The impact of host plant species on the larval development of the large pine weevil Hylobius abietis L.

• 1 The developmental performance of the large pine weevil Hylobius abietis was studied in the laboratory on four species of conifer, Corsican pine Pinus nigra var. maritima, Sitka spruce Picea sitchensis, Douglas‐fir Pseudotsuga menziesii and Japanese larch Larix kaempferi.
• 2 All species supported development, but, there was considerable variation in larval mortality, development time and weight of adults on emergence between host species.
• 3 Levels of mortality were highest in Japanese larch (77%) and lowest in Corsican pine (8.2%), and the heaviest adults emerged from Corsican pine (130 mg) and the smallest from Douglas‐fir (74 mg).
• 4 A constitutive plant defence chemical, lignin, found to vary within a northern provenance of Sitka spruce, also strongly affected larval development.
• 5 The significance of these findings is discussed in relation to the management of H. abietis.

     

Maturation feeding and reproductive development in adult pine weevil, Hylobius abietis (Coleoptera: Curculionidae)

Maturation feeding on conifer bark by newly emerged Hylobius abietis(Linnaeus) is essential for reproductive development. When feeding occurs on young conifer transplants, this weevil causes significant economic damage. Between emergence and oviposition however, weevils feed on bark from different sources but of unknown nutritional 'quality'. The factors influencing the rate of feeding by males and females and female reproductive development were determined in laboratory bioassays using two contrasting food sources - the bark on different species of seedling conifer and on logs of mature trees. The nutritional 'quality' of bark was characterized by the concentration of nitrogen, total sugars, total polyphenols and resin. Regression models were used to show that overall, the rate of feeding on the bark of both seedlings and logs increased with weevil size and was negatively related to nitrogen concentration. The nitrogen concentration in seedling bark (mean 1.1%) was about three times higher than that of logs (mean 0.4%). The rate of reproductive development increased with nitrogen intake during feeding and the preoviposition period for weevils feeding on seedlings and logs was ~ 13 days and 46 days, respectively. Analysis of weevil mortality and of the weight gain of surviving weevils suggests that a nitrogen content of around 0.3% may be limiting for H. abietis. The possibility that nutritionally adequate food resources may be limiting for H. abietis is briefly discussed.     

Preference and performance of the sawfly Diprion pini on host and non-host plants of the genus Pinus

The sawfly, Diprion pini L., is a pest of Pinus in Europe and is mainly found on P. sylvestris L. and P. nigra laricio Poiret. The relative importance of female oviposition capacity and behaviour, egg development, and larval survival on a new host plant was measured on 11 pine species. Five were natural host plants and six non-host plants, five of which are not indigenous to Europe. Oviposition choice tests showed that females discriminated between the pine species. Egg and larval development also differed between pine species. However, the female choice was not linked with hatching rate and larval development. Results of biological tests clearly indicated that there were different response patterns of D. pini life stages in relation to pine species, and these patterns were the same with insects of four different origins. We discuss the importance of each potential barrier to colonisation of a new host.     

Innate preference hierarchies coupled with adult experience,rather than larval imprinting or transgenerational acclimation,determine host plant use in Pieris rapae

The evolution of host range drives diversification in phytophagous insects, and understanding the female oviposition choices is pivotal for understanding host specialization. One controversial mechanism for female host choice is Hopkins’ host selection principle, where females are predicted to increase their preference for the host species they were feeding upon as larvae. A recent hypothesis posits that such larval imprinting is especially adaptive in combination with anticipatory transgenerational acclimation, so that females both allocate and adapt their offspring to their future host. We study the butterfly Pieris rapae, for which previous evidence suggests that females prefer to oviposit on host individuals of similar nitrogen content as the plant they were feeding upon as larvae, and where the offspring show higher performance on the mother's host type. We test the hypothesis that larval experience and anticipatory transgenerational effects influence female host plant acceptance (no‐choice) and preference (choice) of two host plant species (Barbarea vulgaris and Berteroa incana) of varying nitrogen content. We then test the offspring performance on these hosts. We found no evidence of larval imprinting affecting female decision‐making during oviposition, but that an adult female experience of egg laying in no‐choice trials on the less‐preferred host Be. incana slightly increased the P. rapae propensity to oviposit on Be. incana in subsequent choice trials. We found no transgenerational effects on female host acceptance or preference, but negative transgenerational effects on larval performance, because the offspring of P. rapae females that had developed on Be. incana as larvae grew slower on both hosts, and especially on Be. incana. Our results suggest that among host species, preferences are guided by hard‐wired preference hierarchies linked to species‐specific host traits and less affected by larval experience or transgenerational effects, which may be more important for females evaluating different host individuals of the same species.     

