Ants frequently interact with non-myrmecochorous diaspores on the ground in neotropical savannas. Our objective was to quantify the removal rate of these diaspores by ants and vertebrates in order to test the predator avoidance hypothesis, and to test how diaspore traits influence removal by ants and dispersal distance. We also investigated whether seed cleaning (removal of fruit matter simulating ant activity) can influence seed germination. We performed removal experiments with nine diaspore species in a reserve of cerrado savanna in south-east Brazil. Considerable differences in removal rates were found among the nine species. We found a positive linear relationship between lipid content and removal rates for five diaspore species. Vertebrate predation pressure was low for most species, limiting the benefits that ants can provide to plants to escape predators. Ants displace diaspores up to 25 m, which may increase the chance of a seed hitting a safe site. Smaller diaspores attain longer distances of dispersal than large ones. Seed cleaning increased the germination rate for five out of six species tested in greenhouse experiments. Ant activity can have relevant and possibly lasting effects on seed fate of plants adapted for vertebrate dispersal in the cerrado savanna.

Summary 1.A substantial portion of the crop of fruiting trees falls beneath parent plants as a result of dispersal failure. Such diaspores are considered as waste because the likelihood of plant recruitment is usually very small close to parent trees. However, many animals may rescue fallen diaspores and provide them with another chance of dispersal and establishment. 2. We investigated the effectiveness of two broad types of seed dispersal vectors for the regeneration of Xylopia aromatica in the Brazilian cerrado savanna: birds that remove diaspores from plant canopies and ants that harvest diaspores on the ground under the parent plant (as rescuers) or from bird feces (as secondary dispersers). 3. Birds removed a mean of 32% of the crop from plant canopies, but removal was independent of crop size. A large part of the crop (mean of 25%) landed beneath parent plants or was dropped after manipulation by vertebrate frugivores as viable diaspores. Ants from at least five genera removed most fallen diaspores (up to 83%) within 24 h. Ants influenced the fate of a large amount of the crop, and for some trees ants removed as many diaspores as birds. 4. Large ants rescued some diaspores to distances beyond the parent plant crown, but birds may remove diaspores 40-fold farther. However, seedlings of X. aromatica were only found close to nests of large ants, probably due to diaspore rescuing and ⁄ or directed secondary dispersal by certain ant groups following primary dispersal by birds. Although an unknown percentage of seeds was lost to granivorous ants, diaspore removal by ants potentially enhances the likelihood of plant recruitment due to distance-related benefits and directed dispersal to ant nests, while birds play a premier role in long-distance seed dispersal and metapopulation dynamics. 5. Synthesis. Birds and ants provide complementary seed dispersal at different spatial scales to X. aromatica. Since ants remove most fallen diaspores beneath parent plants, the use of diaspore removal rates from plant canopy as a surrogate of plant fitness may be misleading. By acting as secondary dispersers, ants may also provide a fine-tuned dispersal following long-distance dispersal by birds (i.e. diplochory).

The scale at which seed dispersal operates has many implications for the spatial patterns of plant recruitment and diversity. We investigated the effect of short- (ants) and long-distance (birds) seed dispersal of the fleshy-fruited melastome, Miconia rubiginosa, in the Brazilian savanna. We estimated the contribution of dispersal vectors to the removal of the fruit crop from the canopy (birds), and once seeds have reached the cerrado floor (ants) over two fruiting seasons. Birds (13 species) removed up to 23.7% of the fruit crop from the crown, but dropped a substantial proportion of fruits beneath the parent plant. Birds removed a greater proportion of fruits from trees producing large fruit crops, as predicted by the fruit crop size hypothesis. However, up to 18.9% of the fruit crop fell beneath the parent plant as ripe fruit. Most fallen fruits were removed by ants (seven genera), which are likely to play a relatively important role in terms of the quantity of seeds dispersed, especially for plants producing small fruit crops (a conceptual model is presented). Birds and ants did not influence seed germination, but they differ in terms of the spatial scale of dispersal and deposition patterns. Ants probably play an important role in the local population dynamics of Miconia, whereas birds are responsible for long-distance dispersal associated with the colonization of new patches and metapopulation dynamics. By removing seeds from bird droppings, ants may also reshape at a finer scale the seed rain generated by primary dispersers. Indeed, seedlings and saplings of Miconia are more frequently found around leaf-cutter ant nests than in control areas away from ant nests or around large Miconia trees. The quantitative component of dispersal effectiveness by ants acting as "rescuers" of seeds that fail to be dispersed, or fall under parent trees, is probably more important than currently recognized in other systems.

