Dynamics and drivers of fungal communities in a multipartite ant-plant association.

BACKGROUND: Fungi and ants belong to the most important organisms in terrestrial ecosystems on Earth. In nutrient-poor niches of tropical rainforests, they have developed steady ecological relationships as a successful survival strategy. In tropical ant-plant mutualisms worldwide, where resident ants provide the host plants with defense and nutrients in exchange for shelter and food, fungi are regularly found in the ant nesting space, inhabiting ant-made dark-colored piles ("patches"). Unlike the extensively investigated fungus-growing insects, where the fungi serve as the primary food source, the purpose of this ant-fungi association is less clear. To decipher the roles of fungi in these structures within ant nests, it is crucial to first understand the dynamics and drivers that influence fungal patch communities during ant colony development. RESULTS: In this study, we investigated how the ant colony age and the ant-plant species affect the fungal community in the patches. As model we selected one of the most common mutualisms in the Tropics of America, the Azteca-Cecropia complex. By amplicon sequencing of the internal transcribed spacer 2 (ITS2) region, we analyzed the patch fungal communities of 93 Azteca spp. colonies inhabiting Cecropia spp. trees. Our study demonstrates that the fungal diversity in patches increases as the ant colony grows and that a change in the prevalent fungal taxa occurs between initial and established patches. In addition, the ant species significantly influences the composition of the fungal community in established ant colonies, rather than the host plant species. CONCLUSIONS: The fungal patch communities become more complex as the ant colony develops, due to an acquisition of fungi from the environment and a substrate diversification. Our results suggest a successional progression of the fungal communities in the patches during ant colony growth and place the ant colony as the main driver shaping such communities. The findings of this study demonstrate the unexpectedly complex nature of ant-plant mutualisms in tropical regions at a micro scale.

Evolutionary déjà vu? A case of convergent evolution in an ant-plant association.

Obligatory ant-plant symbioses often appear to be single evolutionary shifts within particular ant lineages; however, convergence can be revealed once natural history observations are complemented with molecular phylogenetics. Here, we describe a remarkable example of convergent evolution in an ant-plant symbiotic system. Exclusively arboreal, Myrmelachista species can be generalized opportunists nesting in several plant species or obligately symbiotic, live-stem nesters of a narrow set of plant species. Instances of specialization within Myrmelachista are known from northern South America and throughout Middle America. In Middle America, a diverse radiation of specialists occupies understory treelets of lowland rainforests. The morphological and behavioural uniformity of specialists suggests that they form a monophyletic assemblage, diversifying after a single origin of specialization. Using ultraconserved element phylogenomics and ancestral state reconstructions, we show that shifts from opportunistic to obligately symbiotic evolved independently in South and Middle America. Furthermore, our analyses support a remarkable case of convergence within the Middle American radiation, with two independently evolved specialist clades, arising nearly simultaneously from putative opportunistic ancestors during the late Pliocene. This repeated evolution of a complex phenotype suggests similar mechanisms behind trait shifts from opportunists to specialists, generating further questions about the selective forces driving specialization.

Plant species with larger extrafloral nectaries produce better quality nectar when needed and interact with the best ant partners.

Few studies have explored the phenotypic plasticity of nectar production on plant attractiveness to ants. Here, we investigate the role of extrafloral nectary (EFN) size on the productivity of extrafloral nectar in three sympatric legume species. We hypothesized that plant species with larger EFNs (i) have higher induced nectar secretion after herbivory events, and (ii) are more likely to interact with more protective (i.e. dominant) ant partners. We target 90 plants of three Chamaecrista species in the field. We estimated EFN size and conducted field experiments to evaluate any differences in nectar traits before and after leaf damage to investigate the phenotypic plasticity of nectar production across species. We conducted multiple censuses of ant species feeding on EFNs over time. Plant species increased nectar descriptors after leaf damage, but in different ways. Supporting our hypothesis, C. duckeana, with the largest EFN size, increased all nectar descriptors, with most intense post-herbivory-induced response, taking its place as the most attractive to ants, including dominant species. EFN size variation was an excellent indicator of nectar productivity across species. The higher control over reward production in plants with larger sized EFNs reflects an induction mechanism under damage that reduces costs and increases the potential benefits of indirect biotic defences.

Trees harbouring ants are better defended than con-generic and sympatric ant-free trees.

Plant strategies against herbivores are classically divided into chemical, physical, biotic defences. However, little is known about the relative importance of each type of plant defence, especially in the same species. Using the myrmecophyte Triplaris americana (both with and without ants), and the congeneric non-myrmecophyte T. gardneriana, we tested whether ant defence is more effective than other defences of naturally ant-free myrmecophytes and the non-myrmecophyte congeneric species, all spatially co-occurring. In addition, we investigated how plant traits vary among plant groups, and how these traits modulate herbivory. We sampled data on leaf area loss and plant traits from these tree groups in the Brazilian Pantanal floodplain, and found that herbivory is sixfold lower in plants with ants than in ant-free plants, supporting a major role of biotic defences against herbivory. Whereas ant-free plants had more physical defences (sclerophylly and trichomes), they had little effect on herbivory-only sclerophylly modulated herbivory, but with opposite effects depending on ants' presence and species identity. Despite little variation in the chemicals among plant groups, tannin concentrations and δ13C signatures negatively affected herbivory in T. americana plants with ants and in T. gardneriana, respectively. We showed that ant defence in myrmecophytic systems is the most effective against herbivory, as the studied plants could not fully compensate the lack of this biotic defence. We highlight the importance of positive insect-plant interactions in limiting herbivory, and therefore potentially plant fitness.

Tradeoffs in the evolution of plant farming by ants.

Diverse forms of cultivation have evolved across the tree of life. Efficient farming requires that the farmer deciphers and actively promotes conditions that increase crop yield. For plant cultivation, this can include evaluating tradeoffs among light, nutrients, and protection against herbivores. It is not understood if, or how, nonhuman farmers evaluate local conditions to increase payoffs. Here, we address this question using an obligate farming mutualism between the ant Philidris nagasau and epiphytic plants in the genus Squamellaria that are cultivated for their nesting sites and floral rewards. We focused on the ants' active fertilization of their crops and their protection against herbivory. We found that ants benefited from cultivating plants in full sun, receiving 7.5-fold more floral food rewards compared to shade-cultivated plants. The higher reward levels correlated with higher levels of crop protection provided by the ants. However, while high-light planting yielded the greatest immediate food rewards, sun-grown crops contained less nitrogen compared to shade-grown crops. This was due to lower nitrogen input from ants feeding on floral rewards instead of insect protein gained from predation. Despite this tradeoff, farming ants optimize crop yield by selectively planting their crops in full sun. Ancestral state reconstructions across this ant-plant clade show that a full-sun farming strategy has existed for millions of years, suggesting that nonhuman farmers have evolved the means to evaluate and balance conflicting crop needs to their own benefit.

Nitrogen fixation by diverse diazotrophic communities can support population growth of arboreal ants.

BACKGROUND: Symbiotic ant-plant associations, in which ants live on plants, feed on plant-provided food, and protect host trees against threats, are ubiquitous across the tropics, with the Azteca-Cecropia associations being amongst the most widespread interactions in the Neotropics. Upon colonization of Cecropia's hollow internodes, Azteca queens form small patches with plant parenchyma, which are then used as waste piles when the colony grows. Patches-found in many ant-plant mutualisms-are present throughout the colony life cycle and may supplement larval food. Despite their initial nitrogen (N)-poor substrate, patches in Cecropia accommodate fungi, nematodes, and bacteria. In this study, we investigated the atmospheric N2 fixation as an N source in patches of early and established ant colonies. RESULTS: Via 15N2 tracer assays, N2 fixation was frequently detected in all investigated patch types formed by three Azteca ant species. Quantified fixation rates were similar in early and established ant colonies and higher than in various tropical habitats. Based on amplicon sequencing, the identified microbial functional guild-the diazotrophs-harboring and transcribing the dinitrogenase reductase (nifH) gene was highly diverse and heterogeneous across Azteca colonies. The community composition differed between early and established ant colonies and partly between the ant species. CONCLUSIONS: Our data show that N2 fixation can result in reasonable amounts of N in ant colonies, which might not only enable bacterial, fungal, and nematode growth in the patch ecosystems but according to our calculations can even support the growth of ant populations. The diverse and heterogeneous diazotrophic community implies a functional redundancy, which could provide the ant-plant-patch system with a higher resilience towards changing environmental conditions. Hence, we propose that N2 fixation represents a previously unknown potential to overcome N limitations in arboreal ant colonies.

