to have evolved from ancestral carnivory. Despite recent investigations, the mechanisms underlying this peculiar feeding shift have not been sufficiently elucidated. In particular, no studies have examined the effects of diet composition on male reproductive traits. 2. This study examined male gonad maturation, female fecundity, and larval development under different diets (insect larvae, mixed seeds, and insect larvae + mixed seeds) for Anisodactylus punctatipennis (Coleoptera: Carabidae). 3. Whereas sperm‐bundle length, the durations of larval stages, and adult weight did not differ among diets, the weights of seminal substance‐producing organs, fecundity, and larval survival were higher with diets containing seeds compared to the pure‐animal diet. These findings provide the first conclusive evidence for granivory in Anisodactylus. 4. This study is the first that demonstrates the effects of diet on male reproductive traits in a granivorous carabid. The results were consistent with similar observations for fecundity and larval performance in this and other studies. Thus, in addition to the female and larval traits, the dietary effects on male reproductive traits may also have played an important role in the evolution of granivory in the Carabidae.     

Effect of diet on the fecundity of three carabid beetles

Abstract In the present study, the importance of diet in terms of fecundity is compared for three species of the carabid genus Amara (Coleoptera: Carabidae), using an insect diet, two types of seed diet (Capsella bursa‐pastoris, Stellaria media) and a mixed diet. It is expected that the species of carabid studied have different food requirements for reproduction. Diet affects reproduction performance and egg production significantly. A mixed diet and both single‐seed diets are suitable for reproduction in Amara aenea (DeGeer) because a higher proportion of the females reproduce and lay significantly more eggs than on a purely insect diet. Females of Amara familiaris (Duftschmid) do not reproduce unless provided with seeds of S. media. Seeds of C. bursa‐pastoris or a mixed diet are equally suitable diets for reproduction of Amara similata (Gyllenhal); a diet of insects or seeds of S. media is unsuitable. The results support the hypothesis that the species under investigation have specific food requirements, suggesting that seed feeding has evolved to different degrees in particular species: A. aenea is omnivorous, whereas A. familiaris and A. similata specialize on the seed of a particular plant species or family. This resource partitioning facilitates co‐occurrence of carabid species.     

The effect of GNA lectin on the α-amylase activity of the beet armyworm,Spodoptera exigua Hb. (Lepidoptera: Noctuidae)

Beet armyworm (Spodoptera exigua Hb.) (Lepidoptera: Noctuidae) is the major pest of sugar beet (Beta vulgaris). Pesticide applications are the main method of the insect control. So, alternative method/s is/are needed to control this insect species. So, in the current study, the effect of Galanthus nivalis agglutinin (GNA) (snowdrop lectin) on beet armyworm α-amylase was studied. Measurement of the amylase activity of the larval midgut fed on artificial diet and sugar beet leaves showed that the enzyme activity was higher when the larvae fed on artificial diet. However, in both cases, the fourth instar larvae had the greatest amylase activity. Thus, fourth instar larvae were offered artificial diet containing 1 and 2% GNA. Both treatments of the lectin significantly reduced the α-amylase activity of the insect. For example, amylase activity of the fourth instar larvae in the control (fed only on artificial diet) was 2.62 Uml^?1 whilst the activity of the enzyme in the two treatments including diet containing 2 and 1% lectin was 1.45 and 1.75 Uml^?1, respectively. The achieved data showed that lectin, in addition to have toxic effect on the larval growth and development, affects the α-amylase activity of the insect gut.     

