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Emerald ash borer, Agrilus planipennis (Fairmaire) (Coleoptera: Buprestidae), a phloem-feeding beetle native to Asia, has become one of the most destructive forest pests in North America. Since it was first identified in 2002 in southeast Michigan and Windsor, Ontario, dozens of isolated A. planipennis populations have been discovered throughout Michigan and Ontario, and in 12 other states and the province of Quebec. We assessed realized A. planipennis dispersal at two discrete outlier sites that originated 1 yr and 3 yr earlier from infested nursery trees. We systematically sampled ash trees within an 800 m radius of the origin of each infestation to locate galleries constructed by the progeny of dispersing A. planipennis adults. Our sampling identified eight trees at the 1 yr site infested with a mean ± SE of 11.6 ± 8.4 A. planipennis larvae and 12 trees at the 3 yr site with 25.8 ± 11.1 larvae per meter squared. Dendroentomological analysis indicated that A. planipennis populations were predominantly undergoing a 2 yr (semivoltine) life cycle at both sites. Colonized trees were found out to 638 and 540 m from the epicenters at the 1 yr and 3 yr sites, respectively. Logistic regression was used to determine whether the likelihood of A. planipennis colonization was affected by wind direction, ash phloem abundance, distance from the epicenter, or land-use type (i.e., wooded, residential, agricultural, or urban). Results show that the probability of A. planipennis colonization was significantly affected by ash phloem abundance and decreased with distance from the epicenter.
Onion thrips, Thrips tabaci (Lindeman) (Thysanoptera: Thripidae), can reduce onion bulb yield and transmit iris yellow spot virus (IYSV) (Bunyaviridae: Tospovirus), which can cause additional yield losses. In New York, onions are planted using seeds and imported transplants. IYSV is not seed transmitted, but infected transplants have been found in other U.S. states. Transplants are also larger than seeded onions early in the season, and thrips, some of which may be viruliferous, may preferentially colonize larger plants. Limited information is available on the temporal dynamics of IYSV and its vector in onion fields. In 2007 and 2008, T. tabaci and IYSV levels were monitored in six seeded and six transplanted fields. We found significantly more thrips in transplanted fields early in the season, but by the end of the season seeded fields had higher levels of IYSV. The percentage of sample sites with IYSV-infected plants remained low (<12%) until August, when infection levels increased dramatically in some fields. The densities of adult and larval thrips in August and September were better predictors of final IYSV levels than early season thrips densities. For 2007 and 2008, the time onions were harvested may have been more important in determining IYSV levels than whether the onions were seeded or transplanted. Viruliferous thrips emigrating from harvested onion fields into nonharvested ones may be increasing the primary spread of IYSV in late-harvested onions. Managing T. tabaci populations before harvest, and manipulating the spatial arrangement of fields based on harvest date could mitigate the spread of IYSV.
To characterize habitats of Nannophya pygmaea Rambur (the northern pygmyfly; Odonata: Lilbellulidae), which is endangered in Korea, we analyzed characteristics of surface water and soil, landscape properties, and vegetation types in 22 habitats in eight areas of Korea where nymphs of N. pygmaea have been found since 2005. We divided the habitats into two groups: DS (dwelling site) habitats, where N. pygmaea was observed at the time of the study, and PDS (past dwelling site) habitats, where N. pygmaea recently lived but is no longer found. The habitats were mostly located in former paddy fields on mountain slopes that have been abandoned for 3–7 yr. The main water sources for these habitats were ground water and surface runoff, and the water level was stable at 3–7 cm in depth. The habitats ranged from 300 to 1000 m2 and were dominated by Juncus effusus, which formed tussock mounds. According to the hydrosere model of succession, N. pygmaea appeared mostly in the early stages of plant succession (the period ≈3–7 yr after the initiation of succession in former paddy fields) and N. pygmaea preferred habitats displaying the water and soil characteristics that are typical of the early stages of succession in abandoned paddy fields. These results indicate that the primary habitats of N. pygmaea in Korea are recently abandoned paddy fields that are in an oligotrophic state. As succession proceeds in these habitats, N. pygmaea disappears. A habitat management program should be launched to conserve the habitats and populations of N. pygmaea.
The seasonal abundance of woolly apple aphid (Eriosoma lanigerum Hausmann) was studied from 2005 to 2008 in commercial orchards in central Washington State. There was a single period of crawler migration on the tree trunks each year lasting from 7 to 16 wk. The peak period of crawler activity ranged from early June to late July, although peaks in June were more common. In one orchard, peak densities occurred 7 wk later in 2006 than in 2005. In all orchards where large numbers of crawlers were caught (up to 1,420 per band per week), more crawlers were caught in the lower band (migrating up from root colonies) than in the upper bands (migrating down from the aerial colonies). However, the peak period of crawler migration was similar for root and aerial colonies. In contrast, peaks in population densities of aerial colonies were sometimes bimodal, usually spanning the period from late May through mid-October; however, colonies were detected in mid-April in one orchard. There was no consistent relationship between the crawler density and aerial colony density within an orchard; this raises challenging questions regarding annual reinfestation of the aerial portions of the tree. Nymphs and adults formed 66 and 15%, respectively, of all stages found throughout the season. Alates were only found in September and October in orchards with fall populations, comprising up to 43% of individuals in aerial colonies. Overwintering survival was higher on the root colonies than in the aerial colonies in both years this was studied. Fruit infestation by aphids was relatively rare, and occurred only when aerial colonies were numerous.
The polyphagous drywood termite Kalotermes flavicollis (Fabricius) can establish its colonies inside living trees, causing a progressive debilitation that ends with the death of the plant. For this reason, it is considered one of the major pests of Sherry vineyards in Andalusia, SW Spain. To analyze the trends of colonization of this termite, a detailed investigation was performed during 16 mo in seven districts of the Sherry area, cutting randomly selected grapevine trunks into six sections, from which all individuals were collected and counted according to their developmental instar or caste. The results detected variations in the social composition of the groups colonizing grapevines according to the spatial distribution inside the vine trunk and in the numerical trends over the seasons. A pattern of seasonal migrations of the termites inside the trunk was inferred, with upward movements from spring to autumn and downward movements from autumn to winter. During winter, when termites were mainly located in the medium-basal sections of the trunk, a higher proportion of eggs and larvae was detected. Going from spring to summer, when the majority of the termites was located in the medium-upper sections of the trunk, there was an increase of nymphal instars followed, during autumn, by the appearance of alates, mostly located in the upper sections of the grapevine. Pseudergates and reproductives maintained their proportions and locations almost unchanged during the year.
Two strains of western flower thrips, Frankliniella occidentalis (Pergande), are reputedly found in New Zealand. One strain was recorded in 1934, and it is most common in flowers of Lupinus arboreus outdoors (lupin strain); the other strain was first recorded in New Zealand in 1992 and is found mostly indoors on greenhouse crops (greenhouse strain). Laboratory studies were conducted to compare the life history parameters of these two strains. Thrips from each strain were fed sucrose solution and capsicum or lupin pollen and reared at 25°C, >60% RH, and16 L:8 D photoperiod. Significant differences in life history parameters were found. Preoviposition time was significantly shorter, and oviposition rate and fecundity were markedly higher (four-fold) for the greenhouse than for the lupin strain. The lupin strain performed significantly better on the capsicum pollen, laying more than twice as many eggs than on the lupin pollen over a 14-d period. The greenhouse strain development time from larvae to adult was marginally faster (0.7–1.1 d less) than the lupin strain because of a shorter prepupal and a marginally shorter pupal development time. Females of the greenhouse strain lived on average 69% longer than females from the lupin strain. Large differences in the intrinsic growth rate (rm) were found, with rm being 1.4–1.8 times higher for the greenhouse strain than the lupin strain, depending on pollen source. The results are discussed in relation to different ecological requirements and pest status of the two strains.
An important challenge facing ecologists is to understand how climate change may affect species performance and species interactions. However, predicting how changes in abiotic variables associated with climate change may affect species performance also depends on the biotic context, which can mediate species responses to climatic change. We conducted a 3-yr field experiment to determine how the herbivorous grasshopper Camnula pellucida (Scudder) responds to manipulations of temperature and population density. Grasshopper survival and fecundity decreased with density, indicating the importance of intraspecific competition. Female fecundity tended to increase with temperature, whereas grasshopper survival exhibited a unimodal response to temperature, with highest survival at intermediate temperatures. Grasshopper performance responses to temperature also depended on density. Peak survival in the low-density treatment occurred in warmer conditions than for the high-density treatment, indicating that the intensity of intraspecific competition varies with temperature. Our data show that changes to the temperature regimen can alter grasshopper performance and determine the intensity of intraspecific competition. However, the effects of temperature on grasshopper performance varied with density. Our data indicate the importance of the biotic context in mediating species responses to climatic factors associated with global change.
Peucetia viridans (Hentz) lives almost exclusively on Croton ciliatoglandulifer (Ortega) in the dry forests of western Mexico. This spider is usually found on pubescent shrubs. Within their host plants, P. viridans has been associated with plant height and cover, as well as with number of flowers or inflorescences in anthesis. Flowers can be used as cues of good habitat conditions or because they attract prey detected by the spider. In this study, we assessed the importance of flowers, plant cover, and plant exposure (sun/shade) on the spider distribution in five 50-plant transects. In a field experiment, we also compared the number of spiders between controls, plants from which inflorescences were removed, and plants with artificial inflorescences. The results from the transects indicate that, when the number of flowers per spider is high, spiders were more abundant in exposed locations, which presumably offer better microclimatic conditions; when flowers become scarce, food may be more difficult to find and the spider distribution become strongly associated with the number of flowers, where they are more likely to find prey. Spider abundances on the experimental plot decreased on plants from which flowers were removed in comparison to control plants. Spider abundance increased on those in which artificial inflorescences were added. The similarity between plants with natural and artificial inflorescences suggests that spiders use flowers as cues of good microhabitats instead of prey visitors, which are significantly less abundant on artificial inflorescences.
Many studies have evaluated the effect of forest fragmentation on dung beetle assemblage structure. However, few have analyzed how forest fragmentation affects the processes carried out by these insects in tropical forests where their food sources consist mainly of dung produced by native herbivore mammals. With the conversion of forests to pastures, cattle dung has become an exotic alternative and abundant food for dung beetles. This study compares dung removal rates of native (monkey) and exotic (cow) dung in different-sized fragments of tropical rain forests, during the dry and rainy seasons at the Los Tuxtlas Biosphere Reserve. Dung removal rates were affected by season, dung type, and the interaction between resource type and season. During the dry season, the removal rates of monkey dung were somewhat similar than during the rainy season, whereas the removal rates of cow dung were much higher during the rainy season. Dung beetle biomass and species richness were almost three times greater in monkey dung than in cow dung. Monkey dung attracted species belonging to the dweller, roller, and tunneler guilds; cow dung attracted mostly tunnelers. Therefore, the use of exotic dung may result in a biased misconception of the rates of dung removal in tropical forest and an underestimation of dung beetle diversity. This study highlights the importance of working with natural tropical forest resources when attempting to identify realistic tendencies concerning processes in natural habitats and those modified by fragmentation and by other human activities.
In the Republic of Korea, most denuded forest lands have been restored since the 1960s. In addition, the annual mean temperature in the Republic of Korea has increased ≈1.0°C during the last century, which is higher than the global mean increase of 0.74°C. Such rapid environmental changes may have resulted in changes in the local butterfly fauna. For example, the number of butterflies inhabiting forests may have increased because of reforestation, whereas the number of butterflies inhabiting grasslands may have declined. Furthermore, the number of northern butterflies may have declined, whereas the number of southern butterflies may have increased in response to global warming. Therefore, we compared current data (2002≈2007) regarding the abundance of butterfly species at two sites in the central portion of the Korean Peninsula to data from the late 1950s and early 1970s for the same sites. Changes in the abundance rank of each species between the two periods were evaluated to determine whether any patterns corresponded to the predicted temporal changes. The predicted changes in butterfly abundance were confirmed in this study. In addition, the results showed a different response to habitat change between northern and southern species. In northern butterfly species, butterflies inhabiting forests increased, whereas those inhabiting grasslands declined. However, the opposite was true when southern butterfly species were evaluated. Changes in the abundance indicate that habitat change may be one of the key factors related to the survival of populations that remain around the southern boundary of butterfly species.
Although most North American grasshopper species overwinter as eggs, some species hatch in late summer, overwinter as nymphs, and become adults in late spring. These species periodically reach outbreak densities, but it is unknown if they impact the population dynamics of later developing egg-overwintering grasshopper species. Two experiments were conducted in a northern mixed-grass prairie to examine the effects of herbivory by nymph-overwintering grasshoppers on survival of an egg-overwintering grasshopper, Ageneotettix deorum. When very high densities of nymph-overwintering grasshoppers reduced grass biomass by 80%, survival of A.deorum nymphs was significantly reduced. There was no effect of early season herbivory at a lower density of 16 /m2 on survival of A. deorum nymphs. The effects of early season herbivory appear consistent with exploitative competition. Although a strong reduction in peak grass biomass caused by early season herbivory negatively affected late season grass-feeding grasshopper densities, additional research is needed to examine the potential importance of early summer herbivory on population dynamics of later developing grasshoppers.
Emergence patterns of a natural population of Osmia lignaria subsp. lignaria Say were monitored during a 5-yr period (2003–2007) in relation to air temperature, bloom of eastern redbud (Cercis canadensis L.), and emergence of the parasitoid Chysura kyrae Krombein. Bee cocoons were placed in gelatin capsules and overwintered within shelters in their natural habitat. Mean emergence of female O. l. lignaria was observed at ≈2 d after redbud bloom began; mean male emergence was ≈ 1 wk earlier than for females and was nearly identical to that of C. kyrae. Horticultural models used to predict budburst for several tree species were evaluated with respect to redbud bloom and O. l. lignaria emergence. Each model indicated that both sexes of the bee and redbud had similar chill unit requirements. Differences in emergence and bloom dates were attributed to the different requirements for growing degree hours (GDHs) after chilling requirements were met. The close phenological relationship observed between eastern redbud and bee emergence suggests an important role for this common understory tree in the early nesting success of O. l. lignaria.
The aim of this study was to compare the dung beetle (Coleoptera, Scarabaeidae: Scarabaeinae) community structure at two sites in the Charles Darwin Ecological Refuge in Igarassu, Pernambuco, Brazil. Dung beetles were collected in 2006 using monthly samples from 48 pitfall traps baited with human dung and bovine carrion. The dung beetle communities from the study sites were compared in terms of abundance, species richness, and diversity (Shannon index). Seasonality was analyzed by Spearman correlation between rainfall data and community parameters. In total, 2,560 individuals belonging to 40 species, 16 genera, and 6 tribes were collected. Species richness was higher for the clear-cut area compared with the forest habitat. Estimators of species richness suggested a total richness of 42–47 species in the entire study area. A positive correlation was observed between monthly rainfall and total abundance of individuals for the clear-cut area but not for the forest habitat. This study contributes to a better understanding of Scarabaeinae ecology in the Atlantic rainforest of northeastern Brazil.
Invasive pests, such as the cactus moth, Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae), have not reached equilibrium distributions and present unique opportunities to validate models by comparing predicted distributions with eventual realized geographic ranges. A CLIMEX model was developed for C. cactorum. Model validation was attempted at the global scale by comparing worldwide distribution against known occurrence records and at the field scale by comparing CLIMEX “growth indices” against field measurements of larval growth. Globally, CLIMEX predicted limited potential distribution in North America (from the Caribbean Islands to Florida, Texas, and Mexico), Africa (South Africa and parts of the eastern coast), southern India, parts of Southeast Asia, and the northeastern coast of Australia. Actual records indicate the moth has been found in the Caribbean (Antigua, Barbuda, Montserrat Saint Kitts and Nevis, Cayman Islands, and U.S. Virgin Islands), Cuba, Bahamas, Puerto Rico, southern Africa, Kenya, Mexico, and Australia. However, the model did not predict that distribution would extend from India to the west into Pakistan. In the United States, comparison of the predicted and actual distribution patterns suggests that the moth may be close to its predicted northern range along the Atlantic coast. Parts of Texas and most of Mexico may be vulnerable to geographic range expansion of C. cactorum. Larval growth rates in the field were estimated by measuring differences in head capsules and body lengths of larval cohorts at weekly intervals. Growth indices plotted against measures of larval growth rates compared poorly when CLIMEX was run using the default historical weather data. CLIMEX predicted a single period conducive to insect development, in contrast to the three generations observed in the field. Only time and more complete records will tell whether C. cactorum will extend its geographical distribution to regions predicted by the CLIMEX model. In terms of small scale temporal predictions, this study suggests that CLIMEX indices may agree with field-specific population dynamics, provided an adequate metric for insect growth rate is used and weather data are location and time specific.
The symbiotic relationship of termites and their eukaryotic and prokaryotic gut microbiota is a focal point of research because of the important roles symbionts play in termite nutrition. The use of molecular methods has recently provided valuable insights into the species diversity and the roles of microorganisms in the guts of termites. This paper provides a review of the current knowledge of symbiont species inventories, genome analysis, and gene expression in the guts of subterranean termites. Particular emphasis is given to the termite genera Reticulitermes and Coptotermes (Isoptera: Rhinotermitidae), because they contain pest species of global impact in their native and invasive range.
