Reducing losses inflicted by insect pests on cashew,using weaver ants as a biological control agent

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Effect of supplementary feeding of Oecophylla longinoda on their abundance and predatory activities against cashew insect pests

Many studies have shown the efficiency of using weaver ants (Oecophylla species) as natural biocontrol agents against agricultural pests. Supplementary feeding could promote fast growth of this ant's population and discourage them from moving away. However, such artificial feeding might slow down ants search rates and in this way make them less efficient bio-agents. The experiments were conducted for two consecutive seasons at Naliendele Research Station. Cashew trees planted at a spacing of 12 m × 12 m in 2002 were used to investigate whether supplementary feeding could enhance foraging behaviour of Oecophylla longinoda. Fed O. longinoda colonies fed weaver ants (FWA) were supplemented with a 30% sugar solution and approximately 22 g of finely ground fish meat at two-week intervals while the unfed colonies unfed weaver ants (UWA) had access to only naturally occurring food sources. Weaver ant densities and pest damage were monitored fortnightly on newly damaged shoots, panicles and fruits and nut yields assessed after each harvest season. The results revealed that there was a significant difference (P < 0.05) with higher weaver ant densities in the FWA compared to UWA colonies and significantly lower (P < 0.05) pest damage levels were recorded on weaver ant treatments compared to plots without weaver ants. No significant differences (P > 0.05) in yields and mean damage levels were recorded between the two weaver ant treatments. Highest nut yield (4.22 ± 0.30 kg/tree and 5.37 ± 0.27 kg/tree) was recorded in the fed colonies, followed by non-fed colonies (4.20 ± 0.30 kg/tree and 4.88 ± 0.24 kg/tree) and the least (2.66 ± 0.19 kg/tree and 2.99 ± 0.19 kg/tree) was recorded from the untreated controls in 2012/2013 and 2013/2014, respectively. The studies indicated that supplementary feeding could boost weaver ants to higher population levels without reducing their effectiveness as biocontrol agents.     

Seasonal pattern in food gathering of the weaver ant Oecophylla longinoda (Hymenoptera: Formicidae) in mango orchards in Benin

Prey capture and food scavenging activities of Oecophylla longinoda were monitored through regular weekly samplings during two consecutive years (2009–2010) in a large mango orchard of the Borgou Department of Benin, West Africa, a main mango production area located in the Sudan agro-ecological zone. In both years, interspecific competition with other ants occurred mainly during the dry season (January to March) resulting in increased captures of Formicinae, Myrmicinae and Ponerinae. More prey was caught during the rainy season (end April to end October) than during the rest of the year, with Diptera and Coleoptera prey peaking in May and June, respectively, along with the mango season. As insect prey quickly decreased during November to December weaver ants increasingly collected seeds and plant debris. A total of 241 species of insects were captured including 61 species (25.3%) associated with mango and a few with cashew, among which 48 mango pest species (78.7% of species associated with mango tree). Only five species (2.1%) of beneficial insects were captured. It is concluded that the presence of O. longinoda colonies is beneficial to perennial tree cropping systems such as mango and cashew.     

The effect of weaver ants Oecophylla smaragdina on the shoot borer Hypsipyla robusta on African mahoganies in Australia

• 1 African mahogany Khaya senegalensis is a high‐value timber tree species widely grown in central Africa, south‐east Asia and northern Australia. Pilot plantings show that the tree grows well in the wet‐dry tropical areas of northern Australia, and the shoot borer Hypsipyla robusta (Lepidoptera: Pyralidae) is a potential pest of the tree. The weaver ant Oecophylla smaragdina is an efficient biocontrol agent in some horticulture crops. To investigate whether the ants control shoot borers, field experiments were conducted at two sites near Darwin, Australia from April 2006 to January 2009.
• 2 In the weaver ant treatments, the overall percentage of trees damaged by shoot borers was 0–2.7% at Berrimah Farm and 0–4.2% at Howard Springs, and the damaged trees were attacked once only. In the treatments without weaver ants, however, 9.9–52.1% trees were damaged at Berrimah Farm, and 6.3–64.6% at Howard Springs, and the damaged trees were generally attacked more than once.
• 3 At both sites, significantly fewer trees on each monitoring occasion were damaged in weaver ant treatments than in treatments without weaver ants.
• 4 The mean percentage of overall flushing shoots damaged by the pest at both sites was significantly lower in weaver ant treatments compared with treatments without weaver ants.
• 5 Fewer shoots were damaged per damaged tree in weaver ant treatments compared with treatments without weaver ants.
• 6 The data obtained suggest that weaver ants were effective biological control agents of the shoot borer, and that the ants can be used to manage the pest on African mahogany trees.

     

Using insect traps to increase weaver ant (Oecophylla longinoda) prey capture

Weaver ants (Oecophylla spp.) are managed in plantations to control insect pests and are sometimes harvested as a protein‐rich food source. In both cases, the amount of insect prey caught by the ants is imperative for returns, as more prey leads to more effective biocontrol and to a higher production of ants. Malaise‐like traps placed in trees may catch flying insects without catching ants, as ants may use pheromone trails to navigate in and out of the traps. Thus, ants may increase their prey intake if they are able to extract insects caught in traps. In a mango plantation in Tanzania, we estimated the amount of insects caught by simple traps (cost per trap = 3.9 USD), and whether Oecophylla longinoda was able to collect insects from them. On average, a trap caught 110 insects per month without catching any weaver ants. The number of insects found in traps with ant access was 25% lower than in control traps (ants excluded), showing that ants were able to gather prey from the traps. Ant activity in traps increased over time, showing that prey extraction efficiency may increase as ants customize to the traps. The prey removed from traps by ants constituted 5% of the number of prey items collected by O. longinoda under natural conditions (without traps), potentially increasing to 14% if ants learn to extract all insects. Thus, prey intake may be increased with 5–14% per 3.9 USD invested in traps. These numbers increased to 38 and 78%, respectively, when light was used to attract insects during night time. Combining ant predation with insect trapping is a new approach potentially building increased returns to ant biocontrol and to ant entomophagy.     

At home with the birds: Kalahari tree skinks associate with sociable weaver nests despite African pygmy falcon presence

The way in which animals use habitat can affect their access to key resources or how they are buffered from environmental variables such as the extreme temperatures of deserts. One strategy of animals is to modify the environment or take advantage of structures constructed by other species. The sociable weaver bird (Philetairus socius) constructs enormous colonial nests in trees. These nests are frequented by Kalahari tree skinks (Trachylepis spilogaster) and the two species coexist over a large portion of the Kalahari Desert in South Africa. We tested whether skinks were more abundant in trees containing sociable weaver nests and asked whether the physical features of trees were important predictors of skink abundance. We then focused on potential costs of this association by examining the relationship between skink abundance and the presence of a potential predator, the pygmy falcon (Polihierax semitorquatus), which exclusively uses weaver nest colonies for roosting and nesting. Finally, we simulated a predatory threat to determine if skinks assess risk differently if a weaver nest is present. We found a significant positive association between the presence of weaver nests and skink abundance. In the absence of nests, the type of tree did not influence skink abundance. Skinks used weaver nests and were more likely to perch on the nest than the tree. When threatened with predation, skinks preferred to take refuge in nests. Surprisingly, the presence of nesting pygmy falcons in nests did not influence skink abundance, perhaps because of the abundance of nearby refuges within nests, tree crevices, or in debris at the tree base. We suggest that sociable weaver nests provide multiple benefits to skinks including lowered predation risk, thermal refuges and greater prey availability, although this requires experimental testing. In the current era of global climate change, sociable weaver nests may become a crucial resource for skinks seeking refuge as the Kalahari climate warms.     

Host-ant trail following by myrmecophilous larvae of Liphyrinae (Lepidoptera,Lycaenidae)

In this study we report a case of ant-trail following by lycaenid caterpillars. Euliphyra mirifica and E. leucyana caterpillars are involved in a commensal association with the weaver ant Oecophylla longinoda. The host nests are made with leaves which over the course of time dry out or are broken open by storms, forcing the ants to migrate and build a new nest elsewhere. Euliphyra caterpillars are stimulated by recruitment behaviour which triggers the migration of their host. They then follow the host trails leading to the new nesting site. Laboratory experiments showed that these caterpillars are able to follow host trails under varied conditions: (1) fresh trails actually used by workers, (2) fresh trails in the absence of workers, (3) heterocolonial, 2-month-old trails, and (4) fresh trails washed with water (to simulate the effect of tropical rains). They can also bridge trail gaps of more than 1 cm. Under natural conditions, the trails are frequently situated along thin twigs. The forward progress of the ants in such a situation is not impeded by the presence of large Euliphyra larvae. Workers just climb over the caterpillars, even on larger trails where there is enough room to pass alongside them. This suggests that an allomone is secreted on the dorsal part of the caterpillars. When crawling along heterocolonial trails, the caterpillars are not attacked, even if about 21% of the workers from the new colony spread their mandibles when encountering them. They are then adopted and are admitted to the nest of the new host colony of O. longinoda.     

Comparing different methods for trapping mated queens of weaver ants (Oecophylla longinoda; Hymenoptera: Formicidae)

The predatory efficiency of African weaver ants Oecophylla longinoda and their utilisation in protein production is a function of ant abundance. Reliable control of insect pests in tropical crops is achieved when ant populations are constantly high. Transplanted populations of weaver ant colonies containing egg-laying queens are more stable than those without. Achieving such stability through collection of colonies established in the wild is usually difficult because of uncertainty in locating the nest containing the egg-laying queen. In this study, we investigated four methods that may be used to collect mated queens that subsequently can be used to stock ant nurseries. The catch efficiencies of (1) leaf traps, (2) paper traps (both types providing a refuge for founding queens), (3) random search for queens and (4) light trapping were compared. Light trapping was the most efficient way to collect queens followed by leaf traps, random search and, last, paper traps. Light trapping and random search, though, required the presence of a person throughout the ant's mating season (several months), whereas this was not required when using leaf and paper traps.     

Biological control of the fruit‐spotting bug Amblypelta lutescens using weaver ants Oecophylla smaragdina on African mahoganies in Australia

• 1 African mahogany Khaya senegalensis is a high‐value timber tree. Pilot plantings showed that the fruit‐spotting bug Amblypelta lutescens causes severe damage of the tree in the wet–dry tropics of northern Australia. The weaver ant Oecophylla smaragdina is an efficient biocontrol agent in some horticulture crops. To investigate whether the ants control this pest, field experiments were conducted from April 2006 to January 2009 at two study sites in the Darwin area, Australia. A laboratory experiment was carried out in March 2007 at Berrimah Farm.
• 2 During the experimental period, in the weaver ant treatments, the overall percentage of trees damaged by the pest was 0–8% at both sites, and the damaged trees were attacked once only. In the treatments without weaver ants, however, the damage level was > 80% at Berrimah Farm and 31–100% at Howard Springs, and the damaged trees were attacked more than once.
• 3 The mean percentage of trees damaged per monitoring occasion was 0–2.6% in the weaver ant treatments at both sites, whereas, in the treatments without the ants, the damage percentages were 14.2–27.0% at Howard Springs and 28.2–48.6% at Berrimah Farm.
• 4 Extrafloral nectar of African mahoganies is attractive to weaver ants. Fruit‐spotting bugs only damage the tender parts of flushing shoots and growing tips. Weaver ants live on sugar solution and meat, and they frequently harvest extrafloral nectar on growing shoots, on which they catch nymphs of the pest for their meat supply. The aggressive behaviour of the ants also repels the pest away from flushing shoots.
• 5 The data suggest that weaver ants were effective biocontrol agents of fruit‐spotting bugs, and the ants can be used to manage the pest on African mahoganies.
• 6 The present study demonstrates that the introduced African mahogany comprises another major host of the fruit‐spotting bug.

     

THE BREEDING OF HEUGLIN'S MASKED WEAVER AND ITS NESTING ASSOCIATION WITH THE RED WEAVER ANT

Grimes, L. G. 1973. The breeding of Heuglin's Masked Weaver and its nesting association with the Red Weaver Ant. Ostrich 44: 170–175.

The breeding season of Heuglin “s Masked Weaver Ploceus heuglini at Legon began in the latter half of the major dry season in January and February; continued through the main wet season and ceased in late August and for the rest of the year. The Accra colony had a similar breeding activity but in addition was active for a short period in November which had ended by mid-December. Although the majority of males formed colonies in which the number of males ranged from two to twenty, solitary breeding occurred equally frequently. Most males within the Legon colony, and possibly this is true for all colonies, were polygynous while most solitary males were monogamous. More nests were built by each male than the number of females involved in the polygyny.

A close nesting association with the Red Weaver Ant Oecophylla longinoda existed and the data suggested that the weaver sought the ant. Two cases were found of nesting association with the wasp Belanogaster grisens.     

Territories of the African Weaver Ant (Oecophylla longinoda [Latreille]); A Field Study

The African weaver ant (Oecophylla longinoda) is known to be highly aggressive toward conspecific aliens. In the study area Shimba Hills Reserve (Kenya) individual territories sometimes covered an area of up to approximately 1600 m², comprising 17 major trees. The territorial defense is organized by an elaborate defense recruitment system, which can also be aimed against several other ant species, which are potential competitors of Oecophylla for essential resources. This selective “enemy identification” seems to be the major behavioral mechanism by which the mosaic distribution of ecologically dominant ants is regulated.     

Persistent Intercolonial Trunkroute-marking in the African Weaver Ant Oecophylla longinoda Latreille (Hymenoptera,Formicidae): Tom Thumb's versus Ariadne's Orienting Strategies

The territorial African weaver ant Oecophylla longinoda forages in a 3-dimensional system when attending honeydew secreted by coccid colonies. The orienting strategy allowing workers to reach a static food resource was studied in laboratory conditions. Contrary to other studies devoted to territoriality in O. longinoda, our experiments show that, when moving off the nest, territorial faecal marking by the major workers is not randomly placed all over the home-range area. The chemical trail leading from the nest to a given food site is reinforced by faecal materials both on the horizontal and the vertical planes. In addition to visual cues and chemical trails laid by the major workers (Holldobler & Wilson 1978), foragers use the territorial marking both to localize the food site and to come back to the nest. Thus, anal-drop deposition in O. longinoda also has a clear dual-purpose function: territorial and orientationai marking. The ecological value of such a discrete Tom Thumb's orienting strategy enables workers to quickly reach a food location even after more than 5 months of inactivity on this given site. The persistent marked trail also has an intercolonial effect. This could allow the keeping of exploitation of a definite food site by alien colonies of the same species by reducing time costs for new exploring phases or for learning new spatial cues when foraging on an unknown environment.     

The Antsy Social Network: Determinants of Nest Structure and Arrangement in Asian Weaver Ants

Asian weaver ants (Oecophylla smaragdina) are arboreal ants that are known to form mutualistic complexes with their host trees. They are eusocial ants that build elaborate nests in the canopy in tropical areas. A colony comprises of multiple nests, usually on multiple trees, and the boundaries of the colony may be difficult to identify. However, they provide the ideal model for studying group living in invertebrates since there are a definite number of nests for a given substrate, the tree. Here, we briefly examine the structure of the nests and the processes involved in the construction and maintenance of these nests. We have described the spatial arrangement of weaver ant nests on trees in two distinct tropical clusters, a few hundred kilometres apart in India. Measurements were made for 13 trees with a total of 71 nests in the two field sites. We have considered a host of biotic and abiotic factors that may be crucial in determining the location of the nesting site by Asian weaver ants. Our results indicate that tree characteristics and architecture followed by leaf features help determine nest location in Asian weaver ants. While environmental factors may not be as influential to nest arrangement, they seem to be important determinants of nest structure. The parameters that may be considered in establishing the nests could be crucial in picking the evolutionary drivers for colonial living in social organisms.     

Bioecology of some key cashew insect pests and diseases in diverse habitats and landscapes in Tanzania

Cashew (Anacardium occidentale) is an economically important cash crop for many rural households in Tanzania. However, its production is constrained by some insect pests and diseases. As a prerequisite for the development of a more sustainable integrated insect pest and disease management strategy for cashew, information on the biology and ecology of the key insect pests and diseases in a changing environment, and on influencing biotic and abiotic factors, is needed. Surveys were conducted in the major cashew nut‐producing areas of Tanzania for two seasons: August to December, 2009, and August to December, 2010. Data on number of infested and infected shoots by key insect pests and diseases, natural enemies and associated farmer practices, namely synthetic pesticide use and intercropping systems, were collected from different subzones within agroecological zones. Our data showed that abundance and diversity of key cashew insect pests and diseases were influenced by agroecological zones and subzones. Intercropping was more commonly practised in the northern than in the southern zone. Agrochemicals were most frequently used in the southern agroecological zone and affected the occurrence of natural enemies, notably the weaver ant that was more abundant in the northern zone. Furthermore, our findings revealed that Helopeltis sp. and the powdery mildew remained the major constraints to cashew nut production in Tanzania.     

