Morphological variation and reproductive incompatibility of three coconut-mite-associated populations of predatory mites identified as Neoseiulus paspalivorus (Acari: Phytoseiidae)

Predatory mites identified as Neoseiulus paspalivorus DeLeon (Phytoseiidae) have been considered as agents for classical biological control of the coconut mite, Aceria guerreronis Keifer (Eriophyidae), in Africa and elsewhere. Preliminary identification of geographically distinct populations as belonging to the same species (N. paspalivorus) was based on their morphological similarity. However, laboratory studies recently conducted have shown large differences in feeding behaviors and biological characteristics among individuals collected from three geographic origins: Brazil (South America), Benin and Ghana (West Africa). As morphologically similar specimens do not necessarily belong to the same species, we evaluated under laboratory conditions, reproductive compatibility between the specimens from three geographic locations to ascertain their conspecificity. Morphological measurements were also made to determine whether there is a means of discriminating between them. Inter-population crosses showed complete reproductive isolation between the three geographic populations, but interpopulation discontinuities in morphometric characters were absent. These results indicate that the tested specimens are distinct biological entities despite morphological similarity. Further molecular genetic studies are therefore proposed, including screening for endosymbionts and assessment of genetic differentiation, to determine the cause of reproductive incompatibility and to clarify the taxonomic relationship between those populations.

Life history of Proctolaelaps bulbosus feeding on the coconut mite Aceria guerreronis and other possible food types occurring on coconut fruits

Aceria guerreronis Keifer (Acari: Eriophyidae) is a major pest of coconut fruits (Cocos nucifera L.) in many countries of the Americas, Africa, and parts of Asia. Considerable attention has been given to studies of biological control agents of A. guerreronis. Proctolaelaps bulbosus Moraes, Reis and Gondim Jr. is a predator recently discovered in association with A. guerreronis. Nothing is known about its biology. The aim of this study was to determine suitable food sources for P. bulbosus, among items commonly found on coconut fruits, including A. guerreronis. Food sources evaluated included the mites A. guerreronis, Steneotarsonemus concavuscutum Lofego and Gondim Jr., and Tyrophagus putrescentiae (Schrank), the fungus Rhizopus aff. stolonifer (Ehrenb.) Vuill and coconut pollen; the mite Tetranychus urticae Koch was also included in the assessments, for being a commonly used prey for mass production and laboratory rearing of predatory mites. Proctolaelaps bulbosus was able to develop up to adulthood when fed A. guerreronis, R. aff. stolonifer and T. putrescentiae. It had the highest population growth rates when feeding on the former (R (o) = 17.5; r (m) = 0.392). These results indicate that A. guerreronis is the most suitable food for P. bulbosus among the possible food sources found on coconut fruits and that P. bulbosus can survive in the absence of eriophyid using R. aff. stolonifer as a food source.

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.

Morphological, molecular and cross-breeding analysis of geographic populations of coconut-mite associated predatory mites identified as Neoseiulus baraki: evidence for cryptic species?

Surveys were conducted in Brazil, Benin and Tanzania to collect predatory mites as candidates for control of the coconut mite Aceria guerreronis Keifer, a serious pest of coconut fruits. At all locations surveyed, one of the most dominant predators on infested coconut fruits was identified as Neoseiulus baraki Athias-Henriot, based on morphological similarity with regard to taxonomically relevant characters. However, scrutiny of our own and published descriptions suggests that consistent morphological differences may exist between the Benin population and those from the other geographic origins. In this study, we combined three methods to assess whether these populations belong to one species or a few distinct, yet closely related species. First, multivariate analysis of 32 morphological characters showed that the Benin population differed from the other three populations. Second, DNA sequence analysis based on the mitochondrial cytochrome oxidase subunit I (COI) showed the same difference between these populations. Third, cross-breeding between populations was unsuccessful in all combinations. These data provide evidence for the existence of cryptic species. Subsequent morphological research showed that the Benin population can be distinguished from the others by a new character (not included in the multivariate analysis), viz. the number of teeth on the fixed digit of the female chelicera.

Estimated crop loss due to coconut mite and financial analysis of controlling the pest using the acaricide abamectin.