How do caterpillars cope with xenobiotics? The case of Mythimna unipuncta,a species with low susceptibility to Bt

Mythimna unipuncta is a species with low susceptibility to the Bacillus thuringiensis (Bt) toxin, and this insect occasionally causes devastating damage to maize. In the study region, M. unipuncta‐developed larvae were observed moving from a non‐Bt crop to a nearby Bt crop. Although the first response of many caterpillars to xenobiotics, such as the Bt toxin, is to reduce food intake and prolong development, few studies have focused on the causes and consequences of this response in terms of resistance evolution. To clarify the causes of this response, this work compared changes in the feeding behaviour, cytochrome P450 expression and juvenile hormone titre during the last larval instar of M. unipuncta after Bt ingestion. Four P450 enzymes related to the xenobiotic metabolism of the CYP9 and CYP6 families were identified. Developed larvae fed the Bt diet reduced their food intake and CYP9 expression, experienced prolonged development and presented an altered juvenile hormone balance. The CYP9s were not increased in the larvae that consumed Bt, as previously expected, although their highest expression was observed when larval feeding increased. The high recovery capacity of the larvae contributed to their development when they were fed a non‐Bt diet. The efficiency of responses that act jointly as a defence mechanism against Bt might favour the development of field resistance to the toxin. Therefore, these responses should be further investigated for resistance management programmes.     

Evaluation ofHadena perplexa [Lepidoptera: Phalaenidae] as a biocontrol agent of bladder campionSilene vulgaris [Caryophyllaceae] in Canada: Rearing and host specificity

Rearing techniques and results of preliminary host range tests are reported forHadena perplexa (Denis & Schiffermuller) (Lep.: Phalaenidae) a candidate biocontrol agent against the weed bladder campion,Silene vulgaris (Moench) Garcke, in Manitoba, Canada. In the laboratory, it was necessary to pipette a 15% honey solution in water into the flowers as food for the adult moths. When reared singly to avoid cannibalism, 56% of the 1^st instar larvae developed to pupae. Larvae fed on a natural diet for 10 days can then be reared on either one of 2 artificial diets. Choice oviposition tests and no-choice larval feeding tests were conducted with plant species closely related toS. vulgaris in the generaSilene, Dianthus, Gypsophila, Lychnis, Saponaria. Species in 4 of 5 of these genera were accepted for oviposition, and species in all 5 genera supported the development of 1^st instar larvae to the pupal stage.H. perplexa should not be introduced into Canada.      

Size compensation in moth larvae: attention to larval instars

Environmental perturbations such as starvation and poor diet often prevent animals from attaining their optimal sizes. When the perturbation has a transient character, compensatory responses are expected in terms of faster growth or a prolonged developmental period. In the case of insect larvae, details of such responses are insufficiently known at the proximate level. Attention to responses at the level of particular larval instars should promote an understanding of insect developmental plasticity also in a more general context. To provide an instar‐specific analysis of compensatory growth, larvae of the moth Orgyia antiqua (L.) are reared on inferior diet during one larval instar. Responses in growth parameters are recorded in the course of the manipulated instars, as well as at the level of the entire larval period. The negative relationship between development time and size in response to the inferior food quality, typical of the entire larval periods, is also observed within the manipulated instars taken separately. The manipulated larvae remain smaller than the larvae of the control group (significant in males only), even by the end of the subsequent instar during which all individuals are provided with superior host. In males, close to full size compensation by the time of pupation is achieved only by means of adding an extra larval instar. The inability of larvae to fully compensate during one and even two instars is considered as an indication of the presence of constraints on the within‐instar growth pattern. An alternative, adaptational explanation for the incomplete compensation could be based on the cost of prolonged development period. Given the ecological context of the species' life history, such an explanation appears less likely.     

Effects of food nutritive quality and Bacillus thuringiensis onfeeding behaviour, food utilization and larval growth of spruce budworm Choristoneura fumiferana (Clem.) when exposed as fourth- and sixth-instar larvae

