Ecology of seed and seedling growth for conservation and restoration of tropical dry forest: a review

Abstract

Dry forests are among the most threatened ecosystems and have been extensively converted into grasslands, secondary forest, savanna or agricultural land. Knowledge of seed germination and seedling establishment is required for the success of efforts on restoration of these forests. This review focuses on the ecological requirements at seed and seedling stages, and collates the current knowledge of seed viability, dormancy, germination pattern and seedling behaviour of dry tropical tree species. The spatio-temporal variations within the tropical dry forest biome in soil moisture, light, temperature, nutrients and intensity of predation, significantly affect the seed and seedling traits of component species. The majority of dry tropical species possess orthodox seeds which are characterized by dormancy, while a few have recalcitrant seeds which possess little or no dormancy. Seed coat dormancy, which can be overcome by mechanical or acid scarification or sometimes by transit through animal guts, is most prevalent in the dry tropical forest species. Persistent species dominating the undisturbed portions of the forest have bigger seeds compared to those that mostly occur in disturbed regions and require shade for the survival of their seedlings. Shade demand is associated with drought endurance, and may be absolute in species such as Guettarda parviflora and Coccoloba microstachya, or facultative as in Plumeria alba and Bursera simaruba. The fluctuation in temperature significantly affects seed germination in several species of dry Afromontane forest trees of Ethiopia. Seedling mortality is primarily a function of moisture stress during the dry period. Adaptive responses of seedlings to drought stress include increased chlorophyll content, for example in Acacia catechu, and root biomass, as in several dry forest species (for example Drypetes parvifolia, Teclia verdoornia) of Ghana. Mulching, application of fertilizers, interplanting of leguminous species and mycorrhizal inoculation are useful tools for promoting seedling establishment in nutrient-poor dry tropical soils. Periodic forest fires, and predation affect recruitment and seedling development according to their intensity. Many species experiencing frequent fires have evolved thick seed coats, produce fire-hardy seedlings, or escape the effect by temporal separation of seed dispersal and fire events. Predation may result in abortion of fruits or may enhance germination and recruitment by scarification and dispersal, as in most species of the Guanacaste dry forest. Exposure to elevated CO2 has increased relative growth rate, total leaf area and water use efficiency in most of the dry tropical seedlings tested, but the magnitude of the effect has varied markedly among species. Due to the availability of a large source of energy, large seeds show higher germination percentage, greater seedling survival and increased growth. Seeds originating from different provenances exhibit differences in germination and seedling growth (for example Prosopis cineraria, Albizia lebbeck, Eucalyptus camaldulensis and Acacia mangium), efficiency of nodulation (for example Acacia nilotica, A. auriculiformis), and stress resistance (for example Populus deltoides, Dalbergia sissoo). The review points out the need for coordinated, long-term, field-based studies for identification of multiple cues and niches for germination, on seed and seedling dynamics in response to fire, and on within-species genetic variability for selection of suitable provenances. Field-based studies at species and community levels are also needed to permit manipulations of biotic components to augment the recruitment of desired species and to suppress that of undesirable species.