Better soon than never: climate change induces strong phenological reassembly in the flowering of a Mediterranean shrub community.

● Flowering is a key process in the life cycle of a plant. Climate change is shifting flowering phenologies in the Northern Hemisphere, but studies with long data series at community level are scarce, and even more so those regarding the consequences of phenological changes for emerging ecological interactions. In the Mediterranean region, the effects of climate change are stronger than the global average and there is an urgent need to understand how biodiversity will be affected in this area. ● In this study we investigated how the entire flowering phenology of a community comprising 51 perennial species from the south of the Iberian Peninsula changed from the decade of the 1980s to the 2020s. Furthermore, we have analysed the consequences of these changes for flowering order and co-flowering patterns. ● We have found that the flowering phenology of the community has advanced by about 20 days, and was coherent with the increasing temperatures related to climate change. Individual species have generally advanced their entire flowering phenology (start and end) and increased their flowering duration. The early flowering has resulted in a re-organisation of the flowering order of the community and generated a new co-flowering assemblages of species, with a slight trend towards an increase of shared flowering time among species. ● The advanced flowering phenology and changes on flower duration reported here were of unprecedented magnitude, showcasing the extreme effects of climate change on Mediterranean ecosystems. Furthermore, the effects were not similar among species, which could be attributed to differences in sensitivities of environmental cues for flowering. One consequence of these changes in flowering times is the ecological mismatches indicated by the changes in flower order and co-flowering between decades. The novel scenario may lead to new competition or facilitation interactions and to the loss or gain of pollinators.

Rock n' Seeds: A database of seed functional traits and germination experiments from Brazilian rock outcrop vegetation.

Advancing functional ecology depends fundamentally on the availability of data on reproductive traits, including those from tropical plants, which have been historically underrepresented in global trait databases. Although some valuable databases have been created recently, they are mainly restricted to temperate areas and vegetative traits such as leaf and wood traits. Here, we present Rock n' Seeds, a database of seed functional traits and germination experiments from Brazilian rock outcrop vegetation, recognized as outstanding centers of diversity and endemism. Data were compiled through a systematic literature search, resulting in 103 publications from which seed functional traits were extracted. The database includes information on 16 functional traits for 383 taxa from 148 genera, 50 families, and 25 orders. These 16 traits include two dispersal, six production, four morphological, two biophysical, and two germination traits-the major axes of the seed ecological spectrum. The database also provides raw data for 48 germination experiments, for a total of 10,187 records for 281 taxa. Germination experiments in the database assessed the effect of a wide range of abiotic and biotic factors on germination and different dormancy-breaking treatments. Notably, 8255 of these records include daily germination counts. This input will facilitate synthesizing germination data and using this database for a myriad of ecological questions. Given the variety of seed traits and the extensive germination information made available by this database, we expect it to be a valuable resource advancing comparative functional ecology and guiding seed-based restoration and biodiversity conservation in tropical megadiverse ecosystems. There are no copyright restrictions on the data; please cite this paper when using the current data in publications; also the authors would appreciate notification of how the data are used in publications.

O avanço da ecologia funcional depende fundamentalmente da disponibilidade de dados sobre traços reprodutivos, incluindo dados de plantas tropicais, que têm sido historicamente subrepresentados em bancos de dados de traços funcionais globais. Embora alguns bancos de dados valiosos tenham sido criados recentemente, eles são restritos principalmente a áreas temperadas e a traços vegetativos, como traços de folhas e madeira. Neste artigo apresentamos Rock n' Seeds, um banco de dados de traços funcionais de sementes e experimentos de germinação de vegetações associadas a afloramentos rochosos do Brasil, os quais são reconhecidos como centros notáveis de diversidade e endemismo. Os dados foram compilados através de uma revisão sistemática na literatura, resultando em 103 publicações das quais foram extraídos os traços funcionais das sementes. O banco de dados inclui informações de 16 traços funcionais para 383 taxa de 148 gêneros, 50 famílias e 25 ordens. Estes dezesseis traços incluem dois traços de dispersão, seis de produção, quatro morfológicos, dois biofísicos e dois germinativos; os eixos principais do espectro ecológico da semente. O banco de dados também fornece os dados brutos para 48 experimentos de germinação para um total de 10.187 registros para 281 taxa. Os experimentos de germinação no banco de dados avaliaram o efeito de uma ampla gama de fatores abióticos e bióticos sobre a germinação e diferentes tratamentos de quebra de dormência. Particularmente, 8.255 desses registros incluem a contagem diária da germinação. Estas informações facilitarão a síntese de dados de germinação e a utilização deste banco de dados para uma grande variedade de questões ecológicas. Dada a variedade de traços das sementes e as amplas informações sobre germinação disponibilizadas por este banco de dados, esperamos que ele seja um recurso valioso para o avanço da ecologia funcional comparativa e para orientar a restauração baseada em sementes e a conservação da biodiversidade em ecossistemas tropicais megadiversos. Não há restrições de direitos autorais sobre os dados; favor citar este artigo ao utilizar os dados nas publicações e os autores agradeceriam uma notificação de como os dados são utilizados nas publicações.

