Leaf phenology was monitored for 49 woody species (trees and tall shrubs) each month over a 2.5-year period in a humid, wet-dry tropical eucalypt savanna at Solar Village, near Darwin, Australia. In the 10 most common species, which spanned the range of phenological types, phenology was monitored every two weeks. To investigate the relationships between leaf phenology and plant water status, pre-dawn leaf water potential was monitored in eight common species every 4-6 weeks. Four main phenological types were described: (1) evergreen species, which retained full canopy throughout the year; (2) brevi-or partly deciduous species, in which the amount of canopy fell significantly, but briefly, during at least one dry season during the study period, but to levels not below 50% of full canopy; (3) semideciduous species in which canopy fell to below 50% of full canopy in each of the dry seasons; and (4) fully deciduous species, which lost all leaves during the early-mid dry season, and remained leafless for at least one month. Of these 49 species, 24% were evergreen, 20% were brevideciduous, 29% were semideciduous, and 27% were fully deciduous. Leaf fall occurred 1-2 months earlier in the dry season for the fully deciduous species than for the semideciduous species. Leaf fall in all species was coincident with the attainment of seasonal minima in leaf water potential, which were, on average, about Ϫ1.5 to Ϫ2.0 MPa in the evergreen and semideciduous species, compared with about Ϫ0.5 to Ϫ1.0 MPa in the fully deciduous species. Leaf flushing occurred throughout the dry season in the two evergreen species, with a major peak in the late dry season. In five semideciduous species and one of the fully deciduous species, leaf flushing commenced in the late dry season prior to the occurrence of any rain. In two fully deciduous species, leaf flushing occurred only after the first storms of the early wet season. There was variation in the timing of flushing, both between species within years and between years for some species. However, all species commenced leaf flushing after water potentials rose, following the attainment of seasonal minima in pre-dawn leaf water potential. Soil moisture at 1 m did not fall below permanent wilting point during the dry season; hence, reserves of soil water at the end of the dry season were sufficient to support the whole-plant rehydration that preceded leaf flushing in the absence of rain. These results are consistent with hypotheses, developed in the neotropics, that leaf phenology in trees from the wet-dry tropics is largely controlled by endogenous mechanisms.
Abstract. Variation in structural and compositional attributes of tropical savannas are described in relation to variation in annual rainfall and soil texture along a subcontinental‐scale gradient of rainfall in the wet‐dry tropics of the Northern Territory, Australia. Rainfall varies along the gradient from over 1500 mm p.a. in the Darwin region (c. 12° S) to less than 500 mm in the Tennant Creek region (c. 18° S). Soils are patchy, and sands, loams and clays may occur in all major districts within the region. We utilized a large data set (1657 quadrats ° 291 woody species; with numerous measured and derived sample variables) covering an area of 0.5 million km2. Correlations between floristic composition of woody species and environmental variables were assessed using DCA ordination and vector fitting of environmental variables. Vectors of annual rainfall and soil texture were highly correlated with variation in species composition. Multiple regression analyses incorporating linear and quadratic components of mean annual rainfall and topsoil clay content were performed on three structural attributes (tree height, tree cover, tree basal area) and two compositional attributes (woody species richness, deciduous tree species richness). Tree height declined with decreasing rainfall; cover, basal area, woody species richness and deciduous species richness all declined with decreasing rainfall and increasing soil clay content. Regression models accounted for between 17% and 45% of the variation in the data sets. Variation in other factors such as soil depth, landscape position and recent land‐use practices (for which there were no data on an individual quadrat basis) are likely to have contributed to the large residual variation in the data set.