This study aims at exploring the potential relationship between aphidiine parasitoid development and the primary endosymbiont in aphids by focusing on specific aphid instars and the relative effects on parasitoid oviposition behavior and progeny development. Lysiphlebus ambiguus (Aphidiidae, Hymenoptera) is a solitary parasitoid of several species of aphids, including Aphis fabae. In this study, A. fabae was treated with antibiotic rifampicin to obtain aposymbiotic hosts and exposed to parasitism. L. ambiguus launched significantly more attacks on symbiotic L2 (the second instar), aposymbiotic L3 (the third instar) and L4 (the forth instar) hosts than on the corresponding hosts at the same age. L. ambiguus also parasitized more L1 aphids compared with adults irrespective of whether the aphid was asymbiotic or not. Pupa mortality rate of parasitoid progeny was significantly lower from aposymbiotic hosts than from the corresponding symbiotics at all stages. Female-biased parasitoid progeny was produced from aposymbiotic aphids without respect to host ages, but female progeny increased linearly with host ages at parasitism from symbiotic aphids. Body size of parasitoid progeny increased linearly with host instars at parasitism in symbiotic aphids but did not significantly change across host instars in aposymbiotic aphids. The offspring parasitoids turned out to be generally large in body size from attacking aposymbiotic aphids compared with the symbiotics. Development time of egg-to-adult of parasitoid progeny decreased with host instars in both symbiotic and aposymbiotic aphids but was generally much longer in aposymbiotic aphids than in symbiotic aphids. Our study suggests that age or body size of host aphids may not be the only cue exercised by L. ambiguus to evaluate host quality and that offspring parasitoids may be able to compensate for the nutrition stress associated with disruption of primary endosymbiotc bacteria in aposymbiotic aphids.
We studied the distribution patterns of Wolbachia infection associated with plum curculio strains in eight states of the eastern United States. The presence of the Wolbachia-specific gene wsp identified infections of this endosymbiont in 97.8% of the 93 samples tested. Three distinct Wolbachia strains were identified. The strains wCne1 (593 bp) and wCne2 (593 bp) were 97% identical, and their sequences were both 84% identical with wCne3 (590 bp). BLASTN searches through GenBank showed strong similarities between the wsp sequences of the three strains compared with Wolbachia sequenced from other hosts. Degree of similarity with sequences in other Wolbachia strains is discussed. Polymerase chain reaction-restriction fragment length polymorphism was used for superinfection detection. Of 93 samples, 15 (16.1%), 21 (22.6%), 19 (20.4%), and 36 (38.7%) samples were infected by wCne1, wCne2, wCne1 2, and wCne3, respectively. Only two (2.2%) samples had no infection. The wCne3 strain was always present as a single infection. Wolbachia strains approximate the distribution of plum curculio strains: northern strain infected with wCne1 and wCne2 strains in supergroup B, and southern strain infected with wCne3 strain in supergroup A, with the mid-Atlantic region as the convergence area. Based on haplotype distribution of plum curculio mitochondrial cytochrome oxidase I, there was a closer relation of the mid-southern plum curculio clade to the far-southern clade than to the northern clade. However, Wolbachia symbionts in mid-southern plum curculio are more closely related to those in northern plum curculio than to those in far-southern plum curculio. The relationship of Wolbachia infection with reproductive incompatibility between plum curculio populations was also discussed.
Bacterial communities are known to play important roles in insect life histories, yet their consistency or variation across populations is poorly understood. Bacteria associated with the bark beetle Dendroctonus valens LeConte from eight populations, ranging from Wisconsin to Oregon, were evaluated and compared. We used the culture-independent technique of denaturing gradient gel electrophoresis to visualize bacterial diversity, or individual operational taxonomic units (OTUs), from individual beetles. One-way analysis of similarities was used to test for differences of bacterial communities between sites. Analysis of community profiles showed that individual beetles on average contained 10 OTUs, with frequency of association from 2 to 100% of beetles. OTU sequences most closely matched β- and γ-proteobacteria, and one each matched Bacilli and Actinobacteria. Several OTUs were particularly abundant, most notably an Actinobacterium from 100% and two Proteobacteria from 60% of beetles sampled. Some OTUs were similar to previously described bacteria with known biochemical capabilities and ecological functions, suggesting that some bacterial associates of D. valens may contribute to its ability to exploit a resource low in nutrients and high in defensive compounds. There were significant differences of bacterial communities between sites. The strength of these differences was positively correlated with distance between sites, although additional unexplained factors also contribute to the variation.
Root-feeding beetles, particularly of the curculionid subfamilies Scolytinae and Molytinae, are known to be effective vectors of Ophiostomatoid fungi. Infestation by these insects and subsequent infection by the Ophiostomatoid fungi may play an important role in accelerating symptom progression in pine declines. To examine the relationship between beetles and fungi in longleaf pine stands, root-feeding curculionids were collected in pitfall traps baited with ethanol and turpentine for 62 wk, and Ophiostomatoid fungi were isolated from their body surfaces. The most abundant root-feeding beetles captured were Hylastes tenuis, H. salebrosus, Pachyhbius picivorus, Hylobius pales, and Dendroctonus terebrans. The number of insects captured peaked in spring and fall, although peaks for different insect taxa did not coincide. The most frequently isolated fungi were Grosmannia huntii, Leptographium procerum, L. terebrantis, and L. serpens. Other Ophiostomatoid fungi recovered included Ophiostoma spp. and Pesotum spp. Insect infestation data suggest that Hylastes spp. share an ecological niche, as do Hb. pales and P. picivorus, because the ratios of their fungal symbionts were similar. The fungi associated with D. terebrans suggest that it did not share habitat with the other principle vectors.
Mutualism is a prominent interaction within ecosystems, yet most may actually be conditional. The symbiotic mite, Ensliniella parasitica Vitzthum, ingests the hemolymph of juvenile potter wasps, Allodynerus delphinalis (Giraud), but also protects them from a natural enemy, Melittobia acasta, and is transported to new nests in host pockets specialized for this purpose (i.e., acarinaria). Thus, two different antagonisms from the mite may arise: commensalistic cheating is expected without the natural enemy of the host, and parasitism is predicted with excessive numbers of the mite. However, facultative parasitism mediated by mutualism has rarely been studied in any organism. We found no significant differences in juvenile mortality, nesting rate, or fecundity between mite-free and naturally mite-laden juveniles. However, when overloaded with mites (≈1.5–2.5 times more mites than the maximum number per wasp larva in the field), the developmental period of the male wasp was significantly delayed, and juvenile wasp mortality increased to 30%. These results show that mutualism mediated by parasitism may revert to parasitism, suggesting that either or both organisms in a mutualism mediated by parasitism need population control of the parasite to avoid the risk of parasitism.
The impact of invasive alien species on native organisms is a cause for serious concern. This concern is especially relevant in the Hawaiian archipelago because of its high level of endemicity, severe impacts of accidental introductions of invasive species, and long history of purposeful biological control introductions. Results from a previous study showed that the parasitoid assemblage associated with an endemic moth Udea stellata (Butler) comprised seven species: three adventive species, two purposely introduced species, and two of unknown origin. The objectives of this study were to assess the parasitism levels of alien wasps on populations of U. stellata at different sites and to determine the specific stages that were used by the spectrum of parasitoid species that attack U. stellata. Standardized collections of wild larvae were conducted at eight sites located on the islands of Kauai, Oahu, and Hawaii. In total, 3,531 larvae were collected in a 2-yr survey. Of these, 8.0% were collected as first instar, 23.0% as second instar, 39.0% as third instar, 21.0% as fourth instar, 7.1% as fifth instar, and 1.8% as sixth instar. Of the larvae that survived laboratory rearing, 43.0% were parasitized. Information collected in the surveys was complemented with data from life-table studies to determine stage-specific parasitism. All larval stages were susceptible to parasitism by at least one parasitoid species; second and third instars were susceptible to attack by all seven parasitoid species. Adventive parasitoids rather than purposely introduced ones were responsible for the greater part of the apparent mortality observed. At low and low-medium elevations, the parasitoid assemblage was dominated by adventive species. The two purposely introduced parasitoids were present in remote relatively undisturbed sites on the islands Kauai and Hawaii. The sometimes high parasitism rates by adventive species found in this study were shown to have minimal effect at the population level in our life table study; therefore, care should be taken when interpreting field parasitism data. Carefully addressing current ecological impacts of alien parasitoids on native species is of particular importance for developing more efficient means to quantify the risks of future biological control introductions.
The hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), native to western North America and Asia, was accidentally introduced from Japan to the eastern United States. To potentially establish biological control of A. tsugae, we released a predator endemic to western North America, Laricobius nigrinus Fender (Coleoptera: Derodontidae), from 2003 to 2005, in 22 localities from Georgia to Massachusetts. Release sites spanned the invasive range of the adelgid across five United States Department of Agriculture plant hardiness zones (5a to 7a). Release sizes were 75, 150, 300, 600, or 1,200 adult L. nigrinus per site in the fall, winter, early spring, or sequentially (i.e., fall or winter and early spring). We monitored establishment by annual sampling for L. nigrinus adults with beat sheets and for L. nigrinus larvae by branch clipping. At the end of 3 yr, L. nigrinus was established in 13 of the 22 sites. The following variables were evaluated for their correlation with the numbers of L. nigrinus larvae and adults recovered and for their effect on establishment (scored as F3 presence/absence): (1) Minimum winter temperature at the release site, (2) A. tsugae density at the time of release, (3) release size, and (4) release season. Only minimum winter temperature was correlated with larval recoveries and no variables were correlated with adult recoveries. Logistic regression modeling found that establishment was positively related to minimum winter temperature and release size. We recommend smaller release sizes in warm areas where establishment probability was high (i.e., zones 7a, 6b, and 6a) and larger release sizes in cold areas where establishment probability was low (i.e., zones 5b and 5a). Releases during fall—early spring and across the range of A. tsugae densities tested were successful.
