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Nowadays, in protected vegetable crops, pest management based mainly on biological control represents the most sustainable alternative to pesticide use. The cotton whitefly, Bemisia tabaci, is one of the key pests that negatively impact the yield and quality of such crops in many agricultural systems. The predatory bug Macrolophus pygmaeus is one of the main natural enemies of the whitefly and is widely used for its control. However, the mirid can sometimes behave as a pest itself, causing damage to crops. In this study, we investigated the impact of M. pygmaeus as a plant feeder, by analyzing the combined impact of the whitefly pest and the predator bug on the morphology and physiology of potted eggplants under laboratory conditions. Our results showed no statistical differences between the heights of plants infested by the whitefly or by both insects compared with noninfested control plants. However, indirect chlorophyll content, photosynthetic performance, leaf area, and shoot dry weight were all greatly reduced in plants infested only by B. tabaci, compared with those infested by both pest and predator or with noninfested control plants. Contrarily, root area and dry weight values were more reduced in plants exposed to both of the insect species, compared with those infested only by the whitefly or compared with noninfested control plants, where the latter showed the highest values. These results show how the predator can significantly reduce the negative effects of B. tabaci infestation, limiting the damage it causes to host plants, though the effect of the mirid bug on the underground parts of the eggplant remains unclear. This information might be useful for a better understanding of the role that M. pygmaeus plays in plant growth, as well as for the development of management strategies to successfully control infestations by B. tabaci in cropping environments.

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Despite their importance as biological control agents, zoophytophagous dicyphine mirids can produce economically important damage. We evaluated the phytophagy and potential impact on tomato plants of Dicyphus cerastii and Nesidiocoris tenuis. We developed a study in three parts: (i) a semi-field trial to characterize the type of plant damage produced by these species on caged tomato plants; (ii) a laboratory experiment to assess the effect of fruit ripeness, mirid age, and prey availability on feeding injuries on fruit; and (iii) a laboratory assay to compare the position of both species on either fruit or plants, over time. Both species produced plant damage, however, although both species produced scar punctures on leaves and necrotic patches on petioles, only N. tenuis produced necrotic rings. Both species caused flower abortion at a similar level. Overall, N. tenuis females produced more damage to tomato fruit than D. cerastii. There was an increased frequency of D. cerastii females found on the plants over time, which did not happen with N. tenuis. Our results suggested that, although D. cerastii caused less damage to fruit than N. tenuis, it still fed on them and could cause floral abortion, which requires field evaluation and caution in its use in biological control strategies.

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This study presented biological and economic data for the mass-rearing of Orius minutus in Korea. Simplifying the mass-rearing process through an alternative diet and an artificial oviposition substrate is a prerequisite for enhancing the usability of this insect as a biological control agent. We compare the hatch rate of O. minutus eggs deposited on a plant substrate with that of eggs deposited on two artificial substrates, cork sheets and rubber. The results indicate that cork sheet is the most cost-effective artificial oviposition substrate for the mass-rearing of O. minutus. We also examine five feeding treatments that included two types of brine shrimp eggs and eggs of Ephestia cautella to compare the number of eggs laid in the fifth generation. We found no significant difference between the two treatment groups; 61.3 eggs were laid in the treatment group fed iron-coated brine shrimp and moth eggs, and 67.4 eggs were laid in the control group. The plant-free model developed in our study can reduce rearing costs by 70.5% compared to the conventional mass-rearing model.

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Animals, including herbivores and predators, use diet-mixing to balance their macro- and micronutrient intake. Recent work demonstrated that lady beetles fed only pea aphids from fava beans had reduced fitness caused by a deficiency of dietary sterols. However, beetles redressed this deficit by eating fava bean leaves. In the current study we used Coccinella septempunctata as a model to test the hypotheses that pea aphids are a poor sterol resource independent of their host plant, and that fava beans produce low quality prey regardless of aphid species. Additionally, we tested the reproductive rescue capacity of alfalfa and barley foliage compared to fava, and profiled the sterols of phloem exudates, foliage, and aphids reared on these different hosts. Beetle fecundity and egg viability was significantly better when provided pea aphids reared on alfalfa (compared to fava beans) and green peach aphids reared on fava plants. Alfalfa and barley leaves were not consumed by beetles and did not support beetle reproduction. The sterol profile of aphids largely reflected their host plant phloem. However, green peach aphids from fava acquired 125-times more sterol than pea aphids from fava. Our findings show how the sterol content of different host-plants can affect the third trophic level. Our results suggest that 1) prey quality varies depending on prey species, even when they occur on the same plant, 2) plant species can mediate prey quality, 3) host plant-mediated effects on prey quality partially drive omnivory, and 4) diet-mixing benefits growth and reproduction by redressing micronutrient deficits.

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The generalist predator Euseius stipulatus (Athias-Henriot) and the Tetranychidae-specialist predators Neoseiulus californicus (McGregor) and Phytoseiulus persimilis Athias-Henriot play a key role in the regulation of Tetranychus urticae Koch in Spanish citrus orchards. Previous studies have shown that sour orange (Citrus aurantium L.) and Cleopatra mandarin (Citrus reshni hort. ex Tan.) display extreme resistance and susceptibility to T. urticae, respectively. When offered a choice between these two genotypes infested by T. urticae, E. stipulatus preferred Cleopatra mandarin, whereas the specialists did not show any preference. The present study was undertaken to check whether these preferences could be related to the feeding of E. stipulatus on the host plant and/or to differences in prey feeding on the two plants. Our results demonstrate that E. stipulatus is a zoophytophagous mite, which can engage in direct plant feeding in sour orange and Cleopatra mandarin, whereas neither N. californicus nor P. persimilis do so. Whereas Cleopatra mandarin provided a higher-quality prey/feeding substrate for E. stipulatus, which may be related to its phytophagy, no differences were observed for the two specialists. As higher constitutive and faster inducible defense against T. urticae in sour orange relative to Cleopatra mandarin plants result in sour orange supporting lower T. urticae densities and plant damage, our results demonstrate that pest regulation by specialist natural enemies may be more effective when prey feed on better defended plants.

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Zoophytophagous mirid species, that feed and develop either on prey or plant resources, are often found simultaneously on the same host. Hence, these species can engage in both intraguild predation and cannibalism, which can pose a threat to mirid eggs. Ovipositing females may respond to such risks of predation on their eggs by reducing the number of eggs laid or selecting safer oviposition sites. We tested the oviposition behavior of Macrolophus pygmaeus (Rambur) (Hemiptera: Miridae) females under the risk of cannibalism by M. pygmaeus males and intraguild predation by Nesidiocoris tenuis (Reuter) males (Hemiptera: Miridae) under laboratory conditions. Intraguild predators and cannibals were introduced during or after the oviposition period. The number of eggs laid (using counts of newly hatched nymphs) and their proportion on each part of a tomato plant were both measured. The results reveal that only cannibalism by M. pygmaeus males after the period of oviposition significantly decreased the number of hatched eggs. Cannibalism thus represents a greater risk to mirid eggs than intraguild predation. The M. pygmaeus female responded to the presence of potential intraguild predators (or competitors) by decreasing the number of eggs laid in the upper leaves. The results suggest that M. pygmaeus females avoid competition by N. tenuis, by laying fewer eggs on upper leaves. Cannibalism could regulate zoophytophagous predator populations under prey scarcity conditions and minimize the risk of crop damage associated with those biological control agents.

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While many predatory arthropods consume non-prey foods from lower trophic levels, little is known about what drives the shift from predator to omnivore. Predatory lady beetles often consume non-prey foods like plant foliage and pollen. One species, Coccinella septempunctata, eats foliage to redress sterol deficits caused by eating sterol-deficient prey. Here we explore how omnivory benefits lady beetle fitness. We reared seven species of lady beetles-from five genera distributed across the tribe Coccinellini-on pea aphids in the presence or absence of fava bean foliage; pea aphids have very low sterol content. Foliage supplements lengthened the development times of four species and decreased survival to adulthood of two species; it had no effect on adult mass. We mated beetles in a 2 × 2 factorial design (males with or without foliage paired with females with or without foliage). For each species, we observed a profound paternal effect of foliage supplements on fitness. Females mated to foliage-supplemented males laid more eggs and more viable eggs compared to females mated to non-supplemented males. Foliage-supplemented males produced 2.9-4.6 times more sperm compared to non-supplemented males for the three species we examined. We analysed the sterol profile of four beetle species reared on pea aphids-with or without foliage-and compared their sterol profile to field-collected adults. For two laboratory-reared species, sterols were not detected in adult male beetles, and overall levels were generally low (total ng of sterol/beetle range: 3-33 ng); the exception being Propylea quatuordecimpunctata females (total ng of sterol/beetle range: 50-157 ng). Laboratory-reared lady beetle sterol content was not significantly affected by the presence of foliage. Field-collected beetles had higher levels of sterols compared to laboratory-reared beetles (2,452-145,348 ng per beetle); cholesterol and sitosterol were the dominant sterols in both field-collected and laboratory-reared beetles. Our findings indicate that herbivory benefits lady beetle fitness across the Coccinellini, and that this was entirely a paternal effect. Our data provide a rare example of a nutritional constraint impacting fitness in a sex-specific manner. It also shows, more broadly, how a nutritional constraint can drive predators towards omnivory.

