Contribution of cocoa plantations to the conservation of native ants (Insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic Forest fauna of southern Bahia,Brazil

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests. Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.     

Effects of management intensity on ant diversity in cocoa plantation (Oume,centre west Côte d’Ivoire)

Anthropogenic habitat modifications, including conversion of forest to agricultural production cause losses of native species. In this study we examined the losses suffered by ant communities in relation to the intensity of management in cocoa plantations established in former tropical forest. An extensive sampling protocol consisting of pitfall trapping, leaf litter sampling, soil sampling and hand sampling was used to characterize ant species richness and composition in the native forest and compare it with three cocoa farms differing in their management intensity. Species richness was negatively correlated with management intensity and differed greatly between management practices and the forest. Two subfamilies (Myrmicinae and Amblyoponinae) showed a significant negative correlation with agricultural intensification probably because their species are constrained to living in forest like habitat. The species composition differed greatly between management practices and the forest. Intensively and moderately managed cocoa plantations were most dissimilar to the forest. In contrast, forest ants were well represented in the least intensively managed plantation. Overall, the findings of this study show that only slightly managed cocoa plantations sustain an ant diversity that comes close to that of the forest. This also holds true for cocoa plantations on former agricultural, and thus previously heavily used, land. The findings may help in the conservation of biodiversity as management practices most likely to sustain forest like ant communities have been identified.     

Ant assemblage structure on cocoa trees in smallholder farms in the Centre Region of Cameroon

We investigated the ant community structure in cocoa farms in the Centre Region of Cameroon. Ants were collected on the cocoa trees during the years 2006 and 2007 using chemical knock‐down. We tested the hypothesis of the existence of deterministic factor in the structuration of ant mosaic using C‐Score; we assessed the relationship between the numerical dominant and subdominant ant species using Spearman correlation test and discussed on the influence of vegetation structure and farm management on the ant community structure. A total of 53 ant species belonging to 20 genera and five subfamilies were identified from a set of 51,525 workers collected. C‐score analysis supported the hypothesis that ant community were structured by competition. Negative relationships were found between dominant ant species. Farming practices which were mainly pruning, chemical treatment and habitat structure appeared to influence the ecological status and distribution of dominant ant species.     

An Ant Mosaic Revisited: Dominant Ant Species Disassemble Arboreal Ant Communities but Co-Occur Randomly

The spatial distributions of many tropical arboreal ant species are often arranged in a mosaic such that dominant species have mutually exclusive distributions among trees. These dominant species can also mediate the structure of the rest of the arboreal ant community. Little attention has been paid to how diet might shape the effects of dominant species on one another and the rest of the ant community. Here, we take advantage of new information on the diets of many tropical arboreal ant species to examine the intra- and inter-guild effects of dominant species on the spatial distribution of one another and the rest of the tropical arboreal ant community in a cocoa farm in Bahia, Brazil. Using null model analyses, we found that all ant species, regardless of dominance status or guild membership, co-occur much less than expected by chance. Surprisingly, the suite of five dominant species showed random co-occurrence patterns, suggesting that interspecific competition did not shape their distribution among cocoa trees. Across all species, there was no evidence that competition shaped co-occurrence patterns within guilds. Co-occurrence patterns of subordinant species were random on trees with dominant species, but highly nonrandom on trees without dominant species, suggesting that dominant species disassemble tropical arboreal ant communities. Taken together, our results highlight the often complex nature of interactions that structure species-rich tropical arboreal ant assemblages.     

Changes in ant communities along an age gradient of cocoa cultivation in the Oumé region,central Côte d'Ivoire

We identified the extent to which ant diversity occurs despite conversion of forests into cocoa plantations by examining the communities across four age classes of plantations (classes I–IV with increasing age from 0–5 to 21–40 years) and in their original forests. An extensive sampling protocol consisting of pitfall trapping, leaf litter sampling, soil sampling and hand sampling was used to characterize ant species richness and composition in three replicates of each age class and in the remaining forest patches. A total of one hundred ant species was found in all habitats combined. While the forest was the richest habitat (73 species), species richness in the different plantation age classes varied as follows (sorted in descending order): class IV (69 species) > class III (57 species) > class I (52 species) > class II (43 species). Age gradient was thus significantly positively correlated with mean species richness and with the relative abundance of some subfamilies. The species composition differed greatly between some plantation age classes and the forest. The two youngest cocoa age classes (I and II) were most dissimilar to the forest. In contrast, forest ants were well represented in the old cocoa age classes (III and IV). Three functional guilds (generalist predators, specialist predators and territorially dominant arboreal species) were in their relative abundance significantly correlated to the age gradient. Overall, cocoa cultivations retaining a floristically diverse and structurally complex forest structure are a suitable management system for the conservation of ant species of the formerly forested habitats.     

