Effects of nitrogen and phosphorus availability on the early growth of two congeneric pairs of savanna and forest species / Efeitos da disponibilidade de nitrogênio e fósforo no crescimento inicial de dois pares congenéricos de espécies de savana e floresta

In the tropical region, savannas and seasonal forests, both highly diverse biomes, occur side by side, under the same climate. If so, that mosaic cannot be explained solely by climatic variables, but also by fire, water availability and soil status. Nutrient availability in the soil, especially nitrogen and phosphorus, has been postulated to explain the abrupt transitions between savannas and seasonal forests in tropical regions. Plants from these two biomes may present different nutritional strategies to cope with nitrogen and phosphorus limitation. We used two congeneric pairs of trees — each pair with a species from the savanna and another from the neighboring seasonal forest — to test whether savanna and forest species presented different nutritional strategies during their early development. We cultivated 56 individuals from each of these species in a hydroponics system with four treatments: (1) complete Hoagland solution, (2) Hoagland solution without nitrogen, (3) Hoagland solution without phosphorus, and (4) Hoagland solution without nitrogen and phosphorus. After 45 days, we harvested the plants and measured total biomass, root to shoot ratio, height, leaf area, and specific leaf area. Overall, savanna species were lighter, shorter, with smaller leaves, higher specific leaf areas, and higher root to shoot ratios when compared to the forest species. Nitrogen increased the performance of species from both biomes. Phosphorus improved the performance of the forest species and caused toxicity symptoms in the savanna species. Hence, savanna and forest species presented different demands and were partially distinct already as seedlings concerning their nutritional strategies.

Em regiões tropicais, savanas e florestas estacionais, biomas altamente diversos, ocorrem lado a lado, sob o mesmo clima. Sendo assim, esse mosaico não pode ser explicado somente por variáveis climáticas, devendo ser considerada a frequência e intensidade de incêndios, disponibilidade de água e status do solo. A disponibilidade de nutrientes no solo, especialmente nitrogênio e fósforo, tem sido postulada para explicar as transições abruptas entre savanas e florestas estacionais nos trópicos. Espécies vegetais desses dois biomas podem apresentar estratégias nutricionais diferentes para lidar com a limitação tanto de nitrogênio como de fósforo. Utilizamos dois pares de árvores congenéricas — cada par com uma espécie típica de savana e outra de floresta estacional vizinha — para testar se as espécies da savana e da floresta apresentaram estratégias nutricionais diferentes durante seu desenvolvimento inicial. Cultivamos 56 indivíduos de cada uma dessas espécies em um sistema hidropônico com quatro tratamentos: (1) solução Hoagland completa, (2) solução Hoagland sem nitrogênio, (3) solução Hoagland sem fósforo e (4) solução Hoagland sem nitrogênio e fósforo. Após 45 dias, colhemos as plantas e medimos a biomassa total, a relação raiz / parte aérea, altura, área foliar e área foliar específica. No geral, as espécies savânicas foram mais leves, menores em altura, área foliar e área foliar específica e apresentaram maiores razões entre biomassa radicular por biomassa aérea quando comparadas às espécies florestais. A oferta de nitrogênio aumentou o desempenho das espécies de ambos biomas. O fósforo melhorou o desempenho das espécies florestais e causou sintomas de toxicidade nas espécies savânicas. Concluímos que, já como mudas, espécies congenéricas de savana e floresta apresentaram demandas distintas e foram parcialmente diferentes em relação a suas estratégias nutricionais.

Effects of nitrogen and phosphorus availability on the early growth of two congeneric pairs of savanna and forest species

Abstract In the tropical region, savannas and seasonal forests, both highly diverse biomes, occur side by side, under the same climate. If so, that mosaic cannot be explained solely by climatic variables, but also by fire, water availability and soil status. Nutrient availability in the soil, especially nitrogen and phosphorus, has been postulated to explain the abrupt transitions between savannas and seasonal forests in tropical regions. Plants from these two biomes may present different nutritional strategies to cope with nitrogen and phosphorus limitation. We used two congeneric pairs of trees each pair with a species from the savanna and another from the neighboring seasonal forest to test whether savanna and forest species presented different nutritional strategies during their early development. We cultivated 56 individuals from each of these species in a hydroponics system with four treatments: (1) complete Hoagland solution, (2) Hoagland solution without nitrogen, (3) Hoagland solution without phosphorus, and (4) Hoagland solution without nitrogen and phosphorus. After 45 days, we harvested the plants and measured total biomass, root to shoot ratio, height, leaf area, and specific leaf area. Overall, savanna species were lighter, shorter, with smaller leaves, higher specific leaf areas, and higher root to shoot ratios when compared to the forest species. Nitrogen increased the performance of species from both biomes. Phosphorus improved the performance of the forest species and caused toxicity symptoms in the savanna species. Hence, savanna and forest species presented different demands and were partially distinct already as seedlings concerning their nutritional strategies.

