First trials exploring the potential of phytoseiid mites in managing lychee erinose mite, Aceria litchii (Keifer) (Acari: Eriophyidae), infestations on lychee plants.

Phytoseiid mites have been frequently found in association with the lychee erinose mite, Aceria litchii, on lychee plants in Brazil, suggesting that they are promising candidates as biological control agents against this pest. Here, we investigated whether phytoseiids would suppress A. litchii infestation, i.e. formation of erinea, on lychee plants under field conditions. Four groups of A. litchii-infested plants were randomly distributed in the field, with each group receiving either Phytoseius intermedius, Amblyseius herbicolus, A. herbicolus supplemented with cattail pollen or no predator. During a three-month period, the released predators, along with others present in the surrounding environment, were allowed to freely walk among all plants. In each plant, we evaluated the occurrence of phytoseiid species, their abundance, and the dynamics of erinea formation. A total of 2,097 mites, including 13 other phytoseiid species were identified. The most abundant species were Iphiseiodes zuluagai and Euseius ho, rather than the two predator species that were released. A. herbicolus and P. intermedius failed to establish populations in the majority of the plants, regardless of the presence of pollen, suggesting their ineffectiveness in controlling A. litchii infestations. While there was a significant difference in the proportion of erinea among the four treatments, this contrast was not associated with the presence of phytoseiids, suggesting that other factors might have hindered erinea formation on lychee plants. The reasons behind this outcome are further explored and discussed.

Imperfect diet choice reduces the performance of a predatory mite.

Two mutually unexclusive hypotheses prevail in the theory of nutritional ecology: the balanced diet hypothesis states that consumers feed on different food items because they have complementary nutrient and energy compositions. The toxin-dilution hypothesis poses that consumers feed on different food items to dilute the toxins present in each. Both predict that consumers should not feed on low-quality food when ample high-quality food forming a complete diet is present. We investigated the diet choice of Phytoseiulus persimilis, a predatory mite of web-producing spider mites. It can develop and reproduce on single prey species, for example the spider mite Tetranychus urticae. A closely related prey, T. evansi, is of notorious bad quality for P. persimilis and other predator species. We show that juvenile predators feeding on this prey have low survival and do not develop into adults. Adults stop reproducing and have increased mortality when feeding on it. Feeding on a mixed diet of the two prey decreases predator performance, but short-term effects of feeding on the low-quality prey can be partially reversed by subsequently feeding on the high-quality prey. Yet, predators consume low-quality prey in the presence of high-quality prey, which is in disagreement with both hypotheses. We suggest that it is perhaps not the instantaneous reproduction on single prey or mixtures of prey that matters for the fitness of predators, but that it is the overall reproduction by a female and her offspring on an ephemeral prey patch, which may be increased by including inferior prey in their diet.

Arthropods associated with the lychee erinose mite, Aceria litchii (Acari: Eriophyidae) on lychee trees in Minas Gerais, Brazil.

Lychee fruit production (Litchi chinensis) has been threatened in many regions of the world where the presence has been reported of the litchii erinose mite, Aceria litchii (Keifer). This study aims to identify the arthropod community associated with A. litchii on lychee plants in Minas Gerais, Brazil, and to seek for potential natural enemies associated with this mite pest in this region. We sampled lychee leaves infested with A. litchii in commercial and non-commercial lychee orchards during seven consecutive months, covering the dry and wet seasons of the year. Arthropods found in association with A. litchii on lychee leaves were collected and identified. The results indicate that a great diversity and abundance of mites and hexapods are associated with A. litchii. A total of 985 and 1872 specimens of mites were identified in commercial and non-commercial areas, respectively, belonging to the families Cheyletidae, Cunaxidae, Eupodidae, Iolinidae, Stigmaeidae, Phytoseiidae, Tarsonemidae, Tenuipalpidae, Tetranychidae, Tryophtydeidae, Tuckerellidae, Tydeidae, Winterschmidtiidae and Xenocaligonellidae and the suborder Oribatida. Among them, Phytoseiidae was the most abundant and diverse family with a total of 11 species identified, in which Phytoseius intermedius was the most abundant predatory mite species collected. Minor specimens of hexapods were also been, belonging to the orders Collembola, Blattodea, Coleoptera, Diptera, Hemiptera, Hymenoptera, Psocoptera and Thysanoptera. The presence of a high community of predatory mites in association with A. litchii deserves attention and our results indicate that studies to test the potential of these species and the adoption of management practices that enhance this ecological service must be carried out to achieve satisfactory control of the lychee erinose mite in lychee plants.

Estimating intrinsic growth rates of arthropods from partial life tables using predatory mites as examples.

The intrinsic rate of natural increase of a population (rm) has been in focus as a key parameter in entomology and acarology. It is considered especially important in studies of predators that are potential biological control agents of fast-growing pests such as mites, whiteflies and thrips. Life-table experiments under controlled laboratory conditions are standard procedures to estimate rm. However, such experiments are often time consuming and may critically depend on the precise assessment of the developmental time and the fecundity rate early in the reproductive phase. Using selected studies of predatory mites with suitable life-table data, we investigated whether and how measurements of growth rates can be simplified. We propose a new method for estimating rm from partial life tables, in which the researcher can choose a level of precision based on a stand-in measure of relative error. Based on this choice, the procedure helps the researcher to decide when a life-table experiment can be terminated. Depending on the chosen precision, significant amounts of experimental time can be saved without seriously compromising the reliability of the estimated growth parameter.

Compatibility of two predator species for biological control of the two-spotted spider mite.

Because predators may interfere with each other, an important step towards the implementation of successful release of multiple predators in biocontrol programs requires resolving how predators respond to the presence of heterospecific competitors. Several species of predatory mites are important biocontrol agents and the species Phytoseiulus macropilis and Neoseiulus californicus are used to control the two-spotted spider mite, Tetranychus urticae, an important pest in agriculture worldwide. We investigated their compatibility showing that the two predators do not avoid plants on which the other species is present together with their common prey, and demonstrated that their oviposition rates are not affected by the presence of the other species. However, the distribution of the eggs on leaf discs was affected by the presence of the heterospecific predator. This behaviour might weaken possible interference between these two biocontrol agents, which, in turn, may enable their persistence on plants and favour pest suppression. The increased joint use of several natural enemies for biological control highlights the importance of studies on predator-predator interactions.

High-quality alternative food reduces cannibalism in the predatory mite Amblyseius herbicolus (Acari: Phytoseiidae).

Predatory mites of the Phytoseiidae family are important biological control agents. Many species of this family are omnivores, i.e., besides on prey, they can feed on plant resources such as nectar and pollen. It has been shown that the addition of alternative food for predators to a crop enhances biological control. However, factors such as food availability and quality can also affect interactions such as cannibalism, and thus influence biological control. We investigated the role of quality of the alternative food in the tendency of Amblyseius herbicolus to engage in cannibalism, a common ecological interaction in many phytoseiid mite species. Cannibalism on eggs by A. herbicolus was significantly reduced in the presence of high-quality food (cattail pollen) compared to egg cannibalism without alternative food, whereas this was not the case in the presence of low-quality food (cotton pollen). This suggests that cattail pollen is a high-quality alternative food, not only because it results in increased development and reproduction of predators, but also because it can minimize cannibalism.

Ontogenetic stage-specific reciprocal intraguild predation.

The size or stage of interacting individuals is known to affect the outcome of ecological interactions and can have important consequences for population dynamics. This is also true for intraguild predation (the killing and eating of potential competitors), where the size or ontogenetic stage of an individual determines whether it is the intraguild predator or the intraguild prey. Studying size- or stage-specific interactions is therefore important, but can be challenging in species with complex life histories. Here, we investigated predatory interactions of all feeding stages of the two predatory mite species Neoseiulus californicus and Phytoseiulus macropilis, both of which have complex life cycles, typical for predatory arthropods. Populations of these two species compete for two-spotted spider mites, their prey. We evaluated both the capacity to kill stages of the other predator species and the capacity to benefit from feeding on these stages, both prerequisites for the occurrence of intraguild predation. Ontogeny played a critical role in the occurrence of intraguild predation. Whereas the juveniles of P. macropilis developed from larva until adulthood when feeding on N. californicus eggs, interestingly, adult female P. macropilis did not feed on the smaller stages of the other species. We furthermore show that intraguild predation was reciprocal: both juveniles and adult females of N. californicus preyed on the smallest stages of P. macropilis. These results suggest that a proper analysis of the interactions between pairs of species involved in intraguild predation should start with an inventory of the interactions among all ontogenetic stages of these species.