Host feeding experience affects host plant odour preference of the polyphagous leafminer Liriomyza bryoniae

Herbivorous insects use highly specific volatiles or blends of volatiles characteristic to particular plant species to locate their host plants. Thus, data on olfactory preferences can be valuable in developing integrated pest management tools that deal with manipulation of pest insect behaviour. We examined host plant odour preferences of the tomato leafminer, Liriomyza bryoniae (Kaltenbach) (Diptera: Agromyzidae), which is an economically important agricultural pest widespread throughout Europe. The odour preferences of leafminers were tested in dependence of feeding experiences. We ranked host plant odours by their appeal to L. bryoniae based on two‐choice tests using a Y‐tube olfactometer with five host plants: tomato, Solanum lycopersicum Mill.; bittersweet, Solanum dulcamara L.; downy ground‐cherry, Physalis pubescens L. (all Solanaceae); white goosefoot, Chenopodium album L. (Chenopodiaceae); and dead nettle, Lamium album L. (Lamiaceae). The results imply that ranking of host plant odours by their attractiveness to L. bryoniae is complicated due to the influence of larval and adult feeding experiences. Without any feeding experience as an adult, L. bryoniae males showed a preference for the airflow with host plant odour vs. pure air, whereas females did not display a preference. Further tests revealed that adult feeding experience can alter the odour choice of L. bryoniae females. After feeding experience, females showed a preference for host plant odour vs. pure air. Feeding experience in the larval stage influenced the choice by adults of both sexes: for males as well as females reared on bittersweet the odour of that plant was the most attractive. Thus, host feeding experience both in larval and/or adult stage of polyphagous tomato leafminer L. bryoniae influences host plant odour preference by adults.     

Influence of larval experience on preference of a subterranean insect Delia antiqua on Allium hosts

Numerous studies have reported that larval experience can affect subsequent host plants selection and future oviposition decisions of many different species, but the investigation of pre‐imaginal experiences on host preference of adults has rarely been tested for soil‐dwelling insects. In this study, we present evidence that larval feeding experience can affect adult host preference in the onion maggot, Delia antiqua. By rearing D. antiqua on different host plants, we were able to examine the role of the natal host of different generations and the effect of larval density on host‐choice behaviour. We also performed bioassays by means of switched host treatment to evaluate the host‐selection principle. Choice bioassays among the three host species demonstrated that D. antiqua females preferred to oviposit on their natal host in each generation and host‐switching treatments. Additionally, increasing larval density could intensify this ovipositional preference on the natal host. The overall results showed that host preference of female D. antiqua is determined by larval experience and density. These findings also add support for the controversial Hopkins’ host‐selection principle.     

Reduction of oviposition time and enhanced larval feeding: two potential benefits of aggregative oviposition for the viburnum leaf beetle

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Egg allocation by Bracon hylobii Ratz., the principal parasitoid of the large pine weevil (Hylobius abietis L.), and implications for host suppression

1 The braconid parasitoid Bracon hylobii Ratz. is one of the few specialist natural enemies of the large pine weevil, Hylobius abietis L., a destructive pest of conifer transplants. An assessment of its role as an agent of biological control requires a detailed knowledge of the allocation of its reproductive effort. 2 Parasitoid females were continuously observed in laboratory culture with individually reared host larvae in bark discs. The outcome of sequential parasitoid–host encounters was recorded by subsequent examination of hosts and by rearing all parasitoids. 3 Parasitoids avoided ovipositing on host larvae < 100 mg fresh weight, even though such larvae represented sufficient biomass for complete parasitoid development. All larger larvae were vulnerable to attack, which leaves a window of vulnerability for parasitoids of about 90% of weevil larval life. 4 Parasitoids presented with a range of host sizes showed no preference above 100 mg for the size of host first attacked, but allocated more eggs and a greater total handling time to larger hosts. 5 Most eggs were deposited on the first host attacked, with progressively fewer allocated to subsequent hosts. However, oviposition experience did not affect the time spent on the next host. 6 From these results it is anticipated that when weevil larval size is reduced by less favourable feeding substrates, fewer parasitoid eggs will be allocated to each but more host larvae will ultimately be attacked. 7 Generation time, host finding, oviposition rate, clutch size, life expectancy and diapause induction are strongly affected by temperature. Life expectancy is substantially shorter for parasitoids deprived of non‐host food supplement. At 15 and 20 °C the number of hosts attacked and the number of eggs deposited decreased with female age. 8 Bracon hylobii is inevitably poorly synchronized with a variable life‐cycle host; it is egg‐limited and can enter diapause at a relatively high field temperature. None of these characteristics suggest that it could stabilize the abundance of its host below an economically acceptable threshold density. However, the reproductive potential of the parasitoid suggests that it could make a significant contribution to larval mortality and suppress adult recruitment, thus complementing other control strategies.     

