Field distribution patterns of pests are asymmetrically affected by the presence of other herbivores.

Because plant phenotypes can change in response to attacks by herbivores in highly variable ways, the distribution of herbivores depends on the occurrence of other herbivore species on the same plant. We carried out a field study to evaluate the co-occurrence of three coconut pests, the mites Aceria guerreronis (Acari: Eriophyidae), Steneotarsonemus concavuscutum (Acari: Tarsonemidae) and the moth Atheloca bondari (Lepidoptera: Pyralidae). The eriophyid mite Ac. guerreronis is the most important coconut pest around the world, whereas S. concavuscutum and At. bondari are economically important only in some areas along the Brazilian coast. A previous study suggested that the necrosis caused by Ac. guerreronis facilitates the infestation of At. bondari larvae. Because all three species infest the area under the perianths on coconuts and S. concavuscutum also causes necrosis that could facilitate At. bondari, we evaluated the co-occurrence of all three species. We found that the occurrence of At. bondari was positively associated with Ac. guerreronis, but negatively associated with S. concavuscutum. In addition, the two mite species showed negative co-occurrence. Atheloca bondari was found on nuts of all ages, but more on nuts that had fallen than on those on the trees, suggesting that nuts infested by At. bondari tend to fall more frequently. We discuss the status of At. bondari as a pest and discuss experiments to test the causes of these co-occurrence patterns.

Predator performance is impaired by the presence of a second prey species.

The simultaneous infestation of a plant by several species of herbivores may affect the attractiveness of plants to the natural enemies of one of the herbivores. We studied the effect of coconut fruits infested by the pests Aceria guerreronis and Steneotarsonemus concavuscutum, which are generally found together under the coconut perianth. The predatory mite Neoseiulus baraki produced lower numbers of offspring on fruits infested with S. concavuscutum and on fruits infested with both prey than on fruits with A. guerreronis only. The predators were attracted by odours emanating from coconuts with A. guerreronis, but not by odours from coconuts with S. concavuscutum, even when A. guerreronis were present on the same fruit. Fewer N. baraki were recaptured on fruits with both prey or with S. concavuscutum than on fruits with only A. guerreronis. Furthermore, the quality of A. guerreronis from singly and multiply infested coconuts as food for N. baraki did not differ. Concluding, our results suggest that N. baraki does not perform well when S. concavuscutum is present on the coconuts, and the control of A. guerreronis by N. baraki may be negatively affected by the presence of S. concavuscutum.

Acaricide-impaired functional predation response of the phytoseiid mite Neoseiulus baraki to the coconut mite Aceria guerreronis.

Acaricides may interfere with a myriad of interactions among arthropods, particularly predator-prey interactions. The coconut mite, Aceria guerreronis Keifer (Acari: Eriophyidae), and its phytoseiid predator, Neoseiulus baraki (Athias-Henriot) (Acari: Phytoseiidae), provide an opportunity to explore such interference because the former is a key coconut pest species that requires both predation and acaricide application for its management. The objective of the present study was to assess the effect of the acaricides abamectin, azadirachtin and fenpyroximate on the functional response of N. baraki to A. guerreronis densities. The following prey densities were tested: 5, 10, 20, 40 and 80 preys. The type of functional response and prey handling time (Th) were not altered by the acaricides. However, the attack rate (a') was modified by abamectin and fenpyroximate, and the consumption peak was reduced by abamectin. All of the acaricides allowed for the maintenance of the predator in the field, but exposure to abamectin and fenpyroximate compromised prey consumption.

Evidence of Amblyseius largoensis and Euseius alatus as biological control agent of Aceria guerreronis.

