Moisture enhances the positive effect of leaf‐cutting ant refuse dumps on soil biota activity

Soil biota activity in arid lands is often limited by the availability of water and organic matter. We experimentally explored whether small changes in soil moisture affect the activity of soil biota in external refuse dumps of the leaf‐cutting ant Acromyrmex lobicornis, one of the most important sources of organic matter in a semi‐arid land of north‐western Patagonia. We estimated CO₂ consumption in refuse dumps and in adjacent, non‐nest soil samples at two moisture levels, after 48 and 72 h. Soil biota activity, estimated by respiration rates, was up to 160 times greater in refuse dumps than in adjacent, non‐nest soils. Activity of soil biota in non‐nest soil did not change through time and was not affected by moisture. Conversely, soil biota increased their activity in refuse dump samples only at high moisture condition after 72 h. As the activity of microorganisms is key for soil nutrient generation and availability, refuse dumps may be considered as ‘islands of fertility’ for plants. This effect may be especially important after sporadic spring rainfalls, when the beneficial effect of refuse dumps on soil biota is enhanced. In addition, as refuse dumps generate several times more CO₂ than non‐nest soils, nest areas may be considered also as hot spots of CO₂ emissions. These results illustrate the potential importance of ant nests for nutrient cycling, maintenance of plant cover and carbon balance in arid ecosystems.     

Refuse dumps of leaf‐cutting ants as a deterrent for ant herbivory: does refuse age matter?

Leaf‐cutting ants often avoid contact with their waste because it harbors microorganisms that are dangerous to the ants and their symbiotic fungus. Therefore, the use of ant waste (i.e., refuse dumps) has been proposed as a deterrent method against leafcutter attack. We tested experimentally whether the age of the refuse dump (fresh vs. old) affects the herbivory‐deterrent effect against the leaf‐cutting ant Acromyrmex lobicornis Emery (Hymenoptera: Formicidae). Refuse placed around seedlings significantly delayed the initiation attacks of leaf‐cutting ants, and this deterrent effect decreased gradually over a period of 30 days. The initial strength of this decrease was the same for newly‐discarded ( ‘new’) refuse and refuse from the bottom of the ants’ waste pile (‘old’ refuse). However, the loss of deterrent effect over time was more rapid for new than old refuse. A further experimental manipulation, replacement of refuse every 3 days, had no effect on the deterrent effect for old refuse, but increased this effect for new refuse, although the amount of this increase gradually weakened over the course of the 30‐day experiment. We speculate on the possible causes of these effects, their consequences for the hygienic behavior of leaf‐cutting ants, and on the use of ant debris as short‐term control method against leaf‐cutting ants.     

Influence of nests of leaf‐cutting ants on plant species diversity in road verges of northern Patagonia

Abstract. It has been suggested that ant nests are the most frequent small‐scale disturbance that affect vegetation patterns. However, their effects on plant diversity are little studied. We document effects of nests of the leaf‐cutting ant Acromyrmex lobicornis on physical‐chemical soil properties and their influence on plant diversity near road verges in a desert steppe in NW Patagonia, Argentina. We analysed nest soils and controls for nitrogen, phosphorus, organic matter, moisture retention capacity and texture. We also analysed the vegetation on 42 nests (30 active and 12 abandoned or without life) and 42 areas without nests. Soil around nests had a greater nutrient content and capacity to retain moisture than control soils, which is mainly due to the presence of organic waste that the ants deposit on the soil surface. We found no association between the occurrence of nests and specific groups of plants, but plant diversity was higher at nest‐sites than at nearby non‐nest sites. This increased diversity – which is also found on abandoned nests – is mainly due to the occurrence of a larger number of native and exotic plant species on nest‐sites that are uncommon elsewhere in the study area. The most abundant plant species showed similar cover values at nest and non‐nest sites. This suggests that changes in diversity are associated to edaphic changes caused by nests rather than by changes in competitive balance caused by dominant species exclusion. We propose that the nests of Acromyrmex lobicornis, through increasing the availability of resources, generate favourable microsites that can function both as ‘refuges’ for less frequent native species, and as‘stepping stones’ for less frequent exotic plant species.     

