Seedling Traits Determine Drought Tolerance of Tropical Tree Species

Water availability is the most important factor determining tree species distribution in the tropics, but the underlying mechanisms are still not clear. In this study, we compared functional traits of 38 tropical tree species from dry and moist forest, and quantified their ability to survive drought in a dry-down experiment in which wilting and survival were monitored. We evaluated how seedling traits affect drought survival, and how drought survival determines species distribution along the rainfall gradient. Dry forest species tended to have compound leaves, high stem dry matter content (stem dry mass/fresh mass), and low leaf area ratio, suggesting that reduction of transpiration and avoidance of xylem cavitation are important for their success. Three functional groups were identified based on the seedling traits: (1) drought avoiders with a deciduous leaf habitat and taproots; (2) drought resisters with tough tissues (i.e., a high dry matter content); and (3) light-demanding moist forest species with a large belowground foraging capacity. Dry forest species had a longer drought survival time (62 d) than moist forest species (25 d). Deciduousness explained 69 percent of interspecific variation in drought survival. Among evergreen species, stem density explained 20 percent of the drought survival. Drought survival was not related to species distribution along the rainfall gradient, because it was mainly determined by deciduousness, and species with deciduous seedlings are found in both dry and moist forests. Among evergreen species, drought survival explained 28 percent of the variation in species position along the rainfall gradient. This suggests that, apart from drought tolerance, other factors such as history, dispersal limitation, shade tolerance, and fire shape species distribution patterns along the rainfall gradient.

[1]  S. Díaz,et al.  Plant functional types and ecosystem function in relation to global change , 1997 .

[2]  K. Cao Water relations and gas exchange of tropical saplings during a prolonged drought in a Bornean heath forest, with reference to root architecture , 2000, Journal of Tropical Ecology.

[3]  F. S. Chapin,et al.  Relationship between the structure of root systems and resource use for 11 North American grassland plants , 2003, Plant Ecology.

[4]  E. Veenendaal,et al.  Differences in plant and soil water relations in and around a forest gap in West Africa during the dry season may influence seedling establishment and survival , 1996 .

[5]  D. Bonal,et al.  Contrasting patterns of leaf water potential and gas exchange responses to drought in seedlings of tropical rainforest species , 2001 .

[6]  ALAN P. Smith,et al.  Drought acclimation among tropical forest shrubs (Psychotria, Rubiaceae) , 1992, Oecologia.

[7]  H. Paz Root/Shoot Allocation and Root Architecture in Seedlings: Variation among Forest Sites, Microhabitats, and Ecological Groups1 , 2003 .

[8]  Jeannine Cavender-Bares,et al.  MULTIPLE TRAIT ASSOCIATIONS IN RELATION TO HABITAT DIFFERENTIATION AMONG 17 FLORIDIAN OAK SPECIES , 2004 .

[9]  P. Richards,et al.  The Tropical Rain Forest: An Ecological Study , 1953 .

[10]  M. Swaine,et al.  Distribution and ecology of vascular plants in a tropical rain forest , 1981, Geobotany.

[11]  W. Hoffmann FIRE AND POPULATION DYNAMICS OF WOODY PLANTS IN A NEOTROPICAL SAVANNA: MATRIX MODEL PROJECTIONS , 1999 .

[12]  F. Bongers,et al.  The forests of Upper Guinea: gradients in large species composition , 2004 .

[13]  F. Bongers,et al.  Biodiversity of West African forests: an ecological atlas of woody plant species. , 2004 .

[14]  Derek Eamus,et al.  Ecophysiology of trees of seasonally dry tropics: Comparisons among phenologies , 2001 .

[15]  O. Loucks,et al.  Optimal leaf size in relation to environment , 1972 .

[16]  Bettina M. J. Engelbrecht,et al.  Desiccation Tolerance of Five Tropical Seedlings in Panama. Relationship to a Field Assessment of Drought Performance1 , 2003, Plant Physiology.

[17]  W. Hoffmann,et al.  Comparative growth analysis of tropical forest and savanna woody plants using phylogenetically independent contrasts , 2003 .

[18]  M. Huston,et al.  A theory of the spatial and temporal dynamics of plant communities , 1989, Vegetatio.

[19]  Bettina M. J. Engelbrecht,et al.  Comparative drought-resistance of seedlings of 28 species of co-occurring tropical woody plants , 2003, Oecologia.

[20]  D. Burslem,et al.  Responses to simulated drought and elevated nutrient supply among shade-tolerant tree seedlings of lowland tropical forest in Singapore , 1996 .

[21]  David D. Ackerly,et al.  FUNCTIONAL STRATEGIES OF CHAPARRAL SHRUBS IN RELATION TO SEASONAL WATER DEFICIT AND DISTURBANCE , 2004 .

[22]  Ian J. Wright,et al.  Differences in seedling growth behaviour among species: trait correlations across species, and trait shifts along nutrient compared to rainfall gradients , 1999 .

[23]  A. Gentry,et al.  Changes in Plant Community Diversity and Floristic Composition on Environmental and Geographical Gradients , 1988 .