Food habits of 3 myrmecophilous bug species on myrmecophytic Macaranga (Malpighiales: Euphorbiaceae) vary from herbivory to predation.

Myrmecophytes have mutualistic relationships with symbiotic ants. Although myrmecophytic Macaranga (Malpighiales: Euphorbiaceae) species are well protected by aggressive Crematogaster (Hymenoptera: Formicidae) ants, some bug species occur on the myrmecophytes. To clarify the associations of these bugs with the plants and the ants, we studied the food habits of 3 bug species, Pilophorus lambirensis Nakatani et Komatsu, 2013 (Hemiptera: Miridae: Phylinae), Phylinae sp. 1, and Arbela sp. 1 (Hemiptera: Nabidae). We conducted field observations in a Bornean rainforest. First, we located these bugs and studied their behavioral responses to the ants on Macaranga species; we then conducted stable isotope analyses. All bugs avoided direct contact with ants, but they occurred only on trees with active ants. Pilophorus lambirensis and Phylinae sp. 1 were most commonly observed on the apical parts of host trees, whereas Arbela sp. 1 was mainly in areas distant from the apical parts where ants were sparse. The stable isotope ratios indicated that Phylinae sp. 1 fed on food bodies, which are nutrient-rich spherical bodies produced by Macaranga trees on the apical parts for ants. Although the main diet of the other 2 species remains unclear, nitrogen isotopic signatures demonstrated that P. lambirensis is herbivorous, whereas Arbela sp. 1 is carnivorous. However, the distant location from ants and its isotopic signatures indicated that Arbela sp. 1 rarely fed on the ants. At least 2 mirid bug species might obtain enemy-free space in addition to the food provided by the myrmecophytes.

Mutualism disruption by an invasive ant reduces carbon fixation for a foundational East African ant-plant.

Invasive ants shape assemblages and interactions of native species, but their effect on fundamental ecological processes is poorly understood. In East Africa, Pheidole megacephala ants have invaded monodominant stands of the ant-tree Acacia drepanolobium, extirpating native ant defenders and rendering trees vulnerable to canopy damage by vertebrate herbivores. We used experiments and observations to quantify direct and interactive effects of invasive ants and large herbivores on A. drepanolobium photosynthesis over a 2-year period. Trees that had been invaded for ≥ 5 years exhibited 69% lower whole-tree photosynthesis during key growing seasons, resulting from interaction between invasive ants and vertebrate herbivores that caused leaf- and canopy-level photosynthesis declines. We also surveyed trees shortly before and after invasion, finding that recent invasion induced only minor changes in leaf physiology. Our results from individual trees likely scale up, highlighting the potential of invasive species to alter ecosystem-level carbon fixation and other biogeochemical cycles.

Seeing through the static: the temporal dimension of plant-animal mutualistic interactions.

Most studies of plant-animal mutualistic networks have come from a temporally static perspective. This approach has revealed general patterns in network structure, but limits our ability to understand the ecological and evolutionary processes that shape these networks and to predict the consequences of natural and human-driven disturbance on species interactions. We review the growing literature on temporal dynamics of plant-animal mutualistic networks including pollination, seed dispersal and ant defence mutualisms. We then discuss potential mechanisms underlying such variation in interactions, ranging from behavioural and physiological processes at the finest temporal scales to ecological and evolutionary processes at the broadest. We find that at the finest temporal scales (days, weeks, months) mutualistic interactions are highly dynamic, with considerable variation in network structure. At intermediate scales (years, decades), networks still exhibit high levels of temporal variation, but such variation appears to influence network properties only weakly. At the broadest temporal scales (many decades, centuries and beyond), continued shifts in interactions appear to reshape network structure, leading to dramatic community changes, including loss of species and function. Our review highlights the importance of considering the temporal dimension for understanding the ecology and evolution of complex webs of mutualistic interactions.

