RESUMO

Trade-offs among plant functional traits indicate diversity in plant strategies of growth and survival. The leaf economics spectrum (LES) reflects a trade-off between short-term carbon gain and long-term leaf persistence. A related trade-off, between foliar growth and anti-herbivore defense, occurs among plants growing in contrasting resource regimes, but it is unclear whether this trade-off is maintained within plant communities, where resource gradients are minimized. The LES and the growth-defense trade-off involve related traits, but the extent to which these trade-off dimensions are correlated is poorly understood. We assessed the relationship between leaf economic and anti-herbivore defense traits among sunlit foliage of 345 canopy trees in 83 species on Barro Colorado Island, Panama. We quantified ten traits related to resource allocation and defense, and identified patterns of trait co-variation using multivariate ordination. We tested whether traits and ordination axes were correlated with patterns of phylogenetic relatedness, juvenile demographic trade-offs, or topo-edaphic variation. Two independent axes described ~ 60% of the variation among canopy trees. Axis 1 revealed a trade-off between leaf nutritional and structural investment, consistent with the LES. Physical defense traits were largely oriented along this axis. Axis 2 revealed a trade-off between investments in phenolic defenses versus other foliar defenses, which we term the leaf defense spectrum. Phylogenetic relationships and topo-edaphic variation largely did not explain trait co-variation. Our results suggest that some trade-offs among the growth and defense traits of outer-canopy trees may be captured by the LES, while others may occur along additional resource allocation dimensions.

Assuntos

RESUMO

Leaf aging is a fundamental driver of changes in leaf traits, thereby regulating ecosystem processes and remotely sensed canopy dynamics. We explore leaf reflectance as a tool to monitor leaf age and develop a spectra-based partial least squares regression (PLSR) model to predict age using data from a phenological study of 1099 leaves from 12 lowland Amazonian canopy trees in southern Peru. Results demonstrated monotonic decreases in leaf water (LWC) and phosphorus (Pmass ) contents and an increase in leaf mass per unit area (LMA) with age across trees; leaf nitrogen (Nmass ) and carbon (Cmass ) contents showed monotonic but tree-specific age responses. We observed large age-related variation in leaf spectra across trees. A spectra-based model was more accurate in predicting leaf age (R2  = 0.86; percent root mean square error (%RMSE) = 33) compared with trait-based models using single (R2  = 0.07-0.73; %RMSE = 7-38) and multiple (R2  = 0.76; %RMSE = 28) predictors. Spectra- and trait-based models established a physiochemical basis for the spectral age model. Vegetation indices (VIs) including the normalized difference vegetation index (NDVI), enhanced vegetation index 2 (EVI2), normalized difference water index (NDWI) and photosynthetic reflectance index (PRI) were all age-dependent. This study highlights the importance of leaf age as a mediator of leaf traits, provides evidence of age-related leaf reflectance changes that have important impacts on VIs used to monitor canopy dynamics and productivity and proposes a new approach to predicting and monitoring leaf age with important implications for remote sensing.

Assuntos

RESUMO

Dry forests are common, although highly threatened in the Neotropics. Their ecological processes are mostly influenced by rainfall pattern, hence their cycles exhibit contrasting phases. We studied the phenology of canopy trees in a primary dry forest in Western Brazil in the foothills of the Urucum mountain chain, in order to improve our knowledge on the functioning of these poorly-known forests. Leaf shedding started in the early dry season and was massive in the latter part of this period. Most leaf loss occurred in dry hills, while wet valleys remained evergreen. Anemochorich and autochorich species predominated in dry hills, presumably due to their tolerance to dry conditions and enhanced exposition to winds, which favour diaspores removal and dispersal. Conversely, zoochorich species dominated the wet valleys. Flowering was intense in the late dry season, the driest period of the year, while fruiting was massive just after the onset of rains, as well as flushing. Therefore, most flowering was unrelated to wet conditions, although such an abiotic factor, potentially, triggered the major fruiting episode, widely comprised by zoochorich species. Anemochorich and autochorich species flowered and fruited in the course of the long dry season. The contrasting environmental conditions present in the hills and valleys determine the arrangement of a mosaic in which patches of zoochorich and evergreen trees alternate with patches of non zoochorich and highly deciduous species. Consequently, species with such syndromes exhibited marked flowering and fruiting patterns, accordingly to the pronounced seasonality.

Matas secas neotropicais estão amplamente distribuídas, porém sob elevado risco de desmatamento. Os processos ecológicos nesses ambientes são fortemente influenciados pelo clima, sobretudo o padrão de chuvas, de tal forma que seus ciclos apresentam fases muito contrastantes. Nesse estudo, avaliamos a produção de folhas, flores e frutos em uma mata seca do oeste brasileiro situada no sopé do maciço do Urucum. A perda de folhas teve início no começo da estação seca, mas foi massiva ao final desse período, o mais seco do ano. Espécies decíduas predominaram nas escarpas secas, enquanto as perenifólias foram comuns nos vales úmidos. Nas escarpas, espécies anemo e autocóricas eram muito comuns, potencialmente, por serem mais tolerantes à baixa umidade, bem como favorecidas por ventos mais fortes. Porém, eram raras ou mesmo ausentes nos vales úmidos dominados por espécies zoocóricas. A floração foi intensa, exibindo um pico acentuado ao final da estação seca, seguida de um pronunciado pico de frutificação e produção de folhas com o início das chuvas. Dessa forma, enquanto a floração massiva não foi influenciada pelas chuvas, a frutificação e produção de novas folhas estiveram fortemente relacionadas a esse fator abiótico. As espécies anemo e autocóricas floresceram e frutificaram durante a prolongada estação seca, ao contrário da maioria das zoocóricas. As condições ambientais contrastantes dos vales e escarpas, potencialmente determinam um mosaico em que porções altamente decíduas de mata, com predomínio de espécies anemo e autocóricas, se alternam com outras sempre verdes, dominadas por espécies zoocóricas. Além disso, a forte sazonalidade influencia diferentemente espécies com síndromes de dispersão distintas.