El avance de la ecología funcional depende fundamentalmente de la disponibilidad de datos sobre rasgos reproductivos-incluyendo los de las plantas tropicales-los cuales han estado poco representados en las bases de datos globales de rasgos. Aunque recientemente se han creado algunas bases de datos valiosas, estas se encuentran restringidas principalmente a las zonas templadas y a los rasgos vegetativos, como los de las hojas y la madera. En este artículo presentamos Rock n' Seeds, una base de datos de rasgos funcionales de semillas y experimentos de germinación de la vegetación asociada a afloramientos rocosos de Brasil, los cuales son destacados centros de diversidad y endemismo. Los datos se recopilaron mediante una búsqueda bibliográfica sistemática, que dio como resultado 103 publicaciones de las que se extrajeron los rasgos funcionales de las semillas. La base de datos incluye información de dieciséis rasgos funcionales para 383 taxones de 148 géneros, 50 familias y 25 órdenes. Estos rasgos incluyen dos rasgos de dispersión, seis de producción, cuatro morfológicos, dos biofísicos y dos de germinación; siendo estos los principales ejes del espectro ecológico de las semillas. La base de datos también proporciona los datos brutos de 48 experimentos de germinación, para un total de 10.187 registros de 281 taxones. Dichos experimentos de germinación evaluaron el efecto de una amplia gama de factores abióticos y bióticos sobre la germinación y de diferentes tratamientos para romper la dormancia. En particular, 8.255 de estos registros cuentan con conteos diarios de germinación. Esto facilitará la síntesis de los datos de germinación y el uso de esta base de datos para una gran diversidad de preguntas ecológicas. Dada la variedad de rasgos de las semillas y la amplia información sobre germinación que ofrece esta base de datos, esperamos que sea un recurso valioso para el avance de la ecología funcional comparativa y para orientar la restauración basada en semillas y la conservación de la biodiversidad en ecosistemas tropicales megadiversos. No hay restricciones de derechos de autor sobre los datos; se solicita citar este documento cuando se utilicen los datos en publicaciones y los autores agradecerán ser notificados sobre cómo se utilizan los datos en las publicaciones.

Floristic composition, pollination and seed-dispersal systems in a target cerrado conservation area / Composição florística, modos de polinização e dispersão de sementes de uma área de cerrado relevante para conservação

Abstract: Cerrado remnants can hold an important diversity of plant species of environmental and ecological relevance. We presented a checklist of vascular plants based on 12 years of inventory carried out in 36 plots (10 m x 2 m; 0.18 ha in total) and during unsystematic walks in a remnant area of cerrado sensu stricto located at Itirapina municipality, state of São Paulo, southeastern Brazil. The list comprised 195 plant species, corresponding to 54 families and 131 genera. The richest families were Fabaceae (25 species), Asteraceae (16), Myrtaceae (16), Rubiaceae (11), Bignoniaceae and Malpighiaceae (10 each), Melastomataceae (9), and Erythroxylaceae, Sapindaceae and Annonaceae (6). Predominant life forms included shrubs and trees, with 68% of the species, followed by lianas with 12%, sub-shrub and herbs with 10% each. Bees were the dominant pollinators (67,5%) and the majority of species had seeds dispersed by animals (56.8%), mostly by birds, followed by wind (33.3%) and self-dispersed (11.2%). More than 60% of the total species were classified as "typical" Cerrado species. Bowdichia virgilioides was the only species classified as Near Threatened (NT) and 157 were regarded as Data Deficient (DD). Our dataset provides floristic, structural, and ecological information for one of the targeted areas for Cerrado survey at São Paulo state, contributing to the understanding of diversity patterns and future conservation and restoration actions in this threatened hotspot.

