Frans Bongers

The worldwide leaf economics spectrum

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

The ecology of lianas and their role in forests

Recent studies have demonstrated the increasingly important role of lianas (woody vines) in forest regeneration, species diversity and ecosystem-level processes, particularly in the tropics. Mechanisms responsible for the maintenance of liana species diversity could yield new insights into the maintenance of overall species diversity. Lianas contribute to forest regeneration and competition, not only by competing directly with trees, but also by differentially affecting tree species and thus changing how trees compete among themselves. In addition, they contribute considerably to ecosystem-level processes, such as whole-forest transpiration and carbon sequestration. As the rate of tropical forest disturbance increases, they are likely to increase in relative abundance throughout the tropics and the importance of lianas to many aspects of forest dynamics will grow.

Rates of change in tree communities of secondary Neotropical forests following major disturbances

Rates of change in tree communities following major disturbances are determined by a complex set of interactions between local site factors, landscape history and structure, regional species pools and species life histories. Our analysis focuses on vegetation change following abandonment of agricultural fields or pastures, as this is the most extensive form of major disturbance in Neotropical forests. We consider five tree community attributes: stem density, basal area, species density, species richness and species composition. We describe two case studies, in northeastern Costa Rica and Chiapas, Mexico, where both chronosequence and annual tree dynamics studies are being applied. These case studies show that the rates of change in tree communities often deviate from chronosequence trends. With respect to tree species composition, sites of different ages differ more than a single site followed over time through the same age range. Dynamic changes in basal area within stands, on the other hand, generally followed chronosequence trends. Basal area accumulation was more linked with tree growth rates than with net changes in tree density due to recruitment and mortality. Stem turnover rates were poor predictors of species turnover rates, particularly at longer time-intervals. Effects of the surrounding landscape on tree community dynamics within individual plots are poorly understood, but are likely to be important determinants of species accumulation rates and relative abundance patterns.

ARCHITECTURE OF 54 MOIST-FOREST TREE SPECIES: TRAITS, TRADE-OFFS, AND FUNCTIONAL GROUPS

Tree architecture is an important determinant of the height extension, light capture, and mechanical stability of trees, and it allows species to exploit the vertical height gradient in the forest canopy and horizontal light gradients at the forest floor. Tropical tree species partition these gradients through variation in adult stature (Hmax) and light demand. In this study we compare 22 architectural traits for 54 Bolivian moist-forest tree species. We evaluate how architectural traits related to Hmax vary with tree size, and we present a conceptual scheme in which we combine the two axes into four different functional groups. Interspecific correlations between architecture and Hmax varied strongly from negative to positive, depending on the reference sizes used. Stem height was positively related to Hmax at larger reference diameters (14-80 cm). Species height vs. diameter curves often flattened toward their upper ends in association with reproductive maturity for species of all sizes. Thus, adult understory trees were typically shorter than similar-diameter juveniles of larger species. Crown area was negatively correlated with Hmax at small reference heights and positively correlated at larger reference heights (15-34 m). Wide crowns allow the small understory species to intercept light over a large area at the expense of a reduced height growth. Crown length was negatively correlated with Hmax at intermediate reference heights (4-14 m). A long crown enables small understory species to maximize light interception in a light-limited environment. Light-demanding species were characterized by orthotropic stems and branches, large leaves, and a monolayer leaf arrangement. They realized an efficient height growth through the formation of narrow and shallow crowns. Light demand turned out to be a much stronger predictor of tree architecture than Hmax, probably because of the relatively low, open, and semi-evergreen canopy at the research site. The existence of four functional groups (shade-tolerant, partial-shade-tolerant, and long- and short-lived pioneer) was confirmed by the principal component and discriminant analysis. Both light demand and Hmax capture the major variation in functional traits found among tropical rain forest tree species, and the two-way classification scheme provides a straightforward model to understand niche differentiation in tropical forests.

Seed mass and mast seeding enhance dispersal by a neotropical scatter-hoarding rodent

Many tree species that depend on scatter-hoarding animals for seed dispersal produce massive crops of large seeds at irregular intervals. Mast seeding and large seed size in these species have been explained as adaptations to increase animal dispersal and reduce predation. We studied how seed size and seed abundance simultaneously influenced seed dispersal and predation by scatter-hoarding rodents in the large-seeded rain forest tree Carapa procera (Meliaceae) in French Guiana. We individually tracked the fates of 3000 seeds, using remote video monitoring and thread-marking. Seed size was manipulated by broadly varying intraspecific seed mass, whereas effects of seed abundance were examined by tracking seeds in three seed-rich years and two seed-poor years. The hypotheses, that seed mass and seed abundance both enhance dispersal success and that seed abundance reinforces the effect of seed mass, were supported by the results. Most seeds were removed by the scatter-hoarding rodent red acouchy (Myoprocta acouchy) and subsequently were buried in scattered, single-seeded caches up to distances >100 m. Seeds that were not removed failed to establish seedlings. Seed removal was slower, pre-removal seed predation was greater, and seed dispersal was less far in seed-rich years than in seed-poor years, suggesting poorer dispersal under seed abundance. However, this was more than counterbalanced by a disproportionally greater survival of cached seeds in seed-rich years. The per capita probability of seed survival and seedling establishment was at least 4½ times greater under seed abundance. Large seeds were removed faster, were more likely to be scatter-hoarded, and were dispersed farther away than smaller ones, resulting in a higher probability of seedling establishment for larger seeds. Size discrimination was greater under seed abundance, albeit only during seed removal. Overall, large seeds shed in rich years had the highest probability of seedling establishment. Hence, both larger seed size and greater seed abundance stimulate rodents to act more as dispersers and less as predators of seeds. We conclude that scatter-hoarding rodents can select for both large seed crops and large seeds, which may reinforce mast seeding.

Increasing liana abundance and biomass in tropical forests: