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.

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.

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.

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.

Overall seed dispersal effectiveness is lower in endemic Trillium species than in their widespread congeners.

PREMISE OF THE STUDY: Comparing ecological attributes of endemic species with related, widespread species can reveal differences accounting for rarity. Forests of the southeastern United States are home to many range-restricted endemic and widespread species of Trillium, a genus of ant-dispersed herbs. Evidence suggests that aspects of seed-related life history stages are often correlated with plant rarity, but few studies have tested whether the process of seed dispersal differs for endemic and widespread species. To address this question, we compared aspects of seed dispersal effectiveness (SDE) for three sympatric, widespread endemic Trillium species pairs. METHODS: We observed seed dispersal for Trillium species pairs by ants at eight sites, recorded numbers of seeds dispersed and dispersal distances, and described disperser interactions. To test disperser preference, we presented seeds of each pair to captive colonies of Aphaenogaster picea, a keystone disperser. Seeds were assigned scores based on worker behavior, and we recorded proportions of seeds dispersed after 1 h and 24 h. KEY RESULTS: Field observations yielded some significant within-pair differences. Ants dispersed more seeds of widespread species for all pairs, although dispersal distances did not differ. In laboratory experiments, after 24 h, ants dispersed more seeds of widespread species into nests. CONCLUSIONS: Endemic Trillium species had lower overall SDE than did their widespread congeners. These findings add to the list of ecological and demographic challenges that face endemic plants when compared to common congeners. Lower SDE may negatively impact reproductive rates and the colonization of new habitats, which may contribute to patterns of endemism.

Convergent structure and function of mycelial galleries in two unrelated Neotropical plant-ants.

The construction process and use of galleries by Azteca brevis (Myrmicinae: Dolichoderinae) inhabiting Tetrathylacium macrophyllum (Salicaceae) were compared with Allomerus decemarticulatus (Myrmicinae: Solenopsidini) galleries on Hirtella physophora (Chrysobalanaceae). Though the two ant species are phylogenetically distant, the gallery structure seems to be surprisingly similar and structurally convergent: both are pierced with numerous holes and both ant species use Chaetothyrialean fungi to strengthen the gallery walls. Al. decemarticulatus is known to use the galleries for prey capture and whether this is also the case for Az. brevis was tested in field experiments. We placed Atta workers as potential prey/threat on the galleries and recorded the behaviour of both ant species. We found considerable behavioural differences between them: Al. decemarticulatus was quicker and more efficient at capture than was Az. brevis. While most Atta workers were captured after the first 5 min by Al. decemarticulatus, significantly fewer were captured by Az. brevis even after 20 min. Moreover, the captured Atta were sometimes simply discarded and not taken to the nest by Az. brevis. As a consequence, the major function of the galleries built by Az. brevis may, therefore, be defense against intruders in contrast to Al. decemarticulatus which uses them mainly for prey capture. This may be due to a higher need for protein in Al. decemarticulatus compared to coccid-raising Az. brevis.

Extrafloral nectar at the plant-insect interface: a spotlight on chemical ecology, phenotypic plasticity, and food webs.

Plants secrete extrafloral nectar (EFN) as an induced defense against herbivores. EFN contains not only carbohydrates and amino acids but also pathogenesis-related proteins and other protective enzymes, making EFN an exclusive reward. EFN secretion is commonly induced after wounding, likely owing to a jasmonic acid-induced cell wall invertase, and is limited by phloem sucrose availability: Both factors control EFN secretion according to the optimal defense hypothesis. Non-ant EFN consumers include parasitoids, wasps, spiders, mites, bugs, and predatory beetles. Little is known about the relevance of EFN to the nutrition of its consumers and, hence, to the structuring of arthropod communities. The mutualism can be established quickly among noncoevolved (e.g., invasive) species, indicating its easy assembly is due to ecological fitting. Therefore, increasing efforts are directed toward using EFN in biocontrol. However, documentation of the importance of EFN for the communities of plants and arthropods in natural, invasive, and agricultural ecosystems is still limited.

