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In agricultural settings, examples of effective control strategies using repellent chemicals in integrated pest management (IPM) are relatively scarce compared to those using attractants. This may be partly due to a poor understanding of how repellents affect insect behavior once they are deployed. Here we attempt to identify potential hallmarks of repellent stimuli that are robust enough for practical use in the field. We explore the literature for success stories using repellents in IPM and we investigate the mechanisms of repellency for two chemical oviposition deterrents for controlling Drosophila suzukii Matsumura, a serious pest of small fruit crops. Drosophila suzukii causes injury by laying her eggs in ripening fruit and resulting larvae make fruit unmarketable. In caged choice tests, reduced oviposition was observed in red raspberry fruit treated with volatile 1-octen-3-ol and geosmin at two initial concentrations (10% and 1%) compared to untreated controls. We used video monitoring to observe fly behavior in these caged choice tests and investigate the mode of action for deterrence through the entire behavioral repertoire leading to oviposition. We observed fewer visitors and more time elapsed before flies first landed on 1-octen-3-ol-treated fruits than control fruits and concluded that this odor primarily inhibits behaviors that occur before D. suzukii comes in contact with a potential oviposition substrate (precontact). We observed some qualitative differences in precontact behavior of flies around geosmin-treated fruits; however, we concluded that this odor primarily inhibits behaviors that occur after D. suzukii comes in contact with treated fruits (postcontact). Field trials found reduced oviposition in red raspberry treated with 1-octen-3-ol and a combination of 1-octen-3-ol and geosmin, but no effect of geosmin alone. Recommendations for further study of repellents for practical use in the field are discussed.
For all species, abiotic factors directly affect performance, survival and reproduction, and consequently, their geographic distribution. Species distribution models (SDMs) are important tools to predict the influence of abiotic factors in species distributions and has been more applied over the years. However, these models can be built under different algorithms and using different methods to select environmental predictors, which can lead to different results. Five different algorithms and two sets of environmental predictors were compared to predict the geographic distribution of the blowfly Sarconesia chlorogaster (Wiedemann) (Diptera: Calliphoridae). This species has several occurrence points and a considerable amount of biological data available, which makes S. chlorogaster a good model system to compare environmental predictors. Two sets of environmental predictors (mainly derived from temperature and humidity) were built, and the set based on the influence of abiotic variables on the ecophysiology of S. chlorogaster showed better results than the principal component analysis (PCA) approach using 19 climatic variables. We also employed five modeling algorithms—Envelope Score, Mahalanobis Distance, GARP, Support Vector Machines, and Maxent—and the latter two showed the best performances. The results indicate that temperature is the main factor shaping geographic distribution of S. chlorogaster through its effect on fitness. Furthermore, we showed that this species is mainly distributed in south, southeastern, and some northwestern and southwestern sites of South America. In addition, our results also predicted suitable areas in Ecuador and Colombia, countries without previous records.
Succession in local dung beetle assemblages influences their delivery of ecological functions in natural and modified environments globally. Short-term changes in dung beetle (Coleoptera: Scarabaeidae) species richness, abundance, and biomass were investigated in standardized dung pads in northern, coastal Turkey. For mean tunneling guild abundance, dung deposition time, dung exposure period, and their interaction were significant, and for mean dung dwelling guild abundance, dung exposure period was significant, as was the interaction with dung deposition time, which collectively evidenced temporal resource partitioning, based principally on differences in diel activity. Succession was highly compressed, with maximum abundance at 12 h and maximum species richness at 24 h. A large ball roller and small- to medium-sized tunnelers dominated different periods in the first 24 h but were superseded by dwellers. Regression analysis demonstrated a significant, positive relationship between species richness and the evenness of abundance for both dung deposition times. Correlation analysis generally showed strong, positive correlations between tunneling species, low correlations between tunneling and dwelling species, and low correlations between dwelling species. Niche partitioning based on size difference appears to have acted on the environmental filtering of tunneling species along the temporal gradient of declining moisture, thereby limiting the number of abundant, concurrent species. The aggregation of tunneling species provided opportunities for the less competitive dwelling species to occupy less densely populated zones termed probability refuges. The network of strong, positive correlations between tunneling species may indicate that their collective functionality is vulnerable to loss of efficiency if species are lost.
