Arbuscular mycorrhizal fungi influence life history traits of a lepidopteran herbivore

Results from pot and microcosm studies in the greenhouse have shown that plant growth and foliar chemistry is altered by the presence and species composition of arbuscular mycorrhizal fungi (AMF). The growth and survival of herbivores which feed on plants could, as a consequence, also be affected by these mutualistic soil fungi. Consequently, interactions between AMF, plants and herbivores could occur. To test this, larvae of the common blue butterfly, Polyommatus icarus (Lycaenidae), were fed with sprigs of Lotus corniculatus (Fabaceae) plants which were inoculated with one of two different AMF species, with a mixture of these AMF species or with sprigs of plants which were not inoculated with AMF. Survival and larval weight of third instar larvae fed with plants colonised by AMF were greater than those of larvae fed with non-mycorrhizal plants. Survival of larvae feeding on non-mycorrhizal plants was 1.6 times lower than that of larvae feeding on plants inoculated with a mixture of AMF species and 3.8 times lower than that of larvae feeding on plants inoculated with single AMF species. Furthermore, larvae fed with non-mycorrhizal plants attained only about half the weight of larvae fed with mycorrhizal plants after 11 days of growth. These differences in larval performance might be explained by differences in leaf chemistry, since mycorrhizal plants had a 3 times higher leaf P concentration and a higher C/N-ratio. Our results, thus, show that the presence of belowground mutualistic soil fungi influences the performance of aboveground herbivores by altering their food quality. Larval consumption, larval food use and adult lipid concentrations of the common blue butterfly differed between larvae which were fed with plants inoculated with different AMF species. This suggests that the performance of herbivores is not only influenced by the presence of AMF but also depends on the identity of the AMF species colonising the host plants. Moreover, a significant interaction term between AMF species and maternal identity of the larvae occurred for adult dry weight, indicating that the performance of offspring from different females was differently influenced by AMF species composition. To our knowledge, these results show for the first time that the species composition of AMF communities can influence life-history traits of butterfly larvae and possibly herbivores in general.     

Adult beetles compensate for poor larval food conditions

Life history traits of herbivores are highly influenced by the quality of their hosts, i.e., the composition of primary and secondary plant metabolites. In holometabolous insects, larvae and adults may face different host plants, which differ in quality. It has been hypothesised that adult fitness is either highest when larval and adult environmental conditions match (environmental matching) or it may be mainly determined by optimal larval conditions (silver spoon effect). Alternatively, the adult stage may be most decisive for the actual fitness, independent of larval food exposure, due to adult compensation ability. To determine the influence of constant versus changing larval and adult host plant experiences on growth performance, fitness and feeding preferences, we carried out a match–mismatch experiment using the mustard leaf beetle, Phaedon cochleariae. Larvae and adults were either constantly reared on watercress (natural host) or cabbage (crop plant) or were switched after metamorphosis to the other host. Growth, reproductive traits and feeding preferences were determined repeatedly over lifetime and host plant quality traits analysed. Differences in the host quality led to differences in the development time and female reproduction. Egg numbers were significantly influenced by the host plant species experienced by the adults. Thus, adults were able to compensate for poor larval conditions. Likewise, the current host experience was most decisive for feeding preferences; in adult beetles a feeding preference was shaped regardless of the larval host plant. Larvae or adults reared on the more nutritious host, cabbage, showed a higher preference for this host. Hence, beetles most likely develop a preference when gaining a direct positive feedback in terms of an improved performance, whereby the current experience matters the most. Highly nutritious crop plants may be, in consequence, all the more exploited by potential pests that may show a high plasticity in reproduction and feeding preferences.     

The ‘generalism’ debate: misinterpreting the term in the empirical literature focusing on dietary breadth in insects