Larval food affects oviposition preference,female fecundity and offspring survival in Yponomeuta evonymellus

1. Yponomeuta evonymellus is a monophagous moth that feeds on Prunus padus which is native to Europe. In recent years, larval feeding and egg clusters have also been observed on non‐native Prunus serotina plants; however, survival of larvae on this new host is very low. 2. The objective of the present study was to determine how the feeding of larvae on each of the two host plants impacts oviposition, offspring survival and fecundity in Y. evonymellus. Our hypothesis was that, under controlled conditions, females will lay eggs on the host on which they fed as larvae. We also hypothesised that the lower survival of young larvae feeding on P. serotina was due to the smaller buds and leaves present in this species, relative to those of P. padus. 3. A dual‐choice experiment conducted under laboratory conditions demonstrated that females preferentially chose to oviposit on the plant species on which they fed as larvae. In the experiment, potential fecundity and offspring survival were significantly higher on P. padus than on P. serotina. The reduced performance of Y. evonymellus on P. serotina was correlated with a smaller bud mass and volume, lower leaf mass and surface area, and difficulty in constructing a protective tent against unfavourable weather conditions. 4. In summary, the identity of the host plant species during larval feeding determines adult oviposition preference for that host species. The survival of larvae on P. serotina growing in the nature is low, but for phenology‐related reasons.     

Fecundity, longevity and establishment of Otiorhynchus sulcatus (Fabricius) and Otiorhynchus ovatus (Linnaeus) (Coleoptera: Curculionidae) from the Pacific North-west of the United States of America on selected host plants

1 The fecundity, longevity and establishment of Otiorhynchus sulcatus and Otiorhynchus ovatus from the Pacific North‐west U.S.A. was studied on five selected host plants: Picea abies‘Nidiformis’, Picea glauca‘Conica’, Taxus baccata, Rhododendron catawbiense‘Boursault’ and Fragaria×ananassa‘Totem’. 2 Teneral adults were used to study adult longevity and reproductive success. Leaves of these host plants were used for sustenance for 9 months. Larval establishment was studied by infesting potted host plants with eggs. 3 Fragaria×ananassa‘Totem’ produced the longest survival, shortest preoviposition time, the greatest number of eggs, and the highest fertility for adults of both species. Picea spp. were not good adult hosts for O. sulcatus. Taxus was a good adult host for O. sulcatus, but was a nonhost for adults and larvae of O. ovatus. 4 Adult hosts did not affect preoviposition time or egg viability with O. ovatus adults. With O. sulcatus, preoviposition time was greatly increased and egg viability was < 50% on Picea spp. 5 The best larval host was F.×ananassa‘Totem’ for O. sulcatus and P. glauca‘Conica’ for O. ovatus. Rhododendron was a poor larval host for both species. 6 When all of the studies on these two pests are considered, O. sulcatus appears to have varying host preferences from among its many geographical areas of occurrence whereas O. ovatus has a more universal host selection.     

Host-associated variation in sexual size dimorphism and fitness effects of adult feeding in a bruchid beetle

We studied the effect of larval host (two Convolvulus L. species, Convolvulaceae) on sexual size dimorphism and on the fitness consequences of adult feeding in the bruchid beetle Megacerus eulophus (Erichson) (Coleoptera: Bruchidae). Whereas Convolvulus chilensis Pers. occurs in low density in semiarid habitats, Convolvulus bonariensis Cav. occurs in less stressful environments and exhibits higher population density. Host plants neither differ in seed mass nor in seed nitrogen content, and there were no consistent host‐associated differences in female fitness. Consequently, host quality was considered to be similar. Sexual size dimorphism, expressed as the female:male ratio of body size, was significantly greater in C. chilensis than in C. bonariensis. A greater female size in the C. chilensis population could be selectively advantageous due to higher survival during host‐plant search in stressful environments. Female longevity was affected by food availability (starvation vs. honey‐pollen solution) and by the interaction between food availability and larval host. Fed females lived longer than starved females. This trend was found in both hosts, but it was of greater magnitude in C. chilensis than in C. bonariensis. Fecundity was significantly affected by adult feeding and larval host, and marginally affected by their interaction. Fed females laid more eggs than starved females. However, the increase in fitness of fed females was greater in C. chilensis. Results might be explained by differential selection on plasticity in life‐history traits in contrasting environments. Females in stressful environments should take a greater advantage of the transient availability of resources.     