[24]  G. Parker,et al.  Seasonal balance and vertical pattern of photosynthetically active radiation within canopies of a tropical dry deciduous forest ecosystem in Mexico , 2005, Journal of Tropical Ecology.

[25]  D. Burslem,et al.  Species–habitat associations in a Sri Lankan dipterocarp forest , 2006, Journal of Tropical Ecology.

[26]  Mark G. Tjoelker,et al.  Close association of RGR, leaf and root morphology, seed mass and shade tolerance in seedlings of nine boreal tree species grown in high and low light , 1998 .

[27]  Michael F. Tobin,et al.  Responses of Tropical Understory Plants to a Severe Drought: Tolerance and Avoidance of Water Stress 1 , 1999 .

[28]  Ü. Niinemets,et al.  Tolerance to shade, drought, and waterlogging of temperate northern hemisphere trees and shrubs , 2006 .

[29]  O. Phillips,et al.  Continental-scale patterns of canopy tree composition and function across Amazonia , 2006, Nature.

[30]  L. Poorter,et al.  Effects of seasonal drought on gap and understorey seedlings in a Bolivian moist forest , 2000, Journal of Tropical Ecology.

[31]  J. Cornelissen,et al.  Stem anatomy and relative growth rate in seedlings of a wide range of woody plant species and types , 1998, Oecologia.

[32]  Ü. Niinemets Are compound-leaved woody species inherently shade-intolerant? An analysis of species ecological requirements and foliar support costs , 2004, Plant Ecology.

[33]  L. Poorter Resource capture and use by tropical forest tree seedlings and their consequences for competition , 2003 .

[34]  Bettina M. J. Engelbrecht,et al.  Short dry spells in the wet season increase mortality of tropical pioneer seedlings , 2006, Oecologia.

[35]  L. Sack Responses of temperate woody seedlings to shade and drought: do trade‐offs limit potential niche differentiation? , 2004 .

[36]  R. Borchert,et al.  Soil and Stem Water Storage Determine Phenology and Distribution of Tropical Dry Forest Trees , 1994 .

[37]  P. Reich,et al.  Water stress and tree phenology in a tropical dry forest in the lowlands of Costa Rica. , 1984 .

[38]  Bettina M. J. Engelbrecht,et al.  Drought effects on seedling survival in a tropical moist forest , 2005, Trees.

[39]  o. Prof. em. Dr. h. c. Heinrich Walter,et al.  Vegetation of the Earth and Ecological Systems of the Geobiosphere , 1983, Heidelberg Science Library.

[40]  L. Poorter,et al.  Carbohydrate storage and light requirements of tropical moist and dry forest tree species. , 2007, Ecology.

[41]  W. Hoffmann,et al.  Constraints to seedling success of savanna and forest trees across the savanna-forest boundary , 2004, Oecologia.

[42]  E. Newman,et al.  A METHOD OF ESTIMATING THE TOTAL LENGTH OF ROOT IN A SAMPLE , 1966 .

[43]  W. Carson,et al.  Drought stress and tropical forest woody seedlings: effect on community structure and composition , 2005 .

[44]  Stephen P. Hubbell,et al.  Drought sensitivity shapes species distribution patterns in tropical forests , 2007, Nature.

[45]  Frans Bongers,et al.  Leaf traits are good predictors of plant performance across 53 rain forest species. , 2006, Ecology.

[46]  E. Veenendaal Limits to tree species distributions in lowland tropical rainforest , 1998 .

[47]  L. Poorter,et al.  Diversity of Tropical Tree Seedling Responses to Drought , 2007 .

[48]  Y. Malhi,et al.  Spatial patterns and recent trends in the climate of tropical rainforest regions. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[49]  S. Lavorel,et al.  Predicting changes in community composition and ecosystem functioning from plant traits: revisiting the Holy Grail , 2002 .

[50]  D. Coomes,et al.  IMPACTS OF ROOT COMPETITION IN FORESTS AND WOODLANDS: A THEORETICAL FRAMEWORK AND REVIEW OF EXPERIMENTS , 2000 .

[51]  M. Swaine,et al.  Rainfall and soil fertility as factors limiting forest species distributions in Ghana , 1996 .

[52]  M. Swaine,et al.  Classification and ecology of closed-canopy forest in Ghana. , 1976 .

[53]  T. Killeen,et al.  The Chiquitano Dry Forest, the Transition between Humid and Dry Forest in Eastern Lowland Bolivia , 2006 .

[54]  Peter Ryser,et al.  The importance of tissue density for growth and life span of leaves and roots: a comparison of five ecologically contrasting grasses , 1996 .

[55]  Frans Bongers,et al.  Distribution of twelve moist forest canopy tree species in Liberia and Côte d'Ivoire: response curves to a climatic gradient , 1999 .

[56]  L. Poorter,et al.  Does a ruderal strategy dominate the endemic flora of the West African forests? , 2007 .

[57]  M. Daws,et al.  Effects of topographic position, leaf litter and seed size on seedling demography in a semi-deciduous tropical forest in Panamá , 2005, Plant Ecology.