Slippery flowers as a mechanism of defence against nectar-thieving ants.

BACKGROUND AND AIMS: The great diversity of floral characteristics among animal-pollinated plants is commonly understood to be the result of coevolutionary interactions between plants and pollinators. Floral antagonists, such as nectar thieves, also have the potential to exert an influence upon the selection of floral characteristics, but adaptation against floral antagonists has attracted comparatively little attention. We found that the corollas of hornet-pollinated Codonopsis lanceolata (Campanulaceae) and the tepals of bee-pollinated Fritillaria koidzumiana (Liliaceae) are slippery to nectar-thieving ants living in the plant's habitat; because the flowers of both species have exposed nectaries, slippery perianths may function as a defence against nectar-thieving ants. METHODS: We conducted a behavioural experiment and observed perianth surface microstructure by scanning electron microscopy to investigate the mechanism of slipperiness. Field experiments were conducted to test whether slippery perianths prevent floral entry by ants, and whether ant presence inside flowers affects pollination. KEY RESULTS: Scanning electron microscopy observations indicated that the slippery surfaces were coated with epicuticular wax crystals. The perianths lost their slipperiness when wiped with hexane. Artificial bridging of the slippery surfaces using non-slippery materials allowed ants to enter flowers more frequently. Experimental introduction of live ants to the Codonopsis flowers evicted hornet pollinators and shortened the duration of pollinator visits. However, no statistical differences were found in the fruit or seed sets of flowers with and without ants. CONCLUSIONS: Slippery perianths, most probably based on epicuticular wax crystals, prevent floral entry by ants that negatively affect pollinator behaviour. Experimental evidence of floral defence based on slippery surfaces is rare, but such a mode of defence may be widespread amongst flowering plants.

Short-term plasticity and variation in acacia ant-rewards under different conditions of ant occupancy and herbivory.

In ant-plant defense mutualisms, plants known as myrmecophytes provide food and shelter to ant partners in exchange for defense against herbivores and pathogens. To ensure interaction pay-off, myrmecophytes must regulate their investment in ant-rewards depending on local conditions and herbivore pressure. We investigated how myrmecophyte investment in multiple ant-rewards relates to herbivory, ant defense, and ant occupancy over time. Specifically, we examined the plasticity of ant-rewards produced by swollen-thorn acacias (Vachellia collinsii) under different ant occupancy and herbivory conditions. We compared food rewards (number of extrafloral nectaries and pinnules as a proxy for food bodies) and housing rewards (domatia dimensions) of V. collinsii for three conditions: (1) occupied (defended by the obligate mutualist Pseudomyrmex spinicola) versus unoccupied trees, (2) occupied trees subject to an experimental herbivory manipulation versus control trees, and (3) trees occupied by different ant species varying in their level of defense (P. spinicola, P. simulans, Crematogaster crinosa). We found that food rewards were more likely to vary in time depending on ant occupancy and resident species. Conversely, housing rewards varied with the condition (occupancy or species of partner) and less through time. A one-time herbivory manipulation did not cause any changes to the ant-rewards produced. Our results reveal short-term plasticity in V. collinsii ant-rewards and demonstrate that myrmecophytes with constitutive rewards can adjust their investment in ant-rewards depending on the presence and identity of ant partners.

Gram-negative bacteria associated with a dominant arboreal ant species outcompete phyllosphere-associated bacteria species in a tropical canopy.