Phenotypic plasticity involves adaptive responses to predictable environmental fluctuations and may promote evolutionary change. We studied the regulation of phenotypic plasticity in an important agricultural pollinator, the solitary alfalfa leafcutting bee (Megachile rotundata F.). Specifically, we investigated how larval nutrition affects M. rotundata diapause plasticity and how diapause plasticity affects adult female reproductive behavior. Field surveys and laboratory manipulations of aspects of larval diet demonstrated nutritional regulation of M. rotundata diapause plasticity. Manipulation of larval diet quality through the addition of royal jelly, the caste-determining substance of the honey bee Apis mellifera L., increased the probability of diapause in M. rotundata. We also found that larval nutrition and diapause status affected M. rotundata adult female reproductive behavior. Nutritional effects on larval diapause that also impact adult fitness have intriguing implications for the evolution of developmental plasticity in bees. In particular, as the solitary lifestyle of M. rotundata is considered to be the ancestral condition in bees, nutritionally regulated plasticity may have been an ancestral condition in all bees that facilitated the evolution of other forms of phenotypic plasticity, such as the castes of social bees.
Plant domestication can modify and weaken defensive chemical traits, reducing chemical defenses in plants and consequently their resistance against pests. We characterized and quantified the major defensive flavonols and isoflavonoids present in both wild and cultivated murtilla plants (Ugni molinae Turcz), established in a common garden. We examined their effects on the larvae of Chilesia rudis (Butler) (Lepidoptera: Arctiidae). Insect community and diversity indices were also evaluated. We hypothesized that domestication reduces flavonoid contents and modifies C. rudis preference, the insect community, and diversity. Methanolic extracts were obtained from leaves of U. molinae plants and analyzed by high performance liquid chromatography. Results showed higher insect numbers (86.48%) and damage index (1.72 ± 0.16) in cultivated plants. Four new first records of insects were found associated with U. molinae. Diversity indices, such as Simpson, Shannon, and Margalef, were higher in cultivated plants than in wild plants. Furthermore, eight isoflavonoids were identified in U. molinae leaves for the first time. The five flavonols showed higher concentrations in wild U. molinae leaves (89.8 μg/g) than in cultivated plants (75.2 μg/g); however, no differences were found in isoflavonoids between wild and cultivated plants. The larvae of C. rudis consumed more leaf material of cultivated plants than wild plants in choice (3.8 vs. 0.8 mm2) and no-choice (7.5 vs. 3.0 mm2) assays. Our study demonstrates that domestication in U. molinae reduces the amount of flavonoids in leaves, increasing the preference of C. rudis and the insect community.
In the Asian citrus psyllid (Diaphorina citri Kuwayama), learning facilitates host recognition and mate preference; however, it is unclear whether induced female oviposition preference occurs in this species. We investigated the influence of natal host experience on adult oviposition preference when reared on either ‘Valencia’ orange (Citrus x sinesis) or orange jasmine (Murraya paniculata). Psyllids reared on ‘Valencia’ orange preferred ‘Valencia’ orange as an oviposition host compared with orange jasmine, whereas there was no difference in oviposition between the two hosts in orange jasmine reared psyllids. Nymphs transferred from ‘Valencia’ orange to orange jasmine were smaller in adult size and required more time for development. These findings were reversed in orange jasmine reared psyllids, which increased in size and displayed shorter development times when transferred from orange jasmine to ‘Valencia’ orange. However, mortality increased in nymphs transferred to the non-natal host species in both treatment groups compared with nymphs transferred to the same host. These results indicate an association between host plant preference and performance in this species. Maternal host experience appeared to influence the oviposition preference in this species. Juvenile psyllid performance appeared negatively affected by orange jasmine plants such that fitness was reduced, suggesting benefits for maternal host fidelity in those insects not acclimated to feeding on orange jasmine. Induced oviposition preference may provide an important mechanism of adaptive plasticity in D. citri reproductive strategies, allowing females to discriminate among potential host species in favor of those to which her offspring are best adapted.