Resumo Em regiões tropicais, savanas e florestas estacionais, biomas altamente diversos, ocorrem lado a lado, sob o mesmo clima. Sendo assim, esse mosaico não pode ser explicado somente por variáveis climáticas, devendo ser considerada a frequência e intensidade de incêndios, disponibilidade de água e status do solo. A disponibilidade de nutrientes no solo, especialmente nitrogênio e fósforo, tem sido postulada para explicar as transições abruptas entre savanas e florestas estacionais nos trópicos. Espécies vegetais desses dois biomas podem apresentar estratégias nutricionais diferentes para lidar com a limitação tanto de nitrogênio como de fósforo. Utilizamos dois pares de árvores congenéricas cada par com uma espécie típica de savana e outra de floresta estacional vizinha para testar se as espécies da savana e da floresta apresentaram estratégias nutricionais diferentes durante seu desenvolvimento inicial. Cultivamos 56 indivíduos de cada uma dessas espécies em um sistema hidropônico com quatro tratamentos: (1) solução Hoagland completa, (2) solução Hoagland sem nitrogênio, (3) solução Hoagland sem fósforo e (4) solução Hoagland sem nitrogênio e fósforo. Após 45 dias, colhemos as plantas e medimos a biomassa total, a relação raiz / parte aérea, altura, área foliar e área foliar específica. No geral, as espécies savânicas foram mais leves, menores em altura, área foliar e área foliar específica e apresentaram maiores razões entre biomassa radicular por biomassa aérea quando comparadas às espécies florestais. A oferta de nitrogênio aumentou o desempenho das espécies de ambos biomas. O fósforo melhorou o desempenho das espécies florestais e causou sintomas de toxicidade nas espécies savânicas. Concluímos que, já como mudas, espécies congenéricas de savana e floresta apresentaram demandas distintas e foram parcialmente diferentes em relação a suas estratégias nutricionais.

Effects of nitrogen and phosphorus availability on the early growth of two congeneric pairs of savanna and forest species / Efeitos da disponibilidade de nitrogênio e fósforo no crescimento inicial de dois pares congenéricos de espécies de savana e floresta

In the tropical region, savannas and seasonal forests, both highly diverse biomes, occur side by side, under the same climate. If so, that mosaic cannot be explained solely by climatic variables, but also by fire, water availability and soil status. Nutrient availability in the soil, especially nitrogen and phosphorus, has been postulated to explain the abrupt transitions between savannas and seasonal forests in tropical regions. Plants from these two biomes may present different nutritional strategies to cope with nitrogen and phosphorus limitation. We used two congeneric pairs of trees - each pair with a species from the savanna and another from the neighboring seasonal forest - to test whether savanna and forest species presented different nutritional strategies during their early development. We cultivated 56 individuals from each of these species in a hydroponics system with four treatments: (1) complete Hoagland solution, (2) Hoagland solution without nitrogen, (3) Hoagland solution without phosphorus, and (4) Hoagland solution without nitrogen and phosphorus. After 45 days, we harvested the plants and measured total biomass, root to shoot ratio, height, leaf area, and specific leaf area. Overall, savanna species were lighter, shorter, with smaller leaves, higher specific leaf areas, and higher root to shoot ratios when compared to the forest species. Nitrogen increased the performance of species from both biomes. Phosphorus improved the performance of the forest species and caused toxicity symptoms in the savanna species. Hence, savanna and forest species presented different demands and were partially distinct already as seedlings concerning their nutritional strategies.

Em regiões tropicais, savanas e florestas estacionais, biomas altamente diversos, ocorrem lado a lado, sob o mesmo clima. Sendo assim, esse mosaico não pode ser explicado somente por variáveis climáticas, devendo ser considerada a frequência e intensidade de incêndios, disponibilidade de água e status do solo. A disponibilidade de nutrientes no solo, especialmente nitrogênio e fósforo, tem sido postulada para explicar as transições abruptas entre savanas e florestas estacionais nos trópicos. Espécies vegetais desses dois biomas podem apresentar estratégias nutricionais diferentes para lidar com a limitação tanto de nitrogênio como de fósforo. Utilizamos dois pares de árvores congenéricas - cada par com uma espécie típica de savana e outra de floresta estacional vizinha - para testar se as espécies da savana e da floresta apresentaram estratégias nutricionais diferentes durante seu desenvolvimento inicial. Cultivamos 56 indivíduos de cada uma dessas espécies em um sistema hidropônico com quatro tratamentos: (1) solução Hoagland completa, (2) solução Hoagland sem nitrogênio, (3) solução Hoagland sem fósforo e (4) solução Hoagland sem nitrogênio e fósforo. Após 45 dias, colhemos as plantas e medimos a biomassa total, a relação raiz / parte aérea, altura, área foliar e área foliar específica. No geral, as espécies savânicas foram mais leves, menores em altura, área foliar e área foliar específica e apresentaram maiores razões entre biomassa radicular por biomassa aérea quando comparadas às espécies florestais. A oferta de nitrogênio aumentou o desempenho das espécies de ambos biomas. O fósforo melhorou o desempenho das espécies florestais e causou sintomas de toxicidade nas espécies savânicas. Concluímos que, já como mudas, espécies congenéricas de savana e floresta apresentaram demandas distintas e foram parcialmente diferentes em relação a suas estratégias nutricionais.

