Research Article |
Corresponding author: Wilson Nene ( wilsoninene@gmail.com ) Academic editor: Jack Neff
© 2016 Wilson Nene, Gration M. Rwegasira, Maulid Mwatawala, Mogens Gissel Nielsen, Joachim Offenberg.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Nene W, Rwegasira GM, Mwatawala M, Nielsen MG, Offenberg J (2016) Foraging behavior and Preferences for Alternative Supplementary Feeds by the African Weaver Ant, Oecophylla longinoda Latreille (Hymenoptera, Formicidae). Journal of Hymenoptera Research 50: 117-128. https://doi.org/10.3897/JHR.50.8173
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Weaver ants, Oecophylla spp, are effective predators that control a wide range of insect pests in multiple crops when maintained at high population. Supplementary feeding, particularly during reduced food availability is one of the management practices that maintain and boost weaver ants’ populations. Experiments were conducted between September and October 2013, January and February, 2014 to determine the type of food preferred by weaver ants, O. longinoda. Twenty colonies of O. longinoda were provided with four types of food to determine their feeding preferences. These include anchovy, chicken intestine, fish intestines and earthworms. We examined food preferred by ants based on weight of the food removed and activity of the ants on foods. Furthermore, we examined foraging behavior of ant workers on anchovy food (fresh-ground and dry-ground) in nine O. longinoda colonies. Thereafter, small and large particles of dried-ground anchovy were tested. The results showed that O. longinoda preferred anchovy to other foods provided. However, the results of Analytical Hierarchy Process showed that earthworm and fish intestine were the most accessible food types by farmers, as determined by availability, affordability and applicability. We observed more ants on fresh-ground as opposed to dry anchovy; similarly, large particles were more easily removed than was the case with small particles. Thus, during reduced food availability, farmers in the study area should use earthworms and fish intestines feeds to supplement O. longinoda colonies. Fresh moist anchovy or dry anchovy of large particle sizes can be used where available.
African Weaver Ant, citrus, cashew, food preference, anchovy, Tanzania
Two species of weaver ants, Oecophylla longinoda Latreille and Oecophylla smaragdina Fabricius are generalist predators that protect crops against insect pests (
In addition to biocontrol, Oecophylla is used as a valuable source of food for humans (
Social insect populations are negatively affected when food is scarce (Dusstour and Sympson 2012). Weaver ant colonies may even move among trees in search of forage (
The feeding preferences of Oecophylla longinoda are not well known. In Vietnam, farmers provide fish and chicken intestines to O. smaragdina as supplementary foods during scarcity (
The practice of food supplementation for O. longinoda is limited by inadequate knowledge on food preference (based on cheap local available feeds), forms or states as well as particle sizes. Therefore, the objective of this study was to determine the type of food preferred by weaver ants in terms of form and size. A good understanding of food preferences by weaver ants and their foraging behavior is essential for effective management of O. longinoda during food scarcity.
We conducted experiments at Naliendele Agricultural Research Institute (
The study was approved by the Directorate of Research and Postgraduate Studies (DRPG) of
In each orchard we selected trees with at least 40% of branches occupied by ants (as assessed by as per
Intestines and anchovy were ground by locally made mortar and pestle. Earthworms were dug out of wet soils (close to water ponds and irrigated fields) and chopped into small pieces (approximately 0.5–1 cm in length) by a kitchen knife. About 6 g of each food type were placed in a 0.01×0.1 m bowl, set on a feeding platform. Ants could access the bowls by crawling through a guiding stick. Feed bowls were placed equidistant from the middle of the feeding platform. Feeds and water were provided ad libitum throughout the experimental period. All the food types were tested in 10 colonies in each orchard. Preferences were determined by i) counting all foraging workers observed on food station and inserting their mouths into a food type and ii) weighing the amount of each food removed by the ants. Counting started 60 minutes after more than one forager had discovered each food type. Thereafter, the weight of the remaining food in each bowl was determined. The amount of food type removed by the ants was determined by establishing the difference in weight between the food supplied and the food which remained in the bowl. In each case, the weight loss due to evaporation was deducted. Weight loss due to evaporation was determined in the control food types that were inaccessible by ants.
