Bettina M. J. Engelbrecht

Drought sensitivity shapes species distribution patterns in tropical forests.

Although patterns of tree species distributions along environmental gradients have been amply documented in tropical forests, mechanisms causing these patterns are seldom known. Efforts to evaluate proposed mechanisms have been hampered by a lack of comparative data on species’ reactions to relevant axes of environmental variation. Here we show that differential drought sensitivity shapes plant distributions in tropical forests at both regional and local scales. Our analyses are based on experimental field assessments of drought sensitivity of 48 species of trees and shrubs, and on their local and regional distributions within a network of 122 inventory sites spanning a rainfall gradient across the Isthmus of Panama. Our results suggest that niche differentiation with respect to soil water availability is a direct determinant of both local- and regional-scale distributions of tropical trees. Changes in soil moisture availability caused by global climate change and forest fragmentation are therefore likely to alter tropical species distributions, community composition and diversity.

Functional traits and the growth?mortality trade-off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.

Species distributions in response to individual soil nutrients and seasonal drought across a community of tropical trees

Tropical forest vegetation is shaped by climate and by soil, but understanding how the distributions of individual tree species respond to specific resources has been hindered by high diversity and consequent rarity. To study species over an entire community, we surveyed trees and measured soil chemistry across climatic and geological gradients in central Panama and then used a unique hierarchical model of species occurrence as a function of rainfall and soil chemistry to circumvent analytical difficulties posed by rare species. The results are a quantitative assessment of the responses of 550 tree species to eight environmental factors, providing a measure of the importance of each factor across the entire tree community. Dry-season intensity and soil phosphorus were the strongest predictors, each affecting the distribution of more than half of the species. Although we anticipated clear-cut responses to dry-season intensity, the finding that many species have pronounced associations with either high or low phosphorus reveals a previously unquantified role for this nutrient in limiting tropical tree distributions. The results provide the data necessary for understanding distributional limits of tree species and predicting future changes in forest composition.

Drought effects on seedling survival in a tropical moist forest

The amount and seasonality of rainfall varies strongly in the tropics, and plant species abundance, distribution and diversity are correlated with rainfall. Drought periods leading to plant stress occur not only in dry forests, but also in moist and even wet forests. We quantified experimentally the effect of drought on survival of first year seedlings of 28 co-occurring tropical woody plant species in the understory of a tropical moist forest. The seedlings were transplanted to plots and subjected to a drought and an irrigation treatment for 22 weeks during the dry season. Drought effects on mortality and wilting behavior varied greatly among species, so that relative survival in the dry treatment ranged from 0% to about 100% of that in the irrigated treatment. Drought stress was the main factor in mortality, causing about 90% (median) of the total mortality observed in the dry treatment. In almost half of the species, the difference in survival between treatments was not significant even after 22 weeks, implying that many of the species are well adapted to drought in this forest. Relative drought survival was significantly higher in species associated with dry habitats than in those associated with wet habitats, and in species with higher abundance on the dry side of the Isthmus of Panama, than in those more abundant on the wet side. These data show that differential species survival in response to drought, combined with variation in soil moisture availability, may be important for species distribution at the local and regional scale in many tropical forests.

Desiccation Tolerance of Five Tropical Seedlings in Panama. Relationship to a Field Assessment of Drought Performance

Studies of the desiccation tolerance of the seedlings of five tropical trees were made on potted plants growing in a greenhouse. Pots were watered to field capacity and then dehydrated for 3 to 9 weeks to reach various visual wilting stages, from slightly wilted to dead. Saturated root hydraulic conductance was measured with a high-pressure flowmeter, and whole-stem hydraulic conductance was measured by a vacuum chamber method. Leaf punches (5.6-mm diameter) were harvested for measurement of leaf water potential by a thermocouple psychrometer method and for measurement of fresh and dry weight. In a parallel study, the same five species were studied in a field experiment in the understory of a tropical forest, where these species frequently germinate. Control seedlings were maintained in irrigated plots during a dry season, and experimental plants were grown in similar plots with rain exclusion shelters. Every 2 to 4 weeks, the seedlings were scored for wilt state and survivorship. After a 22-week drought, the dry plots were irrigated for several weeks to verify visual symptoms of death. The field trials were used to rank drought performance of species, and the greenhouse desiccation studies were used to determine the conditions of moribund plants. Our conclusion is that the desiccation tolerance of moribund plants correlated with field assessment of drought-performance for the five species (r2 > 0.94).

Ecological Determinism in Plant Community Structure Across a Tropical Forest Landscape

The ecological mechanisms hypothesized to structure species-rich communities range from strict local determinism to neutral ecological drift. We assessed the degree of ecological determinism in tropical plant community structure by analyses of published demographic data; a broad range of spatial, historical, and environmental variables; and the distributions of 33 herbaceous species (plot size = 0.02 ha) and 61 woody species (plot size = 0.09 ha) among 350 plots in a 16-km2 forest landscape (Barro Colorado Island, Panama). We found a strong degree of cross-landscape dominance by a subset of species whose identities were predictable from sapling survivorship rates under shade. Using canonical ordination we found that spatial and environmental–historical factors were of comparable importance for controlling within-landscape variability in species composition. Past land use had a strong impact on species composition despite ceasing 100–200 years ago. Furthermore, edaphic–hydrological factors, treefall gaps, and an edge effect all had unique impacts on species composition. Hence, ecological determinism was evident in terms of both cross-landscape dominance and within-landscape variability in species composition. However, at the latter scale, the large portion of the explained variance in species composition among plots uniquely attributed to spatial location pointed to an equally important role for neutral processes.