Plant-attracted ants affect arthropod community structure but not necessarily herbivory

Abstract.  1. The effectiveness of ants as plant defenders is equivocal for plants that attract ants via extrafloral nectaries (EFNs).
2. This study focused on the myrmecophilic savannah tree Pseudocedrela kotschyi that attracts ants to EFNs and on the arthropod fauna associated with P. kotschyi . Herbivory and arthropod community composition were compared between trees that were dominated by one of three congeneric ant species, Camponotus acvapimensis , C. rufoglaucus , and C. sericeus , and between trees where ants were experimentally excluded and untreated control trees.
3. Short-term ant-exclusion experiments failed to demonstrate a consistent effect of ants on herbivory.
4. Plants dominated by different ant species differed significantly in leaf damage caused by herbivorous insects. The relative ranking of herbivory levels of the trees dominated by different ant species was persistent in three consecutive years.
5. Ants significantly reduced the abundance of different arthropod groups (Araneae, Blattodea, Coleoptera, Homoptera, non-ant Hymenoptera). Other groups, including important herbivores, seemed not to be affected (Lepidoptera, Orthoptera, Thysanoptera, Heteroptera).
6. The study suggests that the presence of ants only benefits plants when specific ant species are attracted, and protection by these ants is not counterbalanced by their negative effect on other beneficial arthropods.     

Interactions of the ant Crematogaster scutellaris with the fig/fig wasp mutualism

Abstract. 1. A classical example of specialised pollination mutualisms is the relationship between fig trees and their pollinating wasps, in which each partner depends completely on the other for its reproduction; however the fig/fig wasp association is also the target of a great diversity of other species, ranging from specialised parasites to opportunistic foragers, among them ants.
2. The ant community and the sources of ant attraction observed on the Mediterranean fig tree Ficus carica were characterised.
3. A guild of ants attracted by homopterans tended on the plant was distinguished from a second guild composed of two co-dominant ant species ( Crematogaster scutellaris and Pheidole pallidula ) that prey mostly on pollinating wasps, abundant during certain parts of the fig cycle.
4. Foraging workers of C. scutellaris search for prey on the fig inflorescence (syconium), capturing pollinating wasps mostly at the peak of wasp emergence and at a rate estimated to reach 600 prey per day for an entire tree.
5. Detailed study of the predatory sequences displayed under experimental conditions showed that ant workers captured 100% of the pollinating wasps offered, while they captured only 5.5% of the parasitoid wasp specific to the pollinator. The respective impacts of the interaction on ants and on the figs are discussed, as well as several behavioural traits of predation by the dominant ant on pollinators.     

Critical thermal limits in Mediterranean ant species: trade-off between mortality risk and foraging performance

1. In Mediterranean ant communities, a close relationship has been found between activity rhythm in the period of maximum activity and position in the dominance hierarchy: subordinate species are active during the day, when conditions are more severe, while dominants are active during the afternoon and the night.
2. Results obtained in this study confirmed that the species foraging at higher temperatures were closer to their critical thermal limits than the species foraging at lower temperatures.
3. This enabled two extreme strategies of foraging in relation to temperature to be distinguished: (1) heat-intolerant ant species behaved as risk-averse species, foraging at temperatures very far from their critical thermal limits; and (2) heat-tolerant ant species behaved as risk-prone species, foraging very near their critical thermal limits and running a high heat mortality risk.
4. Heat-tolerant species benefited from this strategy by having better foraging performance at high temperatures.
5. This wide range of thermal niches may be one reason why Mediterranean ant faunas are so diverse in the face of limited diversity in vegetation and habitat structure: the daily range of temperature may be sufficiently great to meet the requirement both of heat-adapted and cold-adapted species as well as a spectrum of intermediate forms.     