Ricoseius loxocheles (Acari: Phytoseiidae) is not a predator of false spider mite on coffee crops: What does it eat?

Ricoseius loxocheles (De Leon) (Acari: Phytoseiidae) is often found in coffee crops and is known to feed on coffee leaf rust, Hemileia vastatrix Berkeley and Broome (Uredinales). As the occurrence of coffee leaf rust is limited primarily to the rainy season, the mite may use other food sources to survive during the periods of low pathogen prevalence. It is well known that phytoseiid mites can survive on a variety of food sources, such as herbivorous mites, fungi and pollen. We evaluated the ability of R. loxocheles to survive and reproduce on a diet of Brevipalpus phoenicis Geijskes (Acari: Tenuipalpidae), cattail pollen (Typha spp.), clover rust (Puccinia oxalidis), bee pollen (Santa Bárbara® dehydrated pollen, Santa Bárbara, MG, Brazil) and coffee leaf rust. Ricoseius loxocheles did not survive or reproduce on any B. phoenicis stages tested (egg, larva, adult). The survival and oviposition of R. loxocheles were directly affected by the presence of coffee rust urediniospores, but not by the presence of the prey. Survival and oviposition of the phytoseiid were similar when fed cattail pollen, clover rust and coffee leaf rust but was lower when fed bee pollen. Our results show that R. loxocheles is not a predator of B. phoenicis but it is able to utilize other resources besides coffee leaf rust.

Nematode Root Herbivory in Tomato Increases Leaf Defenses and Reduces Leaf Miner Oviposition and Performance.

The outcome of plant-mediated interactions among herbivores from several feeding guilds has been studied intensively. However, our understanding on the effects of nematode root herbivory on leaf miner oviposition behavior and performance remain limited. In this study, we evaluated whether Meloidogyne incognita root herbivory affects Tuta absoluta oviposition preference on Solanum lycopersicum plants and the development of the resulting offspring. To investigate the M. incognita-herbivory induced plant systemic responses that might explain the observed biological effects, we measured photosynthetic rates, leaf trypsin protease inhibitor activities, and analyzed the profile of volatiles emitted by the leaves of root-infested and non-infested plants. We found that T. absoluta females avoided laying eggs on the leaves of root-infested plants, and that root infestation negatively affected the pupation process of T. absoluta. These effects were accompanied by a strong suppression of leaf volatile emissions, a decrease in photosynthetic rates, and an increase in the activity of leaf trypsin protease inhibitors. Our study reveals that root attack by nematodes can shape leaf physiology, and thereby increases plant resistance.

How to evaluate the potential occurrence of intraguild predation.

Intraguild predation is the combination of exploitative competition and predation among potential competitors that use similar resources. It has the potential to shape population dynamics and community structure. Although there is much empirical evidence for the occurrence of intraguild predation in natural ecosystems, the study of its effects is mainly limited to short-term microcosm experiments. There is, therefore, certain skepticism about its actual significance in nature. A relevant concern is that there is no consensus regarding criteria to evaluate the possible occurrence of intraguild predation in short-term experiments, and methodological differences may therefore underlie apparent inconsistencies among studies. Our purpose here was to evaluate existing criteria to offer guidance for the design of experiments to determine whether two species may potentially engage in intraguild predation. The criteria are based on the condition that intraguild predators need to experience immediate energetic gains when feeding on the intraguild prey. Thus, a relevant experimental design must quantify predation but also fitness benefits of feeding on the other species, i.e. increases in reproduction, somatic growth, or survival.