We here attempt to show, using three broad insect groups – Lepidoptera (mainly leaf‐chewing larval herbivores and nectar‐sucking adults), parasitic Hymenoptera (mainly endoparasitoids, especially of other insects) and aphids (sap‐sucking plant parasites) – how the terms ‘generalist’, usually equated with levels of phagy (oligo‐ and polyphagy), and ‘specialist’ (monophagy), still in widespread parlance, have often been misrepresented. Probably, the reality of generalism, be that herbivorous, predatory and parasitic, can only be demonstrated by detailed empirical field observations plus the use of high‐resolution molecular (DNA) markers, including sequencing, and thereby determining whether the organism in question is really a homogeneous species population over a wide geographical range, or rather comprises a series of morphologically similar/identical cryptic, host‐adapted specialist populations. In the case of insects, the largest group of terrestrial animals on the planet, even if it can be shown that certain species are indeed polyphagous and feed on a variety of plant hosts (herbivores) or prey species (predators and parasitoids), nevertheless, the range of these food items may be highly selective and restricted, depending on morphological–genetical (biochemical/chemical)–behavioural constraints. In the end, while some animals appear to be generalist, this situation may well be illusory. Our present recognition of the term is at best inappropriate, and at worse, inaccurate, as we demonstrate in the examples given, mostly insects. In the meantime, we suggest that the terms used should be re‐defined as four broad classes of specialism–generalism, although the apparent ‘generalism’ is itself conditional on proof following further empirical analyses.     

Studies on host preference and its biological effects on the red palm weevil,Rhynchophorus ferrugineus Olivier in Egypt The fruit trees

A comparative bioassay (choice or non-choice) was developed to evaluate the efficacy of different species of fruit trees as alternative host plants on the morphological and biological aspects of the red palm weevil, Rhynchophorus ferrugineus and also the latent effect of feeding the larvae on pupal and adult stages. Highly significant differences exist between averages of larval body weight of R. ferrugineus larvae fed on fruit trees. The lowest average of larval weight occurred in the larvae fed on the pieces of sour orange, mandarin followed by olive and lime trees. A small increase in body weight appeared in larvae fed on mango, fig and guava in comparison with those fed on date palm trees (control). The larval mortality reached the highest percentages (100%) by obligatory feeding of R. ferrugineus insects on olive, lime, sour orange and mandarin followed by mango (97%) and guava (96%) compared with control (0%), respectively. The latent effect of obligatory feeding on the larvae of R. ferrugineus was highly decreased during pupation (4%, 4% and 8%) when the insect fed on mango, guava and fig, compared with 100% pupation in the case of larvae fed on ornamental palm or date palm, respectively. Most pupae resulting from the treated larvae by R. ferrugineus failed in adult emergence especially those insects fed on olive, lime, sour orange and mandarin trees compared to untreated insects (95.6–96.2%), respectively. Data also proved that if an insect is forced to feed on a non-preferred host plant, this results in death of the insect often during the larval stage, a decrease of pupation or failure of adult emergence. These results led to the conclusion that mango, fig, and guava trees are believed to be vulnerable hosts for red weevil attack. The lowest percentage of food consumption appeared in those larvae fed on mandarin, sour orange, lime and olive trees compared with those fed on palm. The present results emphasise the need to test the efficiency of the compounds extracted from these host plants if it is considered as an insecticide or an alternative host protectant for management of the red palm weevil R. ferrugineus.     

Parasitoids follow herbivorous insects to a novel host plant,generalist predators less so

The ‘enemy‐free space’ hypothesis predicts that herbivorous insects can escape their natural enemies by switching to a novel host plant, with consequences for the evolution of host plant specialisation. However, if natural enemies follow herbivores to their novel host plants, enemy‐free space may only be temporary. We tested this by studying the colonisation of the introduced tree Eucalyptus grandis (Hill) Maiden (Myrtaceae) by insects in Brazil, where various species of herbivores have added eucalyptus to their host plant range, which consists of native myrtaceous species such as guava. Some herbivores, for example, Thyrinteina leucoceraea Ringe (Lepidoptera: Geometridae), cause outbreaks in eucalyptus plantations but not on guava, possibly because eucalyptus offers enemy‐free space. We sampled herbivores (mainly Lepidoptera species) and natural enemies on eucalyptus and guava and assessed parasitism of Lepidoptera larvae on both host plant species during ca. 2 years. Overall, predators were encountered more frequently on guava than on eucalyptus. In contrast, parasitoids were encountered equally and parasitism rates of Lepidoptera larvae were similar on both host plants. This indicates that herbivores may escape some enemies by moving to a novel host plant. However, this escape may be temporary and may vary with time. We argue that studying temporal and spatial patterns of enemy‐free space and the response of natural enemies to host use changes of their herbivorous prey is essential for understanding the role of natural enemies in the evolution of host plant use by herbivorous arthropods.     

Greater impact of host plant species on oviposition of a willow leaf beetle <Emphasis Type="Italic">Plagiodera versicolora</Emphasis> Laicharting (Coleoptera: Chrysomelidae) during the adult stage than in developing larval stage

In some herbivorous insects, such as Coleoptera and aphids, not only the host species of larvae, but also those of adults should be considered as key determinants of potential fecundity because oviposition is affected by the quality of host species during both larval and adult stages. This study examined the relatively greater impact on host species of the larval or adult stage on oviposition of the willow leaf beetle Plagiodera versicolora Laicharting (Coleoptera: Chrysomelidae). We conducted an experiment using a 2 × 2 experimental design, in which either of two different host plant species was fed in larval and adult stages. Females fed on a locally unavailable host Salix eriocarpa in the adult stage did not lay any eggs, but those fed on the locally available host S. babylonica laid 67–75 eggs on average, irrespective of larval host species. Such reproductively inactive females fed S. eriocarpa as an adult host recovered reproductive activity within 3 weeks after changing the host species to S. babylonica. This result indicated that the host species fed in the adult stage had a greater impact on oviposition than in the larval stage.     

Linkages between nectaring and oviposition preferences of Manduca sexta on two co‐blooming Datura species in the Sonoran Desert

1. The oviposition choices of phytophagous insects determine the environment that their offspring will experience, affecting both larval performance and host plant fitness. These choices, however, may be influenced by other activities, such as nectar foraging. 2. In the Sonoran Desert, Manduca sexta (Lepidoptera: Sphingidae) oviposits primarily on the perennial herb Datura wrightii. It has been reported to oviposit on the smaller‐flowered, co‐blooming, sympatric annual Datura discolor as well. Datura is also M. sexta's most important source of nectar in this region. Nectaring and oviposition decisions thus determine M. sexta's reproductive success, as well as the relative benefits (pollination) and costs (herbivory) that each Datura species derives from this interaction. 3. The nectaring and oviposition choices of adult M. sexta on these congeners were studied to investigate how nectar foraging influences oviposition. Larval performance on the two hosts was then assessed. 4. Nectaring and oviposition were behaviourally linked, with M. sexta preferring the perennial D. wrightii as both a nectar source and larval host when given a choice between the two species. This preference disappeared, however, when only D. discolor bore flowers. 5. In the laboratory, larvae developed at equal rates when fed D. wrightii or D. discolor leaves, but survival was higher on D. wrightii when larvae fed on intact plants. 6. These findings suggest that while female preferences match larval performance in most cases, the link between nectaring and oviposition may at times bias moths to lay eggs on inferior larval hosts.     

Shifts in developmental diet breadth of Lymantria xylina (Lepidoptera: Lymantriidae)

The moth Lymantria xylina Swinhoe (1903) (Lepidoptera: Lymantriidae) is a major defoliator of hardwood and fruit trees in Taiwan. Although the plants identified as host plants of L. xylina usually refer to plants used as food or as shelter, most of the host plant identifications have not considered the role of these hosts on larval development. This study investigated various instars feeding on different plants to assess the developmental diet breath of L. xylina. Forty-seven plant species, belonging to 25 families were used in feeding trials. Various bioassays, including first instar survival and long-term feeding trials, indicated the most suitable host plants for the different developmental stages. Results of the first instar survival trial indicated that first instars could survive only on 13 of the tested plant species. In addition, first instars could only successfully grow to pupa on seven of these 13 test plants species. To assess the developmental diet breath shifts of this moth, 38 plant species (excluding those nine plant species that the first instars did not feed upon) were fed to third and fifth instars in long-term feeding trials. Survival to pupa was noted on 12 and 13 test plant species for the third and fifth instars, respectively. In short, we found that the larvae performed differently when fed on various host plants and that the host plant range increased with the larval stage. Therefore, it is necessary to adjust the host plant range of this moth and to consider host plant breadth together with the developmental stages of caterpillars.     

