Host finding behaviour of the coconut mite Aceria guerreronis

For the coconut mite, Aceria guerreronis Keifer, its host plant, the coconut palm, is not merely a source of food, but more generally a habitat to live in for several generations. For these minute organisms, finding a new plant is difficult and risky, especially because their main mode of dispersal is passive drifting with the wind and because they are highly specialized on their host plant. Consequently, the probability of landing on a suitable host is very low, let alone to land in their specific microhabitat within the host. How coconut mites manage to find their microhabitat within a host plant is still underexplored. We tested the hypothesis that they use volatile chemical information emanating from the plant to find a specific site within their host plants and/or use non-volatile plant chemicals to stay at a profitable site on the plant. This was investigated in a Y-tube olfactometer (i.e. under conditions of a directed wind flow) and on cross-shaped arenas (i.e. under conditions of turbulent air) that either allowed contact with odour sources or not. The mites had to choose between odours from specific parts (leaflet, spikelet or fruit) of a non-infested coconut plant and clean air as the alternative. In the olfactometer experiments, no mites were found to reach the upwind end of the Y-tube: <5 % of the mites were able to pass the bifurcation of the “Y”. On the cross-shaped arenas, however, a large number of coconut mites was found only when the arm of the arena contained discs of fruit epidermis and contact with these discs was allowed. The results suggest that coconut mites on palm trees are not attracted to specific sites on the plant by volatile plant chemicals, but that they arrested once they contact the substrate of specific sites. Possibly, they perceive non-volatile chemicals, but these remain to be identified.     

Population dynamics of Aceria guerreronis Keifer (Acari: Eriophyidae) and associated predators on coconut fruits in Northeastern Brazil

Aceria guerreronis Keifer can cause severe damage to coconuts in several countries around the world. Rare studies have been conducted to determine the predatory mites associated with A. guerreronis in Brazil. The study evaluated the prevalence of A. guerreronis and associated predators on the bracts and on the surface of the fruits underneath the bracts, for 12 months, on coconut palms grown along the coast of the States of Alagoas, Paraíba and Pernambuco, Northeastern Brazil. Mites of 10 families were found, but by far the most abundant species was A. guerreronis, corresponding to 99.5% of the mites collected. The prevailing species amongst the predators were the Phytoseiidae mites Neoseiulus baraki (Athias-Henriot) and Neoseiulus paspalivorus (De Leon). These are flat mites that have short limbs, characteristics that allow them to invade the main habitat occupied by A. guerreronis. Other predators were found, but in low numbers, due mainly to their difficulty in reaching the fruit areas most inhabited by A. guerreronis. However, these mites could prey on that pest when it leaves its preferred habitat to disperse. No significant correlations were observed between the levels of abiotic factors and the population levels of A. guerreronis or of the phytoseiids associated with it. This probably occurred due to the simultaneous and conflicting effects of those factors on the mites in the field, or to significant differences between the climatic factors measured in the environment and those prevailing in the habitat occupied by A. guerreronis.     

Plants with spider-mite prey attract more predatory mites than clean plants under greenhouse conditions

Although many predators and parasitoids are known to respond to odours produced by plants infested with their prey under laboratory conditions, there are actually few studies that show that this response leads to higher numbers of predators or parasitoids on the plants under natural conditions. Here we study the response of predatory mites (Phytoseiulus persimilis Athias-Henriot, Acari, Phytoseiidae) to odours from cucumber plants infested with two-spotted spider mites (Tetranychus urticae Koch, Acari, Tetranychidae) in greenhouse release experiments, where predators were released in the centre of a hexagon of cucumber plants. Forty to 57% of all predators released were recaptured on plants within 7 h. Of these, an average of 79.5% were found on infested plants, indicating that these attract about 4 times as many predators as do clean plants. Hence, the blind predatory mites were guided to the plants with prey by herbivore-induced odours produced by the plant, as was indicated by olfactometer experiments, where it was found that P. persimilis preferred odours from infested cucumber plants to odours from clean cucumber plants. The long-range searching behaviour of P. persimilis is discussed.     