Refuse attracts? Effect of refuse dumps of leaf‐cutting ants on floral traits

The nutrient‐rich organic waste generated by ants may affect plant reproductive success directly by enhancing fruit production but also indirectly, by affecting floral traits related with pollinator attraction. Understanding how these soil‐nutrient hot spots influence floral phenotype is relevant to plant–pollination interactions. We experimentally evaluated whether the addition of organic waste from refuse dumps of the leaf‐cutting ant Acromyrmex lobicornis (Hymenoptera: Formicidae: Attini) alters floral traits associated with pollinator attraction in Eschscholzia californica (Ranunculales: Papaveraceae), an entomophilous herb. We analysed flower shape and size using geometric morphometric techniques in plants with and without the addition of refuse‐dumps soil, under greenhouse conditions. We also measured the duration of flowering season, days with new flowers, flower production and floral display size. Plants growing in refuse‐dumps soil showed higher flower shape diversity than those in control soil. Moreover, plants in refuse‐dumps soil showed bigger flower and floral display size, longer flowering season, higher number of flowering days and flower production. As all these variables may potentially increase pollinator visits, plants in refuse‐dumps soil might increase their fitness through enhanced attraction. Our work describes how organic waste from ant nests may enhance floral traits involved in floral attraction, illustrating a novel way of how ants may indirectly benefit plants.     

A meta‐analysis of leaf‐cutting ant nest effects on soil fertility and plant performance

1. Leaf‐cutting ants (LCAs) are considered as one of the most important agents of soil disturbances that affect vegetation patterns, but these assertions are based on isolated studies or anecdotal data. In this study, meta‐analysis techniques were used to quantitatively analyse the generality of these effects and determine some of their sources of variation. 2. The results reveal the following: (i) LCA nest sites showed higher levels of soil fertility than control sites, but the key source of these nutrients is the refuse material rather than the nest soil itself; (ii) refuse material from external piles tended to be richer in nutrient content than refuse material from internal refuse chambers; (iii) nest sites from temperate habitats showed higher cation content than those located in tropical/subtropical habitats; and (iv) nest sites showed higher plant growth than adjacent non‐nest sites (especially if plants have access to the refuse) but similar plant density and plant richness. 3. As LCAs improve nutrient availability in nest sites through the accumulation of refuse material, the location of the refuse will have a relevant role affecting vegetation. LCA species with external refuse dumps could benefit herbs, early vegetation stages and short‐living plants, whereas those with internal refuse chambers could benefit long‐living, large trees. However, the positive effect on individual plants does not extend to population and community levels. The foraging preferences of ants and the changes in microclimatic conditions around nests could act as selective ecological filters. 4. As refuse material from external piles and nest sites in temperate habitats tend to show higher fertility than refuse material from internal nest chambers and nest sites in tropical/subtropical habitats, LCA species with external refuse dumps in temperate regions could be of particular relevance for nutrient cycling and vegetation patterns.     

Performance and fate of tree seedlings on and around nests of the leaf‐cutting ant Atta cephalotes: Ecological filters in a fragmented forest

Habitat fragmentation is currently the most pervasive anthropogenic disturbance in tropical forests and some species of leaf‐cutting ants of the genus Atta (dominant herbivores in the neotropics) have become hyper‐abundant in forest edges where their nests directly impact up to 6% of the forest area. Yet, their impacts on the regeneration dynamics of fragmented forests remain poorly investigated. Here we examine the potential of Atta cephalotes nests to function as ecological filters impacting tree recruitment. Growth, survival and biomass partitioning of experimentally planted seedlings (six tree species) were examined at eight spatially independent A. cephalotes colonies in a large Atlantic Forest fragment. Seedling performance and fate (leaf numbers and damage) were monitored up to 27 months across three habitats (nest centre, nest edge and forest understorey). Plants at illuminated nest centres showed twice the gross leaf gain as understorey individuals. Simultaneously, seedlings of all species lost many more leaves at nests than in the forest understorey, causing a negative net leaf gain. Net leaf gain in the shaded understorey ranged from zero (Licania and Thyrsodium species) to substantial growth for Copaifera and Virola, and intermediate levels little above zero for Protium and Pouteria. Also seedling survival differed across habitats and species, being typically low in the centre and at the edge of nests where seedlings were often completely defoliated by the ants. Lastly, seedling survival increased strongly with seed size at nest edges while there was no such correlation in the forest. Our results suggest that Atta nests operate as ecological filters by creating a specific disturbance regime that differs from other disturbances in tropical forests. Apparently, Atta nests favour large‐seeded tree species with resprouting abilities and the potential to profit from a moderate, nest‐mediated increase in light availability.     