Resumo: Apresentamos uma lista de verificação de plantas vasculares baseada em 12 anos de inventário realizado em 36 parcelas (10 m x 2 m; 0,18 ha no total) e caminhadas assistemáticas em uma área remanescente de cerrado sensu stricto localizada em Itirapina, município do estado de São Paulo, sudeste do Brasil. A lista é composta por 195 espécies de plantas, correspondendo a 54 famílias e 131 gêneros. As famílias mais ricas foram Fabaceae (25 espécies), Asteraceae (16), Myrtaceae (16), Rubiaceae (11), Bignoniaceae e Malpighiaceae (10 cada), Melastomataceae (9) e Erythroxylaceae, Sapindaceae e Annonaceae (6). As formas de vida predominantes incluíram arbustos e árvores (33,7% das espécies), seguidas por lianas (12%), arbustos e ervas (10%). As abelhas foram os polinizadores dominantes (67,5%) e o principal modo de dispersão foi a zoocoria (56,8%), representada principalmente por pássaros, seguida por vento (33.3 %) e auto (11.2 %). Mais de 60% das espécies encontradas foram classificadas como espécies "típicas" de Cerrado. Bowdichia virgilioides foi a única espécie pertencente a uma categoria de ameaça "Quase Ameaçada (NT)", sendo 157 delas classificadas na categoria "Deficiente de Dados (DD)". Nosso conjunto de dados fornece informações florísticas, estruturais e ecológicas para uma das áreas-alvo do levantamento do Cerrado no estado de São Paulo, sudeste do Brasil, contribuindo para a compreensão dos padrões de diversidade e futuras ações de conservação neste hotspot ameaçado.

Temporal organization among pollination systems in a tropical seasonal forest.

Temporal constancy of pollination systems is essential for the maintenance of pollinators through time. Community-level assessment of flowering phenology allows understanding variations across seasons and years and the risks of decoupling flowering and pollinators' activity. We evaluated flowering patterns and temporal diversity of pollination systems in a tropical seasonal forest. We asked whether the temporal organization of flowering times differs among pollination systems; if there is a constancy of pollination systems through the year, since climate and phylogenies constraint flowering time; if there is a prevalent flowering pattern by pollination system, and if the temporal organization of pollination systems by modularity analyses is coherent with grouping by pre-defined seasons. We characterized 10 pollination systems, examined flowering strategies, climate cues and phylogenetic constraints. Pollination by large-to-medium bees dominated (49.2%), followed by diverse insects (22.1%) and flies (14.7%). The remaining systems represented 14% of species. Flowering occurred year-round for most pollination systems, predominating the seasonal flowering strategy. Flowering patterns ranged from aggregated to nested, and random. Climate affected the flowering of most pollination systems, but there was no phylogeny constraint. Modularity grouped pollination systems differently than rainfall seasonality. Contrasting the expectations of reduced temporal constancy, most systems were present year-round, facilitating the exploitation of floral resources by pollinators. Diversity of pollination systems remained constant despite climate seasonality, indicating that several factors influence the optimum flowering time for pollination in seasonally dry vegetations. Global warming may disrupt phenological patterns and the temporal organization of plant communities, a matter for future studies.

Color signals of bee-pollinated flowers: the significance of natural leaf background.

PREMISE: Flower color is a primary pollinator attractant and generally adjusted to the cognitive system of the pollinators. The perception of flower color depends on the visual system of pollinators and also on environmental factors such as light conditions and the background against which flowers are displayed. METHODS: Using bee-pollinated Fabaceae species as a model, we analyzed flower color diversity and compared flower color signals considering both the standard green and the natural leaf background of two tropical seasonally dry vegetations-a mountain rupestrian grassland (campo rupestre) and a woody savanna (cerrado)-compared to a nontropical Mediterranean shrubland. RESULTS: By using natural background, bees discriminated color for 58% of the flowers in the campo rupestre and for only 43% in cerrado. Both vegetations were surpassed by 75% of bee color discrimination in Mediterranean vegetation. Chromatic contrast and purity were similar among the three vegetation types. Green contrast and brightness were similar between the tropical vegetations but differed from the Mediterranean shrubland. Green contrast differences were lost when using a standard green background, and most variables (purity, green contrast, and brightness) differed according to the background (natural or standard green) in all vegetations. CONCLUSIONS: The natural background influenced bee perception of flower color regardless of vegetation. The background of the campo rupestre promoted green contrast for flowers, ensuring flower detection by pollinators and, along with bees, may also act as a selective pressure driving the diversity of flower colors in Fabaceae species. We highlight the importance of considering the natural background coloration when analyzing flower color signals.