Compatibility of Neoseiulus paspalivorus and Proctolaelaps bickleyi, candidate biocontrol agents of the coconut mite Aceria guerreronis: spatial niche use and intraguild predation

The eriophyid mite Aceria guerreronis occurs in most coconut growing regions of the world and causes enormous damage to coconut fruits. The concealed environment of the fruit perianth under which the mite resides renders its control extremely difficult. Recent studies suggest that biological control could mitigate the problems caused by this pest. Neoseiulus paspalivorus and Proctolaelaps bickleyi are two of the most frequently found predatory mites associated with A. guerreronis on coconut fruits. Regarding biological control, the former has an advantage in invading the tight areas under the coconut fruit perianth while the latter is more voracious on the pest mites and has a higher reproductive capacity. Based on the idea of the combined use/release of both predators on coconut fruits, we studied their compatibility in spatial niche use and intraguild predation (IGP). Spatial niche use on coconut fruits was examined on artificial arenas mimicking the area under the coconut fruit perianth and the open fruit surface. Both N. paspalivorus and P. bickleyi preferentially resided and oviposited inside the tight artificial chamber. Oviposition rate of P. bickleyi and residence time of N. paspalivorus inside the chamber were reduced in the presence of a conspecific female. Residence of N. paspalivorus inside the chamber was also influenced by the presence of P. bickleyi. Both N. paspalivorus and P. bickleyi preyed upon each other with relatively moderate IGP rates of adult females on larvae but neither species yielded nutritional benefits from IGP in terms of adult survival and oviposition. We discuss the relevance of our findings for a hypothetic combined use of both predators in biological control of A. guerreronis.     

Population dynamics of Aceria guerreronis (Acari: Eriophyidae) and other mites associated with coconut fruits in Una, state of Bahia, northeastern Brazil

Aceria guerreronis Keifer is a major coconut pest in the Americas, Africa and some Asian countries, and occurs in high population levels in northeastern Brazil. The determination of the climatic conditions that favorably affect its population growth and the prevalence and abundance of predatory mites can promote more efficient control practices. Our objective was to evaluate the pattern of occurrence and population dynamics of A. guerreronis, their associated predators and other mites during a 2?year period in a hybrid coconut plantation in the municipality of Una, state of Bahia, northeastern Brazil. Monthly samples of fruits were taken from June 2008 to May 2010 for qualitative and quantitative evaluation of mites. Aceria guerreronis represented 99.9?% of the mites. An average density of 1,117 mites per fruit and a maximum of 23,596 mites per fruit indicated that the level of infestation can be high in Bahia. Bdella ueckermanni Hernandes, Daud and Feres was the most abundant and frequent predator. Population increase of A. guerreronis was directly related to the temperature rise and inversely related to both the increase of air relative humidity and rainfall. The highest population densities occurred from November to March. The largest A. guerreronis populations occurred in fruits with 32 and 48?% of damaged surface. The relationship between prevailing wind direction and incidence of A. guerreronis could not be corroborated or refuted.     

Attraction of a generalist predator towards herbivore-infested plants

The occurrence and strength of interactions among natural enemies and herbivores depend on their foraging decisions, and several of these decisions are based on odours. To investigate interactions among arthropods in a greenhouse cropping system, we studied the behavioural response of the predatory bug Orius laevigatus (Fieber) (Hemiptera: Anthocoridae) towards cucumber plants infested either with thrips (Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae)) or with spider mites (Tetranychus urticae Koch (Acari: Tetranychidae)). In greenhouse release-recapture experiments, the predatory bug showed a significant preference for both thrips-infested plants and spider mite-infested plants over clean plants. Predatory bugs preferred plants infested with spider mites to plants with thrips. Experience with spider mites on cucumber leaves prior to their release in the greenhouse had no effect on the preference of the predatory bugs. However, this experience did increase the percentage of predators recaptured. Y-tube olfactometer experiments showed that O. laevigatus was more attracted to odours from plants infested with spider mites than to odours from clean plants. Thus, O. laevigatus is able to perceive odours and may use them to find plants with prey in more natural conditions. The consequences of the searching behaviour for pest control are discussed.     