Potential impact of the leaf‐cutting ant Acromyrmex lobicornis on conifer plantations in northern Patagonia,Argentina

• 1 The economic losses associated with crop damage by invasive pests can be minimized by recognizing their potential impact before they spread into new areas or crops.
• 2 We experimentally evaluated the preferences of the leaf‐cutting ant Acromyrmex lobicornis (Hymenoptera: Formicidae) for the most common conifer species commercially planted in northern Patagonia, Argentina. The areas of potential forest interest in this region and the geographical range of this ant overlap. We performed field preference tests and monitored the level of ant herbivory on planted conifer seedlings next to nests.
• 3 Acromyrmex lobicornis preferred some conifer species and avoided foraging on others. Pseudotsuga menziesii and Austrocedrus chilensis were the less preferred species, Pinus ponderosa and Pinus contorta were the most preferred by A. lobicornis.
• 4 The item mostly selected by ants was young needles from P. contorta. This species was also the pine mostly defoliated. Seedlings without ant‐exclusion showed a mean±SE of 60±5% defoliation during the sampling period. Pinus ponderosa was less defoliated; control seedlings showed a mean±SE of 8.5±1% of leaf damage in the sampling period.
• 5 The present study shows how the use of simple field tests of leaf‐cutting ant preferences could allow an improved selection of appropriate conifer species for future plantations in areas where leaf‐cutting ants are present.

     

The Importance of Where to Dump the Refuse: Seed Banks and Fine Roots in Nests of the Leaf‐Cutting Ants Atta cephalotes and A. colombica1

The location of the nutrient‐rich organic refuse produced by a leaf‐cutting ant colony varies among ant species. Atta cephalotes locate their organic refuse in subterranean chambers, whereas A. colombica place their organic refuse on the soil surface near the nest. We studied the effect of the absence or presence of external organic refuse on the abundance of fine roots and seed bank composition in the superficial horizons of ant nests. We sampled soils from ant nests or dumps and adjacent areas of 15 adult nests of A. cephalotes at La Selva (LS), Costa Rica, and of 15 of A. colombica nests on Barro Colorado Island (BCI), Panama. Soils from A. cephalotes nests did not differ from adjacent soils in abundance of fine‐root and seed diversity. In contrast, organic refuse from A. colombica nests was less diverse in seed composition (due to the great abundance of Miconia argentea) and had a greater abundance of fine roots than adjacent areas. Thus the external location of the ant‐nest organic refuse is potentially important in determining the different types of plant recolonization in abandoned or dead ant nests. The relative abundance of these Atta species may influence the structure and/or composition of tropical forests.     

Secondary seed dispersal by ants in Neotropical cerrado savanna: species‐specific effects on seeds and seedlings of Siparuna guianensis (Siparunaceae)

1. Most woody plant species in tropical habitats are primarily vertebrate‐dispersed, but interactions between ants and fallen seeds and fruits are frequent. This study assesses the species‐specific services provided by ants to fallen arillate seeds of Siparuna guianensis, a primarily bird‐dispersed tree in cerrado savanna. The questions of which species interact with fallen seeds, their relative contribution (versus vertebrates) to seed removal, and the potential effects on seedling establishment are investigated. 2. Seeds are removed in similar quantities in caged and control treatments, suggesting that ants are the main dispersers on the ground. Five ant species attended seeds. Pheidole megacephala (≈0.4 cm) cooperatively transported seeds, whereas the smaller Pheidole sp. removed the seed aril on spot. Large (> 1.0 cm) Odontomachus chelifer, Pachycondyla striata, and Ectatomma edentatum individually carried seeds up to 4 m. Bits of aril are fed to larvae and intact seeds are discarded near the nest entrance. 3. Overall, greater numbers of seedlings were recorded near ant nests than in control plots without nests. This effect, however, was only detected near P. megacephala and P. striata nests, where soil penetrability was greater compared with controls. Soil nutrients did not differ between paired plots. 4. This study confirms the prevalence of ant–seed interactions in cerrado and shows that ant‐derived benefits are species‐specific. Ant services range from seed cleaning on the spot to seed displacement promoting non‐random spatial seedling recruitment. Although seed dispersal distances by ants are likely to be shorter than those by birds, our study of S. guianensis shows that fine‐scale ant‐induced seed movements may ultimately enhance plant regeneration in cerrado.     