Abstract 1 Feeding behaviours, and lethal and sublethal (growth, development and food utilization) effects of Foray 48B, a commercial formulation of Bacillus thuringiensis (kurstaki), were investigated on fourth‐ and sixth‐instar spruce budworm larvae according to food nutritive quality. Nitrogen and soluble sugar content of artificial diets were modified to obtain three different qualities of food, simulating variations in nutritive quality of host tree. 2 Larval development times were longer for Bt‐treated larvae and pupal weights were reduced for sixth‐instar larvae only. Bt‐induced mortality levels were influenced by food quality. Ingested dose of Bt and feeding inhibition times were strongly affected by the Bt treatment, but food quality affected only fourth‐instar larvae. Except for food digestibility, nutritional indices were negatively affected by the Bt treatment and by the reduction in food quality. 3 Contrary to early treated larvae (fourth instar), larvae treated at the beginning of the sixth instar were not able to compensate for Bt injury and were consequently more affected by the Bt‐treatment both in terms of lethal and sublethal effects. 4 Bt efficacy was not directly related to the ingested dose. 5 Increase in larval vulnerability to Bt was more likely a consequence of a general stress induced by a less suitable food than a direct interaction between Bt and food nitrogen or sugar compounds. 6 The application of Bt on late‐instar larvae could be a successful operational strategy at low population levels when field sprays target the insect instead of foliage protection.     

Occurrence of Amblycerus species in Cordia trichotoma seeds and their influence on germination

Forest species can have their seeds damaged by granivorous insects, especially by those in their larval stage. In this context, this study aims to report the occurrence of Amblycerus species in Cordia trichotoma seeds, to describe their main damage to seeds and effects on germination, as well as their associated hymenopteran parasitoids. Therefore, seven trees were selected in the municipality of Taquaruçu do Sul, RS, Brazil. Fruits were collected weekly from the medium third of the tree crown, from the beginning of their formation until total dehiscence. To examine the damage caused by granivorous insects within the fruits, 15 fruits from each tree were sectioned with a scalpel. Furthermore, 10 fruits from each tree were stored individually in clear plates to verify the occurrence and identification of granivorous insect species. Evidence of the damage caused to seeds was verified through the germination test by comparing preserved and damaged seeds, with four repetitions of 25 seeds each. The species Amblycerus longesuturalis and Amblycerus profaupar (Chrysomelidae: Bruchinae) were found associated with fruits of C. trichotoma. Female insects predominantly laid eggs on the superior part between the marcescent calyx and the fruit, and larvae perforated the fruit tegument to start consuming seed embryos and reserves. Bruquine larvae are parasitized by Hymenoptera of Bracon, Mirax, Omeganastatus and Triapsis genera. In conclusion, the germination of C. trichotoma seeds is significantly affected by emergence orifices caused by granivorous species.     

How do food quality and larval crowding affect performance of the autumnal moth,Epirrita autumnata?

Outbreak densities of autumnal moth, Epirrita autumnata (Borkhausen) (Lepidoptera: Geometridae), lead to high larval crowding. Phenotypic responses of E. autumnata to larval crowding and to food quality were studied by measuring growth and consumption as well as pupal weight and fecundity. Crowding may trigger increased consumption and faster development to avoid impending food shortage on good quality food. This is suggested by the result that on a good‐quality diet, the growth of crowded larvae was better than that of solitary larvae, though they did not consume more food than solitary larvae. Crowded larvae also completed the last instar earlier than solitary larvae. The fecundity of crowded autumnal moths was not lower than the fecundity of solitarily grown autumnal moths. This may provide conditions for extra rapid population build‐up of E. autumnata. During the population increase phase the crowding effect may facilitate larval performance; however, at peak density the crowding starts to have negative effects on the performance of larvae. On a poor‐quality diet, the performance of crowded and solitary larvae did not differ. The growth of larvae was better on a good‐quality diet than on a poor‐quality diet, due to higher efficiency in food utilization. Larvae feeding on low‐quality diet did not prolong their development time, but pupated at smaller size; this resulted in lower fecundity. A decrease in food quality can be seen as a cue of oncoming food shortage and resource depletion; it may be advantageous to pupate at a smaller size and ensure survival till reproduction, rather than risk prolonging development to achieve larger size and higher fecundity.     