A sample of each food type was analyzed for nutrient compositions at the University of Dar es Salaam. The total carbohydrate, crude protein, total lipids (Fat) and vitamin A were determined according to the procedures described by
Furthermore, we used anchovy food to test for food forms and particle sizes that can be preferred by Oecophylla longinoda. Anchovy was used because it is processed in a standard form. We hypothesized that anchovy type (dry and fresh) affected the foraging behavior of workers. We also hypothesized that the particle size of dry anchovy affects the foraging behavior of workers. The experiments were conducted between May and July 2014. The first experiment involved two different forms of anchovy, dried and fresh. This experiment was conducted for 10 days, with the observation starting around 0900 am each day. The anchovy was sun dried (27–29 °C) for 7 days before grinding. We used fresh-ground anchovy of approximately similar size as the dried one. A Y- shaped feeding arena made up of wood was used as a feeding platform. A feeding bowl was placed on top of a board tied at each end of the Y shaped arena. This gave an equal chance for the workers to access each of the anchovy food types. Nine colonies were used. Each colony occupied at least two citrus trees. One bowl of each anchovy food was supplied per colony. The foraging behavior was assessed by counting workers carrying food particles from the source. The counting was done ten times at an interval of one minute (ten observations) every day per each colony for 10 days consecutively. In the end, we calculated the average number of foraging workers per minute per colony for a given food form.
The second experiment involved dried-ground anchovy of different particle sizes. The particles were measured by using laboratory test sieves (Wagtech International Ltd UK). Two particles sizes were selected; particles ranging from 0.5 to 1 mm in diameter (referred hereinafter as small particles) and; particles of 2 mm d (referred hereinafter as large particles). The experiment was conducted on six weaver ant colonies for 10 days using similar procedures for testing food types (above).
Thereafter, Analytic Hierarchy Process (AHP) (
The analyses were performed using JMP 10.00 software. A non parametric one way ANOVA was used followed by Multiple Comparison-Wilcoxon Each Pair for counting forage workers and the amount of food removed under food preferences. Mann-Whitney tests were performed to compare the number of forage workers on dried and fresh anchovy; similar comparisons were done for small and large particle sizes.
Nutrients composition of the feeds fed to the ants are presented in Table
Food type | Fish intestine | Chicken intestine | Earthworm | |
---|---|---|---|---|
Parameters | Anchovy | |||
Crude Protein (%) | 44.6 | 47.68 | 31.2 | 45.6 |
Total carbohydrate (g/100g) | 0.2 | 3.1 | 6.21 | 0.01 |
Fat content(g/g) | 0.059 | 0.078 | 0.087 | 0.005 |
Moisture content(%) | 89.4 | 52.7 | 78.4 | 92.9 |
Vitamin A(mg/100g) | 4.5 | 3.4 | 5.6 | 0 |
Flavonoids(mg/g) | 0.013 | 0.0002 | 0.001 | 0.00 |
Energy(kj/g | 9.711 | 11.414 | 9.553 | 7.834 |
The numbers of workers foraging on food types were significantly (p=0.05) different in both citrus and cashew orchards, during both dry and rainy seasons (Tables
Food forms (a): Wilcoxon test; N=90; Prob.<0.0001, Food particle sizes (b) Wilcoxon test; N=60; Prob.<0.0001. Number of Weaver Ant counting /10 minutes for 10 days between May and July, 2014 for the given anchovy food forms and different particle sizes at Naliendele Citrus orchard, Tanzania, 2014. Large particle sizes refer to all particles that do not pass on a sieve of less or equal to 1mm, where as small particle sizes are those particle passes on 0.5 or 1 mm sieves.
The p-values for weaver ant counting and amount of food removed (g)/hour in 20 days between September and October 2013 and January and February 2014 in citrus orchard, Naliendele, Tanzania. (Kruskal-Wallis Multiple Comparison-Wilcoxon Each Pair test).
Food types | Citrus dry season | Citrus rainy season | ||
---|---|---|---|---|
Pairwise comparison | Weaver ant counting (P-values) | Amount of food removed (g) (P-values) | Weaver ant counting (P-values) | Amount of food removed (g) (P-values) |
Anchovy versus Chicken intestine | 0.054 | 0.042 | 0.0002 | 0.0011 |
Anchovy versus Earthworm | 0.021 | 0.108 | 0.0002 | 0.0011 |
Anchovy versus Fish intestine | 0.014 | 0.0018 | 0.0003 | 0.0011 |
Chicken intestine versus Fish intestine | 0.121 | 0.0095 | 0.0002 | 0.0011 |
Chicken intestine versus Earthworm | 0.68 | 0.77 | 0.004 | 0.0011 |
Earthworm versus Fish intestine | 0.33 | 0.014 | 0.0155 | 0.0011 |
The p-values for weaver ant counting and amount of food removed (g)/hour in 20 days between September and October 2013 and January and February 2014 in cashew orchard, Naliendele, Tanzania. (Kruskal-Wallis Multiple Comparison-Wilcoxon Each Pair test).