Effects of nitrogen and phosphorus availability on the early growth of two congeneric pairs of savanna and forest species.

In the tropical region, savannas and seasonal forests, both highly diverse biomes, occur side by side, under the same climate. If so, that mosaic cannot be explained solely by climatic variables, but also by fire, water availability and soil status. Nutrient availability in the soil, especially nitrogen and phosphorus, has been postulated to explain the abrupt transitions between savannas and seasonal forests in tropical regions. Plants from these two biomes may present different nutritional strategies to cope with nitrogen and phosphorus limitation. We used two congeneric pairs of trees - each pair with a species from the savanna and another from the neighboring seasonal forest - to test whether savanna and forest species presented different nutritional strategies during their early development. We cultivated 56 individuals from each of these species in a hydroponics system with four treatments: (1) complete Hoagland solution, (2) Hoagland solution without nitrogen, (3) Hoagland solution without phosphorus, and (4) Hoagland solution without nitrogen and phosphorus. After 45 days, we harvested the plants and measured total biomass, root to shoot ratio, height, leaf area, and specific leaf area. Overall, savanna species were lighter, shorter, with smaller leaves, higher specific leaf areas, and higher root to shoot ratios when compared to the forest species. Nitrogen increased the performance of species from both biomes. Phosphorus improved the performance of the forest species and caused toxicity symptoms in the savanna species. Hence, savanna and forest species presented different demands and were partially distinct already as seedlings concerning their nutritional strategies.

Functional diversity, soil features and community functioning: a test in a cerrado site.

Community functioning may be affected by functional diversity, which measures the extent of complementarity in resource use. We tested whether there was a relationship between functional diversity of woody species and community functioning on a fine scale, using FD as a measure of functional diversity and litter decomposition rate as a surrogate for community functioning. We measured eight functional traits from a woodland cerrado community in southeastern Brazil. Then, we tested the correlation between FD and the decomposition rate taking into account differences in soil features and between decomposition rate and each trait separately. The decomposition rate was related to the aluminium and phosphorus concentration in soil, but not to FD, pointing out that functional diversity was not a good predictor of community functioning. There was a non-significant relationship between FD and the decomposition rate even when we considered each trait separately. Most studies in the relationships between biodiversity and community functioning on fine scales were carried out by experimental manipulation of diversity and in temperate regions. We carried out this fine scale study as a mensurative experiment and in a tropical savanna. Our findings indicated that the relationship between biodiversity and community functioning is not as straightforward as usually assumed.

Towards testing the "honeycomb rippling model" in cerrado.

Savannas are tropical formations in which trees and grasses coexist. According to the "honeycomb rippling model", inter-tree competition leads to an effect of trees growing and dying due to competition, which, at fine spatial scale, would resemble honeycomb rippling. The model predicts that the taller the trees, the higher the inter-tree distances and the evenness of inter-tree distances. The model had been corroborated in arid savannas, in what appears to be caused by uneven distribution of rains, but had not yet been tested in seasonal savannas, such as the cerrado, which could be caused by the irregular occurrence of fire.A basic assumption of the model is that strong inter-tree competition affects growth (estimated by height) and mortality (estimated by inter-tree distances). As a first step towards testing this model in the cerrado, we tested this assumption in a single cerrado patch in southeastern Brazil. We placed 80 quadrats, each one with 25 m², in which we sampled all shrubs and trees. For each individual, we measured its height and the distance to its nearest neighbour--the inter-tree distance. We did not find correlations between tree height and both inter-tree distances and evenness of inter-tree distances, refuting the honeycomb rippling model. Inter-tree distances were spatially autocorrelated, but height was not. According to our results, the basic assumption of the model does not apply to seasonal savannas. If, in arid savannas, rainfall events are rare and unpredictable, in seasonal savannas, the rainy season is well-defined and rainfall is considerable. We found horizontal structuring in the community, which may be due to soil nutrient heterogeneity. The absence of vertical structuring suggests that competition for light among adult trees is not as important as competition for nutrients in the soil. We tested the basic assumption of the model in a single patch and at a single moment. To test the model effectively, we suggest this assumption to be tested in many patches over time.

