Interaction engineering: Non-trophic effects modify interactions in an insect galler community.

Theory suggests that non-trophic interactions can be a major mechanism behind community stability and persistence, but community-level empirical data are scarce, particularly for effects on species interactions mediated through changes in the physical environment. Here, we explored how ecosystem engineering effects can feed back to the engineer, not only modulating the engineer's population density (node modulation) but also affecting its interactions with other species (link modulation). Gall induction can be viewed as ecosystem engineering since galls serve as habitat for other species. In a community-level field experiment, we generated treatments with reduced or elevated ecosystem engineering by removing or adding post-emergence galls to different plots of their host plant in the Brazilian Cerrado. We tested the effect of post-emergence galls on the galler, as well as on the galler-parasitoid and galler-aphid interactions. The manipulation of post-emergence galls had little effect on the galler-abundance and survivorship were not affected, and gall volume changed only slightly-but modified interactions involving the galler, parasitoid wasps and inquiline aphids. Aphid inquilines negatively affected density-dependent parasitism rates (interaction modification) likely by killing parasitised galling larvae. Post-emergence galls interfered with aphid inquilinism-likely by the provision of alternative habitat for aphids-and thus interfered with the negative effect of aphids on parasitism (modification of an interaction modification). This work is one of the few studies to demonstrate experimentally the role played by environment-mediated interaction modification at a community level in the field. Moreover, by manipulating a species' ecosystem engineering effect (post-emergence galls) instead of the species itself, we demonstrate the novel result that populations can be regulated by non-trophic effects initiated by their own activities that alter their interaction with other species. This reveals that indirect interactions mediated via the environment offer new pathways of feedback loops for population regulation. Our results indicate that interaction modification has the potential to be a key regulatory mechanism underlying interaction variation in nature, and play a major role in community structure, dynamics and stability.

Experimentally reducing species abundance indirectly affects food web structure and robustness.

Studies on the robustness of ecological communities suggest that the loss or reduction in abundance of individual species can lead to secondary and cascading extinctions. However, most such studies have been simulation-based analyses of the effect of primary extinction on food web structure. In a field experiment we tested the direct and indirect effects of reducing the abundance of a common species, focusing on the diverse and self-contained assemblage of arthropods associated with an abundant Brazilian shrub, Baccharis dracunculifolia D.C. (Asteraceae). Over a 5-month period we experimentally reduced the abundance of Baccharopelma dracunculifoliae (Sternorrhyncha: Psyllidae), the commonest galling species associated with B. dracunculifolia, in 15 replicate plots paired with 15 control plots. We investigated direct effects of the manipulation on parasitoids attacking B. dracunculifoliae, as well as indirect effects (mediated via a third species or through the environment) on 10 other galler species and 50 associated parasitoid species. The experimental manipulation significantly increased parasitism on B. dracunculifoliae in the treatment plots, but did not significantly alter either the species richness or abundance of other galler species. Compared to control plots, food webs in manipulated plots had significantly lower values of weighted connectance, interaction evenness and robustness (measured as simulated tolerance to secondary extinction), even when B. dracunculifoliae was excluded from calculations. Parasitoid species were almost entirely specialized to individual galler species, so the observed effects of the manipulation on food web structure could not have propagated via the documented trophic links. Instead, they must have spread either through trophic links not included in the webs (e.g. shared predators) or non-trophically (e.g. through changes in habitat availability). Our results highlight that the inclusion of both trophic and non-trophic direct and indirect interactions is essential to understand the structure and dynamics of even apparently discrete ecological communities.

Variation of arbuscular mycorrhizal fungal communities along an altitudinal gradient in rupestrian grasslands in Brazil.

Variation in arbuscular mycorrhizal fungi (AMF) communities is described for the first time in rupestrian grasslands in Brazil along an altitudinal gradient of 700 m (800 to 1400 m a.s.l.). Hypotheses tested were that soil properties influence the variation in AMF communities and that the frequency of the most common species of AMF is inversely influenced by the richness of other AMF. Field and laboratory data were collected on AMF community composition, richness, density, and frequency in the altitudinal gradient, and the relationships with several physical-chemical soil properties and altitude were evaluated. Fifty-one species of AMF were recorded, with 14 species being reported as possibly new to science and nine species representing new records for Brazil. This single elevation gradient alone contains 22% of the known world diversity of AMF. Soil properties and AMF community density and richness varied significantly along the elevation (p < 0.05). AMF density and richness were higher at the intermediate altitude, while AMF species composition differed statistically among the altitudes.

Dissimilar forests along the Rio Doce watershed call for multiple restoration references to avoid biotic homogenization.

An environmental disaster caused by the rupture of a mining tailings dam has impacted a large area of the Rio Doce watershed in the Brazilian Atlantic Forest, resulting in unprecedented damage at spatial and temporal scales. The Atlantic Forest is one of the world's most important biodiversity hotspots. A long history of land use conversion has resulted in a highly fragmented landscape. Despite numerous restoration initiatives, these efforts have often biased criteria and use limited species assemblages. We conducted a comprehensive synthesis of the plant community in riparian forests along the Rio Doce watershed. Our work detailed vegetation composition (tree and sapling strata) and examined its relationship with edaphic and landscape factors, aiming to inform restoration projects with scientifically robust knowledge. A total of 4906 individuals from the tree strata and 4565 individuals from the sapling strata were recorded, representing a total of 1192 species from 75 families. Only 0.8% of the tree species and 0.5% of the sapling species occurred in all sampled sectors, with over 84% of the species occurring in a single watershed sector for both strata. We observed a high species heterogeneity modulated by turnover (92.3% in the tree, and 92.7% in the sapling strata) among sites. Overall, our research revealed a gradient of soil fertility influencing species composition across different strata. Additionally, we discovered that preserved landscapes had a positive impact on species diversity within both strata. The species exclusivity in the sampled sites and the high turnover rate imply the need to consider multiple reference ecosystems when restoring the watershed to reduce the risk of biotic homogenization. Finally, the reference ecosystems defined here serve as a basis for the selection of locally particular species in the implementation of restoration projects that aim to improve biodiversity, ecosystem services, and water security.

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.

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation.

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and micro-habitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having a geographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system.

Diversity of Gall-Inducing Insects Associated With a Widely Distributed Tropical Tree Species: Testing the Environmental Stress Hypothesis.

Abiotic factors can affect plant performance and cause stress, which in turn affects plant-herbivore interactions. The Environmental Stress Hypothesis (ESH) predicts that gall-inducing insect diversity will be greater on host plants that grow in stressful habitats. We tested this hypothesis, considering both historical and ecological scales, using the plant Copaifera langsdorffii Desf. (Fabaceae) as a model because it has a wide geographic distribution and is a super-host of gall-inducing insects. According to the ESH, we predicted that 1) on a historical scale, the diversity of gall-inducing insects will be higher in habitats with greater environmental stress and 2) on an ecological scale, gall-inducing insect diversity will be greater on plants that possess greater levels of foliar sclerophylly. We sampled gall-inducing insects on plants of C. langsdorffii in five sites with different levels of water and soil nutrient availability and separated from each other by a distance of up to 470 km. The composition, richness, and abundance of gall-inducing insects varied among study sites. Plants located in more stressful habitats had higher levels of foliar sclerophylly; but richness and abundance of gall-inducing insects were not affected by host plant sclerophylly. Habitat stress was a good predictor of gall-inducing insect diversity on a regional scale, thus corroborating the first prediction of the ESH. No relationship was found between plant sclerophylly and gall-inducing insect diversity within habitats. Therefore, on a local scale, we did not find support for our second prediction related to the ESH.

Ants on plants: a meta-analysis of the role of ants as plant biotic defenses.

We reviewed the evidence on the role of ants as plant biotic defenses, by conducting meta-analyses for the effects of experimental removal of ants on plant herbivory and fitness with data pooled from 81 studies. Effects reviewed were plant herbivory, herbivore abundance, hemipteran abundance, predator abundance, plant biomass and reproduction in studies where ants were experimentally removed (n = 273 independent comparisons). Ant removal exhibited strong effects on herbivory rates, as plants without ants suffered almost twice as much damage and exhibited 50% more herbivores than plants with ants. Ants also influenced several parameters of plant fitness, as plants without ants suffered a reduction in biomass (-23.7%), leaf production (-51.8%), and reproduction (-24.3%). Effects were much stronger in tropical regions compared to temperate ones. Tropical plants suffered almost threefold higher herbivore damage than plants from temperate regions and exhibited three times more herbivores. Ant removal in tropical plants resulted in a decrease in plant fitness of about 59%, whereas in temperate plants this reduction was not statistically significant. Ant removal effects were also more important in obligate ant-plants (=myrmecophytes) compared to plants exhibiting facultative relationships with hemiptera or those plants with extrafloral nectaries and food bodies. When only tropical plants were considered and the strength of the association between ants and plants taken into account, plants with obligate association with ants exhibited almost four times higher herbivory compared to plants with facultative associations with ants, but similar reductions in plant reproduction. The removal of a single ant species increased plant herbivory by almost three times compared to the removal of several ant species. Altogether, these results suggest that ants do act as plant biotic defenses, but the effects of their presence are more pronounced in tropical systems, especially in myrmecophytic plants.

Resilience and restoration of tropical and subtropical grasslands, savannas, and grassy woodlands.

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open-canopy grassy woodlands) remains limited. To incorporate grasslands into large-scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved - frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human-caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species-diverse plant communities, including endemic species, are slow to recover. Complicating plant-community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re-establish. To guide restoration decisions, we draw on the old-growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old-growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old-growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.

Biodiversity recovery of Neotropical secondary forests.

Old-growth tropical forests harbor an immense diversity of tree species but are rapidly being cleared, while secondary forests that regrow on abandoned agricultural lands increase in extent. We assess how tree species richness and composition recover during secondary succession across gradients in environmental conditions and anthropogenic disturbance in an unprecedented multisite analysis for the Neotropics. Secondary forests recover remarkably fast in species richness but slowly in species composition. Secondary forests take a median time of five decades to recover the species richness of old-growth forest (80% recovery after 20 years) based on rarefaction analysis. Full recovery of species composition takes centuries (only 34% recovery after 20 years). A dual strategy that maintains both old-growth forests and species-rich secondary forests is therefore crucial for biodiversity conservation in human-modified tropical landscapes.

Comment on "The global tree restoration potential".

Bastin et al's estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration.

Correction: Effects of a Possible Pollinator Crisis on Food Crop Production in Brazil.

[This corrects the article DOI: 10.1371/journal.pone.0167292.].

Legume abundance along successional and rainfall gradients in Neotropical forests.

The nutrient demands of regrowing tropical forests are partly satisfied by nitrogen-fixing legume trees, but our understanding of the abundance of those species is biased towards wet tropical regions. Here we show how the abundance of Leguminosae is affected by both recovery from disturbance and large-scale rainfall gradients through a synthesis of forest inventory plots from a network of 42 Neotropical forest chronosequences. During the first three decades of natural forest regeneration, legume basal area is twice as high in dry compared with wet secondary forests. The tremendous ecological success of legumes in recently disturbed, water-limited forests is likely to be related to both their reduced leaflet size and ability to fix N2, which together enhance legume drought tolerance and water-use efficiency. Earth system models should incorporate these large-scale successional and climatic patterns of legume dominance to provide more accurate estimates of the maximum potential for natural nitrogen fixation across tropical forests.

Galling Insects of the Brazilian Páramos: Species Richness and Composition Along High-Altitude Grasslands.

In this work, we investigated the factors that determine the distribution of galling insects in high-altitude grasslands, locally called 'campos de altitude' of Mantiqueira Range and tested whether 1) richness of galling insects decreases with altitude, 2) galling insect richness increases with plant richness, 3) variation in galling insect diversity is predominantly a consequence of its ß component, and 4) turnover is the main mechanism driving the beta diversity of both galling insects and plants. Galling insect richness did not exhibit a negative relationship with altitude, but it did increase with plant richness. The additive partition of regional richness (γ) into its local and beta components showed that local diversity (α) of galling insects and plants was relatively low in relation to regional diversity; the ß component incorporated most of the regional diversity. This pattern was also found in the multiscale analysis of the additive partition for galling insects and plants. The beta diversity of galling insects and plants was driven predominantly by the process of turnover and minimally by nesting. The results reported here point out that the spatial distribution of galling insects is best explained by historical factors, such as the distribution of genera and species of key host plants, as well as their relation to habitat, than ecological effects such as hygrothermal stress - here represented by altitude.

Mechanisms Driving Galling Success in a Fragmented Landscape: Synergy of Habitat and Top-Down Factors along Temperate Forest Edges.

Edge effects play key roles in the anthropogenic transformation of forested ecosystems and their biota, and are therefore a prime field of contemporary fragmentation research. We present the first empirical study to address edge effects on the population level of a widespread galling herbivore in a temperate deciduous forest. By analyzing edge effects on abundance and trophic interactions of beech gall midge (Mikiola fagi Htg.), we found 30% higher gall abundance in the edge habitat as well as lower mortality rates due to decreased top-down control, especially by parasitoids. Two GLM models with similar explanatory power (58%) identified habitat specific traits (such as canopy closure and altitude) and parasitism as the best predictors of gall abundance. Further analyses revealed a crucial influence of light exposure (46%) on top-down control by the parasitoid complex. Guided by a conceptual framework synthesizing the key factors driving gall density, we conclude that forest edge proliferation of M. fagi is due to a complex interplay of abiotic changes and trophic control mechanisms. Most prominently, it is caused by the microclimatic regime in forest edges, acting alone or in synergistic concert with top-down pressure by parasitoids. Contrary to the prevailing notion that specialists are edge-sensitive, this turns M. fagi into a winner species in fragmented temperate beech forests. In view of the increasing proportion of edge habitats and the documented benefits from edge microclimate, we call for investigations exploring the pest status of this galling insect and the modulators of its biological control.

Dung Beetles along a Tropical Altitudinal Gradient: Environmental Filtering on Taxonomic and Functional Diversity.

Mountains provide an interesting context in which to study the many facets of biodiversity in response to macroclimate, since environmental conditions change rapidly due to elevation. Although the decrease in biodiversity with increasing elevation is generally accepted, our understanding of the variation of functional diversity along altitudinal gradients is still poorly known. The partitioning of diversity into spatial components can help to understand the processes that influence the distribution of species, and these studies are urgently needed in face of the increasing threats to mountain environments throughout the world. We describe the distribution of dung beetle diversity along an altitudinal gradient on a tropical mountain in southeastern Brazil, including the spatial partitioning of taxonomic and functional diversities. The altitudinal gradient ranged from 800 up to 1400 m a.s.l. and we collected dung beetles at every 100 m of altitude. We used the Rao Index to calculate γ, α and ß diversity for taxonomic and functional diversity of dung beetles. Climatic, soil and vegetation variables were used to explain variation in community attributes along the altitudinal gradient. Dung beetle richness declined with altitude and was related to climatic and vegetation variables, but functional diversity did not follow the same pattern. Over 50% of γ taxonomic diversity was caused by among altitudes diversity (ß), while almost 100% of functional diversity was due to the α component. Contrasting ß taxonomic with ß functional diversity, we suggest that there is ecological redundancy among communities and that the environment is filtering species in terms of the Grinnellian niche, rather than the Eltonian niche. ß taxonomic diversity is caused mainly by the turnover component, reinforcing the hypothesis of environmental filtering. Global warming may have strong effects on mountain communities due to upslope range shifts and extinctions, and these events will lead to an even larger than previously expected loss of diversity as dung beetles γ taxonomic diversity is caused mainly by the ß component.

Effects of a Possible Pollinator Crisis on Food Crop Production in Brazil.

Animal pollinators contribute to human food production and security thereby ensuring an important component of human well-being. The recent decline of these agents in Europe and North America has aroused the concern of a potential global pollinator crisis. In order to prioritize efforts for pollinator conservation, we evaluated the extent to which food production depends on animal pollinators in Brazil-one of the world's agriculture leaders-by comparing cultivated area, produced volume and yield value of major food crops that are pollinator dependent with those that are pollinator non-dependent. In addition, we valued the ecosystem service of pollination based on the degree of pollinator dependence of each crop and the consequence of a decline in food production to the Brazilian Gross Domestic Product and Brazilian food security. A total of 68% of the 53 major food crops in Brazil depend to some degree on animals for pollination. Pollinator non-dependent crops produce a greater volume of food, mainly because of the high production of sugarcane, but the cultivated area and monetary value of pollinator dependent crops are higher (59% of total cultivated area and 68% of monetary value). The loss of pollination services for 29 of the major food crops would reduce production by 16.55-51 million tons, which would amount to 4.86-14.56 billion dollars/year, and reduce the agricultural contribution to the Brazilian GDP by 6.46%- 19.36%. These impacts would be largely absorbed by family farmers, which represent 74.4% of the agricultural labor force in Brazil. The main effects of a pollinator crisis in Brazil would be felt by the poorer and more rural classes due to their lower income and direct or exclusive dependence on this ecosystem service.

Pollinator preferences for Nicotiana alata, N. forgetiana, and their F1 hybrids.

The role of pollinators in plant speciation and maintenance of species boundaries is dubious because most plant species are visited by several types of pollinators, and most pollinator species visit several species of plants. We investigated pollinator preferences and their efficacy as ethological isolation mechanisms between two interfertile species, Nicotiana alata and N. forgetiana and their F1 hybrids. Hawkmoths pollinate N. alata, while primarily hummingbirds and occasionally small hawkmoths visit N. forgetiana. F1 hybrids are easily produced in the greenhouse and although the species grow in similar habitats, hybrids have not been found in nature. In Rio Grande do Sul, Brazil, near where both species are found, experimental plots were studied containing both species, and both species plus F1 hybrids. In the mixed-species plots, hawkmoths showed a strong preference for N. alata. Hummingbirds were less common and only visited N. forgetiana. Hybrid seed was produced but plants made significantly fewer hybrid offspring than predicted by the frequency of interspecific pollinator movements. Nicotiana forgetiana was the seed parent of 97% of the F1 offspring, suggesting an asymmetry in pollen delivery or postpollination processes. In plots containing F1 hybrids plus both parental species, hawkmoths preferred N. alata and undervisited the other two phenotypes, except that in the third plot they visited hybrids in proportion to the hybrid frequency. Hummingbirds strongly preferred N. forgetiana in all plots but also visited F1 hybrids in proportion to their frequency in the third plot. Overall, F1 hybrids were well pollinated and were frequently visited immediately before or after one of the parental species. Thus hybrids could facilitate gene flow between the parental species. We conclude that pollinator discrimination among species is strong but is an imperfect isolation mechanism, especially if hybrids are present.