Reducing the losses caused by Aceria guerreronis Keifer has been an arduous task for farmers. However, there are no detailed studies on losses that simultaneously analyse correlated parameters, and very few studies that address the economic viability of chemical control, the main strategy for managing this pest. In this study the objectives were (1) to estimate the crop loss due to coconut mite and (2) to perform a financial analysis of acaricide application to control the pest. For this, the following parameters were evaluated: number and weight of fruits, liquid albumen volume, and market destination of plants with and without monthly abamectin spraying (three harvests). The costs involved in the chemical control of A. guerreronis were also quantified. Higher A. guerreronis incidence on plants resulted in a 60 % decrease in the mean number of fruits harvested per bunch and a 28 % decrease in liquid albumen volume. Mean fruit weight remained unaffected. The market destination of the harvested fruit was also affected by higher A. guerreronis incidence. Untreated plants, with higher A. guerreronis infestation intensity, produced a lower proportion of fruit intended for fresh market and higher proportions of non-marketable fruit and fruit intended for industrial processing. Despite the costs involved in controlling A. guerreronis, the difference between the profit from the treated site and the untreated site was 18,123.50 Brazilian Real; this value represents 69.1 % higher profit at the treated site. Keywords

Diet-dependent life history, feeding preference and thermal requirements of the predatory mite Neoseiulus baraki (Acari: Phytoseiidae)

Neoseiulus baraki Athias-Henriot (Acari: Phytoseiidae) has been reported from the Americas, Africa and Asia, often in association with Aceria guerreronis Keifer (Acari: Eriophyidae), one of the most important pests of coconut (Cocos nucifera L.) in diVerent parts of the world. That phytoseiid has been considered one of the most common predators associated with A. guerreronis in Brazil. The objective of this study was to evaluate the feeding preference and the eVect of food items commonly present on coconut fruits and several temperature regimes on the life history of a Brazilian population of N. baraki. Completion of immature development was possible when N. baraki was fed A. guerreronis, Steneotarsonemus concavuscutum Lofego and Gondim Jr., and Tyrophagus putrescentiae (Schrank). Fecundity was highest on T. putrescentiae (39.4 eggs), followed by A. guerreronis (24.8 eggs). In choice tests, irrespective of the food on which N. baraki was reared, a larger number of adults of this predator chose leaf discs containing A. guerreronis than discs containing other food items, demonstrating a preference of the former for the latter as food. Egg to adult thermal developmental time was calculated as 84.2 degree-days, above a threshold of 15.8 degrees C. This lower developmental threshold is higher than previously published for phytoseiid species from higher latitudes. Neoseiulus baraki was shown to have higher biotic potential at 30 degrees C (r(m) 0.29). The results suggest N. baraki to be a promising biological control agent of A. guerreronis, well adapted to survive and develop in areas with relatively high temperatures, where that pest prevails.

Distribution of Aceria guerreronis and Neoseiulus baraki among and within coconut bunches in northeast Brazil

Aceria guerreronis Keifer (Acari: Eriophyidae) is considered a major pest of coconut in many countries in the Americas, Africa and parts of Asia. Neoseiulus baraki Athias-Henriot (Acari: Phytoseiidae) is one of the predatory mites most commonly found in association with A. guerreronis in parts of northeast Brazil. The objective of this work was to study the distribution of A. guerreronis and N. baraki among and within coconut bunches. The hypothesis was tested that A. guerreronis and N. baraki are homogenously distributed over the fruits in a bunch, independent of the fruits` age and position. Five collections of bunches, each corresponding to leaves 12-16 from apex (about 2-6 month-old), were conducted in each of three fields in northeastern Brazil, from February to October, 2007. A total of 1,986 fruits were examined. The number of mites, the percentage of fruits hosting them and the level of damage caused by A. guerreronis were evaluated. The highest density of A. guerreronis was observed on fruits of bunch 4 whereas the highest density of N. baraki was observed on bunch 5. Considering all fruits together, no significant differences were observed between densities of either A. guerreronis or N. baraki among the basal, median and apical thirds of the bunches. In younger bunches, fruits of the apical region tend to have lower densities of both mites than fruits of the basal region. This pattern, in association with a similar pattern for the percentage of fruits hosting N. baraki, suggests that the predator initially reaches the basal bunch region, from where it moves to the apical region. The results of the present study suggest that the pest population reduction in bunches older than bunch 4 could be due to (1) an effect of the predator, (2) reduction of the proportion of undamaged tissues amenable to attack, and/or (3) less favorable characteristics of the fruits to attack by A. guerreronis, as indicated by their increasing lignin content as they get older.