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Brontocoris tabidus Signoret (Heteroptera: Pentatomidae) is a zoophytophagous predator of lepidopteran defoliators of eucalyptus in Brazil. This predator complements its diet with plants, which is fundamental for its population maintenance. The objective was to evaluate the B. tabidus development, reproduction, and life table parameters in the field on Eucalyptus cloeziana F. Muell. (T1), Psidium guajava Linn (Myrtales: Myrtaceae) (T2), or without plants (T3). The parameters estimated were: net reproductive rate (Ro); generation duration (DG); time for the population to double in size (TD); intrinsic population growth rate (rm), survival rate (lX), specific fertility (mx), life expectancy (ex), and mortality risk (qx). Ro, DG, TD, and rm were higher in the T1 and T2 than in T3. Plant presence favored the lx, mx, and qx. Ex values were 36.1 and 56.9 in the T3 and T1, respectively. The B. tabidus fertility, longevity, and life table parameters improvement on E. cloeziana and P. guajava plants are due to the water and nutrients obtained from them. B. tabidus can be reared with Tenebrio molitor Linnaeus (Coleoptara: Tenebrionidae) pupae on E. cloeziana or P. guajava plants in the field.

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The proximate forces that create omnivores out of herbivores and predators have long fascinated ecologists, but the causal reasons for a shift to omnivory are poorly understood. Determining what factors influence changes in trophic position are essential as omnivory plays a central role in theoretical and applied ecology. We used sevenspotted lady beetles (Coccinella septempunctata) to test how prey nutrient content affects beetles' propensity to engage in herbivory. We show that beetles consuming an all-prey diet demonstrate normal growth and development, but suffer a complete loss of fitness (spermatogenic failure) that is restored via herbivory and supplementation with phytosterols and cholesterol. Furthermore, we show that lady beetles possess a state-dependent sterol-specific appetite and redressed their sterol deficit by feeding on foliage. These results demonstrate that predators balance their nutrient intake via herbivory when prey quality is low, and reveal a selective force (sterol nutrition) that drives predatory taxa to omnivory.

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Several studies have reported the importance of phytophagy in the history of life from predator stinkbugs. However, knowledge of the nutrients acquired by these insects, from plant materials, is scarce. The phytophagy studies of zoophytophagous predators benefit biological control programs, making them more effective, since the best resources may be offered to these insects, increasing the potential for predation and, thus obtaining, success in pest control. Thus, this study investigated the development and zoophytophagy of predator Podisus nigrispinus (Dallas) (Hemiptera: Pentatomidae) in eucalyptus seedlings, deficient in macronutrients. The experiment was conducted in a greenhouse and the treatments consisted of individual nitrogen omission, phosphorus, potassium, calcium, magnesium and sulfur nutrient solution, where the eucalyptus seedlings were being conducted. The pupae of Tenebrio molitor L. (Coleoptera: Teneobridae) were offered ad libitum. The results of this experiment demonstrated the importance of nutrition in plants for the development and predation of P. nigrispinus. The plants deficient in nitrogen and magnesium reduce the predation efficiency of this zoophytophagous. The omission of calcium, magnesium and sulfur adversely affected the development of this insect predator, extending its immature stages. The proper nutritional balance of the plants is essential for the success in biological control programs that use zoophytophagous predators.

Diversos estudos têm relatado a importância da fitofagia na história de vida dos percevejos predadores. Porém, não se conhece quais são os nutrientes adquiridos por estes insetos nos materiais vegetais. Sabe-se que a fitofagia realizada pelos predadores zoofitófagos beneficia os programas de controle biológico, tornando-os mais eficazes, e que as plantas oferecem recursos que poderão aumentar o potencial de predação, garantindo o sucesso no controle de pragas. Deste modo, este trabalho investigou o consumo alimentar e desenvolvimento do percevejo Podisus nigrispinus (Dallas) (Hemiptera: Pentatomidae), sob mudas de eucalipto, submetidas à solução nutritiva com omissão de macronutrientes. O experimento foi realizado em casa de vegetação e os tratamentos consistiram na omissão individual de nitrogênio, fósforo, potássio, cálcio, magnésio e enxofre da solução nutritiva, onde as mudas de eucalipto se desenvolveram. Pupas de Tenebrio molitor L. (Coleoptera: Teneobridae), foram oferecidas ad libitum. Os resultados deste experimento comprovaram a importância da complementação nutricional em plantas para o consumo alimentar e desenvolvimento do P. nigrispinus. A alimentação nas plantas em que o nitrogênio e magnésio foram omitidos diminuiu o consumo de biomassa animal das presas deste zoofitófago. E a omissão de cálcio, magnésio e enxofre afetou, negativamente, o desenvolvimento deste percevejo predador, prolongando seu estádio imaturo. O balanço nutricional adequado das plantas é fundamental para obter sucesso em programas de controle biológico em que se utilizam predadores zoofitófagos.