The 'Herbivory Uncertainty Principle': application in a cerrado site.

Researchers may alter the ecology of their studied organisms, even carrying out apparently beneficial activities, as in herbivory studies, when they may alter herbivory damage. We tested whether visit frequency altered herbivory damage, as predicted by the 'Herbivory Uncertainty Principle'. In a cerrado site, we established 80 quadrats, in which we sampled all woody individuals. We used four visit frequencies (high, medium, low, and control), quantifying, at the end of three months, herbivory damage for each species in each treatment. We did not corroborate the 'Herbivory Uncertainty Principle', since visiting frequency did not alter herbivory damage, at least when the whole plant community was taken into account. However, when we analysed each species separately, four out of 11 species presented significant differences in herbivory damage, suggesting that the researcher is not independent of its measurements. The principle could be tested in other ecological studies in which it may occur, such as those on animal behaviour, human ecology, population dynamics, and conservation.

Phylogenetic overdispersion of plant species in southern Brazilian savannas.

Ecological communities are the result of not only present ecological processes, such as competition among species and environmental filtering, but also past and continuing evolutionary processes. Based on these assumptions, we may infer mechanisms of contemporary coexistence from the phylogenetic relationships of the species in a community. We studied the phylogenetic structure of plant communities in four cerrado sites, in southeastern Brazil. We calculated two raw phylogenetic distances among the species sampled. We estimated the phylogenetic structure by comparing the observed phylogenetic distances to the distribution of phylogenetic distances in null communities. We obtained null communities by randomizing the phylogenetic relationships of the regional pool of species. We found a phylogenetic overdispersion of the cerrado species. Phylogenetic overdispersion has several explanations, depending on the phylogenetic history of traits and contemporary ecological interactions. However, based on coexistence models between grasses and trees, density-dependent ecological forces, and the evolutionary history of the cerrado flora, we argue that the phylogenetic overdispersion of cerrado species is predominantly due to competitive interactions, herbivores and pathogen attacks, and ecological speciation. Future studies will need to include information on the phylogenetic history of plant traits.

Short-term community dynamics in seasonal and hyperseasonal cerrados.

In South America, the largest seasonal savanna region is the Brazilian cerrado. Our aim was to study temporal changes in some community descriptors, such as floristic composition, richness, species density, plant density, and cylindrical volume, in a seasonal cerrado, comparing it to a nearby hyperseasonal cerrado. In four different seasons, we placed randomly ten 1 m(2) quadrats in each vegetation form and sampled all the vascular plants. Seasonal changes in floristic composition, species density, and plant density were less pronounced in the seasonal than in the hyperseasonal cerrado. Floristic similarity between the vegetation forms was lower when the hyperseasonal cerrado was waterlogged. Richness and species density were higher in the seasonal cerrado, which reached its biomass peak at mid rainy season. The hyperseasonal cerrado, in turn, reached its biomass peak at early rainy season and, despite the waterlogging, maintained it until late rainy season. In the hyperseasonal cerrado, waterlogging acts as an environmental filter restricting the number of cerrado species able to withstand it. The seasonal cerrado community was more stable than the hyperseasonal one. Our results corroborated the idea that changes in the environmental filters will affect floristic composition and community structure in savannas.

Phylogenetic overdispersion of plant species in southern Brazilian savannas / Dispersão filogenética de espécies de plantas em savanas no Sudeste do Brasil

Ecological communities are the result of not only present ecological processes, such as competition among species and environmental filtering, but also past and continuing evolutionary processes. Based on these assumptions, we may infer mechanisms of contemporary coexistence from the phylogenetic relationships of the species in a community. We studied the phylogenetic structure of plant communities in four cerrado sites, in southeastern Brazil. We calculated two raw phylogenetic distances among the species sampled. We estimated the phylogenetic structure by comparing the observed phylogenetic distances to the distribution of phylogenetic distances in null communities. We obtained null communities by randomizing the phylogenetic relationships of the regional pool of species. We found a phylogenetic overdispersion of the cerrado species. Phylogenetic overdispersion has several explanations, depending on the phylogenetic history of traits and contemporary ecological interactions. However, based on coexistence models between grasses and trees, density-dependent ecological forces, and the evolutionary history of the cerrado flora, we argue that the phylogenetic overdispersion of cerrado species is predominantly due to competitive interactions, herbivores and pathogen attacks, and ecological speciation. Future studies will need to include information on the phylogenetic history of plant traits.

Comunidades ecológicas resultam não somente de processos ecológicos atuais, como a competição e os filtros ambientais, mas também de processos evolutivos passados e contínuos. Com base nessas premissas, podemos inferir mecanismos de coexistência contemporânea a partir das relações filogenéticas das espécies em uma comunidade. Estudamos a estrutura filogenética das comunidades de plantas de quatro áreas de cerrado, no Sudeste do Brasil. Calculamos duas medidas das distâncias filogenéticas das espécies amostradas. Estimamos a estrutura filogenética comparando suas distâncias observadas com a distribuição dessas distâncias em comunidades nulas. Obtivemos comunidades nulas aleatorizando as relações filogenéticas do banco regional de espécies. Encontramos uma dispersão filogenética de espécies de cerrado. Há várias explicações para essa dispersão, dependendo da história filogenética dos traços e das interações ecológicas contemporâneas. Entretanto, com base nos modelos de coexistência entre árvores e gramíneas, nas forças ecológicas dependentes da densidade e na história evolutiva da flora do cerrado, argumentamos que a dispersão filogenética das espécies do cerrado é predominantemente devida às interações competitivas, aos ataques de herbívoros e patógenos e à especiação ecológica. Estudos futuros precisarão incluir informações sobre a história filogenética dos traços das plantas.

Short-term community dynamics in seasonal and hyperseasonal cerrados / Seca e alagamento como determinantes da dinâmica da comunidade de cerrados estacional e hiperestacional

In South America, the largest seasonal savanna region is the Brazilian cerrado. Our aim was to study temporal changes in some community descriptors, such as floristic composition, richness, species density, plant density, and cylindrical volume, in a seasonal cerrado, comparing it to a nearby hyperseasonal cerrado. In four different seasons, we placed randomly ten 1 m² quadrats in each vegetation form and sampled all the vascular plants. Seasonal changes in floristic composition, species density, and plant density were less pronounced in the seasonal than in the hyperseasonal cerrado. Floristic similarity between the vegetation forms was lower when the hyperseasonal cerrado was waterlogged. Richness and species density were higher in the seasonal cerrado, which reached its biomass peak at mid rainy season. The hyperseasonal cerrado, in turn, reached its biomass peak at early rainy season and, despite the waterlogging, maintained it until late rainy season. In the hyperseasonal cerrado, waterlogging acts as an environmental filter restricting the number of cerrado species able to withstand it. The seasonal cerrado community was more stable than the hyperseasonal one. Our results corroborated the idea that changes in the environmental filters will affect floristic composition and community structure in savannas.

Na América do Sul, a maior região de savana é o cerrado brasileiro. Estudamos as mudanças na composição florística, riqueza, densidade de espécies, densidade de plantas e volume cilíndrico em um cerrado estacional comparando-o com um cerrado hiperestacional adjacente. Em quatro estações do ano, sorteamos dez parcelas de 1 m² em cada formação vegetal, nas quais amostramos todas as plantas vasculares. Mudanças estacionais na composição florística, densidade de espécies e densidade de plantas foram menos pronunciadas no cerrado estacional. A similaridade entre os cerrados foi menor quando o cerrado hiperestacional estava alagado. A riqueza e a densidade de espécies foram maiores no cerrado estacional, que atingiu o pico de biomassa no meio da estação chuvosa. O cerrado hiperestacional, por sua vez, atingiu o pico de biomassa no começo da estação chuvosa e, apesar do alagamento, o manteve até o final da estação chuvosa. No cerrado hiperestacional, o alagamento atua como um filtro ambiental restringindo o número de espécies de cerrado capazes de tolerá-lo. A comunidade do cerrado estacional foi mais estável que a do cerrado hiperestacional. Nossos resultados corroboraram a idéia de que mudanças de curto prazo nos filtros ambientais das savanas afetam a sua composição florística e estrutura.

Including intraspecific variability in functional diversity.

Linking species and ecosystems often relies on approaches that consider how the traits exhibited by species affect ecosystem processes. One method is to estimate functional diversity (FD) based on the dispersion of species in functional trait space. Individuals within a species also differ, however, and an unresolved challenge is how to include such intraspecific variability in a measure of functional diversity. Our solution is to extend an existing measure to variation among individuals within species. Here, simulations demonstrate how the new measure behaves relative to one that does not include individual variation. Individual-level FD was less well associated with species richness than species-level FD in a single trait dimension, because species differed in their intraspecific variation. However, in multiple trait dimensions, there was a strong association between individual- and species-level FD and richness, because many traits result in a tight relationship between functional diversity and species richness. The correlation between the two FD measures weakened as the amount of intraspecific variability increased. Analyzing natural plant communities we found no relationship between species richness and functional diversity. In these analyses, we did not have to specify the source of intraspecific variation. In fact, the variation was only among individuals. The measure can, however, include differences in the amount of intraspecific variation at different sites, as we demonstrate. Including intraspecific variation should allow a more complete understanding of the processes that link individuals and ecosystems and provide better predictions about the consequences of extinctions for ecosystem processes.

Seasonality of litterfall and leaf decomposition in a cerrado site.

We investigated annual litterfall and leaf decomposition rate in a cerrado site. We collected woody plant litter monthly from April 2001 to March 2002 and from July 2003 to June 2004. We placed systematically 13 litter traps (0.5 x 0.5 m) in a line, 10 m one from the other. We sorted litter into 'leaves', 'stems', 'reproductive structures', and 'miscellanea' fractions, oven-dried them at 80 degrees C until constant mass and weighed the dry material. To assess leaf decomposition rate, we packed leaves recently shed by plants in litter bags. We placed seven sets of nine litter bags in a line, 10 m one from the other, on the soil surface and collected nine bags each time after 1, 2, 3, 4, 6, 9, and 12 months. Total and leaf litter productions showed a seasonal pattern. Leaf litterfall was the phenological attribute that showed the strongest response to seasonality and drought. Decomposition was slower in the cerrado that we studied compared to a more closed cerrado physiognomy, reflecting their structural and environmental differences. Thus, decomposition rates seem to increase from open to closed cerrado physiognomies, probably related to an increase of humidity and nutrients in the soil.

A year in a Cerrado wet grassland: a non-seasonal island in a seasonal savanna environment.

In some Cerrado regions where the water table is superficial and soils are hydromorphic, we may find wet grasslands. We studied temporal changes in some community descriptors, such as species density, plant density, basal area, and cylindrical volume in a Cerrado wet grassland in four different seasons of the year. We also compared the species richness and composition of the wet grassland with a hyperseasonal cerrado, and a seasonal cerrado. We found significant differences among the seasons only for species density. Chao-Sørensen similarity values varied from 0.86 to 0.99 and, in the wet grassland, were not different among the seasons. On the contrary, similarity values between the wet grassland and hyperseasonal and seasonal cerrados were low. Species richness was lower in the wet grassland and higher in the seasonal cerrado. As long as savannas are highly dynamic on all temporal and spatial scales, the wet grassland stability, at least in a short-term view, introduces an important heterogeneity in regional scale. Wet grasslands are also important in the Cerrado domain to increase b-diversity, since they are floristically dissimilar with cerrado vegetation.

Can we predict dispersal guilds based on the leaf-height-seed scheme in a disjunct cerrado woodland?

Although there have been advances in methods for extracting information about dispersal processes, it is still very difficult to measure them. Predicting dispersal groups using single readily-measured traits would facilitate the emergence of instructive comparisons among ecological strategies of plants and offer a path towards improved synthesis across field experiments. The leaf-height-seed scheme consists of three functional traits: specific leaf area, plant canopy height, and seed mass. We tested, applying logistic regression analysis, whether these traits are potential predictors of dispersal guilds in a disjoint cerrado woodland site in southeastern Brazil. According to our results, none of the plant traits studied could predict dispersal guild; this means that abiotically and biotically dispersed species showed similar values of specific leaf area, height, and seed mass. The species of both guilds exhibited sclerophylly, probably a result of the typical soil nutrient deficiency of cerrado, which also may have placed constraints upon plant canopy height regardless of the dispersal mode. In the cerrado, some abiotically dispersed trees might present higher than expected seed mass as support to the investment in high root-to-shoot ratio at the seedling stage. Seeds of bird-dispersed species are limited in size and mass because of the small size of most frugivorous birds. Since soil nutrient quality might contribute to the similarity between the dispersal guilds regarding the three traits of the scheme, other plant traits (e.g., root depth distribution and nutrient uptake strategy) that detail the former should be considered in future predictive studies.

Seasonality of litterfall and leaf decomposition in a cerrado site / Estacionalidade da produção de serapilheira e decomposição foliar em um sítio de cerrado

We investigated annual litterfall and leaf decomposition rate in a cerrado site. We collected woody plant litter monthly from April 2001 to March 2002 and from July 2003 to June 2004. We placed systematically 13 litter traps (0.5 x 0.5 m) in a line, 10 m one from the other. We sorted litter into 'leaves', 'stems', 'reproductive structures', and 'miscellanea' fractions, oven-dried them at 80 °C until constant mass and weighed the dry material. To assess leaf decomposition rate, we packed leaves recently shed by plants in litter bags. We placed seven sets of nine litter bags in a line, 10 m one from the other, on the soil surface and collected nine bags each time after 1, 2, 3, 4, 6, 9, and 12 months. Total and leaf litter productions showed a seasonal pattern. Leaf litterfall was the phenological attribute that showed the strongest response to seasonality and drought. Decomposition was slower in the cerrado that we studied compared to a more closed cerrado physiognomy, reflecting their structural and environmental differences. Thus, decomposition rates seem to increase from open to closed cerrado physiognomies, probably related to an increase of humidity and nutrients in the soil.

Investigamos a produção de serapilheira e a taxa de decomposição foliar em uma área de cerrado sensu stricto. Coletamos mensalmente a serapilheira do componente arbustivo-arbóreo de abril de 2001 a março de 2002 e de julho de 2003 a julho de 2004. Dispusemos sistematicamente 13 coletores (0,5 x 0,5 m) em uma linha, com distância de 10 m entre eles. Separamos a serapilheira nas frações 'folhas', 'galhos', 'estruturas reprodutivas' e 'miscelânea'; as secamos em estufa a 80 °C até atingirem massa constante; e pesamos o material seco. Para analisar a taxa de decomposição foliar, acondicionamos folhas caídas recentemente em sacos de decomposição. Dispusemos sete conjuntos de nove sacos de decomposição em uma linha, distantes 10 m um do outro, sobre a superfície do solo e retiramos nove sacos a cada coleta depois de 1, 2, 3, 4, 6, 9 e 12 meses. As produções totais e de folhas apresentaram um padrão estacional. A queda de folhas foi o atributo fenológico que melhor respondeu à estacionalidade e à seca. A decomposição foi mais lenta no cerrado sensu stricto que estudamos do que em um fragmento de cerradão, o que refletiu em suas diferenças estruturais e ambientais. Portanto, as taxas de decomposição devem aumentar das fisionomias de cerrado abertas para as fechadas, provavelmente devido ao aumento da umidade e dos nutrientes do solo.

A year in a Cerrado wet grassland: a non-seasonal island in a seasonal savanna environment / Um ano em um campo úmido no Cerrado: uma ilha não-estacional em um ambiente de savana estacional

In some Cerrado regions where the water table is superficial and soils are hydromorphic, we may find wet grasslands. We studied temporal changes in some community descriptors, such as species density, plant density, basal area, and cylindrical volume in a Cerrado wet grassland in four different seasons of the year. We also compared the species richness and composition of the wet grassland with a hyperseasonal cerrado, and a seasonal cerrado. We found significant differences among the seasons only for species density. Chao-Sørensen similarity values varied from 0.86 to 0.99 and, in the wet grassland, were not different among the seasons. On the contrary, similarity values between the wet grassland and hyperseasonal and seasonal cerrados were low. Species richness was lower in the wet grassland and higher in the seasonal cerrado. As long as savannas are highly dynamic on all temporal and spatial scales, the wet grassland stability, at least in a short-term view, introduces an important heterogeneity in regional scale. Wet grasslands are also important in the Cerrado domain to increase b-diversity, since they are floristically dissimilar with cerrado vegetation.

Dentro do domínio do Cerrado, os campos úmidos ocorrem em áreas em que o lençol freático é superficial e os solos são hidromórficos. Nosso objetivo foi estudar mudanças temporais em alguns descritores da comunidade, como riqueza de espécies, densidade de espécies, densidade de indivíduos, área basal e volume cilíndrico em um campo úmido em quatro estações do ano. Além disso, comparamos a riqueza e a composição de espécies do campo úmido com um cerrado hiperestacional e outro estacional. Encontramos diferenças significativas entre as estações apenas para a densidade de espécies. Os valores do índice de similaridade Chao-Sørensen variaram entre 0,86 e 0,99 e, no campo úmido, não foram significativamente diferentes entre as estações. Porém, a similaridade foi baixa, quando comparamos o campo úmido com os cerrados hiperestacional e estacional. A riqueza de espécies foi menor no campo úmido e maior no cerrado estacional. A estabilidade do campo úmido introduz uma heterogeneidade importante em escala regional, uma vez que as savanas são bastante dinâmicas em escalas espácio-temporais. Os campos úmidos são importantes no domínio do Cerrado por aumentarem a diversidade b, já que são dissimilares em relação ao cerrado.

Can we predict dispersal guilds based on the leaf-height-seed scheme in a disjunct cerrado woodland? / Podemos prever guildas de dispersão com base no esquema folha-altura-semente em um cerrado sensu stricto disjunto?

Although there have been advances in methods for extracting information about dispersal processes, it is still very difficult to measure them. Predicting dispersal groups using single readily-measured traits would facilitate the emergence of instructive comparisons among ecological strategies of plants and offer a path towards improved synthesis across field experiments. The leaf-height-seed scheme consists of three functional traits: specific leaf area, plant canopy height, and seed mass. We tested, applying logistic regression analysis, whether these traits are potential predictors of dispersal guilds in a disjoint cerrado woodland site in southeastern Brazil. According to our results, none of the plant traits studied could predict dispersal guild; this means that abiotically and biotically dispersed species showed similar values of specific leaf area, height, and seed mass. The species of both guilds exhibited sclerophylly, probably a result of the typical soil nutrient deficiency of cerrado, which also may have placed constraints upon plant canopy height regardless of the dispersal mode. In the cerrado, some abiotically dispersed trees might present higher than expected seed mass as support to the investment in high root-to-shoot ratio at the seedling stage. Seeds of bird-dispersed species are limited in size and mass because of the small size of most frugivorous birds. Since soil nutrient quality might contribute to the similarity between the dispersal guilds regarding the three traits of the scheme, other plant traits (e.g., root depth distribution and nutrient uptake strategy) that detail the former should be considered in future predictive studies.

Embora tenham ocorrido avanços nos métodos de aquisição de informações sobre os processos de dispersão, ainda é difícil medi-los. Predizer grupos de dispersão usando atributos simples e práticos facilitaria o aparecimento de comparações instrutivas entre as estratégias ecológicas das plantas e ofereceria um caminho em direção à síntese efetiva de experimentos de campo. O esquema Folha-Altura-Semente consiste em três atributos funcionais: área foliar específica, altura e massa da semente. Testamos, aplicando análise de regressão logística, se esses atributos são potenciais previsores das guildas de dispersão em uma área disjunta de cerrado sensu stricto no Sudeste brasileiro. Segundo os nossos resultados, nenhum dos atributos estudados pôde prever a guilda de dispersão. Isso significa que espécies abiótica e bioticamente dispersas apresentaram valores similares de área foliar específica, altura e massa da semente. As espécies de ambas as guildas exibiram esclerofilia, provavelmente um resultado da deficiência nutricional típica dos solos do cerrado, o que, por sua vez, pode ter limitado à altura das espécies independentemente do seu modo de dispersão. No cerrado, algumas árvores com dispersão abiótica podem apresentar sementes mais pesadas do que o esperado devido ao investimento em elevada razão raiz-caule no estádio de plântula. Sementes de espécies ornitocóricas são limitadas no seu tamanho e massa por causa do reduzido tamanho da maioria das aves frugívoras. Uma vez que a qualidade nutricional do solo do cerrado pode contribuir com a similaridade entre as guildas de dispersão no que diz respeito aos três atributos do esquema, outros atributos (por ex., distribuição da profundidade da raiz e estratégia de captação de nutrientes) que detalhem a primeira devem ser considerados em estudos futuros.

Soil chemical factors and grassland species density in Emas National Park (central Brazil).

Studies of grasslands on specific soil types suggest that different nutrients can limit biomass production and, hence, species composition and number. The Brazilian cerrado is the major savanna region in America and once covered about 2 million km(2), mainly in the Brazilian Central Plateau, under seasonal climate, with wet summer and dry winter. In view of the importance of soil chemical factors in the distribution of the vegetation forms within the Cerrado domain and which may influence the number of species, we analyzed some soil characteristics in three herbaceous vegetation forms -- hyperseasonal cerrado, seasonal cerrado, and wet grassland -- in Emas National Park, a core cerrado site, to investigate the relationship between number of species and soil characteristics. We collected vegetation and soil samples in these three vegetation forms and submitted the obtained data to multiple linear regression. We found out that aluminum and pH were the best predictors of species density, the former positively related to species density and the latter negatively related. Since the predictable variation in species density is important in determining areas of conservation, we can postulate that these two soil factors are indicators of high species density areas in tropical grasslands, which could be used in selecting priority sites for conservation.

Species convergence into life-forms in a hyperseasonal cerrado in central Brazil.

Whether the functional structure of ecological communities is deterministic or historically contingent is still quite controversial. However, recent experimental tests did not find effects of species composition variation on trait convergence and therefore the environmental constraints should play the major role on community convergence into functional groups. Seasonal cerrados are characterized by a sharp seasonality, in which the water shortage defines the community functioning. Hyperseasonal cerrados experience additionally waterlogging in the rainy season. Here, we asked whether waterlogging modifies species convergences into life-forms in a hyperseasonal cerrado. We studied a hyperseasonal cerrado, comparing it with a nearby seasonal cerrado, never waterlogged, in Emas National Park, central Brazil. In each area, we sampled all vascular plants by placing 40 plots of 1 m(2) plots in four surveys. We analyzed the species convergences into life-forms in both cerrados using the Raunkiaer's life-form spectrum and the index of divergence from species to life-form diversity (IDD). The overall life-form spectra and IDDs were not different, indicating that waterlogging did not affect the composition of functional groups in the hyperseasonal cerrado. However, there was a seasonal variation in IDD values only in the hyperseasonal cerrado. As long as we did not find a seasonal variation in life-form diversity, the seasonal variation of convergence into life-forms in the hyperseasonal cerrado was a consequence of the seasonal variation of species diversity. Because of high functional redundancy of cerrado plants, waterlogging promoted a floristic replacement without major changes in functional groups. Thus, waterlogging in the hyperseasonal cerrado promoted seasonal changes in species convergence into life-forms by reducing species diversity.