Status of Aceria guerreronis Keifer (Acari: Eriophyidae) as a Pest of Coconut in the State of Sao Paulo, Southeastern Brazil

The coconut mite, Aceria guerreronis Keifer, is one of the main pests of coconut palms (Cocos nucifera) in northeastern Brazil. The objective of this study was to evaluate the levels of the coconut mite and other mites on coconut palms in the state of So Paulo and to estimate the possible role of predatory mites in the control of this pest. The effect of cultivated genotypes and sampling dates on the mite populations was also estimated. We sampled attached fruits, leaflets, inflorescences, and fallen fruits. The coconut mite was the main phytophagous mite found on attached and fallen fruits, with average densities of 110.0 and 20.5 mites per fruit, respectively. The prevalent predatory mites on attached and fallen fruits were Proctolaelaps bulbosus Moraes, Reis & Gondim Jr. and Proctolaelaps bickleyi (Bram), both Melicharidae. On leaflets, the tenuipalpids Brevipalpus phoenicis (Geijsks) and Tenuipalpus coyacus De Leon and the tetranychid Oligonychus modestus (Banks) were the predominant phytophagous mites. On both leaflets and inflorescences, the predominant predatory mites belonged to the Phytoseiidae. Neoseiulus baraki (Athias-Henriot) and Neoseiulus paspalivorus (De Leon), predators widely associated with the coconut mite in northeastern Brazil and several other countries, were not found. The low densities of the coconut mite in So Paulo could be related to prevailing climatic conditions, scarcity of coconut plantations (hampering the dispersion of the coconut mite between fields), and to the fact that some of the genotypes cultivated in the region are unfavorable for its development.

Dispersal strategies of Aceria guerreronis (Acari: Eriophyidae), a coconut pest

The dispersal of plant-feeding mites can occur involuntarily, through transportation of infested plant parts, or voluntarily, by walking to new plant parts or to suitable spots where biotic (phoresis) or abiotic (wind, agricultural tools, etc.) factors carry them over long distances. Elucidating the dispersal mechanisms of the coconut mite, Aceria guerreronis Keifer, is important for understanding the process of colonization of new fruits of a same or different plants, essential for the improvement of control strategies of this serious coconut pest. Thus, the objective of this work was to investigate the voluntary dispersal mechanisms of this mite. The hypothesis that the coconut mite disperses by walking, phoresis or wind were tested. The coconut mite was shown to be able to walk short distances between fruits of the same bunch or between bunches of the same plant. Phoresis on insects of the orders Hymenoptera (Apidae), Coleoptera (Curculionidae) and Lepidoptera (Phycitidae) was evaluated in the laboratory and in the field. Although in the laboratory mites were shown to be able to climb onto honeybees, field investigations failed to show these insects as important carriers of the pest, corroborating findings of previous works; however, both laboratory and field investigations suggested the curculionid Parisoschoenus obesulus Casey to be able to transport the coconut mite between plants. Similarly, laboratory and field investigations suggested wind to be important in the dispersal of the coconut mite between plants.

Incidence of tarsonemid mites on Cocos nucifera L. (Arecaceae) from Oman with description of a new species of Nasutitarsonemus Beer and Nucifora (Acari: Tarsonemidae)

Coconut is an important crop grown in the coastal plain of the Dhofar region, south-eastern Oman, on the edge of the Arabian Peninsula desert. It holds a particular place in the landscaping of the region and is also of great interest for the production of coconut drinking water. One of the main pests of coconut in this region is the coconut mite (Aceria guerreronis Keifer). In surveys conducted to understand the dynamics of that mite and its association with other arthropods, the incidence of tarsonemid mites was determined. Steneotarsonemus furcatus de Leon was the most commonly found tarsonemid on fruits as well as on growing tips of coconut seedlings, always at low levels. A few representatives of an undescribed tarsonemid species were also found. That new species is here described as Nasutitarsonemus omani Lofego and Moraes, sp. nov. A key to the species of this little-known genus is provided.

Limitations of Neoseiulus baraki and Proctolaelaps bickleyi as control agents of Aceria guerreronis

Several predatory mites have been found in association with the coconut mite, Aceria guerreronis Keifer, in northeast Brazil. However, the latter still causes damage to coconut in that region. The objectives of this work were to compare the frequencies of occurrence of Neoseiulus (Phytoseiidae) and Proctolaelaps (Melicharidae) species on standing and aborted coconuts in coastal Pernambuco State, northeast Brazil and to analyze their possible limitations as control agents of the coconut mite, based on evaluations of the restrictions they may have to access the microhabitat inhabited by the pest and their functional and reproductive responses to increasing densities of the latter. Neoseiulus baraki (Athias-Henriot) was found mostly on standing coconuts whereas Proctolaelaps bickleyi (Bram) was found mostly on aborted coconuts. Measurements of the entrance to the microhabitat occupied by the coconut mite, between the bracts and the subjacent fruit surface, showed that this different pattern of predator prevalence could be related to predator sizes, although other environmental factors could not be disregarded. Progressively higher predation rate of N. baraki was observed up to an experimental density that corresponded to 1,200 coconut mites per fruit, which is close to the average number determined in northeast Brazil, reducing slightly afterwards. Predation rate of P. bickleyi reduced consistently but slightly with increasing prey densities, but in absolute values, rates were always much higher than determined for N. baraki. The excessively high killing capacity of P. bickleyi, probably related to its high feeding requirement, may be detrimental in terms of stability. In fact, such high requirement for food suggests that P. bickleyi might not have a strong relation with the coconut mite and that the latter may not be its main food source under natural conditions. It is concluded that body sizes of both predators and the exceedingly high feeding requirement of P. bickleyi may limit their performance as control agents of the coconut mite.

