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.

Hybrid zone of a tree in a Cerrado/Atlantic Forest ecotone as a hotspot of genetic diversity and conservation.

The Cerrado, the largest Neotropical savanna, and the Brazilian Atlantic Forest form large ecotonal areas where savanna and forest habitats occupy adjacent patches with closely related species occurring side by side, providing opportunities for hybridization. Here, we investigated the evolutionary divergence between the savanna and forest ecotypes of the widely distributed tree Plathymenia reticulata (n = 233 individuals). Genetic structure analysis of P. reticulata was congruent with the recognition of two ecotypes, whose divergence captured the largest proportion of genetic variance in the data (F CT = 0.222 and F ST = 0.307). The ecotonal areas between the Cerrado and the Atlantic Forest constitute a hybrid zone in which a diversity of hybrid classes was observed, most of them corresponding to second-generation hybrids (F2) or backcrosses. Gene flow occurred mainly toward the forest ecotype. The genetic structure was congruent with isolation by environment, and environmental correlates of divergence were identified. The observed pattern of high genetic divergence between ecotypes may reflect an incipient speciation process in P. reticulata. The low genetic diversity of the P. reticulata forest ecotype indicate that it is threatened in areas with high habitat loss on Atlantic Forest. In addition, the high divergence from the savanna ecotype suggests it should be treated as a different unit of management. The high genetic diversity found in the ecotonal hybrid zone supports the view of ecotones as important areas for the origin and conservation of biodiversity in the Neotropics.

Photosynthetic heat tolerance in plants with different foliar water -uptake strategies.

PREMISE: The distribution and even the survival of plant species are influenced by temperature. In an old climatically buffered infertile landscape (OCBIL) in Brazil, we previously characterized different strategies for foliar water uptake (FWU). It is possible that photosystem II tolerance to heat and excessive light intensity varies among species with different FWU capacities. METHODS: The relationship between FWU, photoinhibition, and thermotolerance was investigated in seven species from this ecosystem. RESULTS: The species with slow water absorption and high water absorption are those that presented less photoinhibition. Contrastingly, the species that have fast and low water absorption presented greater thermotolerance when their leaves are totally hydrated. However, when there is greater leaf dehydration, the most thermotolerant species were those with slow but high water absorption. CONCLUSIONS: Foliar water uptake is an important trait for plants to tolerate excessive light intensity and higher temperatures. Plants in this OCBIL may be differentially affected by future global warming, and the best strategy to deal with this expected climate change is with slow and high absorption of water.

Foliar water-uptake strategies are related to leaf water status and gas exchange in plants from a ferruginous rupestrian field.

PREMISE: Fog is a frequent event in Brazilian rupestrian field and plays an important role in the physiology of several plant species. Foliar water uptake (FWU) of fog may be fast or slow depending on the species. However, fog water may negatively affect CO2 assimilation. Thus, the interference in the water and carbon balance as a result of different strategies of FWU was evaluated to verify whether fog may mitigate possible water deficit in leaves. METHODS: Four plant species with different FWU strategies were studied in a ferruginous rupestrian field with frequent fog. Gas exchange and water potential were measured before dawn and at midday during the dry and rainy seasons, separating foggy from non-foggy days during the dry season. RESULTS: The FWU speed negatively influences CO2 assimilation in the dry season, possibly because of its negative relationship with stomatal conductance, since reduced stomatal aperture impairs carbon entrance. Fog presence increased leaf water potential both in early morning and midday during the dry season. However, during the rainy season, the values of leaf water potential were lower at midday, than during the dry season with fog at midday, which favors leaf gas exchanges. CONCLUSIONS: FWU interferes negatively, but briefly with CO2 assimilation. Nevertheless, FWU prevents water loss through transpiration and increases the water status of plants in the dry season. That is, FWU results in a compensation between CO2 assimilation and foliar hydration, which, in fact, is beneficial to the plants of this ecosystem.

Climatic factors shaping intraspecific leaf trait variation of a neotropical tree along a rainfall gradient.