The population of monarch butterflies east of the Rocky Mountains has experienced a significant decline over the past 20 yr. In order to increase monarch numbers in the breeding range, habitat restoration that includes planting milkweed plants is essential. Milkweeds in the genus Asclepias and Cynanchum are the only host plants for larval monarch butterflies in North America, but larval performance and survival across nine milkweeds native to the Midwest is not well documented. We examined development and survival of monarchs from first-instar larval stages to adulthood on nine milkweed species native to Iowa. The milkweeds included Asclepias exaltata (poke milkweed) (Gentianales: Apocynaceae), Asclepias hirtella (tall green milkweed) (Gentianales: Apocynaceae), Asclepias incarnata (swamp milkweed) (Gentianales: Apocynaceae), Asclepias speciosa (showy milkweed) (Gentianales: Apocynaceae), Asclepias sullivantii (prairie milkweed) (Gentianales: Apocynaceae), Asclepias syriaca (common milkweed) (Gentianales: Apocynaceae), Asclepias tuberosa (butterfly milkweed) (Gentianales: Apocynaceae), Asclepias verticillata (whorled milkweed) (Gentianales: Apocynaceae), and Cynanchum laeve (honey vine milkweed) (Gentianales: Apocynaceae). In greenhouse experiments, fewer larvae that fed on Asclepias hirtella and Asclepias sullivantii reached adulthood compared with larvae that fed on the other milkweed species. Monarch pupal width and adult dry mass differed among milkweeds, but larval duration (days), pupal duration (days), pupal mass, pupal length, and adult wet mass were not significantly different. Both the absolute and relative adult lipids were different among milkweed treatments; these differences are not fully explained by differences in adult dry mass. Monarch butterflies can survive on all nine milkweed species, but the expected survival probability varied from 30 to 75% among the nine milkweed species.
Spotted wing Drosophila, Drosophila suzukii Matsumura (Diptera: Drosophilidae), is an invasive vinegar fly that has become a primary direct pest of berry crops worldwide. We conducted 2 yr of behavioral studies in blueberry plantings to determine how fly activity varied throughout the day. Observations of diurnal activity of adult D. suzukii found the greatest activity in the morning hours between 0600 and 0800 hours, when the majority of flies were on the berries. Flies were also active in the evening mainly between 1800 and 2000 hours; however, this trend was more prominent in 2015, which had cooler and more humid evenings. Experiments examining the effect of irrigation on D. suzukii behavior showed that flies remained active during and after irrigation.The effect of insecticide treatments alone and in combination with irrigation revealed that treatment with spinosad had limited effects on the number of flies per bush, whereas spinetoram reduced the number flying and on the bushes in some cases. Zeta-cypermethrin caused longer and more consistent reduction in D. suzukii flying and on bushes. In all treatments, we observed surviving flies flying near and on treated bushes, indicating that these insecticides do not completely deter fly activity. Irrigation did not influence the effects of zeta-cypermethrin on fly behavior during daily observations up to 3 d after application. Our results highlight that the diurnal patterns of activity of D. suzukii on host plants are flexible and are relatively unaffected by irrigation or insecticide applications.
Herbivore natural enemies base their foraging decision on information cues from different trophic levels but mainly from plant odors. However, the second trophic level (i.e., the herbivorous prey) may also provide reliable infochemical cues for their natural enemies. We have evaluated the role of the aggregation pheromone from Frankliniella occidentalis (Pergande) as a potential kairomone for its natural enemy, the predatory bug Orius laevigatus (Fieber). For this purpose, we have analyzed the response of O. laevigatus to (R)-lavandulyl acetate and neryl (S)-2-methylbutanoate, the two major components of the thrips aggregation pheromone. These compounds have been offered to O. laevigatus adult females and nymphs of the predatory bugs both in separate and as specific (1:1 or 1:2.3) blends, in experiments involving a dual choice Y-tube olfactometer. None of the compounds attracted adults or nymphs when they were individually supplied. Conversely, they were significantly attracted to both adults and nymphs when offered as a blend. A 1:2.3 (R)-lavandulyl acetate:neryl (S)-2-methylbutanoate blend was attractive to both nymphs and adults, while a 1:1 blend elicited response only in nymphs. These results suggest that specific blends of these compounds from the aggregation pheromone may be used as an attractant to O. laevigatus. The results of this work highlight the importance of studying olfactory responses of natural enemies for a better understanding of their foraging behavior. Potential uses of these results in future studies are discussed.
