Eco-Hydrology of a Seasonally Dry Tropical Forest : Tree Growth, Belowground Water Dynamics and Drought-Vulnerability

Abstract

Tropical forests are storehouses of more thanhalf of the world‘s biodiversity and play a key role in global carbon, water and energy cycles. However, as a consequence of rapid anthropogenic climate change, biodiversity and climate functions of these forests are under a threat. Climate is changing not only in mean state but its variability is increasing, with extreme events such as droughts, heat waves and storms also rising. Water is fundamental to plants‘ existence, and in the tropics, is a key determinant of plant species‘richness, composition, growth and survival. There is thus an increasing interest in understanding how changing rainfall may cause functional changes in forests or change their species composition. Therefore, the overarching goal of thisdissertation was to understand the impact of water variability on tropical forest tree growth and vulnerability to drought.Forest tree growth along spatial and temporal rainfall gradientsObservational studies that measure whole forest tree growth along spatial or temporal gradients of rainfall are the most common way of formulating forest growth response curves to water availability, when manipulative experiments are cost-prohibitive or impractical (fire or large mammal disturbance). In the tropics, since very few species show anatomically distinct tree rings, estimating tree growth from trunk diameter is the standard practice to obtain growth patterns across species. However, this method—of equating woody growth to diameter change--is susceptible to bias from water-induced stem flexing. In the absence of bias correction, temporal variability in growth is likely to be overestimated and incorrectly attributed to fluctuations in resource availability, especially in forests with high seasonal and inter-annual variability in water. This problem has been largely ignored in the absence of any corrective measure and due to under-appreciation of the magnitude of error. While diameter re-censuses in permanent sampling plots (PSPs) have been most commonly done at 3-5 year scale (using a graduate tape), increasingly they are done at seasonal and annual scales (using band dendrometers) to closely match variation in rainfall, the scales at which hydrostatic bias may be greater in magnitude relative to woody growth. Besides, along a spatial rainfall gradient, inter-annual variability in water may vary, causing systematic differences in the hydrostatic bias for forests along the gradient. Therefore, one broad objective of this thesis was to evaluate the problem of hydrostatic bias in whole forest growth-rainfall relationship at annual and supra-annual scales, for temporal as well as spatial rainfall gradients and propose and test a novel corrective solution.Further, it also examines if growth-diameter relationship vary along the spatial gradient, which it may arise due to differences in light environments and/or disturbance history and species composition. The missing link of Eco-hydrology Differential responses of tree species in terms of growth and survival to variation in water that they can access, the proximate cause is likely shaped through their life-history strategies, the ultimate cause. However, we neither know the depths at which the diverse tree species in a forest draw water from and its dynamics, nor variation in water at those depths vis-à-vis rainfall patterns—for lack of appropriate methods. This has been a key missing link in understanding how water shapes trees‘ life-history strategies, their demographic trade-offs and co-existence, and also our predictive ability to determine species-specific responses to changing rainfall patterns, especially droughts. Since droughts are highly stochastic events and trees‘ responses to their drought ―experiences‖ may be revealed at decadal scales, long-term evaluations are key. Therefore, the second broad objective of this thesis was to develop a framework to determine trees’ water uptake depths, variation in water availability at those depths and trees’ demographic responses over multiple decades. From this, to understand how belowground hydrology shapes drought-vulnerability, demographic trade-offs and coexistence of forest tree species. This thesis titled—Eco-Hydrology of a Seasonally Dry Tropical Forest: Tree Growth, Belowground Water Dynamics and Drought-Vulnerability—is organized as follows: Chapter 1 lays down an introduction to the thesis, followed by a description of the study site and datasets used in the thesis in Chapter 2. This thesis uses a variety of methods and multiple datasets, all of which are from the protected Seasonally Dry Tropical Forests of the Western Ghats in southern India in the Mudumalai and Bandipur National Parks. It is then followed by three data chapters: Chapter 3 describes the seasonal fluctuations in a five year long (1980-1985) tree diameter time series (using dendrometers) of a Seasonally Dry Tropical Forest in Bandipur National Park to illustrate the issue of hydrostatic stem-flexing. It investigates the possibility that band dendrometers may themselves underestimate stem shrinkage at diurnal or seasonal scale. It also evaluates if there could be a best season and time of the day for undertaking forest diameter censuses that can minimize hydrostatic bias. Chapter 4(published in Forest Ecology and Management)measures the hydrostatic bias in a sample of trees in a 50 ha PSP of a Seasonally Dry Tropical Forest in Mudumalai National Park, and proposes a novel way to correct this bias at the whole community level in the 20 year long 4-year interval growth time series. Chapter 5 (in review with Environmental Research Letters) investigates and presents two new confounding factors in growth-rainfall relationships along a spatial rainfall gradient: hydrostatic bias and size-dependency in growth rates. For this it evaluates forest tree growth estimates in seven 1-ha PSPs (~800 trees, 3-year annual time series 9using dendrometers) along a 1000 mm rainfall gradient spanning a mesic savanna-moist forest transition in Mudumalai National Park. Using the period for which seasonal diameter time series was available (2 yrs), it evaluates if the extent of seasonal fluctuations systematically vary along the gradient—most likely due to hydrostatic stem flexing. It also describes the presence of an anomalous size-diameter relationship in the mesic savanna from a large plots (50 ha PSP, diameter records using graduated tape). These observations are then used to draw insights for ―space for time‖ substitution modeling. Chapter 6 (in prep for Nature Plants) analyses belowground water environments of trees over two decades (1992-2012), a period that includes a prolonged and intense drought, in the 50 ha PSP of a Seasonally Dry Tropical Forest in Mudumalai. It uses a locally parametarised dynamic hydrological model in which site rainfall is also a forcing variable. It then develops a novel dynamic growth model and inversely estimates water uptake depths for adult trees of all common species (include ~9000 trees) in the PSP from their above-ground growth patterns over two decades vis-à-vis belowground water availability at multiple depths. It then examines if species‘ water uptake depth obtained thus is a predictor of their drought-driven mortality. Finally, this is used to evaluate the hydrological niche partitioning tree species operate under and how that drives their water uptake strategies, demographic trade-offs, and drought-vulnerability. Summarizes the thesis and suggests future directions