Floral visitation patterns of two invasive ant species and their effects on other hymenopteran visitors

Abstract.  1. Floral nectar of the native Hawaiian 'ōhi'a tree, Metrosideros polymorpha , is an important food source for several native honeycreepers and yellow-faced bees, Hylaeus spp., but is also attractive to invasive ants.
2. I undertook this study to compare floral visitation patterns of two widespread invasive ants, the Argentine ant, Linepithema humile , and the big-headed ant, Pheidole megacephala , and to determine their effects on nectar volume and floral hymenopteran visitors.
3. In the first year of the study, Argentine ants visited inflorescences more frequently than big-headed ants at mid-day and in the afternoon, but did not occur in higher densities than big-headed ants at any time of day. In the following year, Argentine ants visited inflorescences both more frequently and in higher densities than big-headed ants. Argentine ant density had a stronger association with nectar concentration than big-headed ant density.
4. Nectar volume did not differ between ant-excluded and ant-visited inflorescences for either ant species. However, ant density was negatively associated with nectar volume for both species.
5.  Hylaeus spp. never visited inflorescences with big-headed ants, while non-native honeybees visited inflorescences with and without ants of either species in equal frequency.
6. Most studies of the effects of invasive ants on native arthropods have focused on interactions on the ground. Flowers should not be overlooked as microhabitats from which native arthropods may be displaced by invasive ants.     

The effect of forest disturbance on the leaf litter ant fauna in Ghana

In order to observe the effect of forest loss on the leaf litter ant fauna in Ghana, West Africa, samples were taken in primary forest, secondary forest and in cocoa plots. Ants were extracted from the leaf litter by sieving followed by suspension in Winkler bags. The species composition and species richness in the three different habitats were compared and no significant difference was found between them. It was concluded that most primary forest leaf litter ant species continue to survive in parts of the agricultural landscape which has largely replaced their original habitat.     

Higher productivity at the cost of increased host-specificity when Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation

Abstract .1. Field studies were made of the benefits and costs of two feeding strategies in the genus Maculinea, whose final-instar larvae parasitise Myrmica ant colonies. Maculinea arion is an obligate predator of ant brood, whereas M. rebeli and M. alcon mimic ant larvae and are fed (like cuckoos) directly by the workers.
2. Samples of > 1500 Myrmica nests confirmed laboratory-based predictions that, by feeding at a lower trophic level, many (4.7-fold) more individuals of M. rebeli and M. alcon are supported per ant colony than M. arion.
3. Because of their efficient feeding, cuckoo species often occupied sites where their phytophagous early larval populations coincided to only a small extent (> 10%) with host Myrmica colonies, whereas all sites supporting M. arion had 50–100% of the phytophagous stages within foraging range of the host Myrmica species.
4. Greater host-specificity was identified as another consequence of cuckoo-feeding. The ecological cost of this is discussed .
5. The feeding of other Maculinea species had not been fully described: the data suggest that M. nausithous is a predator of ant brood and confirm that M. teleius is predacious .     

Ant mutualists alter the composition and attack rate of the parasitoid community for the gall wasp Disholcaspis eldoradensis (Cynipidae)

Abstract.  1. The strength or density dependence of pairwise species interactions can depend on the presence or absence of other species, especially potential mutualists.
2. The gall wasp Disholcaspis eldoradensis induces plant galls that secrete a sweet honeydew from their top surfaces while the wasp larvae are active. These galls are actively tended by Argentine ants, which collect the honeydew and drive off parasitoids attempting to attack the gall wasp.
3. When ants were excluded, the total rate of parasitism by seven species of parasitoids increased by 36%, and the rate of gall-wasp emergence decreased by 54%.
4. The total percentage parasitism was affected by gall density when ants were excluded but not when ants were unmanipulated, suggesting a change in parasitoid functional responses due to ant tending.
5. In addition, excluding ants significantly altered the proportions of different parasitoid species that emerged from galls; one parasitoid species increased from 1% to 34%, and another decreased from 46% to 19%.
6. The invasive Argentine ants studied are capable of maintaining the mutualism with the gall wasps that evolved in the presence of different ant species and also act as a selective filter for the local community of generalist parasitoids trying to attack this gall species.     

Microclimatic factors associated with elevational changes in army ant density in tropical montane forest

Abstract.  1. The density (rate of encountering foraging raids) and species richness of army ants (Formicidae: Ecitoninae, and behaviourally convergent Ponerinae) was measured in montane tropical forest. Above-ground and subterranean army ant raids were sampled using standard protocols at four sites across an elevational gradient (1200–1650 m above mean sea level) in and near cloud forest in the area of Monteverde, Costa Rica.
2. Mean ambient temperature differed among sites, and decreased with elevation. For the above-ground foraging army ant species, raid rates also declined with elevation. Surface army ant raid rates, however, were not affected by day to day weather variation within sites (temperature, cloud cover, or precipitation).
3. For the underground foraging army ant species, raid rates did not vary directionally with elevation, and subterranean raid rates were not affected by day to day weather variation within sites.
4. Army ant species richness was not directionally related to elevation, and species sharing among sites was generally high.
5. Army ant community structure changes with elevation in Neotropical montane forest, and the results suggest that the strongest effects are of temperature regimes on the density of raids. These findings provide a baseline against which to detect changes in army ant communities that may accompany directional climate change in tropical cloud forests.     