Assuntos

RESUMO

Dry forests are common, although highly threatened in the Neotropics. Their ecological processes are mostly influenced by rainfall pattern, hence their cycles exhibit contrasting phases. We studied the phenology of canopy trees in a primary dry forest in Western Brazil in the foothills of the Urucum mountain chain, in order to improve our knowledge on the functioning of these poorly-known forests. Leaf shedding started in the early dry season and was massive in the latter part of this period. Most leaf loss occurred in dry hills, while wet valleys remained evergreen. Anemochorich and autochorich species predominated in dry hills, presumably due to their tolerance to dry conditions and enhanced exposition to winds, which favour diaspores removal and dispersal. Conversely, zoochorich species dominated the wet valleys. Flowering was intense in the late dry season, the driest period of the year, while fruiting was massive just after the onset of rains, as well as flushing. Therefore, most flowering was unrelated to wet conditions, although such an abiotic factor, potentially, triggered the major fruiting episode, widely comprised by zoochorich species. Anemochorich and autochorich species flowered and fruited in the course of the long dry season. The contrasting environmental conditions present in the hills and valleys determine the arrangement of a mosaic in which patches of zoochorich and evergreen trees alternate with patches of non zoochorich and highly deciduous species. Consequently, species with such syndromes exhibited marked flowering and fruiting patterns, accordingly to the pronounced seasonality.

Matas secas neotropicais estão amplamente distribuídas, porém sob elevado risco de desmatamento. Os processos ecológicos nesses ambientes são fortemente influenciados pelo clima, sobretudo o padrão de chuvas, de tal forma que seus ciclos apresentam fases muito contrastantes. Nesse estudo, avaliamos a produção de folhas, flores e frutos em uma mata seca do oeste brasileiro situada no sopé do maciço do Urucum. A perda de folhas teve início no começo da estação seca, mas foi massiva ao final desse período, o mais seco do ano. Espécies decíduas predominaram nas escarpas secas, enquanto as perenifólias foram comuns nos vales úmidos. Nas escarpas, espécies anemo e autocóricas eram muito comuns, potencialmente, por serem mais tolerantes à baixa umidade, bem como favorecidas por ventos mais fortes. Porém, eram raras ou mesmo ausentes nos vales úmidos dominados por espécies zoocóricas. A floração foi intensa, exibindo um pico acentuado ao final da estação seca, seguida de um pronunciado pico de frutificação e produção de folhas com o início das chuvas. Dessa forma, enquanto a floração massiva não foi influenciada pelas chuvas, a frutificação e produção de novas folhas estiveram fortemente relacionadas a esse fator abiótico. As espécies anemo e autocóricas floresceram e frutificaram durante a prolongada estação seca, ao contrário da maioria das zoocóricas. As condições ambientais contrastantes dos vales e escarpas, potencialmente determinam um mosaico em que porções altamente decíduas de mata, com predomínio de espécies anemo e autocóricas, se alternam com outras sempre verdes, dominadas por espécies zoocóricas. Além disso, a forte sazonalidade influencia diferentemente espécies com síndromes de dispersão distintas.

RESUMO

We evaluated the hypothesis that photosynthetic traits differ between leaves produced at the beginning (May) and the end (November-December) of the rainy season in the canopy of a seasonally dry forest in Panama. Leaves produced at the end of the wet season were predicted to have higher photosynthetic capacities and higher water-use efficiencies than leaves produced during the early rainy season. Such seasonal phenotypic differentiation may be adaptive, since leaves produced immediately preceding the dry season are likely to experience greater light availability during their lifetime due to reduced cloud cover during the dry season. We used a construction crane for access to the upper canopy and sampled 1- to 2-month-old leaves marked in monthly censuses for six common tree species with various ecological habits and leaf phenologies. Photosynthetic capacity was quantified as light- and CO2-saturated oxygen evolution rates with a leaf-disk oxygen electrode in the laboratory (O2max) and as light-saturated CO2 assimilation rates of intact leaves under ambient CO2 (Amax). In four species, pre-dry season leaves had significantly higher leaf mass per unit area. In these four species, O2max and Amax per unit area and maximum stomatal conductances were significantly greater in pre-dry season leaves than in early wet season leaves. In two species, Amax for a given stomatal conductance was greater in pre-dry season leaves than in early wet season leaves, suggesting a higher photosynthetic water-use efficiency in the former. Photosynthetic capacity per unit mass was not significantly different between seasons of leaf production in any species. In both early wet season and pre-dry season leaves, mean photosynthetic capacity per unit mass was positively correlated with nitrogen content per unit mass both within and among species. Seasonal phenotypic differentiation observed in canopy tree species is achieved through changes in leaf mass per unit area and increased maximum stomatal conductance rather than by changes in nitrogen allocation patterns.