Influence of Dietary Experience on the Induction of Preference of Adult Moths and Larvae for a New Olfactory Cue

In Lepidoptera, host plant selection is first conditioned by oviposition site preference of adult females followed by feeding site preference of larvae. Dietary experience to plant volatile cues can induce larval and adult host plant preference. We investigated how the parent’s and self-experience induce host preference in adult females and larvae of three lepidopteran stem borer species with different host plant ranges, namely the polyphagous Sesamia nonagrioides, the oligophagous Busseola fusca and the monophagous Busseola nairobica, and whether this induction can be linked to a neurophysiological phenotypic plasticity. The three species were conditioned to artificial diet enriched with vanillin from the neonate larvae to the adult stage during two generations. Thereafter, two-choice tests on both larvae and adults using a Y-tube olfactometer and electrophysiological (electroantennography [EAG] recordings) experiments on adults were carried out. In the polyphagous species, the induction of preference for a new olfactory cue (vanillin) by females and 3^rd instar larvae was determined by parents’ and self-experiences, without any modification of the sensitivity of the females antennae. No preference induction was found in the oligophagous and monophagous species. Our results suggest that lepidopteran stem borers may acquire preferences for new olfactory cues from the larval to the adult stage as described by Hopkins’ host selection principle (HHSP), neo-Hopkins’ principle, and the concept of ‘chemical legacy.’     

Pyrrolizidine alkaloids: different acquisition and use patterns in Apocynaceae and Solanaceae feeding ithomiine butterflies (Lepidoptera: Nymphalidae)

Pyrrolizidine alkaloids (PAs) often serve as chemical mediators of plant-herbivore-predator interactions. Butterflies (Danainae and Ithomiinae) and moths (Arctiidae) usually acquire PAs from plant sources (larval host plants, flowers or withered leaves visited by adults—pharmacophagy) and thereby become chemically protected against predators; they also use PAs as pheromone precursors. Study by GC-MS of PAs in three species of Ithomiinae butterflies, their larval host plants and adult alkaloid sources showed three different acquisition patterns: (1) larvae of the primitive Tithorea harmonia sequester PAs from their food plant Prestonia acutifolia (Apocynaceae: Echitoideae), and adults may also acquire these alkaloids from plant sources; (2) larvae of the more derived Aeria olena feed on Prestonia coalita , in whose leaves no PAs were detected, but freshly emerged adults sometimes contain PAs and males intensively seek and sequester these alkaloids in plant sources; and (3) larvae of the still more advanced Mechanitis polymnia feed on several PA-free Solanum species, and adult males sequester the alkaloids from various plant sources. Males and females of all three species contain mostly two PAs, the diastereoisomeric retronecine monoesters lycopsamine and intermedine, stored in the N-oxide form. Larval host plants and adult plant sources showed a large array of PA structures, the most abundant and frequent being lycopsamine and its diastereoisomers intermedine, echinatine, rinderine and indicine, and the deoxy-analogues supinine and amabiline. Bioassays with wild caught and freshly emerged adults suggest that protection against predation by the orb weaving spider Nephila clavipes may be dependent on PA concentration and maybe some spider idiosyncrasies, but freshly emerged Aeria olena without PAs are also liberated by Nephila , suggesting other protective compounds. The role of this spider as a selective pressure for PA acquisition by ithomiines is not clear.     

Oviposition,larval preference,and larval performance in two polyphagous species: does the larva know best?

It is expected that females preferentially oviposit on plant hosts that allow for optimal larval performance. However, this expectation contradicts empirical evidence where adults do not always choose the best host for their descendants. Recent evidence suggests that females’ host selection depends on the number of potential hosts. Females from oligophagous species seem to be able to choose an appropriate host in terms of larval performance, whereas in polyphagous species, adult oviposition preference is not related with larval performance. This suggests that larvae in polyphagous species could be taking a more active role in host selection than their mothers. Here, we evaluated the oviposition preference and the larval preference and performance of two polyphagous species of economic importance, Copitarsia decolora (Guenée) (Lepidoptera: Noctuidae: Cuculliinae) and Peridroma saucia (Hübner) (Lepidoptera: Noctuidae: Noctuinae), on eight species of cultivated plants. In laboratory and greenhouse choice assays, we tested adult preference for oviposition and larval preference at 1 and 24 h. Larval performance was measured in terms of survival to adulthood, length of larval period, and pupal weight. We found that both adult females and larvae actively choose their hosts and that the larval preference toward the hosts is related to the females’ preference in both herbivore species. However, the females and larvae did not preferentially select the host with the best larval performance, indicating that larval performance is not related to female or larval preference and that other selective pressures are influencing the choice of the host plant in these two species.     

