Catching ants with honey: an experimental test of distraction and satiation as alternative modes of escape from flower-damaging ants

According to the distraction hypothesis, extrafloral nectaries (EFN) evolved under selection to entice ants away from floral nectaries, reducing ant-mediated damage to flowers and/or interference with pollinators. Predator-satiation, through production of nectar in either surplus flowers or EFN, provides an alternative mechanism for reducing the impact of ants as flower visitors. I tested these two hypotheses by experimentally adding EFN to flowering plants of the alpine wildflower, Polemonium viscosum, and by surveying the relationship between ant visitation and nectary number in nature. Plants of P. viscosum lack EFN and experience flower damage by ants of Formica neorufibarbus gelida. Ant behavior was compared on plants with five flowers and three experimental EFN and on controls with equal floral display, but no EFN. Addition of EFN increased flower visitation by ants. The effect of EFN on flower visitation did not depend on proximity of EFN to flowers or attractiveness of EFN to ants. Findings suggest that ants perceived patch quality on a whole plant basis, rather than responding to EFN and flowers as distinct nectar patches. Ant visitation did not keep pace with nectary number in nature. The relationship between ant visitation and nectary number per plant was weak and shallow as predicted under satiation. Ant foraging choices on experimental inflorescences showed that ants bypass flowers avoided by earlier ants, enhancing probability of escape via satiation. Results do not support the idea that EFN evolve to reduce flower visitation by ants, but show instead that nectar in surplus flowers can satiate ants and reduce their negative impacts on flower function and integrity.     

Role of extrafloral nectaries of Vicia faba in attraction of ants and herbivore exclusion by ants

We directly evaluated the role of extrafloral nectaries (EFN) in ant attraction and herbivore exclusion by experimental removal of EFN in the laboratory. When EFN of Vicia faba Linnaeus (Leguminosae) were artificially removed, the number of workers of Tetramorium tsushimae Emery (Hymenoptera: Formicidae) visiting the plant decreased, and the efficiency of herbivore exclusion by ants also decreased. Herbivore exclusion by ants was mostly ineffective on a plant when less than four workers visited the plant, but when more than four workers visited, the time a herbivore resided on the plant decreased rapidly with increasing numbers of visiting ants. Therefore, the efficiency of herbivore exclusion from a plant is determined by the number of ants visiting, and EFN play an important role in ant attraction.     

Competition hierarchy and plant defense in a guild of ants on tropical Passiflora

In facultative ant–plant interactions, ants may compete with each other for food provided by extrafloral nectar (EFN) plants. We studied resource competition and plant defense in a guild of ants that use the same EFN resource provided by two species of Passiflora in a seasonal rain forest in tropical China. At least 22 ant species were recorded using the EFN resource, although some of those species were rare. Among these ants, Paratrechina sp.1 and Dolichoderus thoracicus were more aggressive than other species. Ant aggressiveness measured as ant behavioral dominance index (BDI) was positively correlated with ant abundance on the Passiflora species studied. Ant BDI was also positively correlated to the protection that ants provided against herbivory. In Passiflora siamica, the number of workers patrolling on the plants did negatively correlate with average leaf loss per plant. We conclude that in this facultative Passiflora–ant system, plant defense upon herbivore was indeed influenced by the total number of ants present on plant and the aggressiveness of these ants.     

Plant‐derived differences in the composition of aphid honeydew and their effects on colonies of aphid‐tending ants

In plant–ant–hemipteran interactions, ants visit plants to consume the honeydew produced by phloem‐feeding hemipterans. If genetically based differences in plant phloem chemistry change the chemical composition of hemipteran honeydew, then the plant's genetic constitution could have indirect effects on ants via the hemipterans. If such effects change ant behavior, they could feed back to affect the plant itself. We compared the chemical composition of honeydews produced by Aphis nerii aphid clones on two milkweed congeners, Asclepias curassavica and Asclepias incarnata, and we measured the responses of experimental Linepithema humile ant colonies to these honeydews. The compositions of secondary metabolites, sugars, and amino acids differed significantly in the honeydews from the two plant species. Ant colonies feeding on honeydew derived from A. incarnata recruited in higher numbers to artificial diet, maintained higher queen and worker dry weight, and sustained marginally more workers than ants feeding on honeydew derived from A. curassavica. Ants feeding on honeydew from A. incarnata were also more exploratory in behavioral assays than ants feeding from A. curassavica. Despite performing better when feeding on the A. incarnata honeydew, ant workers marginally preferred honeydew from A. curassavica to honeydew from A. incarnata when given a choice. Our results demonstrate that plant congeners can exert strong indirect effects on ant colonies by means of plant‐species‐specific differences in aphid honeydew chemistry. Moreover, these effects changed ant behavior and thus could feed back to affect plant performance in the field.     

Do additional sugar sources affect the degree of attendance of Dysmicoccus brevipes by the fire ant Solenopsis geminata?

Mutualistic interactions between ants and Hemiptera are mediated to a large extent by the amount and quality of sugar‐rich honeydew produced. Throughout the neotropics, the predaceous fire ant Solenopsis geminata (Fabricius) (Hymenoptera: Formicidae) is found in association with colonies of the pineapple mealybug, Dysmicoccus brevipes (Cockerell) (Hemiptera: Pseudococcidae), which they actively tend and protect from attack by natural enemies. In this study, we evaluate the effects of access to a sucrose solution on the mutualistic association between S. geminata and D. brevipes. Ten colonies of either species were established, with D. brevipes maintained on pumpkin, Cucurbita maxima Duchesne (Cucurbitaceae), in screen cages. Five of the S. geminata colonies were permitted access to vials with 20% sucrose solution and a pumpkin with 20 adult mealybugs. The remaining ant colonies were allowed access to mealybug‐infested pumpkins. Ant colonies with access to the sucrose solution attended mealybugs significantly less than those without additional sugar sources. Mealybug survival rates were similar under both treatments. Total body sugars and fructose were nearly twice as high in ants with access to honeydew and sucrose vs. those with access to honeydew and water. Fructose accumulated on the pumpkins over time in both treatments, suggesting that honeydew was not fully exploited by the ants. In conclusion, D. brevipes enjoy lower degrees of ant attendance when S. geminata have alternative sources of carbohydrates. We further discuss the significance of these findings for the conservation of predaceous ants and mealybug biological control.     

Ecological effects of multi‐species,ant–hemipteran mutualisms in citrus

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Ant–plant interaction in the Neotropical savanna: direct beneficial effects of extrafloral nectar on ant colony fitness

Current evidence suggests that ant–plant relationships may influence species composition, abundance, and interactions at the community scale. The main resource that plants offer to ants is extrafloral nectar (EFN) and the major part of published studies shown benefits from ants to plants possessing EFNs. However, the complementary question of whether and how ants benefit from EFNs is rarely addressed. Here, we present the results of a long-term study to demonstrate whether EFN has a positive effect on ant colony fitness. We quantified colony growth rate, survival and the final weight of individuals as measures of benefit derived from EFN. Our results provide clear evidence that EFN can have a significant positive impact on the survivorship, growth and reproduction of the Myrmicinae Cephalotes pusillus. In fact, a diet rich in EFN (providing at least 30 cal per day) resulted in five times more individuals per colony, greater body weights, and more eggs. These results have shed new light on the relationships between ants and EFN-bearing plants such as in tropical and temperate systems. The ant C. pusillus is the first case in which we have firm evidence that EFN improves colony growth and development, corroborating more than 100 years of experimental evidence of benefits to plants in these widespread relationships.     

Consequences of ants and extrafloral nectar for a pollinating seed‐consuming mutualism: ant satiation,floral distraction or plant defense?

Non‐pollinating consumers of floral resources, especially ants, can disrupt pollination and plant reproductive processes. As an alternative food resource to flowers, extrafloral nectar (EFN) may distract and satiate ants from flowers, thereby reducing their antagonistic effects on plants. Yet, EFN may actually attract and increase ant density on plants, thus increasing the disruption of pollination and/or their defense of plants. In this study, we tested the effects of ants and EFN on pollinating seed‐consuming interactions between senita cacti and senita moths in the Sonoran Desert. Prior study of senita showed that EFN can distract ants from flowers, but consequences for plant–pollinator interactions remain unstudied. In our current study, ant exclusion had no effect on pollination or oviposition when moths were abundant (>85% flower visitation). Yet, in an ant by EFN factorial experiment under lower moth abundance (<40% visitation), there was a significant effect of ant exclusion (but not EFN or an ant × EFN) on pollination and oviposition. In contrast with our predictions, ant presence increased rather than decreased pollination (and oviposition) by moths, indicating a beneficial effect of ants on plant reproduction. While ant density on plants showed a saturating response to continuous experimental variation in EFN, in support of ant satiation and distraction, the probability of pollination and oviposition increased and saturated with ant density, again showing a beneficial effect of ants on plant reproduction. Ants showed no significant effect on fruit set, fruit survival, or fruit production of oviposited flowers in the ant exclusion experiment. Ants did not affect the survival of moth larvae, but there was a marginally significant effect of ants in reducing wasp parasitism of moths. We suggest that EFN may not only distract ants from disrupting plant–pollinator interactions, but they may also enhance plant–pollinator interactions by increasing pollination and reducing wasp parasitism. Though often considered dichotomous hypotheses, ant distraction and plant defense may be synergistic, though the mechanism(s) for such positive ant effects on plant–pollinator interactions needs further study.     

Quarantining behaviour in ants: are Myrmica aphid milkers able to detect and get rid of fungus-contaminated aphids?

Fungal infections are highly dangerous for social insects including ants. Close trophobiotic interaction between ants and aphids promotes infection transmission, as aphids can be a disease vector. The ability of ants to detect fungus-infected aphids and get rid of them is important to the prosperity of both symbiotic partners. However, the diversity of quarantining behaviour among ants is still poorly studied. Here, the behaviour of honeydew foragers of two ant species – Myrmica rubra L. and Myrmica scabrinodis Nylander (Hymenoptera: Formicidae, Myrmicinae, Myrmicini) – was studied in laboratory towards Schizaphis graminum (Rondani) (Hemiptera: Aphididae, Aphidini) aphids contaminated with the generalist fungal pathogen Beauveria bassiana (Balsamo-Crivelli) Vuillemin (Hypocreales). This fungus attacks a wide range of hosts including aphids and ants. The removal of conidia-contaminated aphids from the host plant was found not to be typical of the ants studied. Aphid milkers of M. rubra and M. scabrinodis usually displayed non-aggressive behaviour (tolerance, antennation, honeydew collection, grooming) towards the experimental aphids regardless of whether they were covered with conidia or not. Neither ant species, nor the number of milkers had significant effects on their behaviour towards ‘infected’ aphids. However, some individuals were found to demonstrate quarantining behaviour in full. They quickly detected and removed contaminated aphids, placing them at some distance from the plant. Moreover, in addition to the simple carrying of ‘infected’ aphids down, the more effective technique of dropping them from the plant was noted as well. Ants of the genus Myrmica appear to have a tendency to perform a certain sequence of actions to remove conidia-contaminated aphids from the plant. It is likely that in larger colonies or under conditions of increased risk of infection with entomopathogenic fungi, some Myrmica ants are able to deploy and actively use the behavioural pattern of quarantining behaviour to increase their viability.     

Interactions between extrafloral nectaries, aphids and ants: are there competition effects between plant and homopteran sugar sources?