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Ants frequently interact with fleshy fallen diaspores (fruits or seeds) not adapted for ant‐dispersal. Such interactions are usually considered as opportunistic, but recent evidence has indicated that these ants may differ in their effects on diaspore survival and plant recruitment. We investigated if partner choices are recognizable among genera of ants and plants, and if ant and plant traits may influence such preferences in cerrado (savanna‐like vegetation) from southeast Brazil. During a 2‐yr period, 521 ant–diaspore interactions were recorded through various methods, yielding 71 ant species and 38 plant species. Exploitation of fallen diaspores was common among several ant genera, and included carnivorous, omnivorous, and fungivorous ants. Contrary to others areas around the world, where true myrmecochory (seed dispersal by ants) is common among shrubs, ants also exploited diaspores from several cerrado trees. Plant life form, diaspore size, and ant body size did not seem to explain the pattern of interactions observed. Two subsets of preferential interactions, however, segregated fungivorous ants from another group composed of carnivorous and omnivorous ants, probably influenced by the chemical composition of the plant diaspores. Omnivorous ants usually remove the fleshy portion of diaspores on spot and probably provide limited benefits to plants. Carnivorous and fungivorous ants usually remove the whole diaspore to the nest. As each of these ant groups may influence the fitness of diaspores in different ways, there are possible subtle pathways for the evolution of partner choices between ants and these non‐myrmecochorous diaspores.

Edge effects may lead to changes in mutualistic plant-animal interactions, such as seed dispersal, that are critical to plant regeneration. However, research into edge effects is neglected in the Brazilian cerrado, the largest neotropical savanna. We evaluated the consequences of edge effects in the cerrado for the regeneration of Erythroxylum pelleterianum (Erythroxylaceae), a shrub that benefits from seed dispersal by ants. We compared air temperature, relative humidity, and vapor pressure deficit, as well as the frequency and outcome of ant-diaspore interactions between cerrado edges and interiors. The inner portion of cerrado was likely to be moister than its borders, but seed production and germination did not differ between edge and interior of cerrado. Ants removed more seeds near fragment edges than at the interior. However, Myrmicinae ants dominated ant-fruit interactions at edges. These ants are likely to provide few benefits to the seeds. Seedlings of E. pelleterianum growing close to Ponerinae ant nests showed higher survival than seedlings growing away from nests in the interior of cerrado, but such effect disappeared near edges. Widespread seedling mortality due to a higher evaporative demand at edges may partially account for this effect. Furthermore, Ponerinae's nests also showed a lower residence time near edges, decreasing possible benefits derived from ant colony activity such as nutrient enrichment and protection against insect herbivores. Edge effects could change the structure and dynamics of vegetation in cerrado fragments, due in part to the collapse of the mutualistic interactions demonstrated here.

Seed dispersal may involve different vectors of dispersal in two or more sequential phases (i.e., diplochory). However, contributions of each phase to the overall seed dispersal effectiveness (SDE) are poorly understood and hard to evaluate due to post-dispersal processes that affect seed and seedling survival. We investigated the simultaneous bird (phase 1, in plant canopy) and ant (phase 2, on the floor) contributions to SDE with the ornithochoric shrub Erythroxylum ambiguum in a Brazilian Atlantic forest. Twelve species of birds fed on fruit and dispersed approximately 26 % of the seed crop. The remaining seed crop, 90 % of which contained viable seeds, fell to the ground beneath the parental plant. Ants either cleaned seeds in fruits or carried fallen fruit and seeds from bird feces to their nests. Although E. ambiguum has no adaptation for ant dispersal, ants were as quantitatively important as birds. Birds and ants equally increased germination rates compared to controls. However, birds deposited seeds farther from the parent, where seedling survival was higher (78 %) than it was beneath the parent (44 %), whereas ants carried seeds to their nests, where seedling survival was higher (83 %) than in controls away from their nests (63 %). Diplochory allowed a 42 % increase in SDE compared to dispersal in phase 1 alone. High lipid content in the fruit pulp of E. ambiguum may facilitate the inclusion of ants in a second step of dispersal after diaspores reach the floor. Ants can also buffer the dispersal of diplochorous plants against decreases in phase 1 dispersers.

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