Life history and host range of Sauris nr. purpurotincta,an unsuitable biological control agent for Chinese tallowtree

Foreign surveys in China discovered a defoliating insect species feeding on the leaves of Chinese tallowtree (Triadica sebifera), an invasive weed of the southeastern U.S.A. The life history of this species, Sauris nr. purpurotincta (Lepidoptera: Geometridae), was examined and larval no-choice and adult multiple-choice host range tests were conducted in quarantine to evaluate their suitability for biological control of Chinese tallowtree. The results indicated that the larvae have five instars and require approximately 22 days to complete development to the adult stage. Host range tests indicated that the larvae could not feed and complete development on most species tested. However, 40% of the larvae survived when fed leaves of Hippomane mancinella, a state-listed endangered species in Florida, and all larvae survived when fed Morella cerifera, a common native species of the southeastern U.S.A. Multiple-choice oviposition tests indicated eggs were laid on leaves of both a south Florida native plant Gymnanthes lucida and Chinese tallowtree. Considering this broad host range, this species will not be considered further for biological control of Chinese tallowtree in the U.S.A.     

Effects of class-specific,synthetic, and natural proteinase inhibitors on life-history traits of the cotton bollworm Helicoverpa armigera

Herbivorous insects have more difficulty obtaining proteins from their food than do predators and parasites. The scarcity of proteins in their diet requires herbivores to feed voraciously, thus heavily damaging their host plants. Plants respond to herbivory by producing defense compounds, which reduce insect growth, retard development, and increase mortality. Herbivores use both pre- and postdigestive response mechanisms to detect and avoid plant defense compounds. Proteinase inhibitors (PIs) are one example of plant compounds produced as a direct defense against herbivory. Many insects can adapt to PIs when these are incorporated into artificial diets. However, little is known about the effect of PIs on diet choice and feeding behavior. We monitored the diet choice, life-history traits, and gut proteinase activity of Helicoverpa armigera larvae using diets supplemented with synthetic and natural PIs. In choice experiments, both neonates and fourth-instar larvae preferred the control diet over PI-supplemented diets, to varying degrees. Larvae that fed on PI-supplemented diets weighed less than those that fed on the control diet and produced smaller pupae. Trypsin-specific PIs had a stronger effect on mean larval weight than did other PIs. A reduction of trypsin activity but not of chymotrypsin activity was observed in larvae fed on PI-supplemented diets. Therefore, behavioral avoidance of feeding on plant parts high in PIs could be an adaptation to minimize the impact of this plant's defensive strategy.     

Positive effects of cyanogenic glycosides in food plants on larval development of the common blue butterfly

Cyanogenesis is a widespread chemical defence mechanism in plants against herbivory. However, some specialised herbivores overcome this protection by different behavioural or metabolic mechanisms. In the present study, we investigated the effect of presence or absence of cyanogenic glycosides in birdsfoot trefoil (Lotus corniculatus, Fabaceae) on oviposition behaviour, larval preference, larval development, adult weight and nectar preference of the common blue butterfly (Polyommatus icarus, Lycaenidae). For oviposition behaviour there was a female-specific reaction to cyanogenic glycoside content; i.e. some females preferred to oviposit on cyanogenic over acyanogenic plants, while other females behaved in the opposite way. Freshly hatched larvae did not discriminate between the two plant morphs. Since the two plant morphs differed not only in their content of cyanogenic glycoside, but also in N and water content, we expected these differences to affect larval growth. Contrary to our expectations, larvae feeding on cyanogenic plants showed a faster development and stronger weight gain than larvae feeding on acyanogenic plants. Furthermore, female genotype affected development time, larval and pupal weight of the common blue butterfly. However, most effects detected in the larval phase disappeared for adult weight, indicating compensatory feeding of larvae. Adult butterflies reared on the two cyanogenic glycoside plant morphs did not differ in their nectar preference. But a gender-specific effect was found, where females preferred amino acid-rich nectar while males did not discriminate between the two nectar mimics. The presented results indicate that larvae of the common blue butterfly can metabolise the surplus of N in cyanogenic plants for growth. Additionally, the female-specific behaviour to oviposit preferably on cyanogenic or acyanogenic plant morphs and the female-genotype-specific responses in life history traits indicate the genetic flexibility of this butterfly species and its potential for local adaptation.     