Bracon hebetor Say (Hymenoptera: Braconidae) is a gregarious larval ectoparasitoid of several species of Lepidoptera that are associated with stored products. The suitability of 12 potential lepidopteran host species representing four families was studied in this study for the development and reproduction of B. hebetor. The Lepidoptera species used were the Indianmeal moth, Phdia interpunctella (Hübner), Mediterranean flour moth, Ephestia kuehniella (Zeller), almond moth, E. cautella (Walker), rice moth, Corcyra cephalonica (Walker), navel orangeworm, Amyelois transitella (Stainton), greater wax moth (laboratory reared and commercial), Galleria mellonella (Linnaeus) (all Pyralidae); tobacco budworm, Heliothis virescens (Fabricus), corn earworm, Helicoverpa zea (Boddie), beet armyworm, Spodoptera exigua (Hübner) (all Noctuidae); webbing clothes moth, Tineola bisselliella (Hummel) (Tineidae); and Angoumois grain moth, Sitotroga cerealella (Olivier) (Gelichiidae). Experiments were conducted using petri dishes (100 by 15 mm) as experimental arenas at 29 ± 1°C, 65 ± 5% RH, and a photoperiod of 14:10 (L: D) h. B. hebetor females were introduced singly into arenas and given a full-grown host larva every day for 5 consecutive d. Paralysis of the host larvae and oviposition by B. hebetor females were significantly affected by host species. The cumulative fecundity in the 5-d period was highest on A. transitella (106.42 ± 5.19) and lowest on T. bisselliella (9.64 ± 1.28). The egg-to-adult survivorship and progeny sex ratio were also significantly affected by the host species. The highest percentage of parasitoid survival to the adult stage was on A. transitella (84.07 ± 2.26) and zero on T. bisselliella. Egg to adult development time was shortest on E. cautella (9.75 ± 0.25 d) and longest on G. mellonella (12.63 ± 0.28 d). Results from these studies suggest that B. hebetor females can use a wide range of lepidopteran hosts for paralysis and oviposition. However, B. hebetor cannot necessarily develop and reproduce on all host species that it can paralyze and oviposit on, and optimum reproduction is with the stored-product pyralid hosts.
Horizontal transmission of insect viruses is a key factor in their cycling in agroecosystems. Here we study the transmission of the baculovirus HaSNPV among larvae of Helicoverpa armigera (Hübner) in cotton. Transmission of three HaSNPV genotypes was studied from larvae infected with a single virus genotype and from larvae infected with two different genotypes. Genotypes included a wild-type virus, an ecdysteroid UDP-glucosyltransferase (egt) deletion mutant (HaSNPV-LM2) with slightly enhanced speed of kill, and an egt-negative genotype that expresses a neurotoxin gene derived from the scorpion Androctonus australis Hector (HaSNPV-4A). The latter genotype has a substantially increased speed of kill. In three field experiments, the wild-type and egt deletion virus variants and a mixture of the two had similar rates of transmission. Transmission increased with density of infector insects and decreased with time lapsed since the inoculation of the infector larvae. Transmission of the neurotoxin expressing virus was lower than that of the other two genotypes in a glasshouse experiment. The studied genotypes of HaSNPV have significant differences in time to kill and virus yield, but we found no significant differences in rates of virus transmission at the crop level in the case of the egt deletion variant HaSNPV-LM2. Transmission of the transgenic virus genotype HaSNPV-4A was significantly reduced. Overall, differences in transmission between virus genotypes were subtler, and more difficult to detect with statistical significance, than effects of other factors, such as density of infectors and time delay between release of infectors and recipient caterpillars on the plant.
Seventeen isolates of Metarhizium anisopliae (Metschnikoff) Sorokin and three isolates of Beauveria bassiana (Balsamo) Vuillemin (Hypocreales: Clavicipitaceae) were evaluated for their pathogenicity to the adult pea leafminer, Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae), in the laboratory. Flies were contaminated with dry conidia through a velvet material wrapped around the inner side of a cylindrical plastic tube. All the isolates were pathogenic to the pea leafminer, causing mortality between 40 and 100% at 5 d after exposure. The lethal time for 50% mortality (LT50) ranged from 2.6 to 5.4 d, whereas the LT90 values varied between 3.2 and 9.1 d depending on the isolate. An autoinoculation device was evaluated in cage field experiments using only one of the virulent isolates, M. anisopliae ICIPE 20. The device was loaded with 2–3 g of dry conidia. Mortality of up to 100% was observed in flies captured from fungus-treated cages held under laboratory conditions. The average number of spores picked up by a single fly visiting the device increased with days after inoculation. One day after the inoculation, adults picked up an average of 4.1 ± 0.7 × 105 conidia and 39.6 ± 4.0 × 105 conidia 5 d after inoculation. Depending on the sampling date, the LT50 varied between 1.8 and 3.4 d. Results indicate that some isolates of B. bassiana and M. anisopliae are highly pathogenic to L. huidobrensis, suggesting a potential for their use in the control of this pest. They also suggest the possibility of L. huidobrensis suppression with fungi using an autoinoculation device.
Surveys were conducted on regenerating stands of lodgepole pine to determine the relationship between root deformation and susceptibility to attack by the Warren root collar weevil, Hylobius warreni Wood. The total number of trees attacked by H. warreni did not differ between planted and natural trees. A matched case-control logistic regression suggested that root crosssectional area was more important in predicting weevil attack for naturally regenerated trees than for planted trees, but weevils were associated with a larger reduction in height-to-diameter ratios for trees with planted root characteristics than for trees with natural root form. Neither the stability of attacked versus unattacked trees differed significantly and there was no significant interaction of weevil attack and tree type, but weevil-killed trees had different root characteristics than alive, attacked trees. Lateral distribution and root cross-sectional area were significant predictors of alive attacked trees versus weevil-killed trees, suggesting that trees with poor lateral spread or poor root cross-sectional area are more likely to die from weevil attack. We conclude that root deformation does not necessarily increase susceptibility to attack but may increase the likelihood of mortality. Thus, measures to facilitate good root form are needed when planting pine in areas with high risk of Warren root collar weevil attack.
The soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), alternates between a primary overwintering host (buckthorn, Rhamnus sp.) and a secondary summer host (soybean, Glycine max). Selection of soybean cultivars with different maturity groups may provide growers with a management tool for A. glycines, either directly through its effect on summer populations that cause economic damage or indirectly through its effect on the production of migrants that disperse to the primary host in fall. This study investigated the abundance and seasonality of A. glycines on soybean cultivars with different maturity rates in central Indiana. The abscission of soybean foliage occurred earlier for early maturing than late maturing cultivars, but no other consistent difference in development or yield was detected among the cultivars tested in this study. The abundance of aphids did not vary consistently among cultivars when soybean was most susceptible to economic damage. A laboratory assay evaluating the larviposition preference of A. glycines alate females, combined with a 7-yr survey documenting the colonization of buckthorn by winged aphids, indicated that the production of gynoparae on soybean began in mid-September and continued until leaf abscission. The abundance of aphids during this period was higher on late maturing cultivars than on early maturing cultivars in both 2006 and 2008, whereas no significant effect was detected in 2007. Altogether, these results suggest that planting early maturing soybean cultivars has little effect on damage by aphids on the current season crop but may reduce the number of fall migrants to the primary host.
Insect herbivore abundances on host plants are influenced by both plant traits and the physical environment in which that plant grows. This study examined the role of the physical light environment and foliage characteristics in determining abundance of the lacebug Corythuca arcuata Say (Hemiptera: Tingidae) on Quercus alba L. I censused adult C. arcuata across a growing season, quantified leaf characteristics, and measured canopy cover over understory branches of mature Q. alba. Using an information-theoretic approach, a priori hypotheses of the relationship between light, plant traits, and C. arcuata abundance was evaluated. Abundance was best predicted by light environment and carbon content. Adult C. arcuata prefer trees growing under an open canopy and trees with low carbon content; abundance also positively correlated with leaf water content. Although carbon and water did not vary with light in this study, low carbon and high water content are often associated with shadier conditions, suggesting that C. arcuata faces a trade-off between preferences for physical habitat conditions and host plant characteristics.