Amblyseius largoensis (Muma) (Acari: Phytoseiidae) and Euseius alatus De Leon (Acari: Phytoseiidae) are predatory mites that are mostly found on leaves and on the exposed fruit surface of coconut plants. Their morphology hampers the access to the microhabitat occupied by Aceria guerreronis Keifer (Acari: Eriophyidae), the most important pest of coconut fruits throughout the world. However, it was suggested that they can prey on A. guerreronis under natural conditions when this pest leaves its refuge to disperse. Since the trophic interactions between A. largoensis or E. alatus and A. guerreronis are unknown, we compare the frequencies of occurrence of A. largoensis and E. alatus under the bracts of coconut fruits and on coconut leaflets. In addition, because phytoseiids feed by liquid ingestion, we used molecular analysis to confirm the potential role of A. largoensis or E. alatus as predators of A. guerreronis and to assess how fast the A. guerreronis DNA fragment is degradated in the A. largoensis digestive tract. Our study demonstrated that E. alatus was only present on coconut leaflets whereas A. largoensis was found mostly on leaflets and, to a much lesser extent, under the bracts of coconuts. Species-specific ITS primers designed for A. guerreronis were shown to have a high degree of specificity for A. guerreronis DNA and did not produce any PCR product from DNA templates of the other insects and mites associated with the coconut agroecosystem. Based on molecular analysis, we confirmed that the predatory mites, A. largoensis and E. alatus, had preyed on the coconut mite in the field. Overall the predatory mites collected in the field exhibited low levels of predation (26.7% of A. largoensis and 8.9% of E. alatus tested positive for A. guerreronis DNA). The fragment of A. guerreronis DNA remained intact for a very short time (no more than 6 h after feeding) in the digestive tract of A. largoensis.

Limits to ambulatory displacement of coconut mites in absence and presence of food-related cues.

Ambulatory movement of plant-feeding mites sets limits to the distances they can cover to reach a new food source. In absence of food-related cues these limits are determined by survival, walking activity, walking path tortuosity and walking speed, whereas in presence of food the limits are also determined by the ability to orient and direct the path towards the food source location. For eriophyoid mites such limits are even more severe because they are among the smallest mites on earth, because they have only two pairs of legs and because they are very sensitive to desiccation. In this article we test how coconut mites (Aceria guerreronis Keifer) are constrained in their effective displacement by their ability to survive in absence of food (meristematic tissue under the coconut perianth) and by their ability to walk and orient in absence or presence of food-related cues. We found that the mean survival time decreased with increasing temperature and decreasing humidity. Under climatic conditions representative for the Tropics (27 °C and 75 % relative humidity) coconut mites survived on average for 11 h and covered 0.4 m, representing the effective linear displacement away from the origin. Within a period of 5 h, coconut mites collected from old fruits outside the perianth moved further away from the origin than mites collected under the perianth of young fruits. However, in the presence of food-related cues coconut mites traveled over 30 % larger distances than in absence of these cues. These results show that ambulatory movement of eriophyoid mites may well bring them to other coconuts within the same bunch and perhaps also to other bunches on the same coconut palm, but it is unlikely to help them move from palm to palm, given that palms usually do not touch each other.

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.

Behaviour of coconut mites preceding take-off to passive aerial dispersal.

For more than three decades the coconut mite Aceria guerreronis Keifer is one of the most important pests of coconut palms and has recently spread to many coconut production areas worldwide. Colonization of coconut palms is thought to arise from mites dispersing aerially after take-off from other plants within the same plantation or other plantations. The underlying dispersal behaviour of the mite at take-off, in the airborne state and after landing is largely unknown and this is essential to understand how they spread from tree to tree. In this article we studied whether take-off to aerial dispersal of coconut mites is preceded by characteristic behaviour, whether there is a correlation between the body position preceding aerial dispersal and the direction of the wind, and whether the substrate (outer surface of coconut bracts or epidermis) and the wind speed matter to the decision to take-off. We found that take-off can sometimes be preceded by a raised body stance, but more frequently take-off occurs while the mite is walking or resting on its substrate. Coconut mites that become airborne assumed a body stance that had no relation to the wind direction. Take-off was suppressed on a substrate providing food to coconut mites, but occurred significantly more frequently on the outer surface of coconut bracts than on the surface of the fruit. For both substrates, take-off frequency increased with wind speed. We conclude that coconut mites have at least some degree of control over take-off for aerial dispersal and that there is as yet no reason to infer that a raised body stance is necessary to become airborne.