Food types | Cashew dry season | Cashew rainy season | ||
---|---|---|---|---|
Pairwise comparison | Weaver ant counting (P-values) | Amount of food removed (g) (P-values) | Weaver ant counting (P-values) | Amount of food removed (g) (P-values) |
Anchovy versus Chicken intestine | 0.0013 | 0.51 | 0.0002 | 0.0002 |
Anchovy versus Earthworm | 0.0006 | 0.817 | 0.0002 | 0.0002 |
Anchovy versus Fish intestine | 1.00 | 0.86 | 0.0002 | 0.0002 |
Chicken intestine versus Fish intestine | 0.0008 | 0.76 | 0.0002 | 0.0002 |
Chicken intestine versus Earthworm | 0.068 | 0.84 | 0.023 | 0.0002 |
Earthworm versus Fish intestine | 0.0003 | 0.92 | 0.005 | 0.0002 |
Analytical Hierarchy Process results on farmers access to food types for weaver ants.
Alternatives | Weights (Eigen Vector)* | Global score | ||
---|---|---|---|---|
Affordability (0.539) | Availability (0.0845) | Applicability (0.126) | ||
Earthworm | 0.59 | 0.63 | 0.41 | 0.42 |
Fish intestine | 0.25 | 0.19 | 0.27 | 0.18 |
Chicken intestine | 0.13 | 0.28 | 0.22 | 0.12 |
Anchovy | 0.04 | 0.054 | 0.11 | 0.04 |
This study revealed that food preferences and foraging behavior by the Oecophylla longinoda can be influenced by food type, form, as well as particle size. Anchovy was highly preferred by O. longinoda in both citrus and cashew orchards during both dry and rainy seasons. The reason for forage workers’ preference on anchovy is however not clear and could not be confirmed by this study. High preference for anchovy could be due to nutritional composition, particularly flavonoids or proteins. The foraging rate of Pheidole megacephala (Fabricius) depends on the type of protein (
Anchovy was the highest preferred food across orchards and seasons but the order of preference for other food types in both orchards varied between seasons.
Chicken intestine was the least preferred during the rainy season. This is probably due to the fact that individual ant workers more easily remove fish intestine and earthworm than chicken intestine. Chicken intestine became stickier and bound to the food bowl during the rainy season. Thus, the removal of chicken intestine by foragers was difficult. Foragers spent time trying to take sticky-bound food items but they often failed. A temporal change in foraging activities was observed when the food became sticky and bound to the feeding bowl. At 15-30 minutes after food introduction, many ants were recruited and foraging activities increased with more ants observed on chicken intestine. However, as the food became stickier and bound to the feeding bowl, workers shifted to other food sources. It can be concluded that the nature of food at a particular time determines the foraging behavior of Oecophylla longinoda workers, and thereby influences preferences. Previous studies have shown that a large number of nest mates are recruited when ants are facing a non-transportable food items such as shrimps (
More anchovy was removed by ants as opposed to other feeds across seasons and orchards. The probable reason for this could be the form that anchovy assumes after being ground. Anchovy in the field became moist and grainy and could be removed without difficulty unlike the other food types. According to
The quantities of food taken did correspond with the number of foraging workers, except for fish intestine in citrus orchard. We recorded more foraging workers on fish intestine than on chicken intestine and earthworm but the amount of food removed was smaller. Sometimes more ants visit particular feed but remove less (
In this study, we observed higher foraging activities on ground-fresh than on dry anchovy. These results support previous studies whereby three ants, Linepithema humile Mayr, Anoplolepis custodiens F. Smith and Crematogaster peringueyi Emery foraged more on liquid or moist food bait than on dry food bait (Nyamukondiwa and Addison 2014). Similarly, more activities for the ant, L. lumile were recorded on a 25% sugar solutions or honey than was the case with solid based protein foods such as tuna (
However, fresh-ground foods become sticky after some time, making it difficult for ant workers to remove them. Therefore, fresh-ground anchovy should be replenished to avoid stickiness; otherwise dried-ground anchovy should be used.
Forage workers easily collected and took large particles back to their nests in their mouthparts. However, they faced difficulties in collecting small particles and spent more time at the food bowl. A similar finding was reported for the fire ant Solenopsis invicta Buren (
Flavonoids have phytochemical properties against fungal, virus, and bacteria (
The results showed that all four tested feeds were removed by Oecophylla longinoda workers but anchovy was the most preferred. Considering the availability and affordability of the tested food sources, earthworms and fish intestine would be recommended as supplements during scarcity to boost weaver ants colonies on understanding that farmers preferred the less costly and sustainable option. All in all, fresh or dried-ground anchovy with particle sizes greater than 1 mm remains the best choice if availability and affordability are not subjects of concern.
We would like to thank Zonal Director for Naliendele Agricultural Research Institute for granting us permission to conduct this study in Naliendele fields. The funding for this study was provided by Danish International Development Agency (DANIDA) through the project ‘‘Increasing Value of African Mango and Cashew production’’, (DFC No. 10-025AU)