The role of individual variation in flowering and pollination in the reproductive success of a crepuscular buzz-pollinated plant.

BACKGROUND AND AIMS: Plant individuals within a population differ in their phenology and interactions with pollinators. However, it is still unknown how individual differences affect the reproductive success of plants that have functionally specialized pollination systems. Here, we evaluated whether plant individual specialization in phenology (temporal specialization) and in pollination (pollinator specialization) affect the reproductive success of the crepuscular-bee-pollinated plant Trembleya laniflora (Melastomataceae). METHODS: We quantified flowering activity (amplitude, duration and overlap), plant-pollinator interactions (number of flowers visited by pollinators) and reproductive success (fruit set) of T. laniflora individuals from three distinct locations in rupestrian grasslands of southeastern Brazil. We estimated the degree of individual temporal specialization in flowering phenology and of individual specialization in plant-pollinator interactions, and tested their relationship with plant reproductive success. KEY RESULTS: Trembleya laniflora presented overlapping flowering, a temporal generalization and specialized pollinator interactions. Flowering overlap among individuals and populations was higher than expected by chance but did not affect the individual interactions with pollinators and nor their reproductive success. In contrast, higher individual generalization in the interactions with pollinators was related to higher individual reproductive success. CONCLUSIONS: Our findings suggest that individual generalization in plant-pollinator interaction reduces the potential costs of specialization at the species level, ensuring reproductive success. Altogether, our results highlight the complexity of specialization/generalization of plant-pollinator interactions at distinct levels of organization, from individuals to populations, to species.

Flowering Phenology and the Influence of Seasonality in Flower Conspicuousness for Bees.

Flowering patterns are crucial to understand the dynamics of plant reproduction and resource availability for pollinators. Seasonal climate constrains flower and leaf phenology, where leaf and flower colors likely differ between seasons. Color is the main floral trait attracting pollinators; however, seasonal changes in the leaf-background coloration affect the perception of flower color contrasts by pollinators. For a seasonally dry woody cerrado community (Brazilian savanna) mainly pollinated by bees, we verified whether seasonality affects flower color diversity over time and if flower color contrasts of bee-pollinated species differ between seasons due to changes in the leaf-background coloration. For 140 species, we classified flower colors based on human-color vision, and for 99 species, we classified flower colors based on bee-color vision (spectral measurements). We described the community's flowering pattern according to the flower colors using a unique 11 years phenological database. For the 43 bee-pollinated species in which reflectance data were also available, we compared flower color diversity and contrasts against the background between seasons, considering the background coloration of each season. Flowering was markedly seasonal, peaking at the end of the dry season (September), when the highest diversity of flower colors was observed. Yellow flowers were observed all year round, whereas white flowers were seasonal, peaking during the dry season, and pink flowers predominated in the wet season, peaking in March. Bee-bluegreen flowers peaked between September and October. Flowers from the wet and dry seasons were similarly conspicuous against their corresponding background. Regardless of flowering season, the yellowish background of the dry season promoted higher flower color contrast for all flower species, whereas the greener background of the wet season promoted a higher green contrast. Temporal patterns of flower colors and color contrasts were related to the cerrado seasonality, but also to bee's activity, visual system, and behavior. Background coloration affected flower contrasts, favoring flower conspicuousness to bees according to the season. Thus, our results provide new insights regarding the temporal patterns of plant-pollinator interactions.

The diversity and evolution of pollination systems in large plant clades: Apocynaceae as a case study.

Background and Aims: Large clades of angiosperms are often characterized by diverse interactions with pollinators, but how these pollination systems are structured phylogenetically and biogeographically is still uncertain for most families. Apocynaceae is a clade of >5300 species with a worldwide distribution. A database representing >10 % of species in the family was used to explore the diversity of pollinators and evolutionary shifts in pollination systems across major clades and regions. Methods: The database was compiled from published and unpublished reports. Plants were categorized into broad pollination systems and then subdivided to include bimodal systems. These were mapped against the five major divisions of the family, and against the smaller clades. Finally, pollination systems were mapped onto a phylogenetic reconstruction that included those species for which sequence data are available, and transition rates between pollination systems were calculated. Key Results: Most Apocynaceae are insect pollinated with few records of bird pollination. Almost three-quarters of species are pollinated by a single higher taxon (e.g. flies or moths); 7 % have bimodal pollination systems, whilst the remaining approx. 20 % are insect generalists. The less phenotypically specialized flowers of the Rauvolfioids are pollinated by a more restricted set of pollinators than are more complex flowers within the Apocynoids + Periplocoideae + Secamonoideae + Asclepiadoideae (APSA) clade. Certain combinations of bimodal pollination systems are more common than others. Some pollination systems are missing from particular regions, whilst others are over-represented. Conclusions: Within Apocynaceae, interactions with pollinators are highly structured both phylogenetically and biogeographically. Variation in transition rates between pollination systems suggest constraints on their evolution, whereas regional differences point to environmental effects such as filtering of certain pollinators from habitats. This is the most extensive analysis of its type so far attempted and gives important insights into the diversity and evolution of pollination systems in large clades.

How flower colour signals allure bees and hummingbirds: a community-level test of the bee avoidance hypothesis.

Colour signals are the main floral trait for plant-pollinator communication. Owing to visual specificities, flower visitors exert different selective pressures on flower colour signals of plant communities. Although they evolved to attract pollinators, matching their visual sensitivity and colour preferences, floral signals may also evolve to avoid less efficient pollinators and antagonistic flower visitors. We evaluated evidence for the bee avoidance hypothesis in a Neotropical community pollinated mainly by bees and hummingbirds, the campo rupestre. We analysed flower reflectance spectra, compared colour variables of bee-pollinated flowers (bee-flowers; 244 species) and hummingbird-pollinated flowers (hummingbird-flowers; 39 species), and looked for evidence of bee sensorial exclusion in hummingbird-flowers. Flowers were equally contrasting for hummingbirds. Hummingbird-flowers were less conspicuous to bees, reflecting mainly long wavelengths and avoiding red-blind visitors. Bee-flowers reflected more short wavelengths, were more conspicuous to bees (higher contrasts and spectral purity) than hummingbird-flowers and displayed floral guides more frequently, favouring flower attractiveness, discrimination and handling by bees. Along with no phylogenetic signal, the differences in colour signal strategies between bee- and hummingbird-flowers are the first evidence of the bee avoidance hypothesis at a community level and reinforce the role of pollinators as a selective pressure driving flower colour diversity.

Anthropogenic edges, isolation and the flowering time and fruit set of Anadenanthera peregrina, a cerrado savanna tree.

Fragmentation exposes plants to extreme environmental conditions with implications for species phenology and reproduction.We investigated whether isolation and edge effects influence size, flowering time, fruit set, and seedling establishment of Anadenanthera peregrina var. falcata. We compared trees in the interior (n =85), and on the edge (n =74) of a cerrado savanna fragment as well as in a pasture (n =26) with respect to size, flowering phenology, flower and fruit production, fruit and seed set, predispersal seed predation, and seedling establishment. Trees in the pasture were larger and produced a higher number of flowers and fruits than trees on the edge and interior, yet seed set did not differ across environments. The plant size structure explained the flower and fruit production, and the self-compatibility breeding system caused a similar seed set regardless of the environment. First flowering was later and fruit set higher in the interior. We argue that time of first flower influenced the fruit set of Anadenathera. Edge and isolated trees started to flower earlier as a response to microclimatic conditions--mainly temperature--reducing the fruit set. Predispersal seed predation was lower among pasture trees. Conversely, we found seedlings only on the edge and in the interior of cerrado, suggesting that the pasture was of poor quality habitat for Anadenanthera recruitment. Isolation affected the plant size structure and reproduction of Anadenanthera trees. Studies comparing plant phenology under contrasting environmental conditions may offer clues on how global change may affect plant reproduction in the tropics.

Reproductive phenology of coastal plain Atlantic forest vegetation: comparisons from seashore to foothills.

