Seed dispersal is one of the most studied plant–animal mutualisms. It has been proposed that the dispersal of many large-seeded plants from Neotropical forests was primarily conducted by extinct megafauna, and currently by livestock. Parrots can transport large fruits using their beaks, but have been overlooked as seed dispersers. We demonstrate that three macaws (Ara ararauna, A. glaucogularis and A. severus) are the main dispersers of the large-seeded motacú palm Attalea princeps, which is the biomass-dominant tree in the Bolivian Amazonian savannas. Macaws dispersed fruits at high rates (75–100% of fruits) to distant (up to 1200 m) perching trees, where they consumed the pulp and discarded entire seeds, contributing to forest regeneration and connectivity between distant forests islands. The spatial distribution of immature palms was positively associated to the proximity to macaws’ perching trees and negatively to the proximity to cattle paths. The disperser role of livestock, presumably a substitute for extinct megafauna, had little effect due to soil compaction, trampling and herbivory. Our results underscore the importance of macaws as legitimate, primary dispersers of large-seeded plants at long distances and, specifically, their key role in shaping the landscape structure and functioning of this Amazonian biome.

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The extinction of ecological functions is increasingly considered a major component of biodiversity loss, given its pervasive effects on ecosystems, and it may precede the disappearance of the species engaged. Dispersal of many large-fruited (>4 cm diameter) plants is thought to have been handicapped after the extinction of megafauna in the Late Pleistocene and the recent defaunation of large mammals. We recorded the seed dispersal behavior of two macaws (Anodorhynchus hyacinthinus and Anodorhynchus leari) in three Neotropical biomes, totaling >1700 dispersal events from 18 plant species, 98% corresponding to six large-fruited palm species. Dispersal rates varied among palm species (5%–100%). Fruits were moved to perches at varying distances (means: 17–450 m, maximum 1620 m). Macaws also moved nuts after regurgitation by livestock, in an unusual case of tertiary dispersal, to distant perches. A high proportion (11%–75%) of dispersed nuts was found undamaged under perches, and palm recruitment was confirmed under 6%–73% of the perches. Our results showed that these macaws were legitimate, long-distance dispersers, and challenge the prevailing view that dispersal of large-fruited plants was compromised after megafauna extinction. The large range contraction of these threatened macaws, however, meant that these mutualistic interactions are functionally extinct over large areas at a continental scale.

The dispersal of many large-seeded plants is thought to have been handicapped by the extinction of megafauna in the late Pleistocene, and due to the ongoing defaunation of the largest of the extant dispersers. Oversized fruits defined as "megafaunal" provide variable amounts of flesh even though many of them cannot be ingested entirely, nor their seeds defecated, by any extant vertebrate. This apparent mismatch lead to the hypothesis of anachronisms involving extinct megafauna as dispersal-mediated selective agents on fruit traits shaped through endozoochory. It has been suggested that free-ranging livestock partially supply the dispersal functions previously provided by those globally or regionally extinct species. However, there is little knowledge on the role of livestock as a surrogate for megafauna dispersal agents relative to living wild dispersers. Here, we focus on seed dispersal of six palm species (Attalea eichleri, Attalea barreirensis, Attalea speciosa, Attalea princeps, Mauritia flexuosa, Acrocomia totai) with large fruits that conform to the so-called "megafaunal syndrome". Data on seed dispersal were obtained by observations and camera trapping in the Cerrado, Pantanal and Amazonia biomes in Bolivia and Brazil. Rich communities of wild seed dispersers differing among palm species and study areas were recorded, including rodents, monkeys, canids, and a wide variety of birds, especially parrots. Long-distance primary dispersal was mainly conducted by parrots, while multiple species acted as short-and medium-distance secondary dispersers. Among livestock, dispersal was limited to seeds of A. totai and A. princeps moved by several species through stomatochory and endozoochory (mainly regurgitation). These results show that the large seeds can be efficiently dispersed externally by a wide array of present-day vertebrates of variable size but much smaller than extinct megafauna and livestock. A knowledge gap of the natural history of these and other plants with oversized fruits assumed to be maladapted for contemporary dispersal may have been partially favored by neglecting some key disperser guilds (e.g., parrots) and dispersal mechanisms (e.g., ectozoochory). The evaluation of historic and ongoing defaunation of key external dispersers is advocated to understand the influence of actual (rather than putative) dispersers on contemporary frugivore-plant mutualistic interactions.

Rediscovered in the wild twenty years ago, the breeding biology of wild Blue-throated Macaws remains largely unexplored, yet is essential to its effective conservation and recovery. Here, we analyse reproductive parameters in an intensively managed wild population of Blue-throated Macaws, providing the first data on the breeding biology of this critically endangered species. During the six-year study period, 2007–2012, the number of active breeding pairs either remained constant or decreased, depending on the site, and no new breeding pairs were discovered despite extensive searching. We documented nesting attempts in natural cavities in dead palms or live hardwoods, and artificial nest boxes. Egg-laying was concentrated during the end of dry season and the beginning of the wet season, August through December. Hatching failure was the greatest cause of egg losses. Half of the breeding attempts of Blue-throated Macaws produced at least one fledging, on average two, after a 85 days nestling period. An average of 4.3 nestlings per year fledged from all known wild nests combined. Each pair lost roughly 65% of its initial reproductive investment at each nesting attempt. In most successful nesting attempts of individualized pairs, a new nesting attempt was not detected the following year. All monitored breeding pairs showed high nest site fidelity, reusing hardwood-tree cavities and nest boxes. Our findings will aid conservation efforts by refining current actions and prompting new approaches towards the conservation and recovery of the Blue-throated Macaw.

Parrots stand out among birds because of their poor conservation status and the lack of available information on their population sizes and trends. Estimating parrot abundance is complicated by the high mobility, gregariousness, patchy distributions, and rarity of many species. Roadside car surveys can be useful to cover large areas and increase the probability of detecting spatially aggregated species or those occurring at very low densities. However, such surveys may be biased due to their inability to handle differences in detectability among species and habitats. We conducted 98 roadside surveys, covering > 57,000 km across 20 countries and the main world biomes, recording ca. 120,000 parrots from 137 species. We found that larger and more gregarious species are more easily visually detected and at greater distances, with variations among biomes. However, raw estimates of relative parrot abundances (individuals/km) were strongly correlated (r = 0.86–0.93) with parrot densities (individuals/km2) estimated through distance sampling (DS) models, showing that variability in abundances among species (>40 orders of magnitude) overcomes any potential detectability bias. While both methods provide similar results, DS cannot be used to study parrot communities or monitor the population trends of all parrot species as it requires a minimum of encounters that are not reached for most species (64% in our case), mainly the rarest and more threatened. However, DS may be the most suitable choice for some species-specific studies of common species. We summarize the strengths and weaknesses of both methods to guide researchers in choosing the best–fitting option for their particular research hypotheses, characteristics of the species studied, and logistical constraints.

Plant–animal interactions are key to sustaining whole communities and ecosystem function. However, their complexity may limit our understanding of the underlying mechanisms and the species involved. The ecological effects of epizoochory remain little known compared to other seed dispersal mechanisms given the few vectors identified. In addition, epizoochory is mostly considered non-mutualistic since dispersers do not obtain nutritional rewards. Here, we show a widespread but unknown mutualistic interaction between parrots and plants through epizoochory. Combining our observations with photos from web-sources, we recorded nearly 2000 epizoochory events in 48 countries across five continents, involving 116 parrot species and nearly 100 plant species from 35 families, including both native and non-native species. The viscid pulp of fleshy fruits and anemochorous structures facilitate the adherence of tiny seeds (mean 3.7 × 2.56 mm) on the surface of parrots while feeding, allowing the dispersion of these seeds over long distances (mean = 118.5 m). This parrot–plant mutualism could be important in ecosystem functioning across a wide diversity of environments, also facilitating the spread of exotic plants. Future studies should include parrots for a better understanding of plant dispersal processes and for developing effective conservation actions against habitat loss and biological invasions.