Evidence that the fungus cultured by leaf-cutting ants does not metabolize cellulose

Leaf‐cutting ants are a very specialized group of ants that cultivate fungus gardens in their nests, from which they obtain food. The current opinion is that the fungus cultivated by leaf‐cutting ants digests cellulose. Here we reassess the cellulose‐degrading capability of the fungus by using two complementary approaches tested in four Attini species (genera Atta and Acromyrmex): (1) ability of fungus to grow in cellulose; and (2) lignin/cellulose ratio in the refuse material dumped outside the nest, as an indicator of cellulose consumption. We found that (1) the fungus did not grow in cellulose, and (2) the lignin/cellulose ratio was much lower in the ants' refuse than in material digested by cellulose‐digesting organisms, such as brown‐rot fungus, termites, and ruminant mammals. This evidence strongly suggests the inability of the fungus to degrade cellulose. Therefore, the fungus–ant symbiosis and the ecological role of leaf‐cutting ants need to be reconsidered.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Do abandoned nests of leaf‐cutting ants enhance plant recruitment in the Atlantic Forest?

The role played by abandoned nests of leaf‐cutting ants (Atta spp.) as a small‐scale disturbance regime that affects plant recruitment, species coexistence and forest regeneration remains poorly investigated. Here we examine whether abandoned nests of Atta cephalotes serve as regeneration niches and operate as particular plant recruitment habitats, favouring forest regeneration after ant activities cease and leading to the establishment of taxonomically/ecologically distinct plant assemblages. Soil properties, canopy openness, light availability and regenerating plant assemblages were evaluated across 18 nests and adjacent control plots in a large remnant of Atlantic Forest in north‐east Brazil from December 2004 to December 2005. Surprisingly, nests and control plots exhibited very similar light environments irrespective of nest age, but nest soils exhibited substantial reductions in carbon content (1.45 ± 0.24 vs. 1.79 ± 0.13%) and organic matter (2.50 ± 0.41 vs. 3.08 ± 0.23%), and proved to be much more resistant to penetration (30.57 ± 6.08 vs. 39.48 ± 7.53 mm). Functional signature of regenerating plant assemblages exhibited little variation across both habitat types, as they were dominated by pioneer, small‐seeded and vertebrate‐dispersed species. However, abandoned nests exhibited less dense, impoverished and more homogeneous regenerating plant assemblages at local and landscape scale; they clearly lacked nest‐dependent plant species and represented floristic subsets of the flora inhabiting the undisturbed forest. This recruitment bottleneck was transient in the long term because nest‐related effects ameliorated in older nests. Our results suggest that, unlike treefall gaps, abandoned nests represent temporary (relatively long‐lasting) islands of unsuitable substrate that reduce plant recruitment, retard forest regeneration, and fail in providing a special regeneration niche able to promote species coexistence and plant diversity.     

The ‘truck‐driver’ effect in leaf‐cutting ants: how individual load influences the walking speed of nest‐mates

The foraging behaviour of social insects is highly flexible because it depends on the interplay between individual and collective decisions. In ants that use foraging trails, high ant flow may entail traffic problems if different workers vary widely in their walking speed. Slow ants carrying extra‐large loads in the leaf‐cutting ant Atta cephalotes L. (Hymenoptera: Formicidae) are characterized as ‘highly‐laden’ ants, and their effect on delaying other laden ants is analyzed. Highly‐laden ants carry loads that are 100% larger and show a 50% greater load‐carrying capacity (i.e. load size/body size) than ‘ordinary‐laden’ ants. Field manipulations reveal that these slow ants carrying extra‐large loads can reduce the walking speed of the laden ants behind them by up to 50%. Moreover, the percentage of highly‐laden ants decreases at high ant flow. Because the delaying effect of highly‐laden ants on nest‐mates is enhanced at high traffic levels, these results suggest that load size might be adjusted to reduce the negative effect on the rate of foraging input to the colony. Several causes have been proposed to explain why leaf‐cutting ants cut and carry leaf fragments of sizes below their individual capacities. The avoidance of delay in laden nest‐mates is suggested as another novel factor related to traffic flow that also might affect load size selection The results of the presennt study illustrate how leaf‐cutting ants are able to reduce their individual carrying performance to maximize the overall colony performance.     