Efficacy of African weaver ant,Oecophylla longinoda (Hymenoptera: Formicidae) in reducing losses due to frugivorous fruit flies (Diptera: Tephritidae) in smallholder mango production systems in Eastern Tanzania

Fruit flies (Diptera: Tephritidae) are dipteran insects that cause high losses of fruits and vegetables in the tropical and subtropical regions of the world. As quarantine pests, fruit flies affect export markets because trade regulations hamper export of fruits from areas with occurrence of fruit flies. We compared the effectiveness of Oecophylla longinoda with the insecticide; imidacloprid for the control of Bactrocera dorsalis in a 6-year-old orchard of mango cultivar ‘Apple’. Mango samples at different developmental stages were collected between September and November in 2012 and between November 2013 and January 2014 at a three-week interval between sampling dates. Every fruit was cultured in a separate individual plastic container containing approximately 500?g sieved sterilised sand. Three weeks later, larvae and pupae were sieved from sand and counted. In both years, significantly higher (P?<?.0001) incidences and infestation rates were recorded in the control plots when compared to the trees protected with insecticides and O. longinoda. Incidences and infestation rates did not differ significantly (P?>?.05) between insecticide and weaver ants protected fruits, implying that O. longinoda was as effective as imidacloprid in suppressing the population of fruit flies. Peak fruit fly incidences and infestation rates coincided with the ripening stage of mango fruits. Our results have shown that the incidences and infestation rates on incubated mango fruits were lowest in the O. longinoda and the imidacloprid protected trees. Thus, O. longinoda is an effective bio-control agent that could be exploited to serve as a vital Integrated Pest Management component on mango orchards.     

Control of mango seed weevils (Sternochetus mangiferae) using the African Weaver Ant (Oecophylla longinoda Latreille) (Hymenoptera: Formicidae)

The mango seed weevil, Sternochetus mangiferae (Fabricius), is among the major threats to mango production in Tanzania. Sternochetus mangiferae is primarily a quarantine pest whose presence inside the fruits restricts access to new foreign markets and leads to rejections of fruits destined for export. Management options for the pest have largely been dependent on field sanitation and application of synthetic insecticides with some success. Thus, more sustainable methods are needed to substitute insecticides, as this may also open up opportunities for organic markets. We conducted field experiments for two fruiting seasons in a mango plantation at Mlandizi, Kibaha district, along the coastal belt of Tanzania to evaluate and compare the effectiveness of the predaceous ant Oecophylla longinoda Latreille with foliar insecticidal sprays of Dudumida (70 WDG Imidacloprid) in controlling S. mangiferae. Mango seed weevil infestation was assessed fortnightly based on infestation marks on developing fruits starting eight weeks after fruit set to early ripening phase. Between 50 and 64 fruits were sampled, well labelled in jute bags, secured and transported to the laboratory at Kibaha Biological Control Unit (KBCU) for incubation at room temperature using rearing transparent containers. Two weeks later, the fruits were dissected and inspected for the presence of S. mangiferae developmental stages. Field and laboratory results indicated that fruits from trees that were occupied by O. longinoda and from those treated with insecticide showed significantly (P < 0.0001) lower incidences and infestation rates by S. mangiferae than from untreated trees. Furthermore, there were no significant differences between the insecticide and the weaver ant treatments. We conclude that in our experiments, O. longinoda is an efficient biological control agent for a long‐term control programme and is comparable to insecticide (Dudumida) in suppressing S. mangiferae and may be used in Tanzanian mango plantations.     