Ants have efficient and well-studied social immunity mechanisms, which prevent the colony contamination. Little is known about how workers keep their outside territory clear of diseases. We investigated the interactions between Azteca chartifex ants, their associated bacteria and bacteria on the phyllosphere of Byrsonima sericea trees, comparing plants patrolled and not by the ants. The hypothesis is that bacteria associated with the worker's exoskeleton may outcompete the leaf bacteria. Culturable bacteria were isolated from ants, from the main and satellite nests, and from phyllosphere of B. sericea taken from trees that had A. chartifex nests and from trees without nests. The isolates were grouped by Gram guilds and identified at the genus level. There was a higher percentage of Gram-negative isolates in the ants and on the leaves patrolled by them. There was a higher growth rate of ant bacteria from the main nest compared to those found in ants from the satellite nests. The most representative genus among ant isolates was Enterobacter, also found on leaves patrolled by ants. Under favourable in vitro conditions, A. chartifex Gram-negative bacteria outcompete leaf bacteria by overgrowth, showing a greater competition capacity over the Gram-positive bacteria from leaves with no previous interaction with ants in the field. It was demonstrated that ants carry bacteria capable of outcompeting exogenous bacteria associated with their outside territory. The leaf microbiota of a patrolled tree could be shaped by the ant microbiota, suggesting that large ant colonies may have a key role in structuring canopy plant-microbe interactions.

Morphological characterization of domatium development in Callicarpa saccata.

BACKGROUND AND AIMS: Domatia are plant structures within which organisms reside. Callicarpa saccata (Lamiaceae) is the sole myrmecophyte, or 'ant plant', that develops foliar (leaf-borne) myrmeco-domatia in this genus. In this work we examined domatium development in C. saccata to understand the developmental processes behind pouch-like domatia. METHODS: Scanning electron microscopy, sectioning and microcomputed tomography were carried out to compare the leaves of C. saccata with those of the closely related but domatia-less myrmecophyte Callicarpa subaequalis, both under cultivation without ants. KEY RESULTS: Callicarpa saccata domatia are formed as a result of excess cell proliferation at the blade/petiole junctions of leaf primordia. Blade/petiole junctions are important meristematic sites in simple leaf organogenesis. We also found that the mesophyll tissue of domatia does not clearly differentiate into palisade and spongy layers. CONCLUSIONS: Rather than curling of the leaf margins, a perturbation of the normal functioning of the blade/petiole junction results in the formation of domatium tissue. Excess cell proliferation warps the shape of the blade and disturbs the development of the proximal-distal axis. This process leads to the generation of distinct structures that facilitate interaction between C. saccata and ants.

Pollen adaptation to ant pollination: a case study from the Proteaceae.

BACKGROUND AND AIMS: Ant-plant associations are widely diverse and distributed throughout the world, leading to antagonistic and/or mutualistic interactions. Ant pollination is a rare mutualistic association and reports of ants as effective pollinators are limited to a few studies. Conospermum (Proteaceae) is an insect-pollinated genus well represented in the south-western Australia biodiversity hotspot, and here we aimed to evaluate the role of ants as pollinators of C. undulatum. METHODS: Pollen germination after contact with several species of ants and bees was tested for C. undulatum and five co-flowering species for comparison. We then sampled the pollen load of floral visitors of C. undulatum to assess whether ants carried a pollen load sufficient to enable pollination. Lastly, we performed exclusion treatments to assess the relative effect of flying- and non-flying-invertebrate floral visitors on the reproduction of C. undulatum. For this, we measured the seed set under different conditions: ants exclusion, flying-insects exclusion and control. KEY RESULTS: Pollen of C. undulatum, along with the other Conospermum species, had a germination rate after contact with ants of ~80 % which did not differ from the effect of bees; in contrast, the other plant species tested showed a drop in the germination rate to ~10 % following ant treatments. Although ants were generalist visitors, they carried a pollen load with 68-86 % of suitable grains. Moreover, ants significantly contributed to the seed set of C. undulatum. CONCLUSIONS: Our study highlights the complexity of ant-flower interactions and suggests that generalizations neglecting the importance of ants as pollinators cannot be made. Conospermum undulatum has evolved pollen with resistance to the negative effect of ant secretions on pollen grains, with ants providing effective pollination services to this threatened species.

Nest substrate, more than ant activity, drives fungal pathogen community dissimilarity in seed-dispersing ant nests.