Changes in spatial distribution pattern during the larval stage of the fall webworm,Hyphantria cunea drury (Lepidoptera: Arctiidae)

Summary The changes in spatial distribution pattern during larval stage of the fall webworm,Hyphantria cunea were quantitatively investigated in the field experimental populations. The female adult deposits eggs as a cluster and the hatchlings make a compact colonial-web. In this period, the all-or-none type mortality which is characteristic in gregarious insect species was occasionary recognized before spinning a compact colonial-web. Once making a compact colonial-web, the larvae feed the leaves in the colonial-web up to about 5th instar. In this period, the movement of larvae occurred due to the local food shortage in a colonial-web and the expansion of colonial-web. As the larvae developed, the colonial-web was separated into several small groups. These larvae began to disperse about 5th instar. In this period, the local food shortage seems to be an important trigger for the larval dispersal. The mean concentration of larvae on leaves abruptly decreased, and finally the larvae became solitary at the 6th or 7th instars. The dispersal process in later larval stage is not necessarily due to the complete food shortage. The dispersal prior to the occurrence of food shortage may be a safety mechanism to protect the larvae from the food shortage.     

Diversity of birch sawfly responses to seasonally atypical diets

Most insect herbivores are specialised on a particular plant taxon. To have a better understanding of host shift functions and consequences for insect herbivores, it is essential to gather more information on the effects of variation in host quality on specialists across species and environments. We examined the effects of seasonally atypical food on mortality, developmental time, and final body mass of six sawfly species (Hymenoptera: Symphyta) feeding on the foliage of mountain birch (Betula pubescens ssp. czerepanovii), whose pooled larval feeding periods form a gradient and cover the growing season. Insect phenology was manipulated so that the larvae of early-season species would feed on atypically mature leaves and mid- or late-season species would feed on atypically young leaves of their major host plant. Mortality increased dramatically for all species when the larval feeding schedule was advanced or delayed. This indicates a high degree of specialisation not only on a particular host but also to its phenological phases. The main cause of mortality on novel food was a rejection of the diet by the young larvae and their subsequent starvation. An interesting observation was that late-season species showed this response on nutritious young foliage. The effects of seasonally atypical diets on larval development and growth were species-specific and milder than the effects on mortality. Interestingly, for those individuals that accepted it, atypical food seemed to be most beneficial for species appearing at both ends of the seasonal gradient, which might be related to a wider exposure to variable food quality in natural conditions compared with other species. The diversity of responses to atypical food among closely related herbivore species with overlapping feeding periods on the same host plant is the most crucial finding of this study.     

Larval secretions and food odors affect orientation in femalePlodia interpunctella

Substrates contaminated by wandering fifth instar larvae ofPlodia interpunctella (Hübner) (Lepidoptera: Pyralidae) elicit oviposition by conspecific female moths, and larval rearing diet enhances oviposition and also induces upwind flight. Two-choice oviposition assays determined that four-day-old gravid femaleP. interpunctella preferred to lay eggs on dishes containing cornmeal-based rearing diet compared to empty dishes. Pieces of cheesecloth contaminated by fifth instar larvae elicited more oviposition than untreated cheesecloth or dishes with food. The combination of larval contamination and food was preferred over food only or larval contamination only in both two- and four-choice experiments. The factor(s) in larval contamination responsible for eliciting oviposition in female moths was extracted in hexane, confirming that organic semiochemicals are responsible for the effect. The oviposition-eliciting activity of larval contamination was retained on cheesecloth for up to 30 days following treatment with larvae, suggesting the active component(s) is stable and of low relative volatility. In two-choice windtunnel bioassays female moths initiated flight only when rearing food was present in one of the treatments, and they displayed the highest landing responses to a combination of larval contamination and food. Earlier work onP. interpunctella and related pyralid species found that larval contamination due to secretions from the mandibular glands acted as both a spacing pheromone for wandering larvae and as a kairomone for host-seeking parasitoid wasps. The present study suggests that the same or a similar secretion acts as an oviposition-eliciting pheromone for conspecific females.