Herbivores with similar feeding modes interact through the induction of different plant responses.

Plants respond to attacks by herbivores with various defences, which are mounted through the activation of different biochemical pathways that are known to interact. Thus, the attack of a plant by one herbivore species may result in changes in the performances of other species on the same plant. It has been suggested that species with comparable feeding modes induce similar plant defences and such herbivores are therefore expected to have a negative effect on each other's performance. We studied two closely related phytophagous mite species with identical feeding modes. Yet, one of the species (Tetranychus urticae) induces tomato plant defences, whereas the other (T. evansi) reduces them. We found that the "inducing" species benefits from the downregulation of defences by the "reducing" species, which, in turn, suffers from the induction of defences by the inducing species. Moreover, the performances of the two mite species on leaves that were previously attacked by both species simultaneously were intermediate between that on leaves previously attacked by each of the mites separately. The activity of proteinase inhibitor, a defensive compound, was not found to be intermediate in leaves attacked by both species simultaneously--it was almost as high as the activity seen in leaves with defences induced by T. urticae. Oviposition rates of T. urticae showed a nonlinear correlation with inhibitor activity, suggesting that it is potentially problematic to use this activity as an indicator of the level of plant defence. Our results show that herbivores with similar feeding modes have opposite effects on plant defence and differentially affect each other's performance on co-infested plants.

Antipredator behaviours of a spider mite in response to cues of dangerous and harmless predators.

Prey are known to invest in costly antipredator behaviour when perceiving cues of dangerous, but not of relatively harmless predators. Whereas most studies investigate one type of antipredator behaviour, we studied several types (changes in oviposition, in escape and avoidance behaviour) in the spider mite Tetranychus evansi in response to cues from two predatory mites. The predator Phytoseiulus longipes is considered a dangerous predator for T. evansi, whereas Phytoseiulus macropilis has a low predation rate on this prey, thus is a much less dangerous predator. Spider mite females oviposited less on leaf disc halves with predator cues than on clean disc halves, independent of the predator species. On entire leaf discs, they laid fewer eggs in the presence of cues of the dangerous predator than on clean discs, but not in the presence of cues of the harmless predator. Furthermore, the spider mites escaped more often from discs with cues of the dangerous predator than from discs without predator cues, but they did not escape more from discs with cues of the harmless predator. The spider mites did not avoid plants with conspecifics and predators. We conclude that the spider mites displayed several different antipredator responses to the same predator species, and that some of these antipredator responses were stronger with cues of dangerous predators than with cues of harmless predators.

Breaking and entering: predators invade the shelter of their prey and gain protection.

Many herbivorous arthropods construct shelters on their host plant that offer protection against natural enemies. This has resulted in selection on natural enemies to enter these shelters, where they can feed on prey that are inaccessible for competing predators and parasitoids. The spider mite Tetranychus evansi produces a shelter consisting of a dense web that is impenetrable for most predators; the only known natural enemy that can penetrate the web and can forage efficiently on this pest is Phytoseiulus longipes. We show that this predator preferentially foraged and oviposited in the web of its prey. Moreover, intraguild predation on juveniles of these predators was significantly higher outside this web and in the less dense web of a closely related prey species (T. urticae) than inside the web of T. evansi. Although the production of shelters by herbivores may be profitable at first, their adapted natural enemies may reap the benefit in the end.

Ricoseius loxocheles, a phytoseiid mite that feeds on coffee leaf rust.

One of the most important diseases of coffee plants is the coffee leaf rust fungus Hemileia vastatrix Berkeley and Broome (Uredinales). It can cause 30 % yield loss in some varieties of Coffea arabica (L.). Besides fungus, the coffee plants are attacked by phytophagous mites. The most common species is the coffee red mite, Oligonychus ilicis McGregor (Acari: Tetranychidae). Predatory mites of the Phytoseiidae family are well-known for their potential to control herbivorous mites and insects, but they can also develop and reproduce on various other food sources, such as plant pathogenic fungi. In a field survey, we found Ricoseius loxocheles (De Leon) (Acari: Phytoseiidae) on the necrotic areas caused by the coffee leaf rust fungus during the reproductive phase of the pathogen. We therefore assessed the development, survivorship and reproduction of R. loxocheles feeding on coffee leaf rust fungus and measured predation and oviposition of this phytoseiid having coffee red mite as prey under laboratory conditions. The mite fed, survived, developed and reproduced successfully on this pathogen but it was not able to prey on O. ilicis. Survival and oviposition with only prey were the same as without food. This phytoseiid mite does not really use O. ilicis as food. It is suggested that R. loxocheles is one phytoseiid that uses fungi as a main food source.