Competition trade-offs in the organisation of a Mediterranean ant assemblage

Abstract.  1. The organisation of an ant assemblage inhabiting an olive orchard in central Italy was analysed and patterns of dominance among ant species were described in order to assess (i) the relationship between thermal dependency and degree of behavioural dominance, and (ii) the relationship between dominance and discovery ability.
2. Activity patterns of the most abundant species on trees were examined in a sample of 120 trees during spring and summer. The degree of behavioural dominance and the ability of different species to discover new food sources were assessed using tuna baiting on a subset of 80 trees.
3. Different ant species showed contrasting patterns of activity. Some species (such as Lasius lasioides , Camponotus lateralis , and Camponotus piceus ) were most active during the warmer part of the day, while others restricted their activity to the cooler hours ( Camponotus aethiops and Plagiolepis pygmaea ). Some species (such as Crematogaster scutellaris ) were active irrespective of the time of day.
4. No clear relationship was observed between temperature of maximal activity and degree of behavioural dominance. There was, however, a positive relationship between behavioural dominance and thermal range of activity. A positive relationship between dominance and ability to find resources, with the most behaviourally dominant and aggressive species being most efficient in finding food items, was also observed.
5. The results support the idea that the temperature–dominance relationship is much more complex in Mediterranean-type habitats than in other ecosystems. Of particular interest is the positive dominance–discovery relationship. This finding contrasts with previous investigations, which reported a negative relationship between dominance and discovery ability and suggested that this pattern plays a role in promoting the coexistence of species in ant communities.     

Ecological interactions among ants in the genus Linepithema, their phorid parasitoids, and ant competitors

Abstract. 1. Many ant species abandon foraging and retreat underground when parasitoids in the dipteran family Phoridae are present. Although the influence of phorids on ant foraging is well documented, their influence on interspecific competition is less studied. This study examined whether phorids influenced the competitive ability of host ants in the genus Linepithema at two sites in Brazil.
2. The phorid Pseudacteon lontrae attacked Linepithema piliferum at one site, while the phorid Pseudacteon pusillus attacked an unknown Linepithema ( Linepithema sp.) at the other site. Phorid parasitoids of Linepithema were far more common than phorids of other ant species.
3. Despite a high abundance of phorids, it was difficult to conclude that they influenced competition. Captures in pitfall traps indicated that host Linepithema were most active during times of day when phorids were inactive.
4. Camponotus rufipes and Brachymyrmex sp., the most common competitors of Linepithema sp. (60% of all interactions), dominated Linepithema sp. during the day regardless of phorids. Remaining ant species could not be evaluated individually because they interacted with Linepithema sp. infrequently.
5. Ectatomma brunneum was the most common competitor of L. piliferum (58% of all interactions). The high abundance of phorids at this site made it impossible to evaluate interactions between E. brunneum and L. piliferum in the absence of phorids.
6. Phorids seldom influenced exploitative competition by causing host Linepithema to abandon the bait when no ant competitors were present.     

Role of floral repellents in the regulation of flower visits of extrafloral nectary-visiting ants in an Indian crop plant

Abstract.  1. Field investigation of the association between sponge gourd, Luffa cylindrica plants and its ant visitors revealed that five of the six most frequent species: Camponotus compressus, C. paria, Pheidole sp., Pachycondyla tesserinoda and Tetramorium sp. mainly visited the extrafloral (EF) nectaries present on the leaves, bracts, bracteoles and calyx of the plant. Tapinoma melanocephalum was the only ant species observed at the floral as well as the EF nectaries.
2. A bioassay of ant behaviour revealed aversion to young and mature unisexual flowers of sponge gourd in the five predominantly EF nectary-visiting ant species, while floral preference was demonstrated in T. melanocephalum. A significant difference was not found in the number of insect pollinators visiting T. melanocephalum occupied and un-occupied flowers, suggesting the absence of deterrent effect of this tiny ant species on the pollinators.
3. Further behavioural assays showed preference for 2- and 4-day-old leaves and also 2-day-old buds, while the 4-day-old buds induced avoidance in all the species. Androecium and gynoecium had significantly higher repellent effects in comparison to the petals. Thus floral repellents, probably help to reduce nectar theft and prevent loss of pollen function.
4. This aversion was not demonstrated in the case of old flowers. A significantly greater number of insect pollinators visited young and mature flowers compared with old flowers, suggesting that selective exclusion of medium- and large-sized EF nectary-visiting ant species from the flowers, as a result of aversion to floral repellents, serves to avoid the threat of attack to insect pollinators of sponge gourd.     