One of the most damaging insect pests in deciduous forests of North America is the forest tent caterpillar, Malacosoma disstria Hübner. It can feed on a variety of plants, but trembling aspen (Populus tremuloides Michaux) is its preferred host and sugar maple (Acer saccharum Marshall) serves as a secondary one in the northern part of its distribution. Because host plant characteristics influence insect performance and survival, we evaluated the impact of trembling aspen and sugar maple foliage on M. disstria performance. Host effects on insect cold hardiness and overwintering survival of offspring were also studied. Forest tent caterpillar reared on aspen leaves had a shorter development time, higher pupal weights and fecundity, and superior egg parameters (length and weight) compared with those reared on sugar maple leaves. Larvae from the two food treatments had low glucose levels during diapause, whereas glycerol content of insects reared on maple was significantly higher during diapause than larvae fed on aspen. Lower glycerol content may explain the higher overwinter mortality of pharate larvae from aspen-reared parents even though their supercooling points were as low as -36°C. This study shows the influence of host plant on insect life history and the need to consider overwintering success and offspring performance in studies to understand and predict population growth and cycling.
A micro-cage bioassay was developed to test the effect of slash pine pollen (Pinus elliottii Engelm.) supplementation to a whole onion plant (Allium cepa L. variety Pegasus) diet on thrips (Thysanoptera: Thripidae) reproductive parameters. Frankliniella fusca (Hinds) females were placed on two to three-leaf stage onion seedling under a treatment of either slash pine (Pinus elliottii Engelm.) pollen dusting (a pollen supplement) or no pollen treatment. Adult survival, net oviposition, and offspring produced over a series of ten 2-d intervals were recorded. From these values, lx, lxmx, and R0 values were constructed. A trimodal distribution of oviposition was observed with the pollen supplement. Increased oviposition rates led to higher female offspring production per female and to a four-fold increase in F. fusca net reproduction on pollen-treated onions.
In the eastern United States, two invasive specialist insects share a native host plant, Eastern hemlock, Tsuga canadensis. In recent years, much research has focused on the impacts of the hemlock woolly adelgid (Adelges tsugae) because of the detrimental effects it has on hemlock growth and survival. In contrast, the invasive elongate hemlock scale (Fiorinia externa) is thought to have only minor impacts on hemlock. We infested hemlock saplings with each insect and compared them with control (i.e., neither insect herbivore) saplings to assess how early infestations impact Eastern hemlock health (measured using new branch growth, foliar %N, and C:N ratio). Our study showed that, at equal densities, the two insects differed in their effect on Eastern hemlock. F. externa did not impact plant growth or foliar chemistry over the course of the 2-yr experiment. A. tsugae significantly reduced plant growth and caused a reduction of %N in the first year of the experiment. By the end of the experiment, A. tsugae trees had the same %N in their foliage as control and F. externa trees but drastically reduced growth patterns. The most likely explanation for this result is the greater growth in control and F. externa saplings during the second year resulted in the dilution of available foliar N over a larger amount of newly produced plant tissue. For early infestations of both insects, our study suggests that management plans should focus on the more detrimental A. tsugae.
Epidemic populations of Enaphalodes rufulus (Haldeman), red oak borer, a native longhorned wood boring beetle, were implicated as a major contributor to a recent widespread oak mortality event in the Ozark National Forest of Arkansas. We assessed potential factors affecting suitability of a primary host Quercus rubra L., northern red oak, which experienced dieback and mortality throughout two successive borer cohorts. We sampled trees with various E. rufulus infestation levels during 2001–2003 when populations were at outbreak levels and 2003–2005 when borer numbers were declining. We measured phloem thickness and calculated a vigor index; the ratio of the past 5-yr basal area increment to sapwood area. We also counted established first year larval feeding galleries and outer-bark adult emergence holes and measured surface area of feeding galleries on a subset of Q. rubra to assess the importance of host susceptibility versus suitability. Phloem thickness did not exhibit any patterns among hosts of varying infestation levels and was therefore not likely an important factor limiting larval success. Less vigorous Q. rubra appeared to be the most suitable hosts, although it is unclear whether reduced vigor was initially caused by stress of E. rufulus infestation or environmental factors. Host suitability seems to be more important than host susceptibility, as numbers of initiated galleries were not consistently different among host infestation classes and between both cohorts, whereas numbers of emerging adults did differ predictably among host infestation classes.
A 2-yr study was conducted to evaluate the tillering and yield response of rice, Oryza sativa L., whose culms were injured artificially or by larval sugarcane borers, Diatraea saccharalis (F.). Artificially injured plants produced ≈0.49 more tillers than uninjured plants, similar to what has previously been reported for larval injured plants. In contrast, artificial injury did not affect yield per tiller, whereas larval injury did. The proximity of larval injury to the panicle had a negative impact on tiller yield, whereas artificial injury did not. Artificial injury apparently resulted in less injury to vascular tissue than did sugarcane borer larval injury. Until an artificial method of injury is developed that mimics the effects of larval feeding, further injury studies will continue to require sugarcane borer larvae.
Chlorpyrifos is one of the most widely used insecticides worldwide. It has been shown to have deleterious effects on survival of nontarget insects, but its impact on behavior has received less attention. In this study, we investigated the sublethal effects of this insecticide on sexual pheromone discrimination in two Trichogramma species. In these species, sexual pheromones arrest partners belonging to the same species. This specificity is important for reproduction efficacy because interspecific matings are sterile. We used two sympatric and closely related species of Trichogramma to study how two doses (LD 20 and LD 0.1) of chlorpyrifos can alter the discrimination by males and the emission by females of pheromones from both species. When exposed to the LD 20, the males of both species showed a decrease in the discrimination of conspecific sexual pheromones. For one of the two species (T. evanescens Westwood), this decrease even led to a total annihilation of discrimination. A dose as low as the LD 0.1, inducing no apparent mortality, induced the same decrease in pheromone discrimination for T. semblidis Aurivillius males. However, no effect was observed on the discrimination by males of sexual pheromones emitted by females either exposed to an LD 20 or an LD 0.1 of chlorpyrifos. By decreasing the discrimination by males of sexual pheromones, chlorpyrifos may induce interspecific interactions and attempts at copulating that would decrease the fitness of parasitoids. The implications of these results in relation to environmental pollution, the mode of action of the insecticide and the status of natural enemy of Trichogramma are discussed.
Some species of mosquito can detect the presence of larvivorous fish and select against ovipositing in pools supporting them. The effect of kairomones released by the crimson-spotted rainbowfish Melanotaenia duboulayi (Castelnau) on the oviposition behavior of the freshwater mosquitoes Culex annulirostris Skuse, Culex quinquefasciatus Say, and Aedes notoscriptus Skuse was evaluated in both laboratory and outdoor artificial pool experiments. In the laboratory, colony-reared Cx. annulirostris selected against ovipositing in water that had contained M. duboulayi at densities of one fish per 5, 30, 180, and 1,080 liters. In contrast, gravid Ae. notoscriptus showed a preference for water that had contained the highest density of M. duboulayi (1 in 5 liters). Gravid Cx. quinquefasciatus were neither repelled nor attracted to water that had previously contained M. duboulayi. In outdoor artificial pool experiments, wild Cx. annulirostris females selected against ovipositing in pools containing caged M. duboulayi stocked at rates of 1, 4, and 10 g per 1,000 liters. When fish were removed from the pools, the repellent effect persisted for at least 24 h, whereas at the lower stocking rate, an avoidance response was not noted until 48 h after fish were placed in pools. This suggests a volatile substance, however, the exact nature of the kairomone/s has not been identified.
Most adult Lepidoptera feed on nectar, whereas caterpillars consume mainly structural tissue such as leaves, stems, flowers, and/or fruits. This may result in behavioral trade-offs in which search time for high-quality oviposition sites suitable for larval food is restricted by adult foraging needs. Here we report on the preference for and performance on flowering and nonflowering host plants of the generalist herbivore Helicoverpa armigera to explore whether there are such behavioral trade-offs between moth and their caterpillars offpsring. We found that the adult moths have a strong oviposition preference for flowering tobacco and sunflower plants. Young caterpillars preferred to feed on the inflorescences. Adult-realized fecundity was almost 10 times higher when ovipositing on flowering plants. Weight at pupation, which is correlated with potential future fecundity of the caterpillars, was also higher when feeding on flowers. We found no evidence for a behavioral trade-off and conclude that a general preference for flowers by Helicoverpa armigera is highly beneficial from a nutritional perspective for both adults and larvae. The results suggest that the manipulation of flowering plants for the attraction of oviposition is relevant to pest control of this polyphagous species.