Broad bean (Vicia faba), an annual plant bearing extrafloral nectaries (EFN) at the base of the upper leaves, is regularly infested by two aphid species, Aphis fabae and Acyrthosiphon pisum. EFN and A. fabae are commonly attended by the ant, Lasius niger, while Ac. pisum usually remains uninfested. Sugar concentration and sugar composition of extrafloral nectar did not change significantly after aphid infestation. The sugar concentration was significantly higher in EFN (c. 271 µg µl^-1) than in the honeydew of A. fabae (37.5 µg µl^-1). The presence of small A. fabae colonies had no significant effect on ant attendance of EFN, which remained at the same level as that on plants without A. fabae. Obviously, there was no significant competitive effect between the two sugar sources. We suggest that the high sugar concentration in the extrafloral nectar may outweigh the higher quality (due to the presence of melezitose) and quantity of aphid honeydew. Ants and the presence of EFN influenced aphid colony growth. While A. fabae colonies generally grew better in the presence of ants, Ac. pisum colonies declined on plants with EFN or A. fabae colonies. We conclude that EFN may provide some degree of protection for V. faba against those sucking herbivores that are not able to attract ants.     

Extrafloral nectaries as a deterrent mechanism against seed predators in the chemically protected weed Crotalaria pallida (Leguminosae)

Abstract In several plants, extrafloral nectaries (EFN) are located close to the reproductive structures, suggesting that ants may act as a defence against specialized seed predators that overcome chemical defences. Alternatively, ants may also deter herbivores in a generalized manner, thereby protecting the whole plant. In this work, we examined the relationship between the chemically protected weed Crotalaria pallida Ait. (Leguminosae) that bears EFN, its specialized seed predator, the larvae of the arctiid moth Utetheisa ornatrix L. (Arctiidae) and ants. We tested two hypotheses related to the type of deterrence caused by ants. The Seed Predator Deterrence Hypothesis predicts that ant deterrence is directed primarily towards herbivores that destroy seeds and other reproductive structures, without attacking herbivores on vegetative structures. The General Deterrence Hypothesis states that ants are general in their effects, equally deterring herbivores in vegetative and reproductive structures. Our results supported the predictions of the Seed Predator Deterrence Hypothesis, namely, that (i) ant activity on EFN was related to the vulnerability of reproductive structures to attack by U. ornatrix; (ii) ant patrolling was restricted almost entirely to racemes; (iii) ants removed termites used as baits more frequently on racemes than on leaves; and (iv) U. ornatrix larvae were often expulsed from the racemes. These results indicate that EFN can act as another deterrent mechanism in chemically protected plants by promoting the expulsion of specialist seed predators.     

Elevated dominance of extrafloral nectary-bearing plants is associated with increased abundances of an invasive ant and reduced native ant richness

Aim Invasive ants can have substantial and detrimental effects on co‐occurring community members, especially other ants. However, the ecological factors that promote both their population growth and their negative influences remain elusive. Opportunistic associations between invasive ants and extrafloral nectary (EFN)‐bearing plants are common and may fuel population expansion and subsequent impacts of invasive ants on native communities. We examined three predictions of this hypothesis, compared ant assemblages between invaded and uninvaded sites and assessed the extent of this species in Samoa. Location The Samoan Archipelago (six islands and 35 sites). Methods We surveyed abundances of the invasive ant Anoplolepis gracilipes, other ant species and EFN‐bearing plants. Results Anoplolepis gracilipes was significantly more widely distributed in 2006 than in 1962, suggesting that the invasion of A. gracilipes in Samoa has progressed. Furthermore, (non‐A. gracilipes) ant assemblages differed significantly between invaded and uninvaded sites. Anoplolepis gracilipes workers were found more frequently at nectaries than other plant parts, suggesting that nectar resources were important to this species. There was a strong, positive relationship between the dominance of EFN‐bearing plants in the community and A. gracilipes abundance on plants, a relationship that co‐occurring ants did not display. High abundances of A. gracilipes at sites dominated by EFN‐bearing plants were associated with low species richness of native plant‐visiting ant species. Anoplolepis gracilipes did not display any significant relationships with the diversity of other non‐native ants. Main conclusions Together, these data suggest that EFN‐bearing plants may promote negative impacts of A. gracilipes on co‐occurring ants across broad spatial scales. This study underscores the potential importance of positive interactions in the dynamics of species invasions. Furthermore, they suggest that conservation managers may benefit from explicit considerations of potential positive interactions in predicting the identities of problematic invaders or the outcomes of species invasions.     