Knowledge about which bark and ambrosia beetle species are active and at what heights in black walnut canopies is not well understood. Neither is the role of these beetles in spreading Thousand Cankers Disease. To assist with future planned research, which will assess the extent to which these beetle species are associated with Geosmithia morbida Kolařík, Freeland, Utley, and Tisserat (Ascomycota: Hypocreales: Bionectriaceae), experiments were undertaken to monitor bark and ambrosia beetles in urban landscapes and parks in Tennessee between 2011 and 2013. Within mature walnut tree canopies, sticky panel, modified soda bottle, and Lindgren traps were deployed at different heights, with and without ethanol as an attractant and with and without walnut stem sections, or in situ limbs that had been girdled or injection with ethanol to simulate stressed tree tissues. Bark and ambrosia beetle species (Coleoptera: Curculionidae: Scolytinae) collected in greatest abundance included Ambrosiodmus rubricollis (Eichhoff), Ambrosiophilus atratus (Eichhoff), Cnestus mutilatus (Blandford), Dryoxylon onoharaense (Murayama), Euwallacea validus (Eichhoff), Monarthrum fasciatum (Say), Monarthrum mali (Fitch), Xyleborinus saxesenii (Ratzeburg), Xyleborus affinis Eichhoff, Xyleborus ferrugineus (Fabricius), Xylosandrus crassiusculus (Motschulsky), and Xylosandrus germanus (Blandford). C. mutilatus, X. saxesenii, and X. crassiusculus were more active higher in trees than most other species and were strongly attracted to ethanol via all means of lure deployment. C. mutilatus, which were captured from April through October and increased in abundance across the 3-yr study, were most abundant in late May with a second activity period in late August.
Stable flies (Stomoxys calcitrans (L.)) can be a serious pest associated with cattle facilities. In Denmark, they occur most abundantly at organic farms, where they cannot be controlled by means of insecticides. On traditional farms, where chemical control is widely used, development of resistance is of increasing concern. Therefore, interest in biological control or other alternative methods has been growing during the recent years. In order to understand the complex relationships between a pest and its natural enemies in a variable environment, it is necessary to know how temperature affects the dynamics of the involved species. In this paper, we apply data derived from several existing sources to investigate the influence of temperature on development and survival of eggs, larvae, pupae, and adult stable flies, as well as on the fecundity of adult females. We demonstrate that the same modeling framework (called SANDY), previously applied to lifetable data of the pteromalid pupal parasitoid (Spalangia cameroni Perkins), a biological control agent used against stable flies, can also be used to model S. calcitrans. However, the predicted temperature responses depend on the data sources used to parameterize the model, which is reflected by differences in estimated population growth rates obtained from American and non-American studies. Elasticity analysis shows that growth rates are more sensitive to changes in viability, in particular of adult flies, than in fecundity, which may have implications for the management of stable fly populations.
Solar ultraviolet-B (UVB) radiation and radiant heat have lethal effects on plant-dwelling mites, including spider mites, and their natural enemies, such as phytoseiid mites, leading them to reside on lower leaf surfaces. Panonychus spider mites are outcompeted by Tetranychus spider mites and thus exploit upper leaf surfaces, where they are exposed to both UVB radiation and radiant heat. Panonychus spider mites are thought to produce astaxanthin constitutionally. In this study, we compared carotenoid components, antioxidant capacity, lipid peroxidation, survival, and egg production in wild-type (WTS) and albino-type strains (ATS) of Panonychus citri (McGregor). Four carotenoids (neoxanthin, violaxanthin, lutein, and carotene) and their isomers and esters were identified in both strains, but astaxanthin and its esters were present only in WTS. The singlet oxygen scavenging capacity of lipid-soluble ingredients was greater in WTS than in ATS, whereas the oxygen radical absorbance capacities of hydrophilic ingredients were equivalent between them. Lipid peroxide accumulation was clearly higher in ATS than in WTS under both UVB irradiation (25 °C) and high temperature (35 °C) conditions. The findings are consistent with an antioxidant protective function of astaxanthin in this mite. Survival periods at 38 °C were longer in WTS than in ATS, although no difference was shown at 35 °C or under UVB irradiation. Therefore, astaxanthin accumulation was shown to be a major mechanism for survival under radiant heat, although other mechanisms, such as photoreactivation, might play a major role in survival under UVB radiation.
Temperature is one of the most important abiotic factors that influences the development and growth of insects. We investigated the effect of temperature on the development of the weevil Eucryptorrhynchus brandti (Harold). The developmental zeros and effective accumulated temperatures for different developmental stages of E. brandti were estimated under variable temperatures in the laboratory. We observed that the developmental zeros of eggs, larvae, and pupae were 15.27 °C, 20.86 °C, and 14.04 °C, respectively, and the effective accumulated temperatures were 81.46 °Cd, 293.76 °Cd, and 127.76 °Cd, respectively. The rate of development increased as the temperature increased within a particular temperature range, and temperatures from 30 °C to 31 °C were particularly suitable for E. brandti rearing. Based on these results, we may be able to predict the occurrence of E. brandti, providing a reference for physical control measures targeting adults to effectively reduce oviposition and minimize population expansion.
Temperature causes physiological and behavioral changes in egg parasitism. Egg parasitoids are a possible alternative for the control of Lepidoptera pests. The cotton bollworm Helicoverpa armigera (Hübner) is a generalist pest that has recently been introduced into Brazil. We evaluated the suitability of H. armigera eggs as hosts for Trichogramma pretiosum Riley and the effects of temperature (18–33°C) on its survival and development. We assessed the developmental time (egg adult), thermal requirements, age-specific fertility, and parasitism rate upon eggs of H. armigera at six different temperatures. Linear and nonlinear (Brière2) models were fitted to our data in order to describe the relationship between developmental rate and temperature for the immature stages of T. pretiosum and estimate the thermal constant (K), lower developmental temperature threshold (LT), optimum temperature for development (Topt), and upper developmental temperature threshold (Tmax). The estimated K and LT estimated by linear model were 130.38 degree days and 10.3°C, respectively. The T0, Topt, and Tmax by Brière model were 11.52, 32.89, and 41.1°C, respectively. Females of T. pretiosum developed at each temperature were assessed, and parasitism rate was above 75% on the first day, except at 30°C where it took 2 d to reach 75%. At 33°C, the highest rate of parasitism was 38%. The life table parameters predict population increases across the temperature range evaluated. The results demonstrated that T. pretiosum are suitable for the control of H. armigera, and the best performance will be achieved when used in regions with temperatures up to 30°C.
Winter mortality is expected to be a key factor determining the ability of mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), to expand its range in Canada. We determined the mortality rate and supercooling points of eggs from the beetle's historic range in southern British Columbia as well as the recently expanded range in north-central Alberta and tested if eggs require an extended period of chilling to reach their maximum cold tolerance. We found no effect of population source or acclimation time on egg cold tolerance. Although 50% of eggs can survive brief exposure to –20.5 °C (LT50), storage at 0.3 °C and –7.5 °C for 59 d resulted in 50% and 100% mortality, respectively. Our results indicate that eggs suffer significant prefreeze mortality and are not well-adapted to overwintering: eggs are unlikely to survive winter throughout much of the beetle's range. Our results provide information that can be used to help model the climatic suitability of mountain pine beetle, including how changes in seasonality associated with new or changing climates may affect winter survival. In addition to lower lethal temperatures, it is critical that the duration of exposure to sublethal cold temperatures are considered in a comprehensive index of cold tolerance and incorporated into survival and population models.
Harmonia axyridis (Pallas) is a common and abundant predator in China and may be exposed to Cry toxins that are produced in Bt crops either by feeding on plant parts or by feeding on target or nontarget herbivorous insects. A new Bt maize line, expressing the Cry1Ab/Cry2Aj fused protein, has been developed and should be rigorously assessed for the ecological risks on the natural enemy. Laboratory experiments were carried out to study the effects of this Bt maize on nontarget predator H. axyridis via bitrophic interaction of adult H. axyridis feeding on Bt maize pollen and tritrophic interaction of H. axyridis consuming the lepidopteran prey. Spodoptera exigua (Hübner) neonate larvae were used to transfer Bt protein because they could survive after ingesting transgenic cry1Ab/cry2Aj maize kernels in the previous study. ELISA bioassays confirmed that the Bt protein could be transferred, but diluted through Bt maize–prey–predator. Life history parameters such as survival, development, weight, fecundity, and egg hatching rate were not significantly different when H. axyridis consumed prey that had been reared on Bt maize compared with prey reared on a nontransformed parental control. Furthermore, feeding directly on Bt maize pollen also had no detrimental effects on fitness, survival, and weight of female and male adults. In conclusion, our results indicate that transgenic cry1Ab/cry2Aj maize poses no ecological risks on the nontarget predator H. axyridis.
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