Evaluation of different methods to estimate understorey light conditions in tropical forests

The suitability of several methods for estimating light conditions in the understorey of tropical forests, and of different sampling schedules was evalu- ated. Light conditions at 16 understorey sites in a Panamanian lowland forest were continuously measured for 9 mo with quantum sensors and photodiodes. Light conditions at the sites were also assessed indirectly with hemispherical fisheye photographs, plant canopy analysis, 38-mm photographs, 24-mm photographs and a spherical densiometer. Estimates from all indirect methods, except the spherical densiometer, were highly correlated with the direct measurements. Short-term direct light measurements for a day or a week also correlated with long-term light conditions. The indirect measures differed by up to c. 70% from the direct meas- ures relative to single site measurements. Hence, the indirect methods are inad- equate where single site light conditions have to be assessed accurately. However, because light conditions encountered in the understorey varied up to 13-fold, the indirect methods were found to be well suited to rank understorey light conditions among a large number of sites. The results from frequent and infrequent sampling schedules differed only slightly, suggesting that taking indirect measures at the beginning and the end of a study offers a reasonable compromise between accuracy and sampling effort.

Fungal endophytes nearly double minimum leaf conductance in seedlings of a neotropical tree species

Drought strongly influences plant phenology, growth and mortality in tropical forests, thereby shaping plant performance, population dynamics and community structure (Bunker & Carson 2005, Condit et al. 1995). Microbial symbionts of plants profoundly influence host water relations (Losch & Gansert 2002), but are rarely considered in studies of tropical plant physiology. In particular, plant–fungus associations, which are ubiquitous in plant communities and especially common in tropical forests, play important and varied roles in plant water status. Fungal pathogens associated with roots, vascular tissue and foliage may interfere with water uptake and transport, increase rates of foliar transpiration, and induce xylem embolism and tissue death (Agrios 1997). In contrast, rhizosphere mutualists such as ecto- and arbuscular mycorrhizal fungi may benefit hosts by increasing surface area for water uptake, enhancing stomatal regulation of water loss, and increasing root hydraulic conductivity (Auge 2001, Losch & Gansert 2002).

Geological substrates shape tree species and trait distributions in African moist forests.

Background Understanding the factors that shape the distribution of tropical tree species at large scales is a central issue in ecology, conservation and forest management. The aims of this study were to (i) assess the importance of environmental factors relative to historical factors for tree species distributions in the semi-evergreen forests of the northern Congo basin; and to (ii) identify potential mechanisms explaining distribution patterns through a trait-based approach. Methodology/Principal Findings We analyzed the distribution patterns of 31 common tree species in an area of more than 700,000 km2 spanning the borders of Cameroon, the Central African Republic, and the Republic of Congo using forest inventory data from 56,445 0.5-ha plots. Spatial variation of environmental (climate, topography and geology) and historical factors (human disturbance) were quantified from maps and satellite records. Four key functional traits (leaf phenology, shade tolerance, wood density, and maximum growth rate) were extracted from the literature. The geological substrate was of major importance for the distribution of the focal species, while climate and past human disturbances had a significant but lesser impact. Species distribution patterns were significantly related to functional traits. Species associated with sandy soils typical of sandstone and alluvium were characterized by slow growth rates, shade tolerance, evergreen leaves, and high wood density, traits allowing persistence on resource-poor soils. In contrast, fast-growing pioneer species rarely occurred on sandy soils, except for Lophira alata. Conclusions/Significance The results indicate strong environmental filtering due to differential soil resource availability across geological substrates. Additionally, long-term human disturbances in resource-rich areas may have accentuated the observed patterns of species and trait distributions. Trait differences across geological substrates imply pronounced differences in population and ecosystem processes, and call for different conservation and management strategies.

Visual assessment of wilting as a measure of leaf water potential and seedling drought survival

Rainfall and soil moisture availability vary greatly both spatially and temporally. They are prime factors influencing species distribution patterns, diversity and habitat associations in a range of biomes, and limit primary productivity in many natural ecosystems, as well as in forestry and agricultural systems (Hawkins et al. 2003, Kozlowski & Pallardy 1997, Lieth 1975). Projections of drying trends, and increased frequency and intensity of drought events with climate and land-use changes (Hulme & Viner 1998, IPCC 2001) have fuelled an increased interest in the role of drought in determining the structure and function of natural and managed forest systems (Allen & Breshears 1998, Miles et al. 2004). Such projections accentuate the need to assess, understand and predict plant reactions to drought, as well as soil moisture variation at different scales.