Intraspecific trait variation has been singled out as an important mechanism by which individuals can cope with environmental variations and avoid local extinctions. Here we evaluate variation in metamer traits (i.e., traits associated with internodes, petioles and their corresponding leaves) and parameters of chlorophyll fluorescence within and among populations of a neotropical tree, Copaifera langsdorffii. We also evaluated phenotypic plasticity in natural settings comparing traits between shade and sun-exposed metamers. We selected six populations along a climatic gradient ranging from semi-arid to humid and representing three different biomes (Caatinga, Cerrado, and Atlantic Forest). Local climatic conditions significantly affected the morphological and physiological traits of populations. Trait variation among populations was explained mainly by aridity index and evapotranspiration. Individuals from drier regions had lower specific leaf area (SLA), lower investment in leaf area per total dry mass of metamer (LARm), lower specific petiole length (SPL) and lower potential quantum yield (Fv/Fm, only for sun-exposed metamers). Populations from locations with greater environmental heterogeneity (interannual variation) had greater plasticity in response to light for Fv/Fm and electron transport rate (ETR) and morphological traits related to the hydraulic and biomechanical aspects of the leaves (petiole length, internode length and SPL). High intraspecific variation in metamer traits in C. langsdorffii coupled with its ability to modify these traits in response to different climate conditions can explain the success of the species over a range of different habitats and represent important factors for the persistence of this species in the face of climate change.

Genetic and Historical Colonization Analyses of an Endemic Savanna Tree, Qualea grandiflora, Reveal Ancient Connections Between Amazonian Savannas and Cerrado Core.

The evolutionary processes underlying the high diversity and endemism in the Cerrado, the most extensive Neotropical savanna, remain unclear, including the factors promoting the presence and evolution of savanna enclaves in the Amazon forest. In this study, we investigated the effects of past climate changes on genetic diversity, dynamics of species range and the historical connections between the savanna enclaves and Cerrado core for Qualea grandiflora, a tree species widely distributed in the biome. Totally, 40 populations distributed in the Cerrado core and Amazon savannas were analyzed using chloroplast and nuclear DNA sequences. We used phylogeographic, coalescent and ecological niche modeling approaches. Genetic data revealed a phylogeographic structure shaped by Pleistocene climatic oscillations. An eastern-western split in the Cerrado core was observed. The central portion of the Cerrado core harbored most of the sampled diversity for cpDNA. Ecological niche models predicted the presence of a large historical refuge in this region and multiple small refuges in peripheral areas. Relaxed Random Walk (RRW) models indicated the ancestral population in the north-western border of the central portion of the Cerrado core and cyclical dynamics of colonization related to Pleistocene climatic oscillations. Central and western ancient connections between Cerrado core and Amazonian savannas were observed. No evidence of connections among the Amazonian savannas was detected. Our study highlights the importance of Pleistocene climatic oscillations for structuring the genetic diversity of Q. grandiflora and complex evolutionary history of ecotonal areas in the Cerrado. Our results do not support the recent replacement of a large area in the Amazon forest by savanna vegetation. The Amazonian savannas appear to be fragmented and isolated from each other, evolving independently a long ago.

Environmental drivers on leaf phenology of ironstone outcrops species under seasonal climate

ABSTRACT Banded iron formations (BIF) have a particular vegetation type and provide a good model system for testing theories related to leaf phenology, due to unique stressful environmental conditions. As a consequence of the stressful conditions of BIF environment, we hypothesize that most species would retain at least some significant canopy cover, even at the end of the dry season, independently of growth form - trees, shrubs, and sub-shrubs. Considering the strong seasonality, we also hypothesize that photoperiod and rainfall act as triggers for leaf fall and leaf flushing in these environments. The majority of the fifteen studied species had a semi-deciduous behavior and shed their leaves mainly during the dry season, with a recovery at the end of this season. In general, leaf flushing increased around the spring equinox (end of the dry season and start of the rainy season). A trade-off between leaf loss and leaf maintenance is expected in a community in which most plants are naturally selected to be semi-deciduous. Our results suggest photoperiod as a dominant factor in predicting leaf phenology.

Environmental drivers on leaf phenology of ironstone outcrops species under seasonal climate.

Banded iron formations (BIF) have a particular vegetation type and provide a good model system for testing theories related to leaf phenology, due to unique stressful environmental conditions. As a consequence of the stressful conditions of BIF environment, we hypothesize that most species would retain at least some significant canopy cover, even at the end of the dry season, independently of growth form - trees, shrubs, and sub-shrubs. Considering the strong seasonality, we also hypothesize that photoperiod and rainfall act as triggers for leaf fall and leaf flushing in these environments. The majority of the fifteen studied species had a semi-deciduous behavior and shed their leaves mainly during the dry season, with a recovery at the end of this season. In general, leaf flushing increased around the spring equinox (end of the dry season and start of the rainy season). A trade-off between leaf loss and leaf maintenance is expected in a community in which most plants are naturally selected to be semi-deciduous. Our results suggest photoperiod as a dominant factor in predicting leaf phenology.

Climatic drivers of leaf traits and genetic divergence in the tree Annona crassiflora: a broad spatial survey in the Brazilian savannas.

The Cerrado is the largest South American savanna and encompasses substantial species diversity and environmental variation. Nevertheless, little is known regarding the influence of the environment on population divergence of Cerrado species. Here, we searched for climatic drivers of genetic (nuclear microsatellites) and leaf trait divergence in Annona crassiflora, a widespread tree in the Cerrado. The sampling encompassed all phytogeographic provinces of the continuous area of the Cerrado and included 397 individuals belonging to 21 populations. Populations showed substantial genetic and leaf trait divergence across the species' range. Our data revealed three spatially defined genetic groups (eastern, western and southern) and two morphologically distinct groups (eastern and western only). The east-west split in both the morphological and genetic data closely mirrors previously described phylogeographic patterns of Cerrado species. Generalized linear mixed effects models and multiple regression analyses revealed several climatic factors associated with both genetic and leaf trait divergence among populations of A. crassiflora. Isolation by environment (IBE) was mainly due to temperature seasonality and precipitation of the warmest quarter. Populations that experienced lower precipitation summers and hotter winters had heavier leaves and lower specific leaf area. The southwestern area of the Cerrado had the highest genetic diversity of A. crassiflora, suggesting that this region may have been climatically stable. Overall, we demonstrate that a combination of current climate and past climatic changes have shaped the population divergence and spatial structure of A. crassiflora. However, the genetic structure of A. crassiflora reflects the biogeographic history of the species more strongly than leaf traits, which are more related to current climate.

Organ-coordinated response of early post-germination mahogany seedlings to drought.

Water deficit tolerance during post-germination stages is critical for seedling recruitment. In this work, we studied the effect of water deficit on morphological and biochemical responses in different organs of newly germinated mahogany (Swietenia macrophylla King) seedlings, a woody species that occurs in the Amazon rainforest. The root : shoot ratio increased under water deficit. The leaf number and water potential were not altered, although reductions in leaf area and stomatal conductance were observed. Osmotic potential became more negative in leaves of seedlings under severe stress. Water deficit increased fructose, glucose, sucrose and myo-inositol levels in leaves. Stems accumulated fructose, glucose and l-proline. Nitric oxide (NO) levels increased in the vascular cylinder of roots under severe stress while superoxide anion levels decreased due to augmented superoxide dismutase activity in this organ. Water deficit induced glutathione reductase activity in both roots and stems. Upon moderate or severe stress, catalase activity decreased in leaves and remained unaffected in the other seedling organs, allowing for an increase of hydrogen peroxide (H2O2) levels in leaves. Overall, the increase of signaling molecules in distinct organs-NO in roots, l-proline in stems and H2O2 and myo-inositol in leaves-contributed to the response of mahogany seedlings to water deficit by triggering biochemical processes that resulted in the attenuation of oxidative stress and the establishment of osmotic adjustment. Therefore, this body of evidence reveals that the development of newly germinated mahogany seedlings may occur in both natural habitats and crop fields even when water availability is greatly limited.

Testing arbitrary classes of light in a physiognomically heterogeneous area of "campo rupestre" vegetation.

The Brazilian "campo rupestre" vegetation shows high light heterogeneity, characteristic which is often disregarded in ecophysiological studies due the difficult of obtaining accurate measurements in a complex environment. The aim of the present study it was to describe how much heterogeneous is the light distribution in this environment, as well as tried to define classes in relation to the photosynthetically active radiation (PAR) for 30 rupicolous individuals of Bulbophyllum involutum. The individuals were marked and previously classified in light classes visually predetermined ("full sun", "partially shaded" and "shaded"). Once defined these classes, principal component analysis, cluster and dispersal analyses using direct and indirect PAR measures were performed to characterize the environment as well as to validate the inclusion of the individuals in each class. The use of multivariate analysis increased the accuracy of predetermined classes of light, and some individuals from "shaded" and "partially shaded" classes needed to be regrouped. After individuals' reclassification, amplitude of PAR variance was reduced in each class, enhancing homogeneity. Ours results show the possibility to evaluate light environment conditions more precisely in "campo rupestre" vegetation using multivariate analysis, allowing the increase in accuracy of this parameter for ecophysiological researches.

Development of microsatellite markers for Dimorphandra mollis (Leguminosae), a widespread tree from the Brazilian cerrado.

PREMISE OF THE STUDY: Microsatellite markers were developed for Dimorphandra mollis (Leguminosae), a widespread tree in the Brazilian cerrado (a savanna-like vegetation). METHODS AND RESULTS: Microsatellite markers were developed from an enriched library. The analyses of polymorphism were based on 56 individuals from three populations. Nine microsatellite loci were polymorphic, with the number of alleles per locus ranging from three to 10 across populations. The observed and expected heterozygosities per locus and population ranged from 0.062 to 0.850 and from 0.062 to 0.832, respectively. CONCLUSIONS: These microsatellites provide an efficient tool for population genetics studies and will be used to assess the genetic diversity and spatial genetic structure of D. mollis.

Plasticity and stress tolerance override local adaptation in the responses of Mediterranean holm oak seedlings to drought and cold.

Plant populations of widely distributed species experience a broad range of environmental conditions that can be faced by phenotypic plasticity or ecotypic differentiation and local adaptation. The strategy chosen will determine a population's ability to respond to climate change. To explore this, we grew Quercus ilex (L.) seedlings from acorns collected at six selected populations from climatically contrasting localities and evaluated their response to drought and late season cold events. Maximum photosynthetic rate (A(max)), instantaneous water use efficiency (iWUE), and thermal tolerance to freeze and heat (estimated from chlorophyll fluorescence versus temperature curves) were measured in 5-month-old seedlings in control (no stress), drought (water-stressed), and cold (low suboptimal temperature) conditions. The observed responses were similar for the six populations: drought decreased A(max) and increased iWUE, and cold reduced A(max) and iWUE. All the seedlings maintained photosynthetic activity under adverse conditions (drought and cold), and rapidly increased their iWUE by closing stomata when exposed to drought. Heat and freeze tolerances were similarly high for seedlings from all the populations, and they were significantly increased by drought and cold, respectively; and were positively related to each other. Differences in seedling performance across populations were primarily induced by maternal effects mediated by seed size and to a lesser extent by idiosyncratic physiologic responses to drought and low temperatures. Tolerance to multiple stresses together with the capacity to physiologically acclimate to heat waves and cold snaps may allow Q. ilex to cope with the increasingly stressful conditions imposed by climate change. Lack of evidence of physiologic seedling adaptation to local climate may reflect opposing selection pressures to complex, multidimensional environmental conditions operating within the distribution range of this species.