Biology and host range of Heterapoderopsis bicallosicollis: a potential biological control agent for Chinese tallow Triadica sebifera

Chinese tallow, Triadica sebifera, is an invasive weed that infests natural and agricultural areas of the south-eastern USA. A candidate insect for biological control of Chinese tallow has been studied under quarantine conditions. The biology and host range of a primitive leaf-feeding beetle, Heterapoderopsis bicallosicollis, were examined between July 2008 and February 2010. H. bicallosicollis adults fed on 16 of 29 non-target species and survived >10 days on all 29 species, more than the number of days that adults survived when deprived of food. Additionally, adult feeding, oviposition and partial larval development occurred on a Florida endangered species, Heterosavia bahamensis. These data indicated that H. bicallosicollis adult feeding and oviposition may not be limited to the target weed and included several valued North American natives. Therefore, as its release could pose unacceptable risks, testing of this species was discontinued and the quarantine colony destroyed.     

Polyphagy and primary host plants: oviposition preference versus larval performance in the lepidopteran pest Helicoverpa armigera

Oviposition preference and several measures of offspring performance of Helicoverpa armigera (Hübner) were investigated on a subset of its host plants that were selected for their reputed importance in the field in Australia. They included cotton, pigeon pea, sweet corn, mungbean, bean and common sowthistle. Plants were at their flowering stage when presented to gravid female moths. Flowering pigeon pea evoked far more oviposition than did the other plant species and was the most preferred plant for neonate larval feeding. It also supported development of the most robust larvae and pupae, and these produced the most fecund moths. Common sowthistle and cotton were equally suitable to pigeon pea for larval development, but these two species received far fewer H. armigera eggs than did pigeon pea. Mungbean also received relatively few eggs, but it did support intermediate measures of larval growth and survival. Fewest eggs were laid on bean and it was also the least beneficial in terms of larval growth. Among the host plant species tested, only flowering pigeon pea supported a good relationship between oviposition preference of H. armigera and its subsequent offspring performance. Australian H. armigera moths are thus consistent with Indian H. armigera moths in their ovipositional behaviour and larval performance relative to pigeon pea. The results suggest that the host recognition and acceptance behaviour of this species is fixed across its geographical distribution and they support the theory that pigeon pea might be one of the primary host plants of this insect. These insights, together with published results on the sensory responses of the females to volatiles derived from the different host plant species tested here, help to explain why some plant species are primary targets for the ovipositing moths whereas others are only secondary targets of this polyphagous pest, which has a notoriously broad host range. Handling Editor: Joseph Dickens     

Ecophysiological attributes of adult overwintering in insects: insights from a field study of the nut weevil,Curculio nucum

Abstract Diapausing insect species have evolved a great diversity of life cycles, although overwintering occurs at a single development stage within most species. Understanding why diapause has evolved towards a given life stage requires investigation of both the ecological and physiological attributes. Notably, it is suggested that adult overwintering is more energy‐demanding than larval overwintering but it brings fitness gains by allowing adults to be synchronized with their seasonal requisites through an early spring emergence. This hypothesis is tested in field conditions in the nut weevil Curculio nucum, whose life cycle comprises an obligate 2‐year, nonfeeding underground phase, including a larval, followed by an adult, overwintering. In this species, adult wintering leads to an early spring emergence; at first glance, however, this does not enhance synchronization between weevils and their host because adults emerge more than 1 month before starting to breed. It is suggested that adult overwintering ultimately evolved in response to the phenology of the host, by allowing females to oviposit in nuts before their full sclerotinization. Adult overwintering appears to be costly because adults postpone reproduction for 1 year, incur a significant weight loss and require feeding before egg laying. Surprisingly, lipids are unaffected during diapause, lipogenesis even being likely in the summer metamorphosis. These results suggest that the lipids involved in egg production may entirely come from the larval stages, whereas the other nutrients are acquired through adult feeding but this remains to be tested.     

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.     

Foraging behaviour, host plant and host location in the aphid hyperparasitoid Euneura augarus

Euneura augarusis a specialized ectohyperparasitoid of conifer lachnids, which causes considerable mortality in mummies of conifer lachnid parasitoids. For the search on a large scale, E. augarusfemales rely mainly on volatile secondary plant compounds of conifers to locate potential host plants. In the medium-range (i.e., on a given host plant), females detect the presence of hosts either during spiral search flights or during search by walking. Search flights were more effective when mummies were dispersed, while clumped mummies were mainly found by walking. On ant-attended trees, E. augaruswas not able to forage successfully. Choice experiments with Aphidius ervi-mummies showed that E. augarusis able to develop in species that are not attacked in the field. We hypothesize that the proximate mechanism of host plant specificity, and as a consequence, host aphid specificity seems to be the use of typical conifer volatiles for host finding which provide the `search image' for the species.     