Positive ecosystem engineering effects of the ant Atta vollenweideri on the shrub Grabowskia duplicata

Question: How does the ant Atta vollenweideri indirectly affect the shrub Grabowskia duplicata? Does environmental modification induced by nest building affect the life history and population structure of this dominant shrub? Location: Halophytic savanna of western Uruguay, South America. Methods: We compared the density of three shrub size classes, seedlings, saplings and adults, among patches created by ant nest building and in non‐modified areas. We studied key soil properties for plant growth in both live nests and non‐modified areas. We performed a greenhouse experiment to study the effects of soils (live nest versus non‐modified area) on shrub germination probability. We also assessed vegetative propagation by measuring the incidence of root suckers and clonal offshoots in live nests and non‐modified areas through a field survey. Results: Seedling‐size class was more abundant on live nests than in non‐modified areas, suggesting that environmental conditions of this type of habitat (i.e. increased sodium and moisture content of soil, and decreased soil compaction) could facilitate shrub recruitment. We did not find any effects of soil type (live nests versus non‐modified areas) on shrub germination probability. However, the vegetative propagation incidence was higher in the environment with live nests. Conclusions: We found a strong indirect relationship between the ant A. vollenweideri and the shrub G. duplicata. This relationship seems to be mediated by an allogenic engineering process, i.e. soil bioturbation caused by nest building. The environmental conditions of live nests seem to facilitate shrub recruitment, mainly by increasing vegetative propagation.     

Pay it forward: refuse dump from leaf‐cutting ants promotes caterpillar digestive performance by increasing plant nitrogen content

1. The activity of soil‐disturbing animals that increase soil nutrients can affect the carbon : nitrogen (C : N) ratio of plants, which, in turn, may determine the transfer of energy and nutrients through higher trophic levels. However, the strength and sign of this indirect effect depend on whether enhanced nutrient substrates increase plant foliar nutrients and/or plant defensive traits. 2. We investigated how the nutrient‐rich refuse dumps of the leaf‐cutting ant Acromyrmex lobicornis, as a result of their direct effects on thistles, indirectly impact the growth rate and digestive performance of a generalist chewing herbivore. We also included the application of commercial fertilisers to test whether the enhanced soil nutrients comprises the mechanism behind the impact of refuse dumps on the upward cascade effects. 3. Thistles growing on nutrient‐rich substrates (i.e. ant refuse dumps and fertilised soil) had more and larger leaves, up to 80% lower C : N ratios, and higher physical defences compared to plants growing on steppe soil. Caterpillars showed an enhanced digestive performance and growth rate when feed on nutrient‐rich plants and were able to adjust the C : N ratio of their excretion to regulate the relative acquisition of nutrients. 4. The positive effect of nutrient‐rich substrates on caterpillar feeding efficiency suggest that the enhanced nutritional quality of the thistles could compensate for the negative effects of the increased physical defences. The results of the present study indicate how organisms that increase soil nutrient availability may diminish the stoichiometric constraints at the base of food chains, enhancing the development and growth rate of herbivores and, thus, indirectly mediating plant–herbivore interactions.     

Ecosystem Services from Keystone Species: Diversionary Seeding and Seed‐Caching Desert Rodents Can Enhance Indian Ricegrass Seedling Establishment

Seeds of Indian ricegrass (Achnatherum hymenoides), a native bunchgrass common to sandy soils on arid western rangelands, are naturally dispersed by seed‐caching rodent species, particularly Dipodomys spp. (kangaroo rats). These animals cache large quantities of seeds when mature seeds are available on or beneath plants and recover most of their caches for consumption during the remainder of the year. Unrecovered seeds in caches account for the vast majority of Indian ricegrass seedling recruitment. We applied three different densities of white millet (Panicum miliaceum) seeds as “diversionary foods” to plots at three Great Basin study sites in an attempt to reduce rodents' over‐winter cache recovery so that more Indian ricegrass seeds would remain in soil seedbanks and potentially establish new seedlings. One year after diversionary seed application, a moderate level of Indian ricegrass seedling recruitment occurred at two of our study sites in western Nevada, although there was no recruitment at the third site in eastern California. At both Nevada sites, the number of Indian ricegrass seedlings sampled along transects was significantly greater on all plots treated with diversionary seeds than on non‐seeded control plots. However, the density of diversionary seeds applied to plots had a marginally non‐significant effect on seedling recruitment, and it was not correlated with recruitment patterns among plots. Results suggest that application of a diversionary seed type that is preferred by seed‐caching rodents provides a promising passive restoration strategy for target plant species that are dispersed by these rodents.     