The diversity of tropical forest plant phenology has called the attention of researchers for a long time. We continue investigating the factors that drive phenological diversity on a wide scale, but we are unaware of the variation of plant reproductive phenology at a fine spatial scale despite the high spatial variation in species composition and abundance in tropical rainforests. We addressed fine scale variability by investigating the reproductive phenology of three contiguous vegetations across the Atlantic rainforest coastal plain in Southeastern Brazil. We asked whether the vegetations differed in composition and abundance of species, the microenvironmental conditions and the reproductive phenology, and how their phenology is related to regional and local microenvironmental factors. The study was conducted from September 2007 to August 2009 at three contiguous sites: (1) seashore dominated by scrub vegetation, (2) intermediary covered by restinga forest and (3) foothills covered by restinga pre-montane transitional forest. We conducted the microenvironmental, plant and phenological survey within 30 transects of 25 m × 4 m (10 per site). We detected significant differences in floristic, microenvironment and reproductive phenology among the three vegetations. The microenvironment determines the spatial diversity observed in the structure and composition of the flora, which in turn determines the distinctive flowering and fruiting peaks of each vegetation (phenological diversity). There was an exchange of species providing flowers and fruits across the vegetation complex. We conclude that plant reproductive patterns as described in most phenological studies (without concern about the microenvironmental variation) may conceal the fine scale temporal phenological diversity of highly diverse tropical vegetation. This phenological diversity should be taken into account when generating sensor-derived phenologies and when trying to understand tropical vegetation responses to environmental changes.

Diversity of functional traits of fleshy fruits in a species-rich Atlantic rain forest / Diversidade de características funcionais dos frutos carnosos em uma área de Mata Atlântica

Production of vertebrate-dispersed fruits is the most common strategy of tropical woody plants to disperse their seeds. Few studies have documented community-wide variation of fruit morphology and chemistry of vertebrate-dispersed fruits in species-rich tropical communities. We examined the functional diversity of fruit morphological and chemical traits of 186 species representing 57 plant families in an undisturbed lowland plant community in the Atlantic rain forest of SE, Brazil. We were particularly interested in associating morphological and chemical fruit traits to their main seed dispersers, either birds, mammals or 'mixed' (i.e. fruits eaten by birds and mammals). The morphological and chemical traits of fruits at the study site generally resemble the patterns observed in fruits worldwide. Bird fruits tend to be smaller than mammal fruits, being colored black or red, whereas mammal fruits are often yellow or green. Mammal fruits are more variable than bird fruits in relation to morphological traits, while the reverse is true for chemical traits. Mixed fruits resemble bird fruits in the patterns of variation of morphological and chemical traits, suggesting that they are primarily bird-dispersed fruits that are also exploited by mammals. Mixed fruits are common in tropical forests, and represent an excellent opportunity to contrast the effectiveness of different functional groups of frugivores dispersing the same plant species.

A produção de frutos carnosos é a estratégia mais comum adotada por plantas arbóreas tropicais para dispersar suas sementes. Poucos estudos têm documentado variações em nível de comunidade na morfologia e composição química de frutos carnosos em comunidades tropicais ricas em espécie. Nós examinamos a diversidade funcional das características morfológicas e químicas dos frutos de 186 espécies, representando 57 famílias de plantas em uma área de planície coberta por Mata Atlântica bem preservada no sudeste do Brasil. Estávamos particularmente interessados em associar as características morfológicas e químicas dos frutos a seus principais dispersores de sementes: aves, mamíferos ou "misto" (i.e. frutos consumidos por aves e mamíferos). As características morfológicas e químicas dos frutos no geral se assemelharam a padrões observados em outras partes do mundo. Frutos consumidos por aves tendem a ser menores do que os frutos de mamíferos, apresentando predominantemente cor preta ou vermelha, enquanto os frutos de mamíferos são geralmente amarelos ou verdes. Frutos consumidos por mamíferos são mais variáveis do que os frutos de aves em relação às características morfológicas, enquanto o inverso é verdadeiro para as características químicas. Frutos "mistos" assemelham-se aos frutos consumidos exclusivamente por aves em relação aos padrões de variação das características morfológicas e químicas, o que sugere serem eles frutos primariamente ornitocóricos que são também explorados por mamíferos. Frutos "mistos" são comuns em florestas tropicais e representam excelente oportunidade para contrastar a efetividade de diferentes grupos funcionais de frugívoros ao dispersar a mesma espécie de planta.