Ant-caterpillar antagonism at the community level: interhabitat variation of tritrophic interactions in a neotropical savanna.

Ant foraging on foliage can substantially affect how phytophagous insects use host plants and represents a high predation risk for caterpillars, which are important folivores. Ant-plant-herbivore interactions are especially pervasive in cerrado savanna due to continuous ant visitation to liquid food sources on foliage (extrafloral nectaries, insect honeydew). While searching for liquid rewards on plants, aggressive ants frequently attack or kill insect herbivores, decreasing their numbers. Because ants vary in diet and aggressiveness, their effect on herbivores also varies. Additionally, the differential occurrence of ant attractants (plant and insect exudates) on foliage produces variable levels of ant foraging within local floras and among localities. Here, we investigate how variation of ant communities and of traits among host plant species (presence or absence of ant attractants) can change the effect of carnivores (predatory ants) on herbivore communities (caterpillars) in a cerrado savanna landscape. We sampled caterpillars and foliage-foraging ants in four cerrado localities (70-460 km apart). We found that: (i) caterpillar infestation was negatively related with ant visitation to plants; (ii) this relationship depended on local ant abundance and species composition, and on local preference by ants for plants with liquid attractants; (iii) this was not related to local plant richness or plant size; (iv) the relationship between the presence of ant attractants and caterpillar abundance varied among sites from negative to neutral; and (v) caterpillars feeding on plants with ant attractants are more resistant to ant predation than those feeding on plants lacking attractants. Liquid food on foliage mediates host plant quality for lepidopterans by promoting generalized ant-caterpillar antagonism. Our study in cerrado shows that the negative effects of generalist predatory ants on herbivores are detectable at a community level, affecting patterns of abundance and host plant use by lepidopterans. The magnitude of ant-induced effects on caterpillar occurrence across the cerrado landscape may depend on how ants use plants locally and how they respond to liquid food on plants at different habitats. This study enhances the relevance of plant-ant and ant-herbivore interactions in cerrado and highlights the importance of a tritrophic perspective in this ant-rich environment.

Partner manipulation stabilises a horizontally transmitted mutualism.

Mutualisms require protection from non-reciprocating exploiters. Pseudomyrmex workers that engage in an obligate defensive mutualism with Acacia hosts feed exclusively on the sucrose-free extrafloral nectar (EFN) that is secreted by their hosts, a behaviour linking ant energy supply directly to host performance and thus favouring reciprocating behaviour. We tested the hypothesis that Acacia hosts manipulate this digestive specialisation of their ant mutualists. Invertase (sucrose hydrolytic) activity in the ant midguts was inhibited by chitinase, a dominant EFN protein. The inhibition occurred quickly in cell-free gut liquids and in native gels and thus likely results from an enzyme-enzyme interaction. Once a freshly eclosed worker ingests EFN as the first diet available, her invertase becomes inhibited and she, thus, continues feeding on host-derived EFN. Partner manipulation acts at the phenotypic level and means that one partner actively controls the phenotype of the other partner to enhance its dependency on host-derived rewards.

Emerging Trends in Ant-Pollinator Conflict in Extrafloral Nectary-Bearing Plants.

The net outcomes of mutualisms are mediated by the trade-offs between the costs and benefits provided by both partners. Our review proposes the existence of a trade-off in ant protection mutualisms between the benefits generated by the ants' protection against the attack of herbivores and the losses caused by the disruption of pollination processes, which are commonly not quantified. This trade-off has important implications for understanding the evolution of extrafloral nectaries (EFNs), an adaptation that has repeatedly evolved throughout the flowering plant clade. We propose that the outcome of this trade-off is contingent on the specific traits of the organisms involved. We provide evidence that the protective mutualisms between ants and plants mediated by EFNs have optimal protective ant partners, represented by the optimum point of the balance between positive effects on plant protection and negative effects on pollination process. Our review also provides important details about a potential synergism of EFN functionality; that is, these structures can attract ants to protect against herbivores and/or distract them from flowers so as not to disrupt pollination processes. Finally, we argue that generalizations regarding how ants impact plants should be made with caution since ants' effects on plants vary with the identity of the ant species in their overall net outcome.

Exclusive rewards in mutualisms: ant proteases and plant protease inhibitors create a lock-key system to protect Acacia food bodies from exploitation.

Myrmecophytic Acacia species produce food bodies (FBs) to nourish ants of the Pseudomyrmex ferrugineus group, with which they live in an obligate mutualism. We investigated how the FBs are protected from exploiting nonmutualists. Two-dimensional gel electrophoresis of the FB proteomes and consecutive protein sequencing indicated the presence of several Kunitz-type protease inhibitors (PIs). PIs extracted from Acacia FBs were biologically active, as they effectively reduced the trypsin-like and elastase-like proteolytic activity in the guts of seed-feeding beetles (Prostephanus truncatus and Zabrotes subfasciatus), which were used as nonadapted herbivores representing potential exploiters. By contrast, the legitimate mutualistic consumers maintained high proteolytic activity dominated by chymotrypsin 1, which was insensitive to the FB PIs. Larvae of an exploiter ant (Pseudomyrmex gracilis) taken from Acacia hosts exhibited lower overall proteolytic activity than the mutualists. The proteases of this exploiter exhibited mainly elastase-like and to a lower degree chymotrypsin 1-like activity. We conclude that the mutualist ants possess specifically those proteases that are least sensitive to the PIs in their specific food source, whereas the congeneric exploiter ant appears partly, but not completely, adapted to consume Acacia FBs. By contrast, any consumption of the FBs by nonadapted exploiters would effectively inhibit their digestive capacities. We suggest that the term 'exclusive rewards' can be used to describe situations similar to the one that has evolved in myrmecophytic Acacia species, which reward mutualists with FBs but safeguard the reward from exploitation by generalists by making the FBs difficult for the nonadapted consumer to use.

Long-term temporal variation in the organization of an ant-plant network.

BACKGROUND AND AIMS: Functional groups of species interact and coevolve in space and time, forming complex networks of interacting species. A long-term study of temporal variation of an ant-plant network is presented with the aims of: (1) depicting its structural changes over a 20-year period; (2) detailing temporal variation in network topology, as revealed by nestedness and modularity analysis and other parameters (i.e. connectance, niche overlap); and (3) identifying long-term turnover in taxonomic structure (i.e. switches in ant resource use or plant visitor assemblages according to taxa). METHODS: Fieldwork was carried out at La Mancha, Mexico, and ant-plant interactions were observed between 1989 and 1991, between 1998 and 2000, and between May 2010 and 2011. Occurrences of ants on extrafloral nectaries (EFNs) were recorded. The resulting ant-plant networks were constructed from qualitative presence-absence data determined by a species-species matrix defined by the frequency of occurrence of each pairwise ant-plant interaction. KEY RESULTS: Network variation across time was stable and a persistent nested structure may have contributed to the maintenance of resilient and species-rich communities. Modularity was lower than expected, especially in the most recent networks, indicating that the community exhibited high overlap among interacting species (e.g. few species were hubs in the more recent network, being partly responsible for the nested pattern). Structurally, the connections created among modules by super-generalists gave cohesion to subsets of species that otherwise would remain unconnected. This may have allowed an increasing cascade-effect of evolutionary events among modules. Mutualistic ant-plant interactions were structured 20 years ago mainly by the subdominant nectarivorous ant species Camponotus planatus and Crematogaster brevispinosa, which monopolized the best extrafloral nectar resources and out-competed other species with broader feeding habits. Through time, these ants, which are still present, lost their position as network hubs and diminished in their importance in structuring the network; simultaneously, plants gained in importance. CONCLUSIONS: The long-term network analysis reveals a decrease in attended plant species richness, a notable increase in plant species participation from 1990 to 2010 (sustained by less plant taxonomic similarity in the older 1990 network), an increase in the number of ant species and a diminishing dominance of super-generalist ants. The structure of the community has remained highly nested and connected with low modularity, suggesting overall a more participative, homogeneous, cohesive interaction network. Although previous studies have suggested that interactions between ants and EFN-bearing plants are susceptible to seasonality, abiotic factors and perturbation, this cohesive structure appears to be the key for biodiversity and community maintenance.

An Epiphytic Ant-Plant Mutualism Structures Arboreal Ant Communities.

Arboreal ant communities are primarily structured by interactions among ant species, food availability, and physical structures within the environment. Epiphytes are a common feature of tropical forests that can provide ants with both food and nesting space. To date, little work has examined what role epiphytic ant-plants play in structuring arboreal ant communities. We surveyed ant species inhabiting the Australian epiphytic ant-plant Myrmecodia beccarii Hook.f. (Gentianales: Rubiaceae) and how arboreal ant communities are structured in relation to M. beccarii presence on trees. Myrmecodia beccarii was inhabited by the ant Philidris cordata Smith, F. (Hymenoptera: Formicidae) on the majority of Melaleuca viridiflora Sol. Ex Gaertn. (Myrtales: Myrtaceae) trees with ant-occupied ant-plants at our two sites. Dominant arboreal ant species at both study sites exhibited discrete, nonoverlapping distributions, and C-score analysis detected an ant mosaic at one site. The distribution of P. cordata was limited by the distribution of ant-plants for both sites. Philidris cordata dominance on trees was also determined by the presence of M. beccarii occupied by P. cordata at both sites. We suggest that by providing P. cordata with nesting space M. beccarii plays a role in structuring these arboreal ant communities.

Are ants attracted to herbivorized leaves of Caryocar brasiliense Camb. (Caryocaraceae)? / As formigas são atraídas por folhas herbivoradas de Caryocar brasiliense Camb. (Caryocaraceae)?

Abstract Different profiles of secondary compounds are released by plants after herbivore attack. Many of these compounds are used by predators and parasitoids to locate herbivores that are damaging leaves. Such an induced indirect defense was tested with the Caryocar brasiliense-ant system in the dry season, when C. brasiliense has old leaves, and in the rainy season, when C. brasiliense has new leaves. A total of 20 plants were analyzed per season. Two opposite leaves of the same branch were selected for each plant. Approximately 40% of the area of one leaf was removed (treatment leaf) while the other leaf remained intact (control). The number of ants that visited each leaf was counted simultaneously for a period of 15 minutes and the mean difference in ant number was tested by paired t-test. The mean number of ants differed significantly between treatment and control only in the rainy season (t = 3.004, df = 19, p = 0.007). This finding suggests the presence of induced defense in this system only when the leaves are young. The study supports the Optimal Defense Theory since young leaves of C. brasiliense with artificial damage attracted significantly more ants than leaves without damage and represents the first evidence of an induced defense mechanism in the C. brasiliense-ant system.

Resumo Diferentes compostos químicos são liberados pelas plantas após o ataque dos herbívoros. Muitos desses compostos são usados por predadores e parasitoides para localizar os herbívoros que estão injuriando as plantas. Esse tipo de defesa induzida indireta foi testada no sistema Caryocar brasiliense e formigas em duas estações: seca (quando C. brasiliense está com folhas maduras) e chuvosa (quando C. brasiliense está com folhas jovens). Nós analisamos 20 plantas por estação do ano. Em cada planta nós selecionamos duas folhas opostas de um mesmo ramo. Nós removemos cerca de 40% da área de uma das folhas, deixando a outra folha intacta. Nós mensuramos simultaneamente o número de formigas que visitaram cada tipo de folha por 15 min e analisamos a diferença entre o número de formigas em cada tipo de folha através de teste t pareado. Nós observamos diferença significativa no número de formigas que patrulham as folhas tratamento (com herbivoria artificial) e controle (sem hervivoria artificial) apenas na estação chuvosa (t = 3,004, df = 19, p = 0,007). Isso sugere que existe defesa induzida nesse sistema somente quando as folhas são jovens. Nosso estudo corrobora a Teoria de Defesa Ótima já que apenas as folhas jovens de C. brasiliense com dano artificial atraíram significativamente mais formigas do que as folhas sem danos. Esta é a primeira vez que mecanismos de defesa induzida são observados no sistema C. brasiliense-formigas.