Waste Management in the Leaf-Cutting Ant <Emphasis Type="Italic">Acromyrmex lobicornis</Emphasis>: Division of Labour,Aggressive Behaviour,and Location of External Refuse Dumps

In leaf-cutting ants, the handling of waste materials from the fungus culture increases the risk of infection. Consequently, ants should manage their waste in a way that minimizes the spread of diseases. We investigated whether in Acromyrmex lobicornis, waste-worker ants (a) also perform roles in foraging or mound maintenance, (b) are morphologically different than other ant workers, and (c) are aggressively discriminated by other worker ants from the same colony. In addition, we investigated whether the location of external waste piles minimizes the probability that wastes spread to the ant nest. In the field, we (a) marked with different colours waste-workers, foragers and mound-workers and monitored whether these ants interchanged their tasks; (b) measured head width, head length, hind femur length and total length of waste-workers; foragers and mound-workers; (c) forced field encounters between waste-workers and foragers, and (d) measured the cardinal orientation of the waste piles in relation to the colony mound. Waste-worker ants did not perform other function outside the nest; neither foragers nor mound-workers managed the waste. Moreover, waste-workers were smaller than foragers and mound-workers, and were attacked if they tried to enter their nest using foraging entrances. The location of external refuse dumps also appears to reduce contamination risks. Waste piles always were down-slope, and often followed the prevailing wind direction. The importance of behaviours such as the division of labour, aggressions against waste-workers and nest compartmentalization (i.e., the orientation of external waste piles) to minimize the spread of pathogens is discussed.     

Seed handling behaviours of native and invasive seed‐dispersing ants differentially influence seedling emergence in an introduced plant

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Foraging activity of leaf‐cutting ants changes light availability and plant assemblage in Atlantic forest

1. Leaf‐cutting ants (LCAs) have often been denoted as ecosystem engineers because of their multifarious effects on the vegetation, particularly via nest‐driven environmental changes. However, the non‐trophic impacts of LCAs on forest dynamics via foliage harvesting across sizeable foraging zones (so‐called associated ecosystem engineering) are still poorly investigated. 2. Here, light availability and sapling assemblages were assessed within foraging areas and ant‐free control zones of 16 Atta cephalotes colonies located in a large remnant of Atlantic forest in northeastern Brazil. 3. Canopy openness and total light transmission were 1.4 and 1.6 times higher in foraging zones than in control areas. In parallel, sapling density and species richness decreased constantly from control to foraging zone plots. Additionally, shade‐tolerant species exhibited reduced abundance across foraging zones. A non‐metric multidimensional scaling ordination based on taxonomic similarity primarily segregated foraging zone and control plots; foraging zone plots converged to be more similar to each other as well. Finally, some plant species emerged as indicators of LCA‐free zones. 4. These results suggest that LCA foraging activity in the forest canopy directly increases the light availability and indirectly affects the recruitment and the structure of local plant assemblages. 5. Such a biologically significant effect on the light environment and its cascades confirms LCAs as potent ecosystem engineers, particularly as a plant assembly force, which operates beyond the spatial reach of their well‐described nest effects.     

The Multiple Impacts of Leaf‐Cutting Ants and Their Novel Ecological Role in Human‐Modified Neotropical Forests

Herbivory has been identified as a potent evolutionary force, but its ecological impacts have been frequently underestimated. Leaf‐cutting ants represent one of the most important herbivores of the Neotropics and offer an interesting opportunity to address the role played by herbivorous insects through a perspective that embraces population‐ to ecosystem‐level effects. Here we: (1) qualitatively summarize the multiple ways leaf‐cutting ants interact with food plants and their habitats and elucidate the ultimate outcome of such interactions at multiple organization levels; (2) update our understanding of leaf‐cutting ant‐promoted disturbance regimes; and (3) examine potential ecological roles by leaf‐cutting ants within the context of human‐modified landscapes to guide future research agendas. First, we find that leaf‐cutting ants show that some herbivorous insects are able to generate ecologically important disturbance regimes via non‐trophic activities. Second, impacts of leaf‐cutting ants can be observed at multiple spatio‐temporal scales and levels of biological organization. Third, ecosystem‐level effects from leaf‐cutting ants are ecosystem engineering capable not only of altering the abundance of other organisms, but also the successional trajectory of vegetation. Finally, effects of leaf‐cutting ants are context‐dependent, species‐specific, and synergistically modulated by anthropogenic interferences. Future research should examine how leaf‐cutting ants respond to deforestation and influence remaining vegetation in human‐modified landscapes. By promoting either heterogeneity or homogeneity, leaf‐cutting ants operate not only as agricultural pests but also as ecological key players.