Myrmecochory is a widespread mutualism in which plants benefit from seed dispersal services by ants. Ants might also be providing seeds with an additional byproduct benefit via reduced plant pathogen loads in the ant nest environment through their antimicrobial glandular secretions. We investigate this byproduct benefit by identifying fungal communities in ant nests and surrounding environments and quantifying fungal community change (1) through time, (2) between different nest substrates, and (3) as a function of average ant activity levels within nests (based on observed ant activity at nest entrances throughout the summer). We split fungal communities by functional guild to determine seed-dispersing ant-induced changes in the overall fungal community, the animal pathogen fungal community, the plant pathogen fungal community, and the myrmecochore pathogen fungal community. Nest substrate (soil or log) explained much of the variation in fungal community dissimilarity, while substrate occupation (ant nest or control sample) and time had no influence on fungal community composition. Average ant activity had no effect on the community turnover in fungal communities except for the myrmecochore pathogenic fungal community. In this community, higher ant activity throughout the summer resulted in more fluctuation in the pathogenic community in the ant nest. Our results are not consistent with a byproduct benefit framework in myrmecochory, but suggest that nest substrate drives dissimilarity in fungal communities. The influence of nest substrate on fungal communities has important implications for seeds taken into ant nests, as well as ant nest location choice by queens and during nest relocation.

Friend or Foe? Orb-Weaver Spiders Inhabiting Ant-Acacias Capture Both Herbivorous Insects and Acacia Ant Alates.

The orb-weaver spiders Eustala oblonga (Chickering) and Eustala illicita (O. Picard-Cambridge) (Araneae: Araneidae) inhabit the ant-defended acacias Vachellia melanocerus (Beurling) and Vachellia collinsii (Safford) (Fabales: Fabaceae), respectively, in Panama. These spiders do not capture patrolling Pseudomyrmex ants but exploit their plant-protection services to escape predation. What effect the spiders have on the ant-acacia mutualisms is unknown. They may provide an additional layer of plant defense by capturing flying herbivorous insects in their webs. Alternatively, the spiders may disrupt the ant-acacia mutualisms by capturing alate acacia ants during nuptial flights. We evaluated these two hypotheses by sampling insects flying through acacia foliage and by identifying prey remains in webs. The proportions of insects captured on sticky card traps and in webs varied with taxonomic order and ecological role. Herbivorous insects greatly outnumbered other groups captured on sticky cards and were captured in spiders' webs in both acacia species but made up a minority of prey remains in webs. Instead, insect predators and parasitoids made up the majority of prey remains and were comprised primarily by alate ant mutualists of the host acacias. These results provide indirect support for both hypotheses and suggest that the spiders potentially both benefit and harm their host ant-acacia mutualisms. The net effect of spider exploitation, however, is unclear and is likely based on both the effectiveness of plant protection from herbivory provided by the spiders relative to that provided by acacia ants, as well as the overall proportion of the ant reproductive caste the spiders actually capture.

Plant performance response to eight different types of symbiosis.

Almost all plant species interact with one or more symbioses somewhere within their distribution range. Bringing together plant trait data and growth responses to symbioses spanning 552 plant species, we provide for the first time on a large scale (597 studies) a quantitative synthesis on plant performance differences between eight major types of symbiosis, including mycorrhizas, N-fixing bacteria, fungal endophytes and ant-plant interactions. Frequency distributions of plant growth responses varied considerably between different types of symbiosis, in terms of both mean effect and 'risk', defined here as percentage of experiments reporting a negative effect of symbiosis on plants. Contrary to expectation, plant traits were poor predictors of growth response across and within all eight symbiotic associations. Our analysis showed no systematic additive effect when a host plant engaged in two functionally different symbioses. This synthesis suggests that plant species' ecological strategies have little effect in determining the influence of a symbiosis on host plant growth. Reliable quantification of differences in plant performance across symbioses will prove valuable for developing general hypotheses on how species become engaged in mutualisms without a guarantee of net returns.

Ant pollination of Paepalanthus lundii (Eriocaulaceae) in Brazilian savanna.