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Tuta absoluta (Meyrick) has quickly developed into a significant tomato pest worldwide. While the recently found mirid predators Macrolophus basicornis (Stal), Engytatus varians (Distant) and Campyloneuropsis infumatus (Carvalho) of this pest are able to establish and reproduce on tomato, biological knowledge of these mirids is still limited. Here we describe the functional response of the three mirid predators of the tomato pest T. absoluta when offered a range of prey densities (four, eight, 16, 32, 64, 128 and 256 eggs) during a 24 h period inside cylindrical plastic cages in the laboratory. Engytatus varians and M. basicornis showed a type III functional response, whereas C. infumatus showed a type II functional response. At the highest prey densities, C. infumatus consumed an average of 51.0 eggs, E. varians 91.1 eggs, and M. basicornis 100.8 eggs. Taking all information into account that we have collected of these three Neotropical mirid species, we predict that M. basicornis might be the best candidate for control of the tomato borer in Brazil: it has the highest fecundity, the largest maximum predation capacity, and it reacts in a density-dependent way to the widest prey range.

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Podisus distinctus (Hemiptera: Pentatomidae) is a zoophytophagous insect with significant potential for use as a biological control agent in agriculture and forestry because their nymphs and adults actively prey on diverse insect species. The saliva of this insect possesses active substances that cause paralysis and death of the prey. As the first step in identifying compounds of P. distinctus saliva, this study describes the ultrastructure of the salivary glands of this predator. The salivary system of P. distinctus possesses a pair of main salivary glands with a short anterior lobe, a long posterior lobe, and a pair of tubular accessory glands. The main salivary gland of P. distinctus has no associated muscles, suggesting that the saliva-release mechanism occurs with the help of certain thorax muscles. The main salivary gland epithelium has a single layer of cells (varying from cubical to columnar) with cytoplasm rich in rough endoplasmic reticulum, spherical granules of different sizes, a nucleus with a predominance of decondensed chromatin, and nucleolus. The apical cell region has a few short microvilli and the basal region has plasma membrane infoldings. The epithelium of the accessory salivary glands possesses a single-layered epithelium of cubic cells delimiting a narrow lumen. The apical cell region has a high density of microvilli and pleomorphic mitochondria, whereas the central cell region is rich in rough endoplasmic reticulum with a well-developed nucleus and decondensed chromatin. The basal cell region is characterized by the presence of several basal plasma membrane infoldings associated with mitochondria and numerous openings to the hemocoel forming large channels. The ultrastructural characteristics suggest that the main salivary glands and accessory salivary glands play a vital role in protein synthesis for saliva production and that the accessory glands are involved in transport of materials of the hemolymph.

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Effect of gossypol on survival and reproduction of the zoophytophagous stinkbug Podisus nigrispinus (Dallas). Gossypol is a sesquiterpene aldehyde found in cotton plants conferring resistance against herbivory. Although the effect of this sesquiterpenoid on insect pests of cotton is known, the interaction of this compound with zoophytophagous predators such as Podisus nigrispinus (Dallas) (Hemiptera, Pentatomidae) has not been studied so far. Thus, the objective of this study was to evaluate the effect of the purified gossypol on nymphs and adults of P. nigrispinus. Nymphs and adults of this predator were fed on Tenebrio molitor pupae and supplemented with solutions of gossypol at concentrations of 0.00, 0.05, 0.10, and 0.20 percent (w/v) during the nymphal and adult stages or, only during the adult stage of P. nigrispinus. The nymphal stage of the predator was, on average, two days longer when suplemmented with gossypol. Emerged adults had lower fecundity and egg hatching, especially at the highest gossypol concentration (0.20 percent) ingested during the nymphal and adult stages. However, this predator was not affected when it ingested the compound only during the adult stage. P. nigrispinus can have delayed nymphal development and lower reproductive performance when ingesting the gossypol during the nymphal and adult stages, but only at higher concentrations of gossypol than that produced by cotton plants.