Oviposition site selection is crucial for the reproductive success of a herbivore insect species with relatively sedentary larvae. The optimal oviposition theory, i.e., the preference-performance hypothesis, has thus far mainly been tested with a focus on nutritional quality of the host. This study investigates whether female oriental fruit moth Grapholita (Cydia) molesta choose a microhabitat for oviposition characterized by a temperature range within which their offspring perform best. Thermal preferences of females during oviposition were assessed in a circular temperature gradient arena. Offspring performance and survival were assessed under different constant temperature conditions. Females preferred oviposition sites of ∼30°C over lower and higher temperatures. At this temperature, egg, larval, and pupal development was significantly faster than at 22 and 25°C, and larval development was also faster than at 33°C. At 30°C and at the lower temperatures tested, survival of eggs and larvae was significantly higher than at 33°C, whereas development was precluded at 35°C. Furthermore, female pupal weight attained at 30 and 33°C exceeded that reached at the lower temperatures tested. Considering the potentially reduced predation risk caused by the shorter developmental time of eggs and larvae, the laboratory data suggest that this species maximizes its fitness by selecting a thermally optimal environment for its offspring, supporting the optimal oviposition theory. Conversely, it is known that the codling moth (C. pomonella) lacks a mechanism to avoid temperatures lethal to progeny development, which may reflect the differences in geographic ranges of these tortricids.
We studied the dispersal behavior and survival of sterile medfly males either treated or not with ginger root oil (GRO), in field conditions, in Petrolina-PE, northeast Brazil, from May 2006 to December 2007 in a sterile insect technique (SIT) program. The tsl strain Vienna 8 from the Ceratitis capitata Wied. (Diptera: Tephritidae), medfly, mass-rearing facility located in Juazeiro-BA, Brazil, was used. The results showed that sterile males either exposed or not to GRO exhibit similar dispersal behavior and postrelease survival. More than 60% of the sterile males, either treated or not treared wth GRO, were recovered at a 25-m distance from the releasing point, ≈20% at 50 m, and 5% in traps situated 100 m from the releasing point. Around 90% of the sterile males, exposed or not to GRO, were recovered 5 d after release of the sterile male individuals, whereas <1% were recovered after 11 d. Our results imply that ginger root oil can be used to treat sterile medfly males without interfering with their dispersal or survival in the field.
Aphidophagous lady beetles enhance their foraging success in natural settings by consuming other types of food in addition to aphids. Frass analysis was used to examine natural diets of female lady beetles in fields of alfalfa (Medicago sativa L.) in northern Utah. The first (spring) alfalfa crop was censused in 2004 and 2005 to determine the diet of female adults of the introduced Coccinella septempunctata L., and two native species, C. transversoguttata richardsoni Brown, and Hippodamia convergeas Guerin. The proportion of females of the three lady beetle species that fed on pea aphids [Acyrthosiphon pisum (Harris) ] and alfalfa weevil larvae [Hypera postica (Gyllenhal), an abundant alternative prey] increased from early to late season during the first crop. A corresponding seasonal decrease occurred in the proportion of females consuming other types of arthropods (e.g., thrips and collembolans) and nonarthropod food (pollen and fungal spores). Overall, frass analysis indicated that the diets of C. septempunctata and the two native species in alfalfa were similar in their inclusion of a broad variety of foods. The study shows that frass analysis can provide a good overview of the diets of lady beetles in natural settings.
The relative performance of a clear delta trap baited with individual or combination sex pheromone and kairomone lures for codling moth, Cydia pomonella L., was evaluated against three vertical clear pane and colored delta traps in field trials within apple, Malus domestica (Borkhausen). The clear caught significantly more moths of each sex than an orange delta trap when baited with ethyl (E,Z)-2,4-decadienoate (pear ester) and acetic acid (PE AA) and performed similarly to oil-coated pane traps. The clear caught significantly more females than the orange delta trap with pear ester alone, pear ester plus (E,E)-8,10-dodecadien-l-ol (codlemone) (PE-PH), or a combination of PE-PH and acetic acid. Male moth capture was similar in both clear and colored delta traps with all codlemone lures tested. Seasonal moth catches of female moths were higher in clear than either white (second flight, 2008) or orange (first and second flights, 2009) traps baited with PE-PH. Total moth catch was significantly higher in clear than white traps in 2008 and did not differ between clear and orange traps in 2009. Clear traps baited with acetic acid but not with pear ester, PE-PH, or when unbaited caught significantly more nontarget moths than colored traps. These studies suggest that the use of clear traps with their higher captures of female codling moths could improve both monitoring programs and the development of lure and kill strategies.
Testing impacts of large-scale aerial spraying to control public health pests under realistic field conditions are needed to understand impacts on natural populations of nontarget insects. Responses of terrestrial insects to aerial applications of an organophosphate insecticide, naled, used for mosquito and biting midge control were studied on Parris Island Marine Corps Recruit Depot, Parris Island, SC. Aerial applications were made with C-130 aircraft at dusk in 2003 and in 2005. In 2003, we sampled two locations on Parris Island with Malaise traps before and after spraying, and used Before-After analyses to examine changes in terrestrial insect diversity and abundance. In 2005, we sampled insects with yellow pan traps at three locations on Parris Island and at an untreated control site. A Before-After Control-Impact analysis at each location was conducted to compare changes. In 2003, numbers of four of 12 common taxa (Dolichopodidae, Sarcophagidae, Syrphidae, Tachinidae) were lower after sprays. However, there were no significant changes in numbers of common taxa or total numbers in 2005. Shannon diversities (H') were not different in either year indicating that sprays had minimal impact on overall community biodiversity. In contrast, populations of pestiferous biting midges (Culicoides spp.) collected in CDC-style traps were reduced by 94–99% after spraying in both years; mosquito numbers declined by 88.2% in 2003 and 92.5% in 2005, after sprays.
Microplitis mediator Haliday (Hymenoptera: Braconidae) is an important larval endoparasitoid of various lepidopteran pests, including Helicoverpa armigera (Hübner). In China, H. armigera is a key pest of cotton and is currently the focus of several biological control efforts that use M. mediator as principal natural enemy of this pest. To improve the success of biological control efforts, behavioral studies are needed that shed light on the interaction between M. mediator and H. armigera. In this study, we determined M. mediator response to volatile compounds from undamaged, mechanically injured, or H. armigera--damaged plants and identified attractive volatiles. In Y-tube olfactometer assays, we found that mechanically damaged plants and/or plants treated with H. armigera oral secretions did not attract wasps. However, volatiles from H. armigera—damaged plants elicited a strong attraction of both M. mediator sexes. Headspace extracts from H. armigera—damaged cotton were analyzed by coupled gas chromatography—electroantennographic detection (GC-EAD), and a total of seven different compounds were found to elicit electroantennogram (EAG) responses, including an unknown compound. Six different EAD-active volatiles were identified from caterpillar-damaged cotton plants, of which 3, 7-dimethyl-1, 3, 6-octatriene and (Z)-3-hexenyl acetate were the principal compounds. Olfactometer assays indicated that individual synthetic compounds of 3, 7-dimethyl-1, 3, 6-octatriene, (Z)-3-hexenyl acetate, and nonanal were attractive to M. mediator. Field cage studies showed that parasitism of H. armigera larvae by M. mediator was higher on cotton plants to which 3, 7-dimethyl-1, 3, 6-octatriene was applied. Our results show that the combination of terpenoids and green leaf volatiles may not only facilitate host, mate, or food location but may also increase H. armigera parasitism by M. mediator.
Detecting infestations of stink bugs (Heteroptera: Pentatomidae) using pheromones remains problematic, particularly so in the United States for the exotic stink bug, Nezara viridula L., and our native stink bug, Acrosternum hilare (Say). Therefore, we conducted a 2-yr on-farm study to examine the attractiveness and possible cross-attraction of the reported pheromones for N. viridula and A. hilare and those previously discovered for Euschistus servus (Say) and Plautia stall Scott to N. viridula, A. hilare, and E. servus. The attractiveness of selected pentatomid pheromones to tachinid parasitoids of stink bugs was also examined. We showed for the first time under field conditions that N. viridula can be trapped with its reported pheromone, a 3:1 trans- to cis-(Z)-α-bisabolene epoxide blend. In fact, attraction of N. viridula increased with higher pheromone doses. Traps baited with a 5:95 trans- to cis-(Z)-α-bisabolene epoxide blend, the reported male-produced A. hilare attractant pheromone, failed to attract significantly more A. hilare than did unbaited control traps. Instead A. hilare was significantly cross-attracted to the P. stali pheromone [methyl (E,E;,Z) -2,4,6-decatrienoate]. The E. servus pheromone [methyl (E,Z) -2,4-decadienoate], either alone or in combination with P. stali pheromone, was more attractive to E. servus than to N. viridula, P. stali, or A. hilare pheromones. In general, tachinid parasitoids were found responsive to the male-specific volatiles of their known hosts, including the attractiveness of Trichopoda pennipes (F.) to sesquiterpenoid blends characteristic of A. hilare and N. viridula. A tachinid parasitoid of E. servus, Cylindromyia sp., seemed to be attracted to E. servus pheromone. In conclusion, our results indicate that stink bug traps baited with lures containing N. viridula pheromone blend, P. stali pheromone, and E. servus pheromone have the greatest potential for detecting populations of N. viridula, A. hilare, and E. servus, respectively, in diversified agricultural landscapes.