BACKGROUND AND AIMS: Ant-plant associations are widely diverse and distributed throughout the world, leading to complex ecological networks. Regarding ant-plant mutualism, ant pollination is a very rare interaction and few studies have shown the role of ants as pollinators. Therefore, we aimed to evaluate the role of ants as effective pollinators of Paepalanthus lundii (Eriocaulaceae) in a Brazilian savanna. METHODS: Fieldwork with experimental manipulation was conducted to evaluate the fitness of P. lundii, considering potential pollinators. For this, we mainly observed the number of seeds produced in different conditions: control, ant exclusion, exclusion of flying insects, and exclusion (entomophily test) of both ants and flying insects. Furthermore, we evaluated all floral visitors throughout the day, stigma receptivity, the numbers of male and female flowers, and patterns of species co-occurrence, which can indicate the presence of different pollinators in the plants at the same time. KEY RESULTS: We observed a relation between seed production and ant visits; Camponotus crassus was the most frequent floral visitor and the most effective pollinator. Also, we observed a statistical difference between the numbers of male and female flowers produced, with a greater number of male flowers. Furthermore, P. lundii presented flowering asynchrony, with 12 different types of maturation sequence, which indicates a cross-pollination system. Lastly, we observed an overlap of the greatest abundance of C. crassus and the time of plant stigmatic receptivity, and a pattern of non co-occurrence of ants, which shows the pollinator role of this ant. CONCLUSIONS: Our data provide evidence that previous generalizations neglecting the importance of ants as pollinators are wrong. Brazilian savanna can reveal a lot about the ant-pollination syndrome, since this environment presents peculiar characteristics related to this association. Thus, this study has great significance for the understanding of the ant-pollination syndrome, and for the understanding of the complex ecological networks present in these dry arid systems.

Effects of increasing aridity and chronic anthropogenic disturbance on seed dispersal by ants in Brazilian Caatinga.

Anthropogenic disturbance and climate change are the main drivers of biodiversity loss and ecological services around the globe. There is concern that climate change will exacerbate the impacts of disturbance and thereby promote biotic homogenization, but its consequences for ecological services are unknown. We investigated the individual and interactive effects of increasing chronic anthropogenic disturbance (CAD) and aridity on seed dispersal services provided by ants in Caatinga vegetation of north-eastern Brazil. The study was conducted in Catimbau National Park, Pernambuco, Brazil. Within an area of 214 km2 , we established nineteen 50 × 20 m plots that encompassed gradients of both CAD and aridity. We offered diaspores of six plant species, three myrmecochorous diaspores and three fleshy fruits that are secondarily dispersed by ants. We then quantified the number of interactions, seed removal rate and dispersal distances, and noted the identities of interacting ant species. Finally, we used pitfall trap data to quantify the abundances of ant disperser species in each plot. Our results show that overall composition of ant disperser species varied along the gradients of CAD and aridity, but the composition of high-quality dispersers varied only with aridity. The total number of interactions, rates of removal and mean distance of removal all declined with increasing aridity, but they were not related to CAD. These same patterns were found when considering only high-quality disperser species, driven by the responses of the dominant disperser Dinoponera quadriceps. We found little evidence of interactive effects of CAD and aridity on seed dispersal services by ants. Our study indicates that CAD and aridity act independently on ant-mediated seed dispersal services in Caatinga, such that the impacts of anthropogenic disturbance are unlikely to change under the forecast climate of increased aridity. However, our findings highlight the vulnerability of seed dispersal services provided by ants in Caatinga under an increasingly arid climate due to low functional redundancy in high-quality disperser species. Given the large number of plant species dependent on ants for seed dispersal, this has important implications for future plant recruitment and, consequently, for the composition of Caatinga plant communities.

Acacia trees with parasitic ants have fewer and less spacious spines than trees with mutualistic ants.

Obligate ant-defended plants provide food and shelter in exchange for protection against herbivores. Mesoamerican acacia trees have an obligate ant mutualism, but parasitic non-defending ants can also nest on the tree. We assessed whether rewards corresponded to ant defense within a plant species. As we expected, we found that parasite-inhabited trees had fewer swollen spines than ant-defended trees. Spine diameter was smaller in parasite-inhabited plants, but there were no differences in spine length, suggesting that spines serve as mechanical protection against herbivory. Parasite-inhabited plants may have reduced rewards because of plant differences when establishing, a plastic response to limited resources, or differential energy allocation when sensing the lack of defense.