Effect of different ant species on reproductive fitness of Schomburgkia tibicinis (Orchidaceae).

We studied the effect of four ant species on the reproductive fitness (number of fruits produced) of Schomburgkia tibicinis (Orchidaceae), in the coast of Yucatan, Mexico. Ants forage day and night for the nectar produced by the reproductive structures of the orchid. Ant size is: Camponotus planatus (3-4 mm), C. abdominalis (4-6 mm), C. rectangularis (7-9 mm), and Ectatomma tuberculatum (9-12 mm). The results indicate that ant efficiency in disrupting the activities of the main herbivore, (Stethobaris sp./Coleoptera) varies, and that it is apparently related to ant size, three tendencies are clear: (a) lowest fruit production and highest inflorescence damage are significantly associated with the smaller ants and the control; (b) maximum fruit production and minimum inflorescence damage are significantly associated with the larger ant species; and (c) the increase in ant size tends to have a positive effect on the plant's reproductive output (less dead spikes and more matured fruits). We discuss ant effect on the pollination of the orchid, and emphasize that ant presence should not be associated, in general, with benefit to plants.

Studies of a South East Asian ant-plant association: protection of Macaranga trees by Crematogaster borneensis.

In the humid tropics of SE Asia there are some 14 myrmecophytic species of the pioneer tree genus Macaranga (Euphorbiaceae). In Peninsular Malaysia a close association exists between the trees and the small, non-stinging myrmicine Crematogaster borneensis. These ants feed mainly on food bodies provided by the plants and have their colonies inside the hollow internodes. In a ten months field study we were able to demonstrate for four Macaranga species (M. triloba, M. hypoleuca, M. hosei, M. hulletti) that host plants also benefit considerably from ant-occupation. Ants do not contribute to the nutrient demands of their host plant, they do, however, protect it against herbivores and plant competition. Cleaning behaviour of the ants results in the removal of potential hervivores already in their earliest developmental stages. Strong aggressiveness and a mass recruiting system enable the ants to defend the host plant against many herbivorous insects. This results in a significant decrease in leaf damage due to herbivores on ant-occupied compared to ant-free myrmecophytes as well as compared to non-myrmecophytic Macaranga species. Most important is the ants' defense of the host plant against plant competitors, especially vines, which are abundant in the well-lit pioneer habitats where Macaranga grows. Ants bite off any foreign plant part coming into contact with their host plant. Both ant-free myrmecophytes and non-myrmecophytic Macaranga species had a significantly higher incidence of vine growth than specimens with active ant colonies. This may be a factor of considerable importance allowing Macaranga plants to grow at sites of strongest competition.

Azteca protection of Cecropia: ant occupation benefits juvenile trees.

In this 15 month investigation I experimentally demonstrated that sapling Cecropia aff. obtusifolia in lowland western Ecuador grow more vigorously when occupied by the ant Azteca constructor than when the ants have been removed. Thus the interaction is directly beneficial to Cecropia juveniles. The difference in growth is associated with differences in herbivory and vine cover. Removal of ants significantly increases nocturnal Coleoptera herbivory on unoccupied plants. In contrast to the influence on beetle numbers, Azteca are ineffective against Homoptera and cecidomyiid gall flies. Although ant-occupied saplings had less chewing herbivore damage throughout the study, the ants were more effective protectors in the dry season than in the rainy season, when herbivore pressure increased. In addition to reducing herbivory, Azteca efficiently remove vines from occupied saplings.

Benefits conferred by "timid" ants: active anti-herbivore protection of the rainforest tree Leonardoxa africana by the minute ant Petalomyrmex phylax.