Diaphorina citri Kuwayama (Hemiptera: Psyllidae) carries Candidatus liberibacter spp., the putative causal agents of Huanglongbing. D. citri reproduces and develops only on the flushing shoots of its rutaceous host plants. Here we examined whether D. citri is attracted to host plant odors and a mixture of synthetic terpenes. Tests conducted in a vertically oriented Y-tube olfactometer showed that both males and females preferentially entered the Y-tube arm containing the odor from the young shoots of Murraya paniculata (L.) Jack and Citrus limon L. Burm. f. cultivar Eureka. Only males exhibited a preference for the odor of C.sinensis L., whereas the odor of C. × paradisi MacFadyen cultivar Rio Red was not attractive to both sexes. The volatiles emitted by young shoots of grapefruit cultivar Rio Red, Meyer lemon (Citrus × limon L. Burm.f.), and M. paniculata were analyzed by gas chromatograph-mass spectrometry. The samples were comprised of monoterpenes, monoterpene esters, and sesquiterpenes. The number of compounds present varied from 2 to 17, whereas the total amount of sample collected over 6 h ranged from 5.6 to 119.8 ng. The quantitatively dominant constituents were (E)-β-ocimene, linalool, linalyl acetate, and β-caryophyllene. The attractiveness of a mixture of synthetic terpenes, modeled on the volatiles collected from M. paniculata, was evaluated in screened cages in a no-choice test. At three observation intervals, significantly more individuals were trapped on white targets scented with the mixture than on unscented targets. These results indicate the feasibility of developing D. citri attractants patterned on actual host plant volatiles.
The antennae of Grapholita molesta (Busck) are more susceptible to sex pheromone—induced sensory adaptation than the antennae of Choristoneura rosaceana (Harris). Adaptation is detectable in G. molesta at 1/1,000th the aerial concentration of pheromone (i.e., 5.0 × 10-7 ng Z8-12:OAc/ml air) that is required to induce detectable adaptation in C. rosaceana (i.e., 5.0 × 10-4 ng Z11–14:OAc/ml air). In addition, the predicted concentration of pheromone required to induce 50% adaptation in G. molesta (i.e., 1.2 × 10-3 ng Z8–12:OAc/ml air) after 15 min of exposure is only one sixth the estimated aerial concentration required to induce the same level of adaptation in C. rosaceana (i.e., 7.5 × 10-3 ng Z11–14:OAc/ml air) after 15 min of exposure. Sixteen percent and 28% adaptation of G. molesta antennae is predicted after 15 and 30 min of exposure to the equivalent of 1 ng Z8–12:OAc/m3 air (i.e., 1 × 10-6 ng Z8–12:OAc/ml air). The predicted level of adaptation in C. rosaceana antennae, however, is only 1.5 and 9.9% after 15 and 30 min of exposure, respectively, to the equivalent of 1 ng Z11–14:OAc/m3 air (i.e., 1 × 10-6 ng Z11–14:OAc/ml air). The approximately three-fold greater level of sensory adaptation in G. molesta antennae after 30 min of exposure to a pheromone concentration measured in pheromone-treated orchards (i.e., 1 ng/m3) may be one reason why this species is more readily controlled than C. rosaceana using mating disruption.
We conducted laboratory and field bioassays to characterize the pheromone system of an ash bark beetle, Hylesinus pruinosus Eichhoff (Coleoptera: Curculionidae: Scolytinae). Solitary females in newly initiated galleries in ash logs produced ( )-exo-brevicomin, whereas male beetles paired with females produced (-)-endo-brevicomin, lesser quantities of ( )-exo-brevicomin, and a third compound that could not be identified. Beetles produced these compounds also after exposure to juvenile hormone III, and they were the sole volatile chemicals isolated from beetles or aerations of infested logs that elicited electrophysiological responses from antennae of either sex. In the field, both sexes were strongly attracted to traps baited solely with either racemic or pure ( )-endo-brevicomin. Racemic exo-brevicomin was much less attractive to both sexes than racemic endo-brevicomin, and it did not increase attraction of endo-brevicomin when released in combination. Host odors (volatiles from mechanically damaged ash branches) failed to attract beetles or increase attractiveness of racemic exo-brevicomin. Our evidence suggests that male-produced ( )-endo-brevicomin is the major component of an aggregation pheromone for H. pruinosus, with ( )-exo-brevicomin and the unidentified male compound playing an indeterminate role in the chemical ecology of this species. Our data thus show an instance in which the major aggregation pheromone component of a bark beetle is produced by the secondarily arriving sex, a rare occurrence in bark beetles but one which has been reported previously for the Hylesini.
A seven-state survey showed that the recently detected invasive Asian banded elm bark beetle, Scolytus schevyrewi Semenov, was abundant in areas of Colorado and Wyoming, whereas the long-established European elm bark beetle, S. multistriatus (Marsham), was not as abundant. In one of a series of studies to evaluate whether S. schevyrewi is competitively displacing S. multistriatus in their North American zone of sympatry, we characterized long-range flight responses infested or uninfested to small cut logs of American, Chinese, and Siberian elm, Ulmus americana, U. parvifolia, and U. pumila. Trials were conducted in Colorado and Wyoming to test the flight response of S. schevyrewi; in California to test the response of S. multistriatus; and in Nevada to test the responses of both species simultaneously. Studies with S. schevyrewi showed that males and females aggregated toward Ulmus spp. host volatiles but provided no evidence of a putative aggregation pheromone during the 0- to 48- or 48- to 96-h period of infestation. In contrast, S. multistriatus was attracted to U. pumila over unbaited controls, more to U. pumila infested with conspecific females than without, and more during the 48- to 96- versus 0- to 48-h period of infestation. This confirmed that male and female S. multistriatus aggregated toward host volatiles and that females produced an aggregation pheromone. In a cross-attraction study, S. schevyrewi displayed neither flight preference nor interruption to 17. pumila infested with conspecifics, heterospecifics, or a mix of both species. Response of S. multistriatus was too low to draw conclusions. Although S. multistriatus aggregates moderately to host volatiles and strongly to female-derived pheromones emitted after a few days, S. multistriatus may have a relative disadvantage by selecting elm hosts more slowly than S. schevyrewi, which aggregates very strongly to host volatiles. The differential long-range host location strategy may be one factor in a chain of behavioral events that leads to advantageous host colonization and development by S. schevyrewi.
Methyl salicylate (MeSA) is a common herbivore-induced plant volatile that, when applied to crops, has the potential to enhance natural enemy abundance and pest control. The impacts of MeSA in strawberry were unknown and examined in the spring and midsummer period. Strawberry plots contained no lures (control) or two 30-d MeSA lures (Predalure) in the center: one lure 0.61 m aboveground over a sticky trap, and one lure on a plant near the ground. Arthropod abundance was monitored at the point source, 5 m and 10 m away from lures over 31 d with white sticky traps, pitfall traps, and leaf inspection. Twenty-seven and nine comparisons were made among beneficial and pest arthropods, respectively. Overall positive responses were found among Chrysopidae in July–August 2008 and Orius tristicolor (White) in May–June 2009 to MeSA based on sticky traps. Chrysopidae showed attraction to the point source, but not at 5 m and 10 m. Ground-dwelling predators collected in pitfall traps such as Araneae, the carabid beetles, Pterostichus melanarius (Illiger), and Nebria brevicollis (Fabricius) did not respond. Increased abundance of six natural enemy groups appeared on various dates between 3 and 24 d after placement of lures in the field based on leaf inspection and sticky traps. Conversely, fewer Coccinellidae were captured on sticky traps on days 0–3, and fewer natural enemies were observed on leaves on day 28 in MeSA plots. MeSA did not increase nor decrease pest abundance.
When invasive herbivorous insects encounter novel plant species, they must determine whether the novel plants are hosts. The Mediterranean pine engraver, Orthotomicus erosus (Wollaston), an exotic bark beetle poised to expand its range in North America, accepts hosts after contacting the bark. To test the hypothesis that O. erosus accepts hosts on the basis of gustatory cues, we prepared bark and phloem extracts from logs of four North American tree species that we had used in previous host acceptance experiments. Water, methanol, and hexane extracts of red pine, tamarack, balsam fir, and paper birch were presented alone and in combination on a neutral filter paper substrate in a section of a plastic drinking straw. Boring behavior in response to the three-extract combinations differed from the pattern of acceptance previously observed among species when the beetles were in contact with the bark surface. Only the aqueous extracts of tamarack, Larix laricina, increased the initiation and the extent of boring by O. erosus on the filter paper substrate. We conclude that the effects of extracted chemicals do not match the behavior of the beetles observed when penetrating excised bark and phloem discs, indicating that host selection by O. erosus may not be predictable from bark and phloem chemistry alone. Instead, host acceptance may be determined by nongustatory stimuli or by a combination of stimuli including gustatory and nongustatory cues.