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 Para 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 ill moderately tempered and humid areas. We discuss the significance of our findings for natural and biological control of A. guerreronis.

Population trends of the red palm mite, raoiella indica hirst (acari: tenuipalpidae) and associated entomopathogenic fungi in trinidad, antigua, st kitts and nevis and dominica

A survey to determine population trends and entomopathogenic fungi associated with the red palm mite (RPM), Raoiella indica, was conducted in Trinidad, Antigua, St. Kitts and Nevis and Dominica. RPM population density was evaluated by sampling a total of ten coconut palms per site in Antigua, St. Kitts and Nevis, Dominica, and Trinidad (Manzanilla and Icacos). Mites from the four islands were either surface sterilized or left unsterilized before being cultured on Tap Water Agar (TWA). A total of 318 fungal colonies were retrieved. A further 96 mites from Dominica were kept on sterile moist filter paper in a humidity chamber and a further 85 colonies were isolated. Based on morphological observations of all 403 isolates, a sample consisting of 32 colonies (8 %) was sent for identification at CABI-UK. Of the 27 fungi positively identified, 15 isolates belonged to the genera Cladosporium, three to Simplicillium spp., and one to Penicillium. Other fungi genera with limited or no entomopathogenic potential included: Aspergillus, Cochliobolus, Fusarium, Pestalotiopsis and Pithomyces. The results show a potential use of entomopathogenic fungi for population management of the red palm mite in the Caribbean region.

Can the red palm mite threaten the Amazon vegetation?

The red palm mite Raoiella indica Hirst (Tenuipalpidae) was first reported in the New World in 2004, dispersing quickly and widely while adopting new plant species as hosts. Since then, it has caused severe damage in this region, especially to coconut (Cocos nucifera L.). It was first found in Brazil in 2009, in the northern Amazonian state of Roraima. In the present study, native and introduced plants were sampled between March 2010 and February 2011 in sites of the 15 Roraima municipalities, to estimate its distribution and the associated mite fauna. In addition, monthly samples were taken from a coconut plantation in Mucajai throughout the same period, for an initial appraisal of the levels R. indica could reach. It was found in 10 municipalities, on 19 plant species of four families. Six species are reported for the first time as hosts. Among the associated predators, 89.1% were Phytoseiidae, most commonly Amblyseius largoensis (Muma), Iphiseiodes zuluagai Denmark & Muma and Euseius concordis (Chant). The highest densities of R. indica, 1.5 and 0.35 mites/cm2 of leaflet (approx total of 331 and 77 mites/leaflet), were reached respectively in March 2010 and February 2011. The highest density of phytoseiids on coconut (0.009 mites/cm2 or about 2 mites/leaflet) was reached in November 2010. The average densities of R. indica recorded for Roraima were comparable to those reported for countries in which the mite is reportedly economically damaging. The dispersal of R. indica through the Amazon forest may result in damage to cultivated and native palms, and plants of other families, if the projected increase in both the frequency and the severity of drought events occurs. Parts of the Amazon have undergone periods of low rainfall, a condition that appears to favour the biology of this mite. Its eventual arrival to northeastern Brazil may result in heavy economic and ecological losses.

Amblyseius largoensis in controlling red palm mite under semi-field conditions.

The objective of this work was to evaluate the efficiency of two Amblyseius largoensis (Acari:Phytoseiidae) populations in controlling Raoiella indica (Acari: Tenuipalpidae). The treatments were: release of A. largoensis from the island of La Réunion; release of A. largoensis from the state of Roraima, Brazil; and a control, without predator release. Initially, 20 predators were released per plant; three other releases were done at a rate of ten adults per plant, at 46, 135, and 156 days after the first release. The population densities were estimated every 20 days, during six months. Both A. largoensis populations evaluated are not sufficiently efficient to control the R. indica population.