Joachim Offenberg

Balancing between mutualism and exploitation: the symbiotic interaction between Lasius ants and aphids

Abstract Nests of Lasius niger (L.) ants were given varied food regimens to test whether their behaviour towards an aphid partner, Aphis fabae (Scop.), changed with alternative food supplies. Honeydew collection and predation on aphids were measured by video monitoring the movement of ants between their nest and an aphid aggregation. Data collected from the aphid aggregations enabled comparisons between remaining aphid biomass and between the tending intensities of the ants. I tested how ant behaviour was influenced by their access to alternative prey and sugar. The results showed that ants accepted a honey solution as a substitute for the honeydew produced by aphids. Ants not only attended their aphid partners, but also preyed on them. The average predation rate increased eightfold when ants were offered the alternative of sugar, whereas alternative prey had no significant effect. In contrast, ant-tending intensity decreased with alternative sugar whereas alternative prey elicited no effect.

Evidence that insect herbivores are deterred by ant pheromones

It is well documented that ants can protect plants against insect herbivores, but the underlying mechanisms remain almost undocumented. We propose and test the pheromone avoidance hypothesis—an indirect mechanism where insect herbivores are repelled not only by ants but also by ant pheromones. Herbivores subjected to ant predation will experience a selective advantage if they evolve mechanisms enabling them to avoid feeding within ant territories. Such a mechanism could be based on the ability to detect and evade ant pheromones. Field observations and data from the literature showed that the ant Oecophylla smaragdina distributes persistent pheromones throughout its territory. In addition, a laboratory test showed that the beetle Rhyparida wallacei, which this ant preys on, was reluctant to feed on leaves sampled within ant territories compared with leaves sampled outside territories. Thus, this study provides an example of an ant–herbivore system conforming to the pheromone avoidance hypothesis.

REVIEW: Ants as tools in sustainable agriculture

Summary With an expanding human population placing increasing pressure on the environment, agriculture needs sustainable production that can match conventional methods. Integrated pest management (IPM) is more sustainable, but not necessarily as efficient as conventional non-sustainable measures. Being predatory and organized as superorganisms, ants possess traits making them suitable agents in IPM. Recent works on weaver ants Oecophylla spp. showcase ants as highly efficient pest controllers. A synthesis shows that weaver ants can reduce pest numbers and their damage and increase yields in multiple crops. Their efficiency is comparable to chemical pesticides or higher, while at lower costs. They provide a rare example of documented efficient conservation biological control. Weaver ants share beneficial traits with almost 13 000 other ant species and are unlikely to be unique in their properties as control agents. A synthesis of applied work on other ant species illustrates potentials for control of arthropod pests, weeds and plant diseases in orchards, forestry and arable crops. Synthesis and applications. By showing that ant biocontrol can match synthetic pesticides in a wide setting of agricultural systems, this review emphasizes the potential of managing ants to achieve sustainable pest management solutions. The synthesis suggests future directions and may catalyse a research agenda on the utilization of ants, not only against arthropod pests, but also against weeds and plant diseases. Lastly, it may stimulate implementation of already proven ant-based integrated pest management techniques.

The distribution of weaver ant pheromones on host trees

Abstract.The visible anal spots deposited by Oecophylla smaragdina ants have been suggested to deter ant prey, affect interspecific competition and facilitate mutualists and parasites in tracking down Oecophylla ants. I measured the density of anal spots on host trees with and without ants and tested for correlations between spot density, ant activity and the likelihood of being detected by an ant. Spots were only found on trees with ants. On ant-trees, spots were distributed throughout the trees but with higher densities in areas with high ant activity and pheromone densities were higher on twigs compared to leaves. Also there was a positive correlation between spot density and the likelihood of being detected by ants. Anal spots may thus function as reliable cues to interacting species and be an important factor in shaping the community around Oecophylla colonies.

