Actinobacteria that live mutualistically with leaf-cutter ants secrete antibiotics that may induce antibiotic resistance in nearby soil bacteria. We tested for the first time whether soil bacteria near and inside Atta cephalotes nests in Costa Rica show higher levels of antibiotic resistance than bacteria collected farther away. We collected soil samples 0 m to 50 m away from ant nests and grew bacteria from them on agar with paper discs treated with antibiotics of common veterinary use. As a proxy for antibiotic resistance, we measured the distance from the edge of each disc to the closest bacterial colonies. In general, resistance to oxytetracycline increased with proximity to leaf-cutter ant nests. Antibiotic resistance to oxytetracycline was also higher in samples collected inside the nest than in samples from the nest mound; not all antibiotics demonstrated the same trend. A preliminary exploratory morphological analysis suggests bacterial communities between 0 m and 50 m from ant nests were similar in diversity and abundance, indicating the pattern of antibiotic resistance described above may not be caused by differences in community composition. We conclude that actinobacteria living mutualistically with A. cephalotes drive natural antibiotic resistance to tetracycline in proximal bacterial communities.

Flower and leaf herbivory might cause relevant and negative impacts on plant fitness. While flower removal or damage by florivores produces direct negative effects on plant fitness, folivores affect plant fitness by reducing resource allocation to reproduction. In this study, we examine the effects of both flower and leaf herbivory by leaf-cutting ants on the reproductive success of the shrub species Miconia nervosa (Smith) Triana (Family Melastomataceae) in a fragment of Atlantic Forest in Northeast Brazil. We conducted a randomized block-designed field experiment with nine replicates (blocks), in which three plants per block were assigned to one of the three following treatments: undamaged plants (ant exclusion), leaf-damaged plants (ant exclusion from reproductive organs, but not from leaves), and flower + leaf-damaged plants (no exclusion of ants). We then measured flower production, fruit set, and fruit production. Our results showed that flower + leaf-damaged plants reduced flower production nearly twofold in relation to undamaged plants, while flower set in leaf-damaged plants remained constant. The number of flowers that turned into fruits (i.e., fruit set), however, increased by 15% in flower + leaf-damaged plants, while it slightly decreased in leaf-damaged compared to undamaged plants. Contrastingly, fruit production was similar between all treatments. Taken together, our results suggest a prominent role of ant floral herbivory across different stages of the reproductive cycle in M. nervosa, with no consequences on final fruit production. The tolerance of M. nervosa to leaf-cutting ant herbivory might explain its high abundance in human-modified landscapes where leaf-cutting ants are hyper-abundant.

Successional state of forest and availability of pioneer plants are recognized factors affecting densities of leaf-cutting ants. However little is known about how abiotic factors can shape nest distributions. We investigated the effect of topography, soil, forest successional state and inundation risk on nest density and size of Atta cephalotes colonies along streams in a tropical wet forest in Costa Rica. In each forest type, we surveyed 12 sites, each site comprising five transects (10 × 100 m) varying in topography and proximity to streambeds. We found no difference regarding nest size or density between forest types or soil consociation. Nest density varied with topographic environment, with significantly higher nest density on slope tops (farther from streambeds) and without colonies in valley bottoms (closer to streambeds). Nests found in areas affected by the last great inundation before our study were scarcer and smaller than nests in non-flooded areas. We showed that inundation events favour an accumulation of Atta colonies towards higher sites, where they are also allowed to become larger and may survive longer. Inundation risk may be a strong force shaping the distribution of leaf-cutting ant nests in tropical floodplain forests, even concealing the relevance of successional state of forest.

This study examined whether high nutrient concentrations associated with leaf-cutting ant nests influence plant growth and plant water relations in Amazon rain forests. Three nests of Atta cephalotes were selected along with 31 Amaioua guianensis and Protium sp. trees that were grouped into trees near and distant (>10 m) from nests. A 15N leaf-labelling experiment confirmed that trees located near nests accessed nutrients from nests. Trees near nests exhibited higher relative growth rates (based on stem diameter increases) on average compared with trees further away; however this was significant for A. guianensis (near nest 0.224 y−1 and far from nest 0.036 y−1) but not so for Protium sp. (0.146 y−1 and 0.114 y−1 respectively). Water relations were similarly species-specific; for A. guianensis, near-nest individuals showed significantly higher sap flow rates (16 vs. 5 cm h−1), higher predawn/midday water potentials (−0.66 vs. −0.98 MPa) and lower foliar δ13C than trees further away indicating greater water uptake in proximity to the nests while the Protium sp. showed no significant difference except for carbon isotopes. This study thus shows that plant response to high nutrient concentrations in an oligotrophic ecosystem varies with species. Lower seedling abundance and species richness on nests as compared with further away suggests that while adult plants access subterranean nutrient pools, the nest surfaces themselves do not encourage plant establishment and growth.

Habitat fragmentation is gradually altering tropical forests all around the globe (Wright 2005), as human populations convert large blocks of pristine forest habitats into permanently fragmented or human-modified landscapes (Tabarelli et al. 2004). Fragmentation processes apparently create opportunities for small sets of species, which proliferate across modified landscapes, while others are driven to extinction or remain as minimal populations (Tabarelli et al. 2010). This ‘winners vs. losers paradigm’ (sensu McKinney & Lockwood 1999) probably describes one of the most conspicuous rearrangements of tropical biotas in response to human disturbances (in addition to biomass collapse), with unanticipated consequences for ecosystem functioning and long-term biodiversity persistence (Lôbo et al. 2011). Despite such appeal, the ecological identity of proliferating organisms and the mechanisms underlying their high abundance levels remains poorly appreciated.