Birds adopting weaver nests for breeding in Africa

Weavers build domed, long-lasting nests that may also be adopted by other species for breeding, probably reducing the energetic costs of nest building to varying degrees. In an extensive literature search, 57 species were found to have at least one record of adopting a weaver nest. There is one known obligate nest user, the Pygmy Falcon Polihierax semitorquatus. Four species were classed as near obligate nest adopters. Four species were listed as common, 10 as occasional, and 38 species as rare nest adopters. Other than the falcon and lovebirds Agapornis species, these nest adopters are passerines. Of the 57 species of nest adopters, 35 species had confirmed eggs and/or chicks found in the weaver nests. Most nest adopter species were in the Estrildidae family (20 species), with three estrildids classed as near obligates. This was followed by the Muscicapidae family, species that build cup nests. By building their cup nests inside weaver nests, there is likely protection from adverse weather and predators. Overall, there appeared to be a large diversity of nest adopter species for the traditional savanna living weavers (Ploceus, Bubalornis, Anaplectes, Plocepasser and Philetairus).     

Raptors breeding on weaver nests in trees and on man-made structures

Several raptor species nest on top of large weaver nests. These weaver nests are usually sited in trees, but 11.7% of Red-billed Buffalo-Weaver Bubalornis niger and 25.7% of Sociable Weaver Philetairus socius nests occur on man-made structures. In an extensive literature search, a total of 16 raptor species were recorded as nesting on top of weaver nests. At least 10 raptor species used weaver nests built in trees. Seven raptor species used weaver nests on man-made sites and four raptor species only used weaver nests built on man-made sites. No owls have been recorded as using weaver nests on man-made sites. There are historical records of raptors nesting on top of weaver nests in trees, while nesting on top of weaver nests sited on man-made structures appears to be a more recent adaptation. Costs and benefits of nesting on man-made sites are briefly listed. Nesting on man-made sites may increase (by both weavers and raptors) and raptor researchers are encouraged to document cases of raptors nesting on weaver nests where these are placed in trees or artificial sites, so that there is a record of changes in frequency of nest site usage by raptors.     

Weaving resistance: silk and disease resistance in the weaver ant <Emphasis Type="Italic">Polyrhachis dives</Emphasis>

Social insects are at risk from a diverse range of parasites. The antibiotic-producing metapleural gland is an ancestral trait in ants which is thought to be one of their primary mechanisms of resistance. However, the metapleural gland has been lost secondarily in three ant genera, which include weaver ants that are characterised by the remarkable construction of their nests using larval silk. Silken nests may have allowed reduced investment in costly disease resistance mechanisms like the metapleural gland if the silk has antimicrobial properties, as in other insects, or is a hygienic substrate. Here we examine this hypothesis in the weaver ant Polyrhachis dives. We found no evidence of a beneficial effect of silk. The presence of silk did not improve the already high resistance of ants to the entomopathogenic fungus Metarhizium, the ants only rarely interacted with the silk regardless of whether they were exposed to Metarhizium or not, and silk also did not inhibit the in vitro germination or growth of Metarhizium. Furthermore, silk was found in vitro to be heavily contaminated with the facultative entomopathogenic fungus Aspergillus flavus, and many more ants sporulated with this fungus when kept with silk in vivo than when they were kept without silk. Further work is needed to examine the effects of silk on other parasites and of silk from other weaver ants. However, the results in combination suggest that silk in P. dives is unlikely to provide protection against parasites and that it is also not a hygienic substrate. Alternative explanations may therefore be needed for the loss of the metapleural gland in weaver ants.     

Ecology of the fruit spotting bug, Amblypelta lutescens lutescens Distant (Hemiptera: Coreidae) in cashew plantations, with particular reference to the potential for its biological control

Abstract  The fruit spotting bug, Amblypelta lutescens lutescens, is one of the principal insect pests of cashews in Australia. Its population dynamics were studied using field observations and long-term monitoring to find suitable management methods. Observations of bugs reared in netting bags showed a sequence of change in bug-damage symptoms after 12 h and up to 3 d. Field observations revealed that adults preferred to feed and rest on the shady side of the tree. The number of bugs observed accounted for only 17–35% of the total variability in the number of damaged shoots, suggesting that the number of flushing shoots (leaf, flower or young nuts) with fresh damage symptoms was a more reliable parameter for determining the presence and level of activity of bugs than was a direct estimate of the number of bugs. The green tree ant, Oecophylla smaragdina (F), was the most important factor regulating bug populations. When predation was excluded as a factor, the number of flushing shoots and maximum temperature accounted for 80% of the total variability in the bug damage. Green tree ants should be considered as an important biological control agent for fruit spotting bug, and monitoring should be commenced when cashew trees start to flush (using damaged shoots as indicator).