Economic Injury Levels and Economic Thresholds for Leucoptera coffeella as a Function of Insecticide Application Technology in Organic and Conventional Coffee (Coffea arabica), Farms.

Leucoptera coffeella (Lepidoptera: Lyonetiidae) is one of the main pests in coffee crops. The economic injury level (EIL) is the lowest density of the pest at which economic damages match the costs of control measures. The economic threshold (ET) is the density of the pest at which control measures must be taken so that this population does not reach the EIL. These are the main indices used for pest control decision-making. Control of L. coffeella is carried out by manual, tractor, airplane or drone applications. This work aimed to determine EILs and ETs for L. coffeella as a function of insecticide application technology in conventional and organic Coffea arabica crops. Data were collected over five years in commercial C. arabica crops on seven 100 ha central pivots. The cost of control in organic crops was 16.98% higher than conventional. The decreasing order of control cost was manual > drone > airplane > tractor application. Coffee plants were tolerant to low densities (up to 15% mined leaves) of the pest that caused losses of up to 6.56%. At high pest densities (54.20% mined leaves), losses were high (85.62%). In organic and conventional crops and with the use of different insecticide application technologies, EIL and ET were similar. The EIL and ET were 14% and 11% of mined leaves, respectively. Therefore, these indices can be incorporated in integrated pest management programs in C. arabica crops. The indices determined as a function of insecticide application technology in organic and conventional coffee are important as they serve producers with different technological levels. Additionally, EILs and ETs can contribute to more sustainable production, as control methods will only be employed when the pest density reaches these indices.

Survival and behavioural response to acaricides of the coconut mite predator Neoseiulus baraki.

The coconut mite, Aceria guerreronis Keifer, is a major pest of coconut palm in the world. The control of this pest species is done through acaricide applications at short time intervals. However, the predators of this pest may also be affected by acaricides. Among the predators of A. guerreronis, Neoseiulus baraki (Athias-Henriot) has potential for biological control. The objective of this study was to assess the effect of acaricides on the survival and behavior of N. baraki. The survivorship of N. baraki was recorded in surface-impregnated arenas. Choice and no-choice behavioral bioassays were carried out using a video tracking system to assess the walking behavior of the predator under acaricide exposure. Although all acaricides negatively affected the survival of N. baraki, chlorfenapyr and azadirachtin caused lower effect than the other acaricides. No significant differences in walking behavior were observed under exposure to fenpyroximate, chlorfenapyr and chlorpyrifos on fully-contaminated arenas. Azadirachtin and chlorpyrifos caused repellence. Irritability was observed for all acaricides, except for abamectin. Chlorfenapyr was the most suitable product for managing the coconut mite because of its low effect on survival and behavior of N. baraki.

A herbivore that manipulates plant defence.

Phytopathogens and herbivores induce plant defences. Whereas there is evidence that some pathogens suppress these defences by interfering with signalling pathways involved in the defence, such evidence is scarce for herbivores. We found that the invasive spider mite Tetranychus evansi suppresses the induction of the salicylic acid and jasmonic acid signalling routes involved in induced plant defences in tomato. This was reflected in the levels of inducible defence compounds, such as proteinase inhibitors, which in mite-infested plants were reduced to even lower levels than the constitutive levels in herbivore-free plants. Additionally, the spider mite suppressed the release of inducible volatiles, which are implicated in plant defence. Consequently, the mites performed much better on previously attacked plants than on non-attacked plants. These findings provide a new perspective on plant-herbivore interactions, plant protection and plant resistance to invasive species.

Olfactory response of predatory mites to vegetative and reproductive parts of coconut palm infested by Aceria guerreronis.

The phytophagous mite Aceria guerreronis Keifer is an important pest of coconut worldwide. A promising method of control for this pest is the use of predatory mites. Neoseiulus baraki (Athias-Henriot) and Proctolaelaps bickleyi Bram are predatory mites found in association with A. guerreronis in the field. To understand how these predators respond to olfactory cues from A. guerreronis and its host plant, the foraging behavior of the predatory mites was investigated in a Y-tube olfactometer and on T-shaped arenas. The predators were subjected to choose in an olfactometer: (1) isolated parts (leaflet, spikelet or fruit) of infested coconut plant or clean air stream; (2) isolated parts of non-infested or infested coconut plant; and (3) two different plant parts previously shown to be attractive. Using T-shaped arenas the predators were offered all possible binary combinations of discs of coconut fruit epidermis infested with A. guerreronis, non-infested discs or coconut pollen. The results showed that both predators were preferred (the volatile cues from) the infested plant parts over clean air. When subjected to odours from different infested or non-infested plant parts, predators preferred the infested parts. Among the infested plant parts, the spikelets induced the greatest attraction to predators. On the arenas, both predators preferred discs of coconut fruits infested with A. guerreronis over every other alternative. The results show that both predators are able to locate A. guerreronis by olfactory stimuli. Foraging strategies and implications for biological control are discussed.

Suitability of the predatory mites Iphiseiodes zuluagai and Euseius concordis in controlling Polyphagotarsonemus latus and Tetranychus bastosi on Jatropha curcas plants in Brazil.

One of the most promising plant species for biofuel production in Brazil is the physic nut Jatropha curcas. Major phytosanitary problems include the attack of two pest mite species, the broad mite Polyphagotarsonemus latus and the spider mite Tetranychus bastosi. Owing to pesticide-related problems, there is an increasing demand for sustainable environmental-friendly control methods such as biological control. In this study we evaluated the suitability of the predatory mite species Iphiseiodes zuluagai and Euseius concordis in controlling P. latus and T. bastosi on J. curcas. The number of T. bastosi killed by I. zuluagai was lower than the number of P. latus consumed. Euseius concordis preyed upon both T. bastosi and P. latus but the number of prey killed was always lower in comparison with I. zuluagai. However, P. latus and T. bastosi are suitable for the development of I. zuluagai and E. concordis as oviposition of both predators did not differ in relation to prey species. The preference of I. zuluagai for leaves of plants infested by either P. latus or T. bastosi, combined with the higher values for predation obtained by this predatory mite when fed on P. latus, compared to those values obtained by E. concordis, suggests that I. zuluagai can be more efficient than E. concordis in reducing populations of P. latus and T. bastosi under field conditions. Furthermore, we report here on the first record of predatory mites associated with P. latus and T. bastosi on native J. curcas plants in Brazil. In conclusion, we emphasize the crucial importance of predatory mites as agents of natural biological control of mite pests on J. curcas in small farms.

When treatment increases the contaminant's ecotoxicity: A study of the Fenton process in the degradation of methylene blue.

The degradation of dyes can generate harmful by-products, thereby requiring the need to evaluate the toxicity to aquatic organisms. This study aims to evaluate the chronic ecotoxicity of methylene blue dye degraded by the Fenton process using the non-target planarian Girardia tigrina as a sensitive bioindicator of environmental contamination. The bioassays evaluated the lethality of several concentrations of the untreated and degraded dye methylene blue (MB), as well as, their sub-lethal effects on locomotion, feeding, regeneration, and reproduction. In both acute and chronic tests, the degraded dye had a stronger toxic effect when compared to the untreated dye. This negative effect after treatment was mainly associated with the presence of residual hydrogen peroxide and iron (and consequently the hydroxyl radical formed). We conclude that the utilization of the Fenton process using less oxidizing agents should be considered as important alternatives for the protection of aquatic ecosystems, without compromising the efficient removal of MB.