Much of the ecological alteration faced by human-modified Neotropical forests can be assigned to edge effects, including the proliferation of some voracious herbivores such as leaf-cutting ants. However, the underlying mechanisms/impacts of tropical forest edge on herbivores performance and their foraging behaviour (e.g. dietary diversity) have rarely been investigated. The goal of this study was, therefore, to determine whether and how the annual diet (i.e. species richness, diversity and the relative proportion of pioneer versus non-pioneer species of plant materials) of Atta cephalotes colonies differs in the forest edge versus the interior zone of a large remnant of Atlantic forest in northeastern Brazil. Among the key results was a strong habitat effect on dietary diversity (explaining ca. 40–50% of the variation), which, in edge colonies, decreased approximately by one fourth compared to interior colonies (inverse of Simpson's index: 3.7±0.84 versus 4.99±0.95). There was a predominance of leaf fragments collected from pioneer species in the diet in both habitat (86% in edge and 80.4% in interior). Edge colonies collected proportionally more fragments from pioneer species than colonies located in the forest interior. Our results are the first to demonstrate an edge-mediated relaxation of dietary restrictions in leaf-cutting ants. These findings render robust support to previous evidence indicating the reduction of bottom-up forces as a key factor explaining both edge-induced hyper-abundance and increased herbivory of leaf-cutting ants in human-modified Neotropical landscapes.

Leaf-cutting ants (species of Atta and Acromyrmex) are dominant herbivores and play a key role as ecosystem engineers of tropical and subtropical America (Fowler et al. 1989, Weber 1972). Not only are they among the most polyphagous and voracious herbivorous insects, cutting up to 15% y−1 of the leaf standing crop (Urbas et al. 2007, Wirth et al. 2003), but also they strongly affect the light environment and the nature of plant assemblages via ant-nest-mediated disturbances (Farji-Brener & Illes 2000, Hull-Sanders & Howard 2003, Moutinho et al. 2003). Some leaf-cutting ant species have turned into an omnipresent feature of present-day neotropical landscapes and a wealth of studies has documented their abundance to drastically increase with increasing agricultural land use, disturbance and deforestation/fragmentation (Fowler et al. 1986, Jaffe 1986, Terborgh et al. 2001, Vasconcelos & Cherrett 1995, Wirth et al. 2007). In view of their ecosystem engineering capacity and the ever-increasing conversion of tropical forests into agricultural landscapes (Wright 2005), it has been concluded that disturbance-driven accumulation of Atta colonies leads to far-reaching and deleterious consequences in present-day neotropical landscapes (Wirth et al. 2008). But what about the opposite scenario of regenerating forests? Is disturbance-mediated hyper-abundance of leaf-cutting ants a reversible phenomenon? We believe that this question is highly relevant because (1) knowledge of the dynamics of leaf-cutting ant populations during forest regeneration is lacking and (2) natural secondary succession has become a widespread phenomenon after land is abandoned or temporarily fallowed (Wright 2005). In the Brazilian Amazon during the 1990s, for example, secondary forests have reclaimed 31% of the once deforested land (Perz & Skole 2003).

Leaf-cutting ants (genera Atta and Acromyrmex) have been denoted key species of American rain-forest ecosystems (Fowler et al. 1989) because of their multifarious effects on the vegetation. Being dominant herbivores, cutting up to 13% of the standing leaf crop in a colony's territory per year, they affect directly and significantly individual plants, plant communities and ecosystems (Wirth et al. 2003). The considerable ecological impact of these ants is paralleled by the well-known fact that some species strongly benefit from human-driven habitat alterations and represent prime pests throughout Latin America (Cherrett 1986). Numerous studies have documented populations of leaf-cutting ant to increase with increasing agricultural land use, deforestation and/or disturbance (Fowler et al. 1986, Jaffe & Vilela 1989, Jonkman 1979). Specifically, elevated colony densities have been recorded in (1) transformed vegetation such as pastures (Fowler 1983) and plantations (Jaffe 1986, Oliveira et al. 1998), (2) early successional forests (Vasconcelos & Cherrett 1995), and recently (3) isolated forest remnants (Terborgh et al. 2001).

Nests of leaf-cutting ants (Hymenoptera: Formicidae: Attini) are abundant disturbances in Neotropical rain forests, and could affect the plant community both while the nests are active and after they are abandoned. We measured the diversity and abundance of understorey plants (>1 m in height) in the area around active and abandoned nests of leaf-cutting ants (Atta cephalotes) at the La Selva Biological Station in Costa Rica. Sampel quadrats on active nests had reduced diversity (number of morphospecies) and abundance of both small (height >10 cm) and large (10 cm–1 m) understorey plants, when compared to the nearby forest floor (3 and 13 m from the nest edge). Abandoned nests had greater diversity and marginally greater abundance of small understorey plants relative to nearby forest; there was no difference in diversity or abundance of large understorey plants. Leaf-cutting ant nests create gaps in the plant understorey when active, but serve as centres of recruitment for small plants after they are abandoned. Thus, like canopy gaps, ant nests could play an important role in recruitment of new individuals and maintenance of plant species diversity in tropical forests.