The parasitoid of a fruit moth caterpillar utilizes fruit components as nutrient source to increase its longevity and fertility

Longevity and fecundity of female wasps are two decisive factors for the effectiveness of parasitoid species as biological control agents. Accessibility and suitability of nutrient sources determine parasitoid survival and reproduction. Host, nectar and honeydew feeding are frequent adult parasitoid behaviors to cover nutritional needs. Here we postulate that especially parasitoid species of endophytic herbivores might use plant tissue as a nutrient source that becomes accessible upon herbivory. We investigated the influence of plant consumption and host feeding on longevity and fecundity of Hyssopus pallidus, a gregarious ecto-parasitoid of caterpillars of the codling moth that feed inside apple fruits. Longevity of unmated and mated ovipositing female parasitoids was highest in treatments with fruit pulp. While longevity in this treatment was not significantly different from that with honey, it was significantly higher than in treatments without food, with water or with a host alone.Reproduction was significantly increased by these sugar-rich nutrient sources compared to the control with a host alone. In contrast, host feeding did not yield any significant contribution to longevity and fecundity in a series of bioassays with different host–parasitoid ratios and with differently aged and sized hosts, compared to controls without food.We conclude that in this synovigenic species host feeding does not contribute to longevity and fecundity, but females can increase survival and reproduction in the field relying solely on the plant tissue damaged by their host caterpillar.     

Flower nectar of an autogamous perennialRorippa indica as an indirect defense mechanism against herbivorous insects

This report shows that one of the most important roles of the flower nectar of an autogamous perennialRorippa indica (L.) Hieron is as an attractant for employing some ant species as a defense against herbivorous insects. The plant has flowers from spring to early winter. Its flower nectar is frequently stolen by some ant species (hereafter cited as ants) which also feed on small herbivorous insects on the plant. Internations among the tritrophic levels (R. indica, herbivores, ants) were experimentally examined and the followings became clear. (1) Ants were attracted toR. indica in search of its flower nectar. (2) The gradual secretion of flower nectar seemed to detain ants on the plant. (3)Pieris butterfly lavae were the major herbivores onR. indica and were potentially harmful to the plant. (4) The presence of ants reduced the survival rate ofP. rapae larvae onR. indica. (5) The presence of ants reduced the feeding damage toR. indica. (6) The disadvantage of nectar use by ants seemed to be minimal for the plant since the ants did not disturb the other flower visitors. These facts suggest a mutualistic relationship betweenR. indica and ants. That is, the flower nectar serves as an indirect defense against herbivorous insects.     

The effect of β-aminobutyric acid on the growth of herbivorous insects feeding on Brassicaceae

dl ‐β‐Aminobutyric acid (BABA) is a nonprotein amino acid that can enhance defences in a variety of plants against a wide range of pathogens. BABA can also reduce infestation by phytopathogenic nematodes and has recently been shown to suppress the growth of aphids feeding on legumes. This investigation examined the effect of applying BABA as a root drench to a range of Brassicaceae, including Arabidopsis thaliana, on the performance of two species of aphid (Myzus persicae and Brevicoryne brassicae) and the larvae of two species of Lepidoptera (Trichoplusia ni and Plutella xylostella). Application of BABA reduced the performance of all four insect species, and inhibition of insects occurred on all the plants tested. The results illustrate that BABA‐induced resistance (BABA‐IR) can affect generalist and specialist insect herbivores and inhibit insects feeding with mandibulate as well as sap‐feeding mouthparts. The BABA‐induced suppression of B. brassicae and P. xylostella feeding on A. thaliana provides a means to further examine the mechanisms of BABA‐IR to insects using this model plant.     

