Abstract
Brazilian savanna is a seasonally dry biome, highly diverse and distributed mainly on nutrient-limited soils. Interactions between water stress and nutrient availability are important evolutionary filters in these environments. Previous evidence indicated that reducing the nutritional limitation increases growth rate, optimizes water transport and decreases stomatal conductance in woody plants. However, the anatomical mechanisms that explain such responses are not well-understood. We studied the effects of long-term (20 years) nutrient addition (N, NP and P) on soil chemistry and hydraulic morphological and anatomical traits in six dominant woody savanna species. Nutrient addition and decrease in soil moisture, probably related to changes in grass cover, influenced the plant hydraulic traits at the anatomical level, namely increasing the xylem vessels’ diameter. Consequently, the specific theoretical xylem conductivity (K tx) increased in five species under NP and P addition. Additionally, the stomatal pore index (SPI) decreased with species-specific responses regarding the intrinsic water use efficiency (iWUE). Four species had higher vulnerability to cavitation (l vul) under NP and P addition. Using community-weighted mean and structural equation modeling approaches, we observed that nutrient enrichment at the community level did not affect iWUE, while the vulnerability to cavitation (l vul) strongly increased. The K tx and SPI were positively and negatively affected by nutrient addition, respectively, but the effects were not as strong as expected due to contrasting species responses. These changes optimized water transport with a hydraulic safety cost and reduced water loss. In comparison with responses to N addition, the greater P-limitation in Cerrado vegetation explains the inter-specific convergence in the responses of P-fertilized individuals. We showed that long-term responses to increased nutrient availability in dystrophic soils include anatomical changes in savanna woody vegetation with relevant interactions with soil-plant–atmosphere water relations.
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