Efeito do gossipol na sobrevivência e reprodução do percevejo zoofitófago Podisus nigrispinus (Dallas). O gossipol é um aldeído sesquiterpeno produzido pelo algodoeiro que confere resistência contra a herbivoria. A interação deste sesquiterpeno com predadores zoofitófagos, como Podisus nigrispinus (Dallas) (Hemiptera, Pentatomidae), é inexistente apesar do reconhecido efeito do gossipol sobre insetos pragas do algodoeiro. Assim, este estudo avaliou o efeito do extrato de gossipol sobre ninfas e adultos de P. nigrispinus. O predador foi alimentado com pupas de Tenebrio molitor L. (Coleoptera, Tenebrionidae) e suplementado com soluções do gossipol nas concentrações 0, 0,05, 0,10 e 0,20 por cento (peso/volume) durante os estágios ninfal e adulto e somente durante o estágio adulto. A duração do estágio ninfal do predador foi em média dois dias mais longo quando suplementado com gossipol. Adultos oriundos de ninfas suplementadas com gossipol apresentaram menor fecundidade e viabilidade dos ovos, em especial na maior concentração (0,20 por cento). Contudo, adultos não foram afetados quando suplementados com gossipol somente durante este estágio. Os resultados demonstram que a ingestão prolongada do gossipol ocasiona efeito no desenvolvimento e reprodução do predador P. nigrispinus, porém somente em concentração superior àquela produzida naturalmente pela planta de algodão.

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Este trabalho investigou o efeito de seis plantas daninhas [Ageratum conyzoides L., Amaranthus hybridus L., Bidens pilosa L., Desmodium tortuosum (Sw.), Euphorbia heterophylla L., Ricinus communis L.] e do algodoeiro Gossypium hirsutum L cv. CNPA Precoce 1, na biologia de Podisus nigrispinus (Dallas). Ovosdo predador foram fixados às plantas e, posteriormente, os insetos foram alimentados com pupas de Tenebrio molitor (L.) (Coleoptera: Tenebrionidae) até a morte. Para avaliar a preferência para oviposição, fêmeas acasaladas foram confinadas em gaiolas contendo plantas daninhas e algodoeiro, observando-se a seleção da planta para oviposição através do número de posturas e ovos. D. tortuosum propiciou o mais rápido desenvolvimento ninfal de P. nigrispinus. A viabilidade da fase ninfal não foi afetada em relação à espécie de planta e fêmeas produzidas em R. communis foram mais leves em relação às produzidas em G. hirsutum. Fêmeas mantidas em A. conyzoides tiveram maior fecundidade, enquanto aquelas mantidas em D. tortuosum, E. heterophylla e R. communis apresentaram a menor fecundidade. A taxa líquida de reprodução (Ro) e o tempo médio da geração (T) de P. nigrispinus foram maiores em A. conyzoides e menores em R. communis. A taxa intrínseca de crescimento populacional (r m) e a razão finita de crescimento (lambda) foram maiores em B. pilosa e menores em G. hirsutum. Dentre as plantas estudadas P. nigrispinus preferiu ovipositar em A. hybridus e B. pilosa. Os resultados sugerem que plantas daninhas como A. conyzoides, B. pilosa e A. hybridus podem beneficiar a colonização e manutenção de P. nigrispinus no agroecossistema algodoeiro por fitofagia.

This study investigated the effects of six weeds [Amaranthus hybridus L., Desmodium tortuosum (Sw.), Euphorbia heterophylla L., Ageratum conyzoides L., Bidens pilosa L., Ricinus communis L.] and the cotton plant Gossypium hirsutum L. CNPA Precocious 1 on Podisus nigrispinus (Dallas) life history characteristics and oviposition behavior. The study begun with P. nigrispinus eggs placed on plants and caged throughout up to female death. Egg production and selected plant for oviposition by P. nigrispinus were investigated caging mated females on weeds and cotton plants simultaneously. Nymphal development of P. nigrispinus was shorter on D. tortuosum and longer on A. hybridus. Nymphal viability was not affected by any species of plant studied. Nymphs reared on R. communis produced females weighting less compared to the other plant species. The weed A. conyzoides promoted higher female fecundity compared to those females reared on D. tortuosum, E. heterophylla and R. communis. The demographic parameters net reproductive rate (Ro) and mean generation time (T) of P. nigrispinus were higher for females reared on A. conyzoides, while the intrinsic rate of population increase (r m) and finite rate of increase (lambda) suggest that better performance of the predator will be yielded on B. pilosa. The weed plants A. hybridus and B. pilosa were significantly preferred by P. nigrispinus females to lay eggs. These results suggest that the weeds A. conyzoides, B. pilosa and A. hybridus can benefit the colonization and maintenance of P. nigrispinus in the cotton ecosystem by phytophagy.