Mouthparts of flower-visiting insects

This review deals with the morphology and function of adult insect mouthparts modified to feed on nectar, pollen or petals. Specialization to nectar-feeding is evident in formation of proboscides of various lengths and designs. Proboscides of many Hymenoptera and Diptera function according to adhesion mechanisms that load nectar onto extensible apical mouthpart regions before fluid is conveyed along the food canal to the mouth by capillarity and suction. Predominantly suctorial proboscides evolved once in Lepidoptera, probably twice in Coleoptera, variously in some Hymenoptera and several times with similar design in Diptera. Many of them are particularly long and have sealed food tubes, specialized apical regions, new proboscis resting positions and modified feeding movements. Mouthparts of obligate pollen-feeding insects can be characterized by modified mandibles, specialized bristles for pollen manipulation and elaborate feeding movements. Often saliva is crucial for pollen retention and ingestion. In Coleoptera, intact pollen is gathered by sweeping movements of mouthparts; in Diptera, it is suspended in saliva prior to suction. Pollen is crushed by asymmetrical mandibles in aglossatan Lepidoptera and one group of basal Hymenoptera. Pollen-piercing mouthparts occur in Thysanoptera and one group of Diptera. Some butterflies and few Diptera extract nutrients from pollen by mixing it externally with saliva on their mouthparts. No mouthpart specializations to petal-feeding are reported in flower-visiting insects.     

Family matters: effect of host plant variation in chemical and mechanical defenses on a sequestering specialist herbivore

Insect herbivores contend with various plant traits that are presumed to function as feeding deterrents. Paradoxically, some specialist insect herbivores might benefit from some of these plant traits, for example by sequestering plant chemical defenses that herbivores then use as their own defense against natural enemies. Larvae of the butterfly species Battus philenor (L.) (Papilionidae) sequester toxic alkaloids (aristolochic acids) from their Aristolochia host plants, rendering larvae and adults unpalatable to a broad range of predators. We studied the importance of two putative defensive traits in Aristolochia erecta: leaf toughness and aristolochic acid content, and we examined the effect of intra- and interplant chemical variation on the chemical phenotype of B. philenor larvae. It has been proposed that genetic variation for sequestration ability is ??invisible to natural selection?? because intra- and interindividual variation in host-plant chemistry will largely eliminate a role for herbivore genetic variation in determining an herbivore??s chemical phenotype. We found substantial intra- and interplant variation in leaf toughness and in the aristolochic acid chemistry in A. erecta. Based on field observations and laboratory experiments, we showed that first-instar larvae preferentially fed on less tough, younger leaves and avoided tougher, older leaves, and we found no evidence that aristolochic acid content influenced first-instar larval foraging. We found that the majority of variation in the amount of aristolochic acid sequestered by larvae was explained by larval family, not by host-plant aristolochic acid content. Heritable variation for sequestration is the predominant determinant of larval, and likely adult, chemical phenotype. This study shows that for these highly specialized herbivores that sequester chemical defenses, traits that offer mechanical resistance, such as leaf toughness, might be more important determinants of early-instar larval foraging behavior and development compared to plant chemical defenses.     

A trade-off between female lifespan and larval diet breadth at the interspecific level in Lepidoptera

A prediction arising from several evolutionary diet breadth models is that, in insect herbivores whose adults practise adaptive host plant selection based on larval performance, female adult lifespan should be negatively correlated with larval diet breadth. In one category of models, female adult lifespan drives evolutionary changes in larval diet breadth; in the other category, larval diet breadth drives evolutionary changes in female adult lifespan. Applying the method of independent contrasts to a biologically and phylogenetically diverse array of Lepidoptera, we ask whether larval diet breadth—as measured by the number of larval food plant species reported in the literature—is negatively correlated with female adult lifespan at the interspecific level. We show that these two life history variables are indeed inversely related. Next, we relax the assumption, common to all of the models, that the female adult is the life stage responsible for the distribution of progeny among different host plants. By introducing into our data set three species whose females are incapable of flight (due to either aptery or brachyptery), and whose larvae are the dispersive stage, the negative correlation between female adult lifespan and larval diet breadth is lost, when using the independent contrasts method. We interpret this effect as supporting the models’ common prediction. Ours is the first reported evidence of a lifespan/diet breadth trade-off at the interspecific level among insects, and it confirms the findings of a previous study in which the degree of habitat specialisation among arthropods was inversely related to proxy measures of the degree of search time constraint. In one of our “diet breadth drives changes in lifespan” models, the females’ type of egg maturation strategy (as measured by the ovigeny index) is predicted to be positively correlated with larval diet breadth, and it mediates a female adult lifespan/larval diet breadth trade-off; however, we found no convincing support for such a role.