In this study, we demonstrate that an important benefit provided by the small host-specific ant Petalomyrmex phylax to its host plant Leonardoxa africana is efficient protection against herbivores. We estimate that in the absence of ants, insect herbivory would reduce the leaf area by about one-third. This contributes considerably to the fitness of the plant. Our estimates take into account not only direct damage, such as removal of leaf surface by chewing insects, but also the effects of sucking insects on leaf growth and expansion. Sucking insects are numerically predominant in this system, and the hitherto cryptic effects of ant protection against the growth-reducing effects of sucking insects accounted for half of the total estimated benefit of ant protection. We propose that the small size of workers confers a distinct advantage in this system. Assuming that resource limitation implies a trade off between size and number of ants, and given the small size of phytophagous insects that attack Leonardoxa, we conclude that fine-grained patrolling by a large number of small workers maximises protection of young leaves of this plant. Since herbivores are small and must complete their development on the young leaves of Leonardoxa, and since a high patrolling density is required for a fine-grained search for these enemies, numerous small ants should provide the most effective protection of young leaves of Leonardoxa. We also discuss other factors that may have influenced worker size in this ant.

The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree.

One possible function of extrafloral nectaries is to attract insects, particularly ants, which defend plants from herbivores. We determined whether ants visiting saplings of the tree Stryphnodendronmicrostachyum (Leguminosae) provide protection (decreased plant damage due to ant molestation or killing of herbivores) and benefit (increased plant growth and reproduction associated with ant presence) to the plant. We compared ant and herbivore abundance, herbivore damage and growth of ant-visited plants and ant-excluded plants grown in sun and shade microhabitats of a 6-ha plantation in Costa Rica over a 7-month period. Results show that ants provided protection to plants not by reducing herbivore numbers but by molesting herbivores. Ants also reduced the incidence of pathogen attack on leaves. Protection was greater in the shade than in the sun, probably due to lower herbivore attack in the sun. Protection was also variable within sun and shade habitats, and this variability appeared to be related to variable ant visitation. Results also indicate that ant presence benefits the plant: ant-visited plants grew significantly more in height than ant-excluded plants. The cultivation of ants may serve as an important natural biological control in tropical forestry and agroforestry systems, where increased plant density can otherwise lead to increased herbivore attack.

O bambu Merostachys fischeriana (Bambusoideae: Bambuseae) como habitat para formigas de Floresta Tropical Montana / The bamboo Merostachys fischeriana (Bambusoideae: Bambuseae) as a canopy habitat for ants of Neotropical Montane Forest

Although Merostachys fischeriana is very abundant in the Brazilian Atlantic Rainforest, little attention has been paid to the biological interactions with other animals. The present study describes some of the interactions between ants and this bamboo species. The experiment was carried out in a fragment of a montane tropical forest in the Parque Estadual do Itacolomi, near Ouro Preto, MG, Brazil. Thirty culms of bamboo were randomly collected. The ants were obtained by direct collection from nodes and internodes. Morphometric variables of the bamboo were recorded for characterization of potential ant habitat. Merostachys fischeriana grows in rosettes as a thin bamboo (average = 1,0 cm; se = 0,27; n = 20) and is tall enough to reach the upper canopy of this low forest (average = 9,1 m; se = 2,72; n = 20). Fifteen ant species were sampled. Brachymyrmex heeri Forel was the most abundant in the nodes, while Camponotus crassus Mayr (Hymenoptera: Formicidae) was the most abundant in the internodes. The composition of the species that inhabit the internodes was different from the composition in the node (Q-test: Q = 3,76; P = 0,05). The level of occupation was defined by the number of holes (F = 10,33; P < 0,01), the number of internodes in the canopy (F = 6,84; P = 0,01) and the length of the culm (F = 7,52; P = 0,01). The plant's morphology allowed the occurrence of additional species of ants in the canopy and influenced the composition of the entire ant assemblage.