Fitness costs associated with low-level dimethoate resistance in Phytoseiulus macropilis.

Phytoseiulus macropilis Banks (Acari: Phytoseiidae) is an effective predator of tetranychid mites, but there are no data on its response to pesticides. We investigated the resistance of the predatory mite P. macropilis to the acaricides abamectin and dimethoate, and we examined the fitness costs associated with resistance. Two populations were tested: one from conventional cultivation and another from an area not commercially exploited. After the application of acaricides to the predator, we determined the lethal effects of the acaricides, the instantaneous rate of population increase (r(i)), the predation on Tetranychus urticae Koch (Acari: Tetranychidae) and its ability to locate prey in an olfactometer. P. macropilis exhibited resistance to dimethoate only. The low level of resistance (9.4x) of the predator did not affect their ability to locate prey. However, the dimethoate resistant population was not as effective in contatining prey population when in lower density and exhibited a more pronounced decrease of r(i) in the presence of this acaricide, due to the reduced oviposition of the predator, a likely consequence of the different genetic background of this population.

Effects of Acaricides on the Functional and Numerical Responses of the Phytoseid Predator Neoseiulus idaeus (Acari: Phytoseiidae) to Spider Mite Eggs.

Integrated control tactics are often necessary for pest management. This is especially true for organisms such as the two-spotted spider mite, Tetranychus urticae Koch. The management of this mite pest species relies on pesticide use, but its short life cycle associated with high selection pressure results in frequent problems of acaricide resistance and population outbreaks. Therefore, combining acaricides and natural enemies is an appealing strategy for managing this pest species. The predatory mite Neoseiulus idaeus Denmark & Muma (Phytoseiidae) is important in arid environments, where other natural enemies show low efficacy. Thus, we investigated the effects of representative acaricides used for managing spider mites around the world in several crops (i.e., abamectin, fenpyroximate, and azadirachtin), on the functional and numerical responses of the phytoseid predator N. idaeus to increasing egg densities of its prey. Acaricide exposure did not affect the type of N. idaeus functional response or attack rate (a). However, acaricide exposure decreased the amount of consumed prey and increased prey handling time (Th). All acaricides affected the numerical response of the predator, which reduced oviposition rates. Therefore, caution is required in attempts to integrate the control methods.

Which Species of Coconut Moth Occurs in Brazil: Atheloca subrufella vs. Atheloca bondari (Lepidoptera: Pyralidae)?

The moths, Atheloca subrufella (Hulst 1887) and A. bondari, (Heinrich 1956) are species known for their economic impact on coconut production, which Brazil is the fourth largest global producer. The first record of Atheloca in Brazil was reported by Bondar in 1940, where the author registered it being A. subrufella. The studies performed by C. Heinrich in 1956 related the existence of divergence in specimens of Brazilian Atheloca suggesting the presence of morphological differences between the males of A. bondari and A. subrufella. In this study, Atheloca specimens from the five states of northeastern Brazil were used. Samples from Pernambuco state were sent to taxonomist Dr. V. O. Becker (Uiraçu Institute-BA) for identification. Male individuals from the other states were mounted for photographic documentation, highlighting the characteristics that differentiate the two species. A fragment of the cytochrome c oxidase subunit 1 gene was sequenced and then compared with that of the Atheloca spp. deposited in GenBank. An analysis was conducted to evaluate the genetic distance between A. bondari and A. subrufella. The results indicate greater interspecific (0.030-0.034) than intraspecific (0.000-0.002) genetic variation between the groups, reinforcing the hypothesis of two distinct species. A geographic distribution map and a table with the host plants were constructed based on a literature review. This study concluded that the species occurring in Brazil is A. bondari, as suggested by C. Heinrich. Atheloca bondari and A. subrufella have only been reported in plants of the family Arecaceae, but only the coconut tree (Cocos nucifera L.) is shared by the two species.

Alternative food sources for the ladybird Brumoides foudrasii (Mulsant) (Coleoptera: Coccinellidae).