Tuned protection of aphids by ants against a predatory hoverfly

1. Aphid‐tending ants that feed on honeydew have evolved strategies against aphidophagous insects and tune their aggressive behaviour according to the level of danger for their trophobionts. Here we investigate how Lasius niger Linnaeus (Hymenoptera: Formicidae) ants react to different instars of Episyrphus balteatus De Geer (Diptera: Syrphidae) hoverflies which vary in their voracity and defensive abilities. 2. During pairwise encounters, early syrphid instars (eggs, L1, and L2 larvae) elicited lower aggression scores compared to third larval instars (L3), which was intensively bitten by ants. L3 tried to escape from ants by releasing a sticky and toxic secretion over biting ants that died or underwent severe morbidity. 3. In a standardised system including the host plant, aphid, tending ant, and hoverfly, the ability of ants to protect an Aphis fabae Scopoli (Hemiptera: Aphididae) colony was evaluated. Early E. balteatus instars placed onto the plant elicited no mobilisation of ants, which often removed the hoverfly successfully. Eggs and early instars appeared as the weak links for integrated pest management by hoverfly auxiliaries. 4. In contrast, L3 induced the number of ant patrollers to increase at a local scale without any further recruitment from inside the ant nest. L3 syrphids were quite efficient at gluing ants with defensive secretions and at resisting to removal attempts by ants. 5. While supporting the assumption that ants tune their defensive response to the aphidophagous predator, the present results also showed a lack of efficient protection of their trophobionts from the most voracious late syrphid instar.     

Increased Host Investment in Extrafloral Nectar (EFN) Improves the Efficiency of a Mutualistic Defensive Service

Extrafloral nectar (EFN) plays an important role as plant indirect defence through the attraction of defending ants. Like all rewards produced in the context of a mutualism, however, EFN is in danger of being exploited by non-ant consumers that do not defend the plant against herbivores. Here we asked whether plants, by investing more in EFN, can improve their indirect defence, or rather increase the risk of losing this investment to EFN thieves. We used the obligate plant-ant Acacia-Pseudomyrmex system and examined experimentally in the field during the dry and the rainy seasons how variations in EFN secretion are related to (i) ant activity, to (ii) the ant-mediated defence against herbivores and (iii) the exploitation of EFN by non-ant consumers. Extrafloral investment enhanced ant recruitment and was positively related to the ant mediated defence against herbivores. The ant-mediated protection from exploiters also increased in proportion to the nectar sugar concentration. Although the daily peak of EFN production coincided with the highest activity of EFN thieves, Pseudomyrmex ferrugineus ants protected this resource effectively from exploiters. Nevertheless, the defensive effects by ants differed among seasons. During the dry season, plants grew slower and secreted more EFN than in the rainy season, and thus, experienced a higher level of ant-mediated indirect defence. Our results show that an increased plant investment in an indirect defence trait can improve the resulting defensive service against both herbivores and exploiters. EFN secretion by obligate ant-plants represents a defensive trait for which the level of investment correlates positively with the beneficial effects obtained.     

Arboreal Ants Use the “Velcro? Principle” to Capture Very Large Prey

Plant-ants live in a mutualistic association with host plants known as “myrmecophytes” that provide them with a nesting place and sometimes with extra-floral nectar (EFN) and/or food bodies (FBs); the ants can also attend sap-sucking Hemiptera for their honeydew. In return, plant-ants, like most other arboreal ants, protect their host plants from defoliators. To satisfy their nitrogen requirements, however, some have optimized their ability to capture prey in the restricted environment represented by the crowns of trees by using elaborate hunting techniques. In this study, we investigated the predatory behavior of the ant Azteca andreae which is associated with the myrmecophyte Cecropia obtusa. We noted that up to 8350 ant workers per tree hide side-by-side beneath the leaf margins of their host plant with their mandibles open, waiting for insects to alight. The latter are immediately seized by their extremities, and then spread-eagled; nestmates are recruited to help stretch, carve up and transport prey. This group ambush hunting technique is particularly effective when the underside of the leaves is downy, as is the case for C. obtusa. In this case, the hook-shaped claws of the A. andreae workers and the velvet-like structure of the underside of the leaves combine to act like natural Velcro® that is reinforced by the group ambush strategy of the workers, allowing them to capture prey of up to 13,350 times the mean weight of a single worker.     

The function of extrafloral nectaries in the pollination and reproduction of Sambucus javanica

Sambucus javanica is a perennial herb with extrafloral nectaries (EFNs) on its inflorescences. To explore the ecological functions of EFNs, a factorial combination experiment of ant (access or exclusion) and EFNs (with or without) at the plant level was created in two populations. The role of EFNs in the attraction of ants and flying pollinators, the defensive role of ants against foliar herbivores, the effects of ants on pollinator visitation and the effects of ant–pollinator interactions on fruit production in one or both populations were assessed. Ants were common on the ant-access plants with EFNs, but absent from the ant-access plants without EFNs. Foliar herbivory was independent of ant and EFN treatments and their interactions. The visitation frequency of flying pollinators (honeybees and syrphid flies) and fruit set were significantly higher for plants with EFNs than plants without EFNs, but were not affected by ant treatments or their interactions with EFN treatments. These results suggest that EFNs in S. javanica attracted both ants and flying pollinators, but ants did not present a defensive role against herbivores, did not deter flying pollinators from visiting inflorescences and had no effects on fruit production. In addition, ants were not significant pollen vectors.     

Effectiveness of protein-rich pea flour for the control of stored-product beetles

Abstract The association between visiting ants and the extrafloral nectaries (EFN)‐bearing shrub Hibiscus pernambucensis Arruda (Malvaceae) was investigated in two different coastal habitats – a permanently dry sandy forest and a regularly inundated mangrove forest. In both habitats the frequency of plants with ants and the mean number of ants per plant were much higher on H. pernambucensis than on non‐nectariferous neighbouring plants. In the sandy forest the proportion of live termite baits attacked by ants on H. pernambucensis was much higher than on plants lacking EFNs. In the mangrove, however, ants attacked equal numbers of termites on either plant class. Ant attendance to tuna/honey baits revealed that overall ant activity in the sandy forest is higher than in the mangrove area. The vertical distribution (ground vs. foliage) of ant activity also differed between habitats. While in the mangrove foraging ants were more frequent at baits placed on foliage, in the sandy forest ant attendance was higher at ground baits. Plants housing ant colonies were more common in the mangrove than in the sandy forest. Frequent flooding in the mangrove may have resulted in increased numbers of ant nests on vegetation and scattered ant activity across plant foliage, irrespective of possession of EFNs. Thus plants with EFNs in the mangrove may not experience increased ant aggression towards potential herbivores relative to plants lacking EFNs. The study suggests that the vertical distribution of ant activity, as related to different nest site distribution (ground vs. foliage) through a spatial scale, can mediate ant foraging patterns on plant foliage and probably affect the ants’ potential for herbivore deterrence on an EFN‐bearing plant species.     

Does the availability of arboreal honeydew determine the prevalence of ecologically dominant ants in restored habitats?

Ants are extensive users of arboreal sugars, but little is known about how ecological dominance or habitat succession influences this interaction. We investigated how the availability and use of arboreal sugar resources by ants changes across a restoration chronosequence. We surveyed the use and availability of hemipteran honeydew and floral nectar on the two dominant plant genera, Eucalyptus and Acacia, in study sites in south eastern Australia. Sugars used by ants are likely to drive their role as ecosystem engineers, while sugars not used by ants remain available to other organisms. We also tested whether the use of sugars differed between ecologically dominant and non-dominant ants; taxa likely to perform different functions in ecosystems. No floral nectar was available on Acacia, but later successional eucalypts supported more floral resources and fewer mutualist hemiptera. Successional stage significantly affected how much sugar remained unexploited by ants, with similar trends for ant use of sugars. Non-dominant ants used mainly floral nectar, while hemipteran honeydew resources were used disproportionately by dominant ants, consistent with the prediction that this group monopolises persistent carbohydrate resources. This pattern was similar across successional stages, but the difference was least in habitats with the greatest availability of floral nectar, suggesting that high sugar availability may reduce the incentive to defend honeydew. Across habitat types, the proportion of dominant ants increased with the availability of hemipteran honeydew. This suggests that honeydew availability may regulate ecological dominance, thus affecting ant-driven ecosystem processes.     

Ants have a negative rather than a positive effect on extrafloral nectaried Crotalaria pallida performance

Crotalaria pallida (Fabaceae) is a pantropical plant with extrafloral nectaries (EFNs) near the reproductive structures. EFN-visiting ants attack and remove arctiid moth Utetheisa ornatrix larvae, the main pre-dispersal seed predator, but the impact of ants on C. pallida fitness is unknown. To assess this impact, we controlled ant presence on plants and evaluated the reproductive output of C. pallida with and without ants. Predatory wasps also visit EFNs, prey upon U. ornatrix larvae, and may be driven out by ants during EFN feeding. Does this agonistic interaction affect the multitrophic interaction outcome? We found it difficult to evaluate the effect of both visitors because cages excluding wasps affect plant growth and do not allow U. ornatrix oviposition. Therefore, we verified whether ant presence inhibited wasp EFN visitation and predicted that (1) if ants confer a benefit for C. pallida, any negative effect of ants on wasps would be negligible for the plant because ants would be the best guardians, and (2) if ants are poor guardians, they would negatively affect wasps and negatively impact the fitness of C. pallida. Surprisingly, we found that the number of seeds/pods significantly increased, ca. 4.7 times, after ant removal. Additionally, we unexpectedly verified that controls showed a higher percentage of herbivore bored pods than ant-excluded plants. We found that wasps spent less time visiting EFNs patrolled by ants (ca. 299 s less). These results support our second prediction and suggest that the outcome of multitrophic interactions may vary with natural enemy actors.     

Extrafloral nectar as a driver of arboreal ant communities at the site‐scale in Brazilian savanna

Despite years of study, it remains unclear if and to what extent the effects of extra‐floral nectaries (EFNs) on arboreal ants observed on individual trees scale up to larger spatial scales. Here, we address this issue in Brazilian savanna and tested three predictions: (i) Trees with EFN have higher richness of arboreal ant species than trees without; (ii) Arboreal ant species richness increases with the proportion of total EFN‐bearing trees at the site scale, due to a higher occurrence of non‐core ant species; (iii) Ant species composition changes with the proportion of EFN‐bearing trees at the site scale. We sampled arboreal ants in 32 plots with EFN‐bearing trees ranging from 0% to 60% of all trees. We sampled 72 ant species, from which 17 (mostly belonging to Camponotus, Cephalotes and Crematogaster) were identified as core species in at least one of the ant‐EFN networks in the 32 plots. Ant species richness was significantly higher on EFN‐bearing trees. We identified 11 ant species that preferentially occurred on EFN‐bearing trees, all of which were core partners in networks. Species richness at the site scales increased with the proportion of EFN‐bearing trees, regardless of tree density and richness; this pattern was due to a higher occurrence of non‐core ant species. Finally, species composition also varied with the proportion of EFN‐bearing trees. Therefore, we found that the presence of EFNs not only influences arboreal ants on individual trees but also has a substantial effect on the ant‐EFN network on a broader community scale. The increase in non‐core species site scale reveals that this interaction is unlikely to result in substantially enhanced protection services for EFN‐bearing plants.