Leaf phenology was monitored for 49 woody species (trees and tall shrubs) each month over a 2.5‐year period in a humid, wet–dry tropical eucalypt savanna at Solar Village, near Darwin, Australia. In the 10 most common species, which spanned the range of phenological types, phenology was monitored every two weeks. To investigate the relationships between leaf phenology and plant water status, pre‐dawn leaf water potential was monitored in eight common species every 4–6 weeks. Four main phenological types were described: (1) evergreen species, which retained full canopy throughout the year; (2) brevi‐ or partly deciduous species, in which the amount of canopy fell significantly, but briefly, during at least one dry season during the study period, but to levels not below 50% of full canopy; (3) semideciduous species in which canopy fell to below 50% of full canopy in each of the dry seasons; and (4) fully deciduous species, which lost all leaves during the early‐mid dry season, and remained leafless for at least one month. Of these 49 species, 24% were evergreen, 20% were brevideciduous, 29% were semideciduous, and 27% were fully deciduous. Leaf fall occurred 1–2 months earlier in the dry season for the fully deciduous species than for the semideciduous species. Leaf fall in all species was coincident with the attainment of seasonal minima in leaf water potential, which were, on average, about −1.5 to −2.0 MPa in the evergreen and semideciduous species, compared with about −0.5 to −1.0 MPa in the fully deciduous species. Leaf flushing occurred throughout the dry season in the two evergreen species, with a major peak in the late dry season. In five semideciduous species and one of the fully deciduous species, leaf flushing commenced in the late dry season prior to the occurrence of any rain. In two fully deciduous species, leaf flushing occurred only after the first storms of the early wet season. There was variation in the timing of flushing, both between species within years and between years for some species. However, all species commenced leaf flushing after water potentials rose, following the attainment of seasonal minima in pre‐dawn leaf water potential. Soil moisture at 1 m did not fall below permanent wilting point during the dry season; hence, reserves of soil water at the end of the dry season were sufficient to support the whole‐plant rehydration that preceded leaf flushing in the absence of rain. These results are consistent with hypotheses, developed in the neotropics, that leaf phenology in trees from the wet–dry tropics is largely controlled by endogenous mechanisms.
Seasonal variations in carbon assimilation of eight tree species of a north Australian tropical savanna were examined over two wet seasons and one dry season (18 months). Assimilation rates (A) in the two evergreen species, Eucalyptus tetrodonta F. Muell. and E. miniata A. Cunn. ex Schauer, were high throughout the study although there was a 10-20% decline in the dry season compared with the wet season. The three semi-deciduous species (Erythrophleum chlorostachys (F. Muell.) Baillon, Eucalyptus clavigera A. Cunn. ex Schauer, and Xanthostemon paradoxus F. Muell.) showed a 25-75% decline in A in the dry season compared with the wet season, and the deciduous species (Terminalia ferdinandiana Excell, Planchonia careya (F. Muell.) Kunth, and Cochlospermum fraseri Planchon) were leafless for several months in the dry season. Generally, the ratio of intercellular CO(2) concentration to ambient CO(2) concentration (C(i):C(a)) was larger in the wet season than in the dry season, indicating a smaller stomatal limitation of photosynthesis in the wet season compared with the dry season. In all species, the C(i):C(a) ratio and A were essentially independent of leaf-to-air vapor pressure difference (LAVPD) during the wet season, but both parameters generally declined with increasing LAVPD in the dry season. The slope of the positive correlation between A and transpiration rate (E) was less in the wet season than in the dry season. There was no evidence that high E inhibited A. Instantaneous transpiration efficiency was lowest in the wet season and highest during the dry season. Nitrogen-use efficiency (NUE) was higher in the wet season than in the dry season because the decline in A in the dry season was proportionally larger than the decline in foliar nitrogen content. In the wet season, evergreen species exhibited higher NUE than semi-deciduous and deciduous species. In all species, A was linearly correlated with specific leaf area (SLA) and foliar N content. Foliar N content increased with increasing SLA. All species showed a decline in midday leaf water potential as the dry season progressed. Dry season midday water potentials were lowest in semi-deciduous species and highest in the deciduous species, with evergreen species exhibiting intermediate values.
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