Ant Exclusion Research Articles

Cover crops dismantle keystone ant/aphid mutualisms to enhance insect pest suppression and weed biocontrol

Abstract Cover crops are multifunctional tools that mitigate environmental impacts of agriculture, enhance resilience to weather extremes and suppress weeds and arthropod pests. Cover crops provide non‐crop food and habitat resources that attract natural enemies of pests, but their outcomes for pest management are less clear in regions where keystone mutualisms between red imported fire ants and aphids dominate. Here, we manipulate ant exclusion treatments and cover crop treatments (living mulches and terminated cover crops) that vary in food/habitat resources and examine responses of ants, aphids, other herbivores and predators in a cotton agroecosystem. Living mulches reduced both ants and aphids in the crop canopy by 97% and 93%, respectively, relative to bare soil treatments, and terminated cover crops reduced them as well by a lesser degree (~50%). Non‐aphid herbivores occurred in low densities system‐wide and increased in living mulches, whilst native predators had variable responses to cover crops and ant exclusion. Cover crops had no effect on prey removal in the crop canopy, but living mulches tripled rates of weed seed biocontrol, relative to bare soils. Cover crops elicited a shift in fire ant foraging from cotton foliage downward to the soil‐surface, preventing competitive exclusion by keystone ant/aphid mutualists that dominate crop monocultures. Cover crops altered the system‐wide impacts of fire ants: reducing ecosystem disservices (i.e. aphid tending) and enhancing ecosystem services (i.e. weed seed biocontrol). These results provide incentives for cover crop adoption as a regenerative practice in large‐scale commercial agriculture.

Urbanization drives partner switching and loss of mutualism in an ant-plant symbiosis.

Mutualistic interactions between species underpin biodiversity and ecosystem function, but may be lost when partners respond differently to abiotic conditions. Except for a few prominent examples, effects of global anthropogenic change on mutualisms are poorly understood. Here we assess the effects of urbanization on a symbiosis in which the plant Cordia nodosa house ants in hollow structures (domatia) in exchange for defense against herbivores. We expected to find that mutualist ants would be replaced in the city by heat-tolerant opportunists, leaving urban plants vulnerable to herbivory. In five protected forest sites and five urban forest fragments in southeast Perú, we recorded the identity and heat tolerance (CTmax) of ant residents of C. nodosa. We also assayed their plant-defensive behaviors and their effects on herbivory. We characterized the urban heat-island effect in ambient temperatures and within domatia. Forest plants housed a consistent ant community dominated by three specialized plant ants, whereas urban plants housed a suite of 10 opportunistic taxa that were, collectively, about 13 times less likely than forest ants to respond defensively to plant disturbance. In the forest, ant exclusion had the expected effect of increasing herbivory, but in urban sites, exclusion reduced herbivory. Despite poor ant defense in urban sites, we detected no difference in total standing herbivory, perhaps because herbivores themselves also declined in the city. Urban sites were warmer than forest sites (daily maxima in urban domatia averaged 1.6°C hotter), and the urban ant community as a whole was slightly more heat tolerant. These results illustrate a case of mutualism loss associated with anthropogenic disturbance. If urbanization is representative of increasing anthropogenic stressors more broadly, we might expect to see destabilization of myrmecophytic mutualisms in forest ecosystems in the future.

Insectivorous birds and bats outperform ants in the top-down regulation of arthropods across strata of a Japanese temperate forest.

Birds, bats and ants are recognised as significant arthropod predators. However, empirical studies reveal inconsistent trends in their relative roles in top-down control across strata. Here, we describe the differences between forest strata in the separate effects of birds, bats and ants on arthropod densities and their cascading effects on plant damage. We implemented a factorial design to exclude vertebrates and ants in both the canopy and understorey. Additionally, we separately excluded birds and bats from the understorey using diurnal and nocturnal exclosures. At the end of the experiments, we collected all arthropods and assessed herbivory damage. Arthropods responded similarly to predator exclusion across forest strata, with a density increase of 81% on trees without vertebrates and 53% without both vertebrates and ants. Additionally, bird exclusion alone led to an 89% increase in arthropod density, while bat exclusion resulted in a 63% increase. Herbivory increased by 42% when vertebrates were excluded and by 35% when both vertebrates and ants were excluded. Bird exclusion alone increased herbivory damage by 28%, while the exclusion of bats showed a detectable but non-significant increase (by 22%). In contrast, ant exclusion had no significant effect on arthropod density or herbivory damage across strata. Our results reveal that the effects of birds and bats on arthropod density and herbivory damage are similar between the forest canopy and understorey in this temperate forest. In addition, ants were not found to be significant predators in our system. Furthermore, birds, bats and ants appeared to exhibit antagonistic relationships in influencing arthropod density. These findings highlight, unprecedentedly, the equal importance of birds and bats in maintaining ecological balance across different strata of a temperate forest.

Foraging ants affect community composition and diversity of phyllosphere fungi on a myrmecophilous plants, Mallotus japonicus.

Many microorganisms inhabit the aboveground parts of plants (i.e. the phyllosphere), which mainly comprise leaves. Understanding the structure of phyllosphere microbial communities and their drivers is important because they influence host plant fitness and ecosystem functions. Despite the high prevalence of ant-plant associations, few studies have used quantitative community data to investigate the effects of ants on phyllosphere microbial communities. In the present study, we investigated the effects of ants on the phyllosphere fungal communities of Mallotus japonicus using high-throughput sequencing. Mallotus japonicus is a myrmecophilous plants that bears extrafloral nectaries, attracting several ant species, but does not provide specific ant species with nest sites like myrmecophytes do. We experimentally excluded ants with sticky resins from the target plants and collected leaf discs to extract fungal DNA. The ribosomal DNA internal transcribed spacer 1 (ITS1) regions of the phyllosphere fungi were amplified and sequenced to obtain fungal community data. Our results showed that the exclusion of ants changed the phyllosphere fungal community composition; however, the effect of ants on OTU richness was not clear. These results indicate that ants can change the community of phyllosphere fungi, even if the plant is not a myrmecophyte.

Interspecific competition between ants and African honeybees (Apis mellifera scutellata) may undermine the effectiveness of elephant beehive–deterrents in Africa

AbstractBeehive deterrents are commonly used to mitigate human–elephant conflict and protect woody vegetation. To ensure hive activity, reduce abscondment risks, and maintain deterrent effectiveness, resident bee colonies require supplementary feeding during periods of low resource availability. However, our study found that ants frequently consume the supplementary feed in open feeders intended for bees. Anoplolepis custodiens was the most numerically dominant species that excluded bees from the feeders, followed by Camponotus and Crematogaster spp. With higher ant abundance, the predicted probability of zero bees being present at feeders increased up to 82%. This competition may undermine the efficacy of beehive deterrents as a conflict mitigation tool. We developed a simple and effective ant exclusion method that raised the overall predicted probability of bees' presence at supplementary feeding stations from 32% to 68%. Our findings suggest that innovative solutions to exclude ants from supplementary feed may improve the implementation and success of this conflict mitigation method across Africa.

Vertebrates, but not ants, protect rainforest from herbivorous insects across elevations in Papua New Guinea

AbstractAimThe effects of insectivorous predators on herbivorous prey should have a cascading effect releasing herbivory pressure and favouring plant's biomass. However, it remains unclear whether different types of predators regulate herbivores to the same degree across seasons, and how their interactions affect lower trophic levels across elevations where predator communities differ significantly. Therefore, we investigated the impact of excluding flying vertebrate predators and ants (individually and in combination) on arthropods and herbivory across tropical seasons along a rainforest gradient spanning 3500 m a.s.l.LocationPapua New Guinea.TaxonMulti‐taxon.MethodsWe excluded predators from 560 saplings in two 6‐month‐long predator exclusion experiments, controlling for seasonality. Saplings were spread across eight sites, evenly spaced at 500 m elevational increments from 200 to 3700 m a.s.l. We measured change in the abundance of arthropods and herbivory damage and analysed them by linear and generalized linear mixed models.ResultsExclusion of flying vertebrate predators, but not ants, led to a significant increase in both arthropod density and herbivory damage. The density of arthropods increased significantly by 37% when flying vertebrates were excluded and by 33% when both flying vertebrates and ants were excluded. Both season and elevation significantly influenced this effect. Leaf damage increased significantly by 50% in exclosures of flying vertebrates alone and by 36% in combined exclosures of flying vertebrates and ants. In contrast, the exclusion of ants alone had no significant effect on arthropod density or leaf damage, which increased by 12% and 9%, respectively, although the effect decreased with increasing elevation.Main conclusionsThe overall effect of flying vertebrate predators on arthropod density and leaf damage remains consistently strong along the whole elevational gradient. In contrast, ant‐driven trophic cascades were detected only in lowland. Disappearance of insectivorous flying vertebrates could lead to substantial negative consequences for plants.

Impacts of invasive ants on pollination of native plants are similar in invaded and restored plant communities

Ants are amongst the most successful invaders worldwide and can markedly modify invaded communities through biotic interactions. Invasive ants, for example, can compete with native pollinators for resources, act as legitimate pollinators, or disrupt plant-pollinator mutualistic interactions, thereby affecting native plant reproduction. Ecological restoration aims to mitigate the impact of invasive species and restore ecological communities and ecosystem functioning. Here we investigated the combined effects of two invasive ant species, the yellow crazy ant (Anoplolepis gracilipes) and white-footed ant (Technomyrmex albipes), on the pollination and reproduction of native plants in the context of a large-scale plant community restoration experiment. We conducted ant exclusion experiments and pollinator observations to obtain data on plant reproduction and pollination of four plant species. The experiments were conducted across eight granitic inselbergs on the island of Mahé, Seychelles, four of which are undergoing restoration since 2011 and four remained as invaded control sites (unrestored). Our findings indicate that native plants on Seychelles’ inselbergs relied primarily on flying flower visitors for pollination, considering that the contribution of ants to fruit set was generally small and substantially lower than that of flying pollinators. Moreover, while flying pollinators approached flowering plants irrespective of invasive ant presence, the number of probed flowers per visit was reduced, in both restored and unrestored communities. Therefore, native plant reproduction was negatively affected by the presence of invasive ants and this effect was similar in restored and unrestored plant communities. Our research suggests that invasive ants may contribute to changes in plant communities by deterring legitimate pollinators of native plants, an impact that is unlikely to be alleviated through early stages of ecological restoration.

Exclusion of Mediterranean ant species enhances biological control of the invasive mealybug Delottococcus aberiae in citrus.

Delottococcus aberiae is an invasive mealybug that produces severe damage in Spanish citrus. This mealybug has established a mutualistic relationship with native Mediterranean ant species that may limit biological control of this pest. Herein, we evaluated the effect of tending ants on the biological control of D. aberiae. To do this, we compared: (i) the density of D. aberiae, (ii) the density of its natural enemies, and (iii) the damage produced by the mealybug in trees with (control) and without ants (ants excluded with sticky barriers) in two citrus orchards across two consecutive years. Lasius grandis was the most abundant ant species in both orchards and represented more than 95% of the ants tending D. aberiae in control trees. Spiders and lacewings were the most abundant predators observed in mealybug colonies, and the exclusion of mutualistic ants increased their abundance. Moreover, in control trees, ant activity throughout the year was negatively correlated with the relative abundance of predators (number of predators per mealybug). No parasitoids were recovered during field experiments. Ant exclusion reduced the density of D. aberiae and the ratio of damaged fruit at harvest across years and orchards. This work corroborates the previous finding that D. aberiae benefits from its mutualistic relationship with L. grandis, probably because the presence of ants reduced the abundance of generalist predators. This mutualism can be disrupted using physical barriers on on the trunk. Further research should assess other methods of ant control that are more economic and feasible for citrus producers. © 2023 Society of Chemical Industry.

Structural and functional organization of ant communities in secondary meadow ecosystems

Introduction. Due to a significant biomass and diverse ecological niches, ants (Formicidae, Hymenoptera) are crucial in establishing the structure and functioning of the co-adaptive species complexes, i.e. myrmecocomplexes, to which they belong. The majority of ant species, which build their nests with visible anthills, create humpy micro-relief changing the habitats of numerous plant and animal species. The number of active ant families and the number of their nests indicate the functional power of every species in an ant community. According to the dynamics of nest numbers, it is possible to analyze the changes in the ecosystem as a whole. Materials and methods. The research territory is located in the SE suburb of Lviv on the edge of Davydiv and Holohory ridges. It includes two former agricultural sites: fallow arable land and fallow grazing land. The material was collected by the method of ant exclusion with its following conservation, laboratory analysis and determination. Anthill mapping was conducted on 200 sq. m sites. Results and discussion. The results suggest that the more structured a habitat is, the higher ant species diversity is observed: seven species were found in the fallow grazing land (Formica pratensis – dominant, F. cunicularia – subdominant, Tetramorium caespitum, Lasius niger, L. flavus, Myrmica rubra, Solenopsis fugax – all are influents). It is typical of the three influent species (L. flavus, T. caespitum, M. rubra) to distribute within the territory of a dominant’s foraging area. However, the influent ant species avoid contacting each other. The less structured habitat (fallow arable land) is presented by only two of the most adaptive species, which are in high numbers (L. niger – 95 % of the site nests, and L. flavus – only 5 % of them). Conclusions. Thus, each ant community is a system of „dominant–subdominant–influent” species, which is under permanent transformation due to anthropogenic habitat changes; only well-adapted species can stay in the community under the influence of the mentioned changes. The research on the spatial distribution of ant nests testifies to the prospects and need for the mapping method used to establish the type and level of previous anthropogenic impact on the ecosystem (grazing, ploughing, grass burning, etc.).

Resilience of native ant community against invasion of exotic ants after anthropogenic disturbances of forest habitats.

The positive association between disturbances and biological invasions is a widely observed ecological pattern in the Anthropocene. Such patterns have been hypothesized to be driven by the superior competitive ability of invaders or by modified environments, as well as by the interaction of these factors. An experimental study that tests these hypotheses is usually less feasible, especially in protected nature areas. An alternative approach is to focus on community resilience over time after the anthropogenic disturbance of habitats. Here, we focused on ant communities within a forest to examine their responses after disturbance over time. We selected the Yanbaru region of northern Okinawa Island, which is a biodiversity hotspot in East Asia. We compared ant communities among roadside environments in forests where the road age differed from 5 to 25 years. We also monitored the ant communities before and after disturbance from forest thinning. We found that the species richness and abundance of exotic ants were higher in recently disturbed environments (roadsides of 5–15 years old roads), where the physical environment was warmer and drier. In contrast, the roadsides of 25‐year‐old roads indicated the potential recovery of the physical environment with cooler and moister conditions, likely owing to regrowth of roadside vegetation. At these sites, there were few exotic ants, except for those immediately adjacent to the road. The population density of the invasive species Technoymex brunneus substantially increased 1–2 years after forest thinning. There was no evidence of the exclusion of native ants by exotic ants that were recorded after disturbance. Our results suggest that local ant communities in the Yanbaru forests have some resilience to disturbance. We suggest that restoration of environmental components is a better strategy for maintaining native ant communities, rather than removing exotic ants after anthropogenic disturbance.

Long‐term quantification of leaf‐cutting ant damage in willow forestations in the lower delta of the Paraná River, Argentina

Abstract Leaf‐cutting ants are major pests of Neotropics forest plantations. The lower delta of the Paraná River contains the main Argentine Salicaceae production, strongly attacked by Acromyrmex lundii and Acromyrmex ambiguus. Nevertheless, there is no damage quantification in willow plantations attributed to leaf‐cutting ant species. In an area without leaf‐cutting ant control, we installed 15 blocks with eight willow stakes each: four clones × two treatments; with and without leaf‐cutting ant exclusion. We used two traditional (Americano, Nigra 4) and two new (Géminis, Yaguareté) commercial clones. During 2014–2018, we measured the damaged foliage, height and diameter of each tree. Foliage was damaged intensely during the first 2 years. After 1537 days, the loss in height and diameter was greater in Americano (70%), followed by Géminis (50%–60%), Yaguareté (40%–50%) and Nigra 4 (45%–40%). Stake survival with exclusion was greater (>80%) than stakes without exclusion (<50%). Total loss of wood volume was 93% for Americano, followed by Géminis (77%), Yaguareté (66%) and Nigra 4 (51%). Although the new clones were heavily attacked, they produced two to three times more wood volume than Americano; replacing Americano with the new clones would help to reduce leaf‐cutting ants impact on plantations and pesticides released into the environment.

Weak effects of birds, bats, and ants on their arthropod prey on pioneering tropical forest gap vegetation.

The relative roles of plants competing for resources versus top-down control of vegetation by herbivores, in turn impacted by predators, during early stages of tropical forest succession remain poorly understood. Here we examine the impact of insectivorous birds, bats, and ants exclusion on arthropods communities on replicated 5 × 5 m of pioneering early successional vegetation plots in lowland tropical forest gaps in Papua New Guinea. In plots from which focal taxa of predators were excluded we observed increased biomass of herbivorous and predatory arthropods, and increased density, and decreased diversity of herbivorous insects. However, changes in the biomass of plants, herbivores, and arthropod predators were positively correlated or uncorrelated between these three trophic levels and also between individual arthropod orders. Arthropod abundance and biomass correlated strongly with the plant biomass irrespective of the arthropods' trophic position, a signal of bottom-up control. Patterns in herbivore specialization confirm lack of a strong top-down control and were largely unaffected by the exclusion of insectivorous birds, bats, and ants. No changes of plant-herbivore interaction networks were detected except for decrease in modularity of the exclosure plots. Our results suggest weak top-down control of herbivores, limited compensation between arthropod and vertebrate predators, and limited intra-guild predation by birds, bats, and ants. Possible explanations are strong bottom-up control, a low activity of the higher order predators, especially birds, possibly also bats, in gaps, and continuous influx of herbivores from surrounding mature forest matrix.

Functional roles of ants in a temperate grassland.

Ants in temperate grasslands are consumers and ecosystem engineers, influencing biodiversity and potentially grassland productivity. However, the effects of ant exclusion or suppression on resource removal and the biological community in temperate grasslands have yet to be fully explored. We conducted ant-suppression experiments and evaluated the effects of ants on ground-dwelling arthropod communities in the field by using pitfall and bait traps. In the laboratory, we evaluated the effects of ants on the ant-attended aphid Aphis rumicis, which is a honeydew resource for ants, and the slug (Deroceras laeve), an aphid predator. Aboveground arthropod communities were not affected by the ant-suppression treatment. However, slugs (D. laeve and Ambigolimax valentianus) visited bait resources more frequently in the ant-suppression treatment area. In the ant-absence condition in the laboratory experiment, there were fewer aphids on the plants compared to the ant-presence condition owing to predation by D. laeve. Our results suggest that ant abundance in temperate grasslands influences the predation activity of slugs toward honeydew sources such as aphids.

Density thresholds and the incorporation of biocontrol into decision-making to enhance the control of Cacopsylla pyri in pear (cv. Ercolini) orchards.

Economic injury level (EIL) and economic threshold (ET) are customary tools for integrated pest management. Cacopsylla pyri L. is a major pest in pear orchards. The aim of this work was to establish EIL and ET for the optimization of the use of insecticides to control this psyllid, considering biocontrol and two spraying strategies (low-toxicity versus broad spectrum chemicals). This research was conducted over 4 years in five commercial pear, cv. Ercolini, orchards in south-eastern Spain. Psyllids and ant populations were followed using periodic sampling, and the russet on fruits was quantified. The effect of spray intensity and ant exclusion on psyllid abundance and yield were also tested: both had a significant effect on the cumulative number of C. pyri (CNP), yield and fruit weight. Yield was found to be negatively correlated with CNP. The russet index (RI) increased in a sigmoidal fashion as a function of CNP, being significantly higher with than without ant exclusion. The commercial categorization of fruits was explained satisfactorily as a function of CNP and the cumulative number of ants (CNA). The quantitative EIL was established at a CNP of 427.2 for spraying with paraffinic oil and 425.7 for abamectin. As for the cosmetic EIL, when CNA was zero, this EIL was 24.2, at a CNP of 16.6 for spraying with paraffinic oil or abamectin. The use of products of low toxicity, for the conservation of ants, is expected to increase ET and, thus, reduce the intensity of spraying. © 2021 Society of Chemical Industry.

Ground cover and native ant predation influence survival of metamorphic amphibians in a southeastern pine savanna undergoing restoration

Longleaf pine savannas historically supported abundant ground cover maintained by frequent fire but little other disturbance. Ground cover creates microclimates with lower temperatures, higher humidity, and increased soil moisture that may benefit wildlife, particularly small vertebrates such as amphibians. Today, most historical pine savannas have had extensive soil disturbance and altered fire regimes resulting in reduced ground cover and altered soil fauna communities including predatory invertebrates. We used a factorial terrestrial cage study to test the effects of native wiregrass (Aristida spp.) cover and the exclusion of a native predatory ant (Dorymyrmex smithi) on the survival of post‐metamorphic Ornate chorus frogs (Pseudacris ornata) and Gopher frogs (Rana capito). Although we were unable to achieve full ant exclusion, ant reduction in exclusion treatments and plant cover had an interactive effect on metamorph survival. Ant exclusion tended to increase Gopher frog survival and this effect was more pronounced when wiregrass was present. Within ant treatments, survival of Gopher frogs increased slightly with increasing wiregrass cover. Ornate chorus frogs had a high probability of survival (>95%) in all ant exclusion treatments regardless of wiregrass cover; however, in treatments without ant exclusion, survival increased with increasing wiregrass cover. Our results demonstrate that high abundances of a native ant species and low coverage of native wiregrass, which are legacies of historical soil disturbance and altered fire regimes, interact to elevate mortality of juvenile amphibians. Minimizing soil disturbance and restoring native ground cover are likely important for amphibian habitat management within historical southeastern pine savannas.

Intraspecific competition favors ant–plant protective mutualism

AbstractThe dependency of the anti‐herbivore defense on ant–plant protective mutualism often varies depending on abiotic and biotic conditions. Although intraspecific competition is a primary interaction between neighboring plants, its effects on ant–plant mutualisms have yet to be sufficiently elucidated. In order to determine the effects of intraspecific competition and competitor genotype on ant–plant mutualisms, I conducted competition and ant‐removal experiments and examined their effects on damage to the leaves of Urena lobata var. tomentosa plants. I found that larger numbers of worker ants visited the plants growing with non‐siblings than plants growing alone and that plants growing with non‐siblings had a higher shoot to root ratio and secreted greater volumes of extrafloral nectar than plants growing alone and/or with siblings. Under the presence of both sibling and non‐sibling competitors, I observed that when ants were removed from plants, those grown with conspecific neighbors were characterized by a higher percentage of damaged leaf area than plants harboring ants. The effect of ant exclusion on leaf damage was more pronounced in plants grown with non‐siblings than those grown near siblings. However, when the plants were grown alone, I detected no significant difference in percentage leaf damage between the ant‐excluded and ant‐harboring plants. The results indicate that neighboring plants can exert strong effects on ant–plant protective mutualisms, thereby highlighting the need to take into consideration plant–plant interactions in studies on these mutualistic associations.

Ants reduce fruit damage caused by psyllids in Mediterranean pear orchards

AbstractBACKGROUNDPsyllids are principal pests of pear orchards. Pest control based on the intensive use of chemicals is facing difficulties and strategies with a low impact on natural enemies are considered as the most suitable for psyllid control. Ants have been reported as the main predators of Cacopsylla pyri (Hemiptera: Psyllidae) in the southern Mediterranean area. The aim of this work was to determine the contribution of ants to the reduction of the fruit damage caused by psyllids at different intensities of spraying. Five levels of spraying intensity and ant exclusion were tested in a pear orchard with three blocks of five plots each.RESULTSThe incidence of russet on fruits increased with the abundance of psyllids and with ant exclusion. In the absence of ants, the impact of russet increased progressively from the highest to lowest intensity of spraying. By contrast, when ants were present no significant differences were found between the lowest (two application of paraffinic oil) and the highest (six applications of abamectin plus paraffinic oil) spraying levels. In the presence of ants, the incidence of russet in the plots with three applications of paraffinic oil was similar to those with three or six applications of paraffinic oil plus abamectin).CONCLUSIONSThe importance of ants for the control of psyllids and the reduction of fruit damage is shown. Reduction in the frequency of spraying and the use of low toxic insecticides for natural enemies is presented as the most sustainable strategy for psyllid control in pear orchards. © 2020 Society of Chemical Industry

Extranuptial nectaries in flowers: ants increase the reproductive success of the ant-plant Miconia tococa (Melastomataceae).

Although the production of extranuptial nectar is a common strategy of indirect defence against herbivores among tropical plants, the presence of extranuptial nectaries in reproductive structures is rare, especially in ant-plants. This is because the presence of ants in reproductive organs can generate conflicts between the partners, as ants can inhibit the activity of pollinators or even castrate their host plants. Here we evaluate the hypothesis that the ant-plant Miconia tococa produces nectar in its petals which attracts ants and affects fruit set. Floral buds were analysed using anatomical and histochemical techniques. The frequency and behaviour of floral visitors were recorded in field observations. Finally, an ant exclusion experiment was conducted to evaluate the effect of ant presence on fruit production. The petals of M. tococa have a secretory epidermis that produces sugary compounds. Nectar production occurred during the floral bud stage and attracted 17 species of non-obligate ants (i.e. have a facultative association with ant-plants). Ants foraged only on floral buds, and thus did not affect the activity of pollinators in the neighbouring open flowers. The presence of ants in the inflorescences increased fruit production by 15%. To our knowledge, the production of extranuptial nectar in the reproductive structures of a myrmecophyte is very rare, with few records in the literature. Although studies show conflicts between the partners in the ant-plant interaction, ants that forage on M. tococa floral buds protect the plant against floral herbivores without affecting bee pollination.

Defence variation within a guild of aphid‐tending ants explains aphid population growth

Mutualism studies often focus on the service provided by single species, while variation in benefits provided by multiple partners is less understood. Ant‐aphid food‐for‐protection mutualisms are suitable to study the implications of intra‐guild service variation because they often involve several ant species with varying levels of aggressiveness. We studied an aphid species and its associated ant guild to address whether intra‐guild defence variation against aphid natural enemies explains aphid performance on plants (thistles). We surveyed plants with natural abundances of aphids associated with different ant species and estimated aphid population growth. We conducted confrontation experiments between ant species and aphid natural enemies (ladybugs and hoverfly larvae). In plants patrolled by the most aggressive ant species, we determined the ant's probability of expelling aphid natural enemies and tested whether ant exclusion affects the abundance of aphids and their natural enemies. The ant Dorymyrmex tener was the most abundant and frequent species on plants and the most aggressive against aphid natural enemies. Aphid populations grew faster on plants patrolled by D. tener compared to plants patrolled by Camponotus distinguendus or D. richteri. Field experiments confirmed that D. tener effectively expels aphid natural enemies from plants. When D. tener was excluded, the density of aphids decreased, while the abundance of aphid natural enemies increased. The disruption of aphid predation by aggressive and numerically dominant ant species is a determinant of aphid population dynamics. This study illustrates the importance of considering intra‐guild service variation to better understand multi‐partner mutualisms.

Nectar Secretion of Floral Buds of Tococa guianensis Mediates Interactions With Generalist Ants That Reduce Florivory.

The specialised mutualism between Tococa guianensis and ants housed in its leaf domatia is a well-known example of myrmecophily. A pollination study on this species revealed that flowers in the bud stage exude a sugary solution that is collected by ants. Given the presence of this unexpected nectar secretion, we investigated how, where, and when floral buds of T. guianensis secret nectar and what function it serves. We studied a population of T. guianensis occurring in a swampy area in the Cerrado of Brazil by analyzing the chemical composition and secretion dynamics of the floral-bud nectar and the distribution and ultrastructure of secretory tissues. We also measured flower damage using ant-exclusion experiments. Floral bud nectar was secreted at the tip of the petals, which lack a typical glandular structure but possess distinctive mesophyll due to the presence of numerous calcium oxalate crystals. The nectar, the production of which ceased after flower opening, was composed mainly of sucrose and low amounts of glucose and fructose. Nectar was consumed by generalist ants and sporadically by stingless bees. Ant exclusion experiments resulted in significantly increased flower damage. The floral nectar of T. guianensis is produced during the bud stage. This bud-nectar has the extranuptial function of attracting generalist ants that reduce florivory. Pollen is the unique floral resource attracting pollinators during anthesis. Tococa guianensis, thus, establishes relationships with two functional groups of ant species: specialist ants acting against herbivory and generalist ants acting against florivory.