What determines conditionality in ant–Hemiptera interactions? Hemiptera habitat preference and the role of local ant activity

Abstract.  1. This paper describes spatial variation in density of a mutualist Hemiptera, and attempts to elucidate an understanding of the spatial variation in conditionality of its mutualism with ants.
2. Aggregations of the membracid treehopper, Campylenchia sp., occurred more frequently, and with higher number of individuals, in isolated trees as compared with their occurrence in the same tree species in wooded patches.
3. Using treehopper aggregations as the level of replication, there was a habitat × ant presence interaction associated with colony survival time. However, when the median survival time of all aggregations within each tree was used, only an additive effect of both habitat type and ant presence were apparent. This suggests that treehopper aggregations in some isolated trees experienced more benefit from ant presence than other trees.
4. The ant:treehopper ratio averaged over each tree was correlated with effect of ant presence on survivorship (survival time without ants/survival time with ants). Therefore, conditionality in the ant–membracid mutualism appears related to ant attendance level at the scale of individual trees.
5. Ant attendance at treehopper aggregations in isolated and matorral trees did not differ, and therefore there is no clear mechanism by which ants create the contrast in treehopper densities between the two habitat types.
6. The main force shaping initial treehopper densities in the two habitats is likely plant quality. Results suggest that growing situation (or more specifically plant health) is likely a good predictor of membracid density, while general ant activity within a tree is the best indicator of interaction strength.     

Invertebrate vectors of Phytophthora palmivora, causing black pod disease of cocoa in Ghana

Evidence is presented for the involvement of a range of invertebrates in the epidemiology of black pod disease of cocoa. Tent-building ant species, notably Crematogaster striatula, Camponotus acvapimensis and Pheidole megacephala , are considered to contribute to vertical spread of the pathogen within the cocoa tree. The building material used in tent construction is a potential source of inoculum. Certain phytophagous invertebrates and sugar-feeding Diptera colonize black pods in the field and may carry viable inoculum externally, on their bodies and mouthparts, or internally, in their faeces. Two species, Brachypeplus pilosellus (Coleoptera) and Chaetonerius latifemur (Diptera), stand out as important vectors because of their abundance, their habit of visiting wounded pods and their proven efficiency in transmission experiments. These vector species quickly colonize the lesions of pod-wounding insects (mirids, pod borers) and are considered to be the main means of local and long-distance horizontal spread of the pathogen. Radioactive tracers were extensively used to determine the movements of potential vectors.     

The influence of temperature and fine-scale resource distribution on resource sharing and domination in an ant community

Abstract.  1. In order to coexist in sympatry, subordinate species must somehow obtain resources that dominant taxa may generally control. This study examines the response of an ant community to fine-scale variation in resource distribution, as a mechanism enabling resource acquisition by subordinates in the presence of dominant taxa. Food (6 g) was portioned as one, eight, or 64 items in a 0.4-m² area, centred on nests of Monomorium sydneyense Forel, considered the most dominant ant in the area. This ant is a newly established exotic species in our study site of Tauranga, New Zealand.
2. As the number of food items increased, a significant increase in the number of species utilising the food was observed, associated with an increase in the non-utilised proportion of food items.
3. The changing occupation rates of food items by M. sydneyense and three other species were modelled against soil surface temperature and the varying dispersion rates of the food resource. Significant main effects of food density, site, species, and temperature were observed on the probability of food being occupied by a species. However, there were also significant interaction effects, making it impossible to interpret the main effects in isolation. Monomorium sydneyense dominated a smaller proportion of resources in trials with increased resource distribution and cooler temperatures, allowing more species to access resources.
4. There was considerable variation between species in their responses to variation in temperature and food distribution. Although most species were able to respond to increasing resource dispersion, species that were in low abundance apparently could not. However, even species that could respond to increasing resource dispersion were limited in the number of resources they could secure.