The predator Brumoides foudrasii (Mulsant) (Coleoptera, Coccinellidae) has been naturally found in plants infested by mealybugs. In this study, the striped mealybug Ferrisia dasylirii (Cockerell) (Hemiptera, Pseudococcidae) and Anagasta kuehniella Zeller (Lepidoptera, Pyralidae) eggs were evaluated as diets for the development and reproduction of B. foudrasii. Brumoides foudrasii immatures developed faster when fed with A. kuehniella eggs than when fed with F. dasylirii. However, the survival and longevity of B. foudrasii adults were not affected by the diets. Oviposition was more frequent when B. foudrasii females were fed with F. dasylirii (95%) than those fed with A. kuehniella eggs (65%). Brumoides foudrasii females fed with F. dasylirii were 2.5 times more fecund than those fed with A. kuehniella eggs. Although both diets may be considered proper for B. foudrasii, A. kuehniella eggs were more suitable for immature development, while F. darsyrili provided more nutritious resources for adult biological parameters as oviposition period, egg viability and fecundity. These results suggests that this predator can play an important role in regulating populations of the stripped mealybug F. dasylirii in the field.

Alternative food sources for the ladybird Brumoides foudrasii (Mulsant) (Coleoptera: Coccinellidae) / Fontes de alimento alternativo para a joaninha Brumoides foudrasii (Mulsant) (Coleoptera: Coccinellidae)

Abstract The predator Brumoides foudrasii (Mulsant) (Coleoptera, Coccinellidae) has been naturally found in plants infested by mealybugs. In this study, the striped mealybug Ferrisia dasylirii (Cockerell) (Hemiptera, Pseudococcidae) and Anagasta kuehniella Zeller (Lepidoptera, Pyralidae) eggs were evaluated as diets for the development and reproduction of B. foudrasii. Brumoides foudrasii immatures developed faster when fed with A. kuehniella eggs than when fed with F. dasylirii. However, the survival and longevity of B. foudrasii adults were not affected by the diets. Oviposition was more frequent when B. foudrasii females were fed with F. dasylirii (95%) than those fed with A. kuehniella eggs (65%). Brumoides foudrasii females fed with F. dasylirii were 2.5 times more fecund than those fed with A. kuehniella eggs. Although both diets may be considered proper for B. foudrasii, A. kuehniella eggs were more suitable for immature development, while F. darsyrili provided more nutritious resources for adult biological parameters as oviposition period, egg viability and fecundity. These results suggests that this predator can play an important role in regulating populations of the stripped mealybug F. dasylirii in the field.

Resumo O predador Brumoides foudrasii (Mulsant) (Coleoptera, Coccinellidae) tem sido encontrado naturalmente sobre plantas infestadas por cochonilhas. Neste estudo, a cochonilha listrada Ferrisia dasyliri (Cockerell) (Hemiptera, Pseudococcidae) e ovos de Anagasta kuehniella Zeller (Lepidoptera, Pyralidae) foram avaliados como dietas para o desenvolvimento e reprodução de B. foudrasii. Formas amaturos de B. foudrasii desenvolveram-se mais rapidamente quando alimentados com ovos de A. kuehniella do que quando alimentados com F. dasyliri. No entanto, a sobrevivência e longevidade de adultos de B. foudrasii não foram afetadas pelas dietas. A oviposição foi mais freqüente em B. foudrasii alimentadas com F. dasylirii (95%) do que aqueles alimentados com ovos de A. kuehniella (65%). Fêmeas de B. foudrasii alimentadas com F. dasylirii foram 2,5 vezes mais fecundas do que aquelas alimentados com ovos de A. kuehniella. Embora ambas as dietas possam ser consideradas apropriadas para B. foudrasii, ovos de A. kuehniella foram mais adequadas para o desenvolvimento de imaturos, enquanto F. dasylirii proporcionou mais recursos nutritivos para os parâmetros biológicos do adulto, tais como período de oviposição, viabilidade de ovos e fecundidade. Estes resultados sugerem que este predador pode ser potencialmente eficiente na regulação populacional da cochonilha F. dasylirii em campo.