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.     

Preference–performance relationship and influence of plant relatedness on host use by Pityogenes chalcographus L.

• 1 Pityogenes chalcographus L. (Coleoptera: Scolytinae) causes damage in European coniferous forests, primarily on Picea abies L. Karst., but is also recorded on other native and exotic Pinaceae species. Estimating the adequacy between adult preference and larval performance of this beetle among its host‐range, as well as the influence of plant taxonomic relatedness on these parameters, would provide useful information on the beetle's ability to shift onto novel hosts.
• 2 Choice and no‐choice assays were conducted under laboratory conditions. Adult preference and larval performance parameters among two native (Pinus sylvestris L. and Picea abies) and three exotic north American [Pinus contorta Dougl., Picea sitchensis (Bong.) Carr. and Pseudotsuga menziesii Mirbel (Franco)] conifer species were measured.
• 3 Pityogenes chalcographus exhibited a significant positive relationship between preference and performance. Picea abies was both the preferred and the most suitable host species for larval development. The closest relative, P. sitchensis, was the second best choice in terms of preference and performance. Pseudotsuga menziesii occupied an intermediate position for both beetle preference and performance, and Pinus spp. were the least suitable hosts for beetle development.
• 4 Adult preference and larval performance ranking among hosts provides little support to the plant taxonomic relatedness hypothesis. Taxonomic relatedness could play a role on the diet breadth, although only at a limited scale, within the genus Picea. At higher taxonomic levels, other factors such as bark thickness might be decisive.

     

The metazoan parasites of swordfish from Atlantic tropical–equatorial waters

The metazoan parasite fauna composition of the swordfish Xiphias gladius is reported from 145 (72 females and 73 males) specimens caught in Atlantic tropical–equatorial waters during June to July 2006 and August 2007. Seventeen metazoan taxa (two monogeneans, two digeneans, one acanthocephalan, four cestodes, seven nematodes and one copepod) were identified by morphological and genetic methodologies. Among those identified, Anisakis brevispiculata, Anisakis paggiae and the acanthocephalan Bolbosoma vasculosum are new host records. Dominant species were: the anisakid nematode larvae Anisakis physeteris, Anisakis brevispiculata, Hysterothylacium incurvum and Hysterothylacium corrugatum (s.l.), the adult cestode Fistulicola plicatus, the plerocercoid Hepatoxylon squali and the monogeneans Tristoma integrum and Tristoma coccineum, which showed high prevalences and intensities. The relationship between metazoan parasites and host size and sex was investigated for the first time. No statistically significant correlation was found between host sex and parasite burden. Infection levels with T. integrum and T. coccineum were observed to be significantly negatively correlated to host size. Infection levels by endoparasites observed in X.gladius from Atlantic tropical‐equatorial area may be related to host feeding behaviour and the availability of the parasites’ intermediate and final hosts in the area studied. Anisakids of the genus Anisakis, were identified genetically to species level, and were considered to be promising biological tags for the characterization of X.gladius stocks.     

Reproductive development and maternal effects in the pine weevil Hylobius abietis

1. In a laboratory study of maturation feeding of female pine weevil Hylobius abietis on current and 1‐year‐old stem bark of transplants of Scots and Corsican pine, Norway and Sitka spruce, Douglas fir, and hybrid larch, the length of the pre‐oviposition period was influenced by the species on which weevils fed. The shortest pre‐oviposition period was on hybrid larch (11.8 days) and the longest on Douglas fir (15.5 days). 2. The species on which weevils fed also affected fecundity but there was evidence of a species–year interaction. Over a period of 36 days, most eggs were laid by weevils feeding on current stem of Norway spruce and Corsican and Scots pine and fewest on current stem of Sitka spruce. 3. Significant maternal effects on egg size were observed both in relation to female size and conifer species. The largest eggs were laid on Corsican pine and the smallest on Douglas fir, with no evidence of a trade‐off between number of eggs laid and their size. 4. There was a positive relationship between egg and larval size and between larval size and survival on logs of four conifer species. Residual resistance mechanisms in the bark of recently cut stumps and larval competition are discussed briefly in relation to the importance of the observed maternal effects on weevil population dynamics.