The root weevil Diaprepes abbreviatus L. has been continuously reared since 1992 on an artificial diet (F1675) first reported in 1982. The weight of weevils reared on F1675 was more than twice as great as field-collected individuals. Recently, several ingredients included in the original F1675 diet were shown to have little or no effect on insect performance. We examined the effects of varying three ingredients of a simplified diet on the weight of adult D. abbreviatus. We used a geometric design combined with response surface models to evaluate the effects of the amount and proportion of diet ingredients to identify optimal diet recipes for normative weight gain defined in relation to field-collected D. abbreviatus from locations in Florida. Our results showed that it is possible to produce adults of normative weight or, for that matter, any mean weight within a wide range by varying the proportions of cottonseed meal, wheat germ and cellulose, a non-nutritive filler. Although wheat germ contributed to greater weight gain, survival of larvae to adult was lower on diets containing only wheat germ compared with diets containing only cottonseed meal as the principal nutritive components. The analyses of all the variables measured indicate that cottonseed meal is the only major nutritive component, in addition to standard vitamin and salt mixes, required to produce adult D. abbreviatus of normative weight.
Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is the vector of the bacteria that causes citrus greening and is considered one of the world's most important citrus diseases. We examined how host, geographic region, and gender affect the thermal requirements of D. citri. The insects were reared in climatic chambers at constant temperatures of 18, 20, 22, 25, 28, 30, and 32 ± 1°C, 70 ± 10% RH, and a 14 h photophase. Host plants for D. citri included orange (Citrus sinensis [Rutaceae]) varieties Pêra and Natal, the rootstock, Rungpur lime (C. limonia [Rutaceae]) and the natural host, Orange jessamine (Murraya paniculata [Rutaceae]). To study the influence of geographic origin on thermal requirements, we studied D. citri populations from Piracicaba, SP (warmer region) and Itapetininga, SP (cooler region). The duration and survival of the development stages and the duration of the total development (egg-adult) did not differ significantly on the different hosts, but it did vary with temperature. Nymphs of D. citri created on the different hosts have the same thermal requirements. The thermal requirements for this species collected from the two climate regions were identical; males and females also had the same thermal requirements.
Factors that influence thysanopteran wing diphenism are not well known. In these studies, the impact of food quality, mediated through nitrogen addition, and conspecific density was explored on the wing diphenism of an herbivorous thrips species (Anaphothrips obscurus Müller) (Thysanoptera: Thripidae). In the first study, nitrogen was added to timothy grass (Phleum pretense L.) (Poales: Poaceae) transplants, and naturally occurring thrips populations were caged on the plants. Thrips abundance and foliar nutrients were assessed every 2 wk. A separate factorial experiment in growth chambers explored the impact of both plant nitrogen addition and thrips abundance on wing diphenism. Thrips density was manipulated by adding either 3 or 40 thrips to potted and caged timothy. Thrips abundance and foliar nutrients were measured 58 d after treatment placement. Plant quality directly affected thrips wing diphenism independent of thrips density in both experiments. Near the end of the field cage experiment, density may have indirectly impacted wing diphenism. In both experiments, plant quality and thrips density interacted to affect thrips population abundance. Plant quality alone can affect thrips wing diphenism, but it remains unclear whether density alone can affect thrips wing diphenism. This is a unique and understudied system that will be useful to examine generalized theories on the negative interaction between reproduction and dispersal.
Most studies on ecological impact of solar ultraviolet (UV) radiation generally focus on plants. However, UV radiation can also affect organisms at other trophic levels. Protection against mortality induced by solar UV has, therefore, been hypothesized as one of the reasons why Typhlodromalus aripo hides in the apex of cassava plants during the day and comes out at night to prey on spider mites on leaves. In laboratory experiments using UV lamps, we determined the impact of UVA and UVB radiation on survival and oviposition of two leaf-inhabiting mites (Amblydromalus manihoti, Euseius fustis) and the apex-inhabiting mite (T. aripo), all three species being predators used for controlling the cassava green mite Mononychellus tanajoa in Africa. Whereas on leaf discs UVA has no negative impact on survival of the three predators, UVB is lethal to all of them. In contrast, nearly 85% of T. aripo survived after exposure to UVB inside apex of cassava plants. Exposure of A. manihoti and E. fustis to UVB radiation on the lower surface of a cassava leaf resulted in 36% survival. Oviposition and hatching of eggs laid after exposure to UVB were not affected, but eggs directly exposed to UVB did not hatch. Although caution should be exercised to extrapolate laboratory studies to the field, our results support the hypothesis that lower side of leaves, but especially plant apices, represent refuges that protect predatory mites from UVB. This might explain why T. aripo moves out of the apex to forage on leaves only during the night.
The lepidopteran stemborers Busseola fusca (Fuller) (Lepidoptera: Noctuidae), Sesamia calamistis (Hampson) (Lepidoptera: Noctuidae), and Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) are effectively controlled by Bt maize that expresses the Cry1Ab insecticidal protein. Another noctuid species, the cutworm Agrotis segetum (Denis and Schiffermüller) (Lepidoptera: Noctuidae), which is the most common and injurious pest of maize seedlings in South Africa, is exposed to Bt toxin for a part of its life cycle. The effect of this exposure to Bt maize has not been studied yet. The aims of this study were to determine the effects of Bt maize (events MON810 and Bt11) on larval mass, development time, survival, and fecundity of A. segetum. Laboratory studies were conducted with first- and fourth-instar larvae and moths. Results showed that the effect of Cry1Ab toxin on the biology of A. segetum larvae and moths were largely insignificant. The effects of the two Bt maize events on the different parameters measured in this study was not similar between the Bt events and their respective iso-hybrids. Compared with larvae that fed on conventional (non-Bt) maize, Bt maize did not affect survival of first-instar larvae. However, mean mass of larvae that fed on Bt maize (Bt11) was significantly lower. Feeding on Bt maize did not have a significant effect on development and survival of fourth-instar larvae or moth longevity. It did, however, delay the development period to pupa formation. Fewer eggs were laid by moths fed as larvae on maize event Bt11 compared with MON810. This study indicates that Bt maize will most likely not have any significant effect on the control of A. segetum under field conditions.
Corn (Zea mays L.) transformed with a gene from the bacterium Bacillus thuringiensis (Bt) comprises 49% of all corn in the United States. The input of senesced corn tissue expressing the Bt gene may impact stream-inhabiting invertebrates that process plant debris, especially trichopteran species related to the target group of lepidopteran pests. Our goal was to assess risk associated with transgenic corn debris entering streams. First, we show the input of corn tissue after harvest was extended over months in a stream. Second, using laboratory bioassays based on European corn borer [Ostrinia nubilalis (Hübner)], we found no bioactivity of Cry1Ab protein in senesced corn tissue after 2 wk of exposure to terrestrial or aquatic environments. Third, we show that Bt near-isolines modify growth and survivorship of some species of invertebrates. Of the four nontarget invertebrate species fed Bt near-isolines, growth of two closely related trichopterans was not negatively affected, whereas a tipulid crane fly exhibited reduced growth rates, and an isopod exhibited reduced growth and survivorship on the Cry1Ab near-isoline but not on the stacked Cry1Ab Cry3Bb1 near-isoline. Because of lack of evidence of bioactivity of Bt after 2 wk and because of lack of nontarget effects on the stacked near-isoline, we suggest that tissue-mediated differences, and not the presence of the Cry1Ab protein, caused the different responses among the species. Overall, our results provide evidence that adverse effects to aquatic nontarget shredders involve complex interactions arising from plant genetics and environment that cannot be ascribed to the presence of Cry1Ab proteins.
Nontarget impacts of six transgenic Bt rice lines (expressing the Cry1Ab or Cry1Ab/Cry1Ac protein) on the thrips, Stenchaetothrips biformis (Bagnall), attacking the rice seedling and tillering stages, were evaluated under laboratory and field conditions. Laboratory results showed relatively longer larval, pupal development and preoviposition durations of S. biformis. Although it had a shorter oviposition period, female adult longevity and less total laid eggs were found when fed on some tested Bt rice in comparison to non-Bt controls. S. biformis population dynamics in Bt and non-Bt plots were monitored using the plastic bag and beat plate methods. In the field, the temporal patterns of S. biformis population changes were similar between tested Bt rice lines and their respective control; however, the total number of S. biformis individuals collected from the Bt plots were significantly less or the same, varying from variety to variety, compared with those from the non-Bt plots. ELISA results showed that the Bt insecticidal protein could be transferred from Bt rice to the thrips, and the concentrations of the protein in rice leaves and thrips were not significantly correlated with some important biological parameters of the thrip. In addition, the potential effects of Bt rice on the abundance of S. biformis candidate predators are also discussed. In conclusion, our results show that the six Bt rice lines assessed may be less preferable host plants to S. biformis at the individual and population levels in comparison to the non-Bt rice plants.
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