Multiple sources of local knowledge: a global review of ways to reduce nuisance from the beneficial weaver ant Oecophylla

The weaver ants Oecophylla smaragdina and O. longinoda are abundant in tropical Asia, Australia and Africa. Although local people and a handful of scientists appreciate the benefits these tree-dwelling ants bring in terms of medicinal and food source, protection of tree crops, and enhancement of fruit and nut quality, Oecophylla has one major drawback: it also bites people. Perception of this nuisance, and the response to it, is influenced by the frequency of encounters and the perceived benefits gained from Oecophylla. We used a range of methods in more than ten countries to document how people reduce weaver ant nuisance, including interactive rural radio programs. Apart from growers, also ant brood collectors, fruit-pickers and intermediaries in the value chain hold in-depth ecological knowledge on weaver ants. Ways to make Oecophylla more widely accepted and build ecological literacy at various levels of the society are discussed.

Occurrence of sexuals of African weaver ant ( Oecophylla longinoda Latreille) (Hymenoptera: Formicidae) under a bimodal rainfall pattern in eastern Tanzania

The African weaver ant, Oecophylla longinoda, is being utilized as a biocontrol agent and may also be targeted for future protein production. Rearing of mated queens in nurseries for colony production is needed to cater for such demands. Thus, newly mated queens must be collected for use as seed stocks in the nurseries. To collect mated queens efficiently it is important to identify when sexuals occur in mature colonies. We studied the occurrence of sexuals in O. longinoda colonies for 2 years in Tanga, Tanzania, a region characterized by a bimodal rainfall pattern. We found that O. longinoda sexuals occurred almost throughout the year with abundance peaks from January to April. Production of sexuals appeared to be triggered by rainfall, suggesting that populations in areas with long rainy periods may show prolonged mating periods compared to populations experiencing extended dry periods. The bimodal rain pattern may thus cause a low production over a long period. The average yearly production of queens per tree and per colony was estimated to be 449 and 2753, respectively. The average number of queens per nest was 17. Worker abundance declined from January to March with minimum by the end of this period, being inversely proportional to the production of sexuals. In conclusion, mated queens may be collected almost throughout the year, but most efficiently by the onset of the long rainy season when the majority disperse.

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

In Benin cashew plantations, yields and nut quality are lost mainly as a result of insect pests. In the present study, we investigated the effectiveness of the African weaver ant Oecophylla longinoda as a biocontrol agent against Beninese cashew pests. In a 2‐year study, nut yield and quality were compared among: (i) trees with weaver ants O. longinoda; (ii) trees where weaver ants were sugar‐fed; (iii) integrated pest management (IPM) trees with weaver ants combined with fruit fly bait spray; and (iv) control trees receiving no control measures. All treatments with ants showed significantly higher yields than the control, with the IPM treatment leading to the highest yield. Compared with the control trees, the ants, ant sugar‐fed and the IPM trees produced 78%, 122% and 151% more nuts, respectively. Nuts produced on control trees were of a higher quality on average because they were less damaged by thrips (probably because the fruit fly bait worked as a contact poison on thrips); this was also the case for the IPM treatment. In absolute numbers, however, trees in ant treatments produced more first‐quality nuts. To achieve a broader and effective control of both coreid bugs and thrips, a combination of weaver ants and supplementary compatible control measures is recommended.

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 ants mating season (several months), whereas this was not required when using leaf and paper traps.

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 ants 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.

The Effect of Queen and Worker Adoption on Weaver Ant (Oecophylla smaragdina F.) Queen Fecundity

Incipient ant colonies are often under fierce competition, making fast growth crucial for survival. To increase production, colonies can adopt multiple queens (pleometrosis), fuse with other colonies or rob brood from neighboring colonies. However, different adoption strategies might have different impacts such as future queen fecundity or future colony size. O. smaragdina queen production was measured in incipient colonies with 2, 3 or 4 founding queens, following the transplantation of 0, 30 or 60 pupae from a donor colony. Pupae developed into mature workers, resulting in increased worker/queen ratios in pupae transplanted treatments and leading to increases in the per capita queen production. Conversely, more queens did not induce increased per capita fecundity. Thus, brood robbing added individuals to the worker force and increased future production of resident queens, whereas queen adoption increased the colony’s future production, but not the production of individual queens.