How predatory mites find plants with whitefly prey

We investigated the searching behaviour of two species of predatory mites, Typhlodromips swirskii (Athias-Henriot) and Euseius scutalis (Athias-Henriot), both known to feed on immature stages of the whitefly Bemisia tabaci Gennadius. When released in a greenhouse inside a circle of cucumber plants that were alternatingly clean or infested with immature whiteflies, the mites took several days to find plants. Both species were recaptured significantly more on plants with whiteflies. This suggests that the mites are able to discriminate between plants with and without whiteflies. The predators may either have been attracted to plants with whiteflies from a distance or arrested on plants with whiteflies. Typhlodromips swirskii that had previously fed on whitefly immatures on cucumber leaves were significantly attracted by volatiles from cucumber plants with whiteflies in a Y-tube olfactometer. This suggests that the mites use volatile cues to discriminate between infested and clean plants. However, this response waned rapidly; if predators, experienced as above, were starved for 3–4 h in absence of cucumber leaves, they no longer preferred volatiles of infested plants to clean plants. Furthermore, T. swirskii that had no experience with immature whiteflies on cucumber plants also did not prefer odours of infested plants to those of clean plants. Because the release experiment with this species in the greenhouse was done with inexperienced predators, this suggests that the aggregation of mites on plants with whiteflies was mainly caused by differential arrestment of mites on plants with prey and clean plants. For T. swirskii, this was in agreement with the finding that the fraction of predators on plants with prey increased with time to levels higher than 70%. A less clear trend was found for E. scutalis, for which the fraction of predators on plants with prey stabilized soon after release to levels from 54–70%. Hence, the predatory mites may find plants with prey by random searching, but they are subsequently arrested on these plants. An earlier study showed that 87% of all whiteflies released in a set-up as used here were recaptured within 1 day. Hence, the effectiveness with which predatory mites locate plants with whiteflies is low compared with that of their prey. We expect this to generate spatial patterns in the dynamics of predator and prey and this may have consequences for biological control of whiteflies with predatory mites.     

Below-ground plant parts emit herbivore-induced volatiles: olfactory responses of a predatory mite to tulip bulbs infested by rust mites

Although odour-mediated interactions among plants, spider mites and predatory mites have been extensively studied above-ground, belowground studies are in their infancy. In this paper, we investigate whether feeding by rust mites (Aceria tulipae) cause tulip bulbs to produce odours that attract predatory mites (Neoseiulus cucumeris). Since our aim was to demonstrate such odours and not their relevance under soil conditions, the experiments were carried out using a classic Y-tube olfactometer in which the predators moved on a Y-shaped wire in open air. We found that food-deprived female predators can discriminate between odours from infested bulbs and odours from uninfested bulbs or artificially wounded bulbs. No significant difference in attractiveness to predators was found between clean bulbs and bulbs either wounded 30 min or 3 h before the experiment. These results indicate that it may not be simply the wounding of the bulbs, but rather the feeding by rust mites, which causes the bulb to release odours that attract N. cucumeris. Since bulbs are belowground plant structures, the olfactometer results demonstrate the potential for odour-mediated interactions in the soil. However, their importance in the actual soil medium remains to be demonstrated.     

Odour-mediated responses of a predatory mirid bug and its prey, the two-spotted spider mite

It has been shown that many natural enemies of herbivorous arthropods use herbivore induced plant volatiles (HIPVs) to locate their prey. Herbivores can also exploit cues emitted by plants infested with heterospecifics or conspecifics. A study was conducted to test whether green bean HIPVs as well as odours emitted directly by spider mites influenced the orientation behaviour of the predatory mirid bug, Macrolophus caliginosus and its prey, Tetranychus urticae in a Y-tube olfactometer. Our results show that both spider mites and M. caliginosus preferred spider mite infested green bean plants to uninfested plants. For M. caliginosus this response was mediated by HIPVs whereas for T. urticae it was mediated through a composite response to both HIPVs and odours emitted directly by the conspecifics (and their associated products). The results may be of use in practical biocontrol situations, through e.g., plant breeding for improved HIPV production, conditioning of mass-reared predators to appropriate cues, and employment of “push–pull-strategies” by using HIPVs.     

Induced Response of Tomato Plants to Injury by Green and Red Strains of Tetranychus Urticae

We studied the induced response of tomato plants to the green strain and the red strain of the spider mite Tetranychus urticae. We focused on the olfactory response of the predatory mite Phytoseiulus persimilis to volatiles from T. urticae-infested tomato leaves in a Y-tube olfactometer. Tomato leaves attracted the predatory mites when slightly infested with the red strain, or moderately or heavily infested with the green strain. In contrast, neither leaves that were slightly infested with green-strain mites, nor leaves that were moderately or heavily infested with the red strain attracted the predators. We discuss the specific defensive responses of tomato plants to each of the two strains.     

Distribution patterns of coconut mite, Aceria guerreronis, and its predator Neoseiulus aff. paspalivorus in coconut palms

Distribution patterns and numerical variability of the coconut mite Aceria guerreronis Keifer (Acari: Eriophyidae) and its predator Neoseiulus aff. paspalivorus DeLeon (Phytoseiidae) on the nuts of 3- to 7-month-old bunches of coconut palms were studied at two sites in Sri Lanka. At the two sites, coconut mites were present on 88 and 75% of the nuts but no more than three-quarters of those nuts showed damage symptoms. N. aff. paspalivorus was found more on mature nuts than on immature nuts. Spatial and temporal distribution of coconut mites and predatory mites differed significantly. The mean number of coconut mites per nut increased until 5-month-old bunches and declined thereafter. The densities of predatory mites followed a similar trend but peaked 1 month later. Variability in the numbers of mites among palms and bunches of the same age was great, but was relatively low on 6-month-old bunches. The results indicate that assessment of infestation levels by damage symptoms alone is not reliable. Sampling of coconut and/or predatory mite numbers could be improved by using several nuts of 6-month-old bunches. The effect of predatory mites on coconut mites over time suggests that N. aff. paspalivorus could be a prospective biological control agent of A. guerreronis.     

Variation in the olfactory response of 13 populations of the predatory mite Amblyseius womersleyi to Tetranychus urticae-infested plant volatiles (Acari: Phytoseiidae, Tetranychidae)

We studied the response of the predatory mite Amblyseius womersleyi collected in 13 different sites in Japan toward Tetranychus urticae-infested kidney bean leaf volatiles in a Y-tube olfactometer. The predatory mites were collected from eight plant species infested by one of three tetranychid mite species. The predators' responses to the infested-leaf volatiles varied from 33% to 97% among the populations. The predators collected at 10 sites showed a significant preference for infested-leaf volatiles, whereas those collected at three tea plantations did not distinguish between the infested- and uninfested-leaf volatiles. We discussed the possible factors that affected the olfactory response of A. womersleyi towards the infested leaf volatiles.     

Exploration of the acarine fauna on coconut palm in Brazil with emphasis on Aceria guerreronis (Acari: Eriophyidae) and its natural enemies

Coconut is an important crop in tropical and subtropical regions. Among the mites that infest coconut palms, Aceria guerreronis Keifer is economically the most important. We conducted surveys throughout the coconut growing areas of Brazil. Samples were taken from attached coconuts, leaflets, fallen coconuts and inflorescences of coconut palms in 112 localities aiming to determine the occurrence and the distribution of phytophagous mites, particularly A. guerreronis, and associated natural enemies. Aceria guerreronis was the most abundant phytophagous mite followed by Steneotarsonemus concavuscutum Lofego & Gondim Jr. and Steneotarsonemus furcatus De Leon (Tarsonemidae). Infestation by A. guerreronis was recorded in 87% of the visited localities. About 81% of all predatory mites belonged to the family Phytoseiidae, mainly represented by Neoseiulus paspalivorus De Leon, Neoseiulus baraki Athias-Henriot and Amblyseius largoensis Muma; 12% were Ascidae, mainly Proctolaelaps bickleyi Bram, Proctolaelaps sp nov and Lasioseius subterraneus Chant. Neoseiulus paspalivorus and N. baraki were the most abundant predators on attached coconuts. Ascidae were predominant on fallen coconuts, while A. largoensis was predominant on leaflets; no mites were found on branches of inflorescences. Leaflets harboured higher mite diversity than the attached coconuts. Mite diversity was the highest in the state Pará and on palms surrounded by seasonal forests and Amazonian rain-forests. Neoseiulus paspalivorus, N. baraki and P. bickleyi were identified as the most promising predators of A. guerreronis. Analyses of the influence of climatic factors revealed that dry ambient conditions favour the establishment of A. guerreronis. Neoseiulus paspalivorus and N. baraki have differing climatic requirements; the former being more abundant in warm and dry areas, the latter prevailing in moderately tempered and humid areas. We discuss the significance of our findings for natural and biological control of A. guerreronis.     

Residual bioassay to assess the toxicity of Acaricides against Aceria guerreronis (Acari: Eriophyidae) under laboratory conditions

Aceria guerreronis Keifer (Acari: Eriophyidae) is considered a major pest of the coconut (Cocos nucifera L.), and the use of pesticides is the current method to control it. However, no standard toxicological tests exist to select and assess the efficiency of molecules against the coconut mite. The aim of this study was to develop a methodology that allows for the evaluation of the relative toxicity of acaricides to A. guerreronis through rapid laboratory procedures. We confined A. guerreronis on arenas made out of coconut leaflets and tested two application methods: immersing the leaf fragments in acaricides and spraying acaricides on the leaf fragments under a Potter spray tower. In the latter application method, we sprayed leaf fragments both populated with and devoid of mites. We evaluated the comparative toxicity of two populations (Itamaracá and Petrolina, Pernambuco, Brazil) by spraying on leaflets without mites and submitted the mortality data to probit analysis after 24 h of exposure. No difference was observed in the LC50, regardless of whether the leaflets were immersed or sprayed with acaricide (abamectin, chlorfenapyr or fenpyroximate). The toxicity of chlorfenapyr and fenpyroximate did not differ, irrespective of whether it was applied directly to the leaflet or to the mite; however, the toxicity of abamectin was higher when applied directly to the mite. Chlorpyrifos and abamectin toxicities were lower for the Petrolina population than for the Itamaracá population. Immersing and spraying coconut leaflets can be used to assess the mortality of A. guerreronis under laboratory conditions.     

Acaricides impair prey location in a predatory phytoseiid mite

The use of predatory mites as the sole management tactic in biological control programmes frequently does not fully and reliably prevents damage of phytophagous mites on plants. Therefore, as an alternative, the integration of predatory mites with acaricides can provide more effective control of phytophagous mites than that of the predators only. However, for such integration, acaricides minimal negative impacts on predatory mites are required. In this study, we evaluated the sublethal effects of three acaricides on the foraging behaviour of Neoseiulus baraki (Athias‐Henriot) (Acari: Phytoseiidae) in a coconut production system. The acaricides were assessed for interference with the location of prey habitat using a Y‐tube olfactometer and for interference with the location of the prey colony within the habitat using a video‐tracking system. In addition to the choice of odour source, the time required and the distance walked to make the choice were assessed. The acaricides tested were abamectin, azadirachtin and fenpyroximate. The predatory mite preferred coconuts infested with the coconut mite Aceria guerreronis Keifer (Acari: Eriophyidae) over uninfested coconuts when not exposed to acaricides. However, when exposed to acaricides, the predator did not distinguish between infested and uninfested fruits. When exposed to abamectin, N. baraki spent more time resting and walked greater distances before making the choice of an odour source. Thus, the acaricides impair the ability of the predatory mite N. baraki to locate a prey habitat and to locate a prey within that habitat. The acaricides differentially affected prey foraging by interfering with odour perception.     

Attraction of the predatory mites Typhlodromalus manihoti and Typhlodromalus aripo to cassava plants infested by cassava green mite

The attraction of the predatory mites, Typhlodromalus manihoti and Typhlodromalus aripo, to the host plant-spider mite complex, Manihot esculenta – Mononychellus tanajoa, was investigated with a Y-tube olfactometer. Factors examined included predator starvation period, several combinations of cassava leaf biomass and initial M. tanajoa infestations, M. tanajoa-damaged leaves with mites and/or their residues removed, M. tanajoa alone, and mechanically damaged cassava leaves. We found that females of T. manihoti and T. aripo were significantly attracted to M. tanajoa-infested cassava leaves when the predators were starved for 2, 6, or 10 h. Satiated T. aripo was significantly attracted to infested cassava leaves whereas satiated T. manihoti did not discriminate between infested and non-infested leaves. When a choice was given between either two or four leaves infested with 200 female M. tanajoa and an equivalent number of non-infested leaves, 2 h-starved T. manihoti and T. aripo were significantly attracted to each of the infested groups of cassava leaves. At a density of 12 female M. tanajoa per leaf on four leaves, 2 h-starved T. manihoti was still attracted to M. tanajoa-infested leaves whereas 2 h-starved T. aripo was not attracted. When a choice was given between non-infested cassava leaves and either infested leaves from which only M. tanajoa females had been removed, or infested leaves from which all M. tanajoa and their visible products (web, feces) had been wiped off, T. aripo preferred odors from both types of previously infested leaves. Typhlodromalus manihoti was only attracted to infested leaves from which the M. tanajoa females only had been removed. Finally, the two predators were not attracted to 400 female M. tanajoa on clean cotton wool or to mechanically wounded leaves. This supports the hypothesis that M. tanajoa damage induces volatile cues in cassava leaves that attract T. manihoti and T. aripo to M. tanajoa-infested leaves.     

捕食螨化学生态研究进展

捕食螨是重要的生物防治因子。早在20世纪70年代就发现了捕食螨的性信息素,许多研究证明植物挥发物在捕食螨向猎物定位过程中发挥着至关重要的作用,影响捕食螨寻找猎物的植物挥发物来源于未受害植物、机械损伤植物、猎物危害植物、非猎物危害植物。人工合成的植物挥发物组分对捕食螨具有引诱作用,但引诱活性低于虫害诱导植物释放的挥发性混合物。捕食螨的饲养条件、饥饿程度、学习与经验行为等会影响捕食螨对植物挥发物的反应。介绍了信息素与植物挥发物对捕食螨的作用,并讨论了目前存在的问题和研究前景。     

Sub-lethal effects of fenbutatin oxide on prey location by the predatory mite <Emphasis Type="Italic">Iphiseiodes zuluagai</Emphasis> (Acari: Phytoseiidae)

We used a Y-tube olfactometer to assess the sub-lethal effects of the acaricide fenbutatin oxide on the olfactory response of the predatory mite Iphiseiodes zuluagai towards odours from: (1) air or undamaged coffee plants; (2) undamaged or red spider mite Oligonychus ilicis-infested coffee plants; (3) undamaged or false spider mite Brevipalpus phoenicis-infested coffee plants. Predatory mite adult females were exposed to residues of fenbutatin oxide or distilled water on leaf discs during a period of 72 h prior experiments. When exposed to distilled water (control treatments), predatory mites significantly preferred undamaged plants over air, O. ilicis-infested plants over undamaged plants, and they did not prefer B. phoenicis-infested plants over undamaged plants. However, predatory mites that had been exposed to residues of fenbutatin oxide were neither attracted towards undamaged plants nor to O. ilicis-infested plants. Thus, fenbutatin oxide affected negatively the olfactory response of I. zuluagai. We conclude that sub-lethal-effect studies should be considered in pesticide selectivity programs since the ability of predatory mites to locate their prey may be negatively affected by non-lethal concentrations of pesticides.     

A key volatile infochemical that elicits a strong olfactory response of the predatory mite <Emphasis Type="Italic">Neoseiulus californicus</Emphasis>, an important natural enemy of the two-spotted spider mite <Emphasis Type="Italic">Tetranychus urticae</Emphasis>

Herbivore-induced plant volatiles (HIPVs) emitted from lima bean leaves infested with the two-spotted spider mites Tetranychus urticae strongly attract the predatory mites Neoseiulus californicus. Among these HIPVs, methyl salicylate and linalool can attract the predators. Three green-leaf volatiles (GLVs) of (Z)-3-hexen-1-ol, (Z)-3-hexenyl acetate and (E)-2-hexenal, found in the odor blends from T. urticae-infested leaves and physically damaged leaves, can also attract the predators. To search for a strong predator attractant, the olfactory responses of N. californicus to each synthetic compound or their combinations were investigated in a Y-tube olfactometer. When presented a choice between a mixture of the five compounds (i.e. the two HIPVs and the three GLVs) and T. urticae-infested leaves, N. californicus did not discriminate between these odor sources. The same trend was observed when either a mixture of the two HIPVs or methyl salicylate vs. T. urticae-infested leaves were compared. In contrast, the predators preferred T. urticae-infested leaves to linalool, each of the three GLVs, or a mixture of the three GLVs. These results indicated that methyl salicylate is a strong predator attractant, and its potential attractiveness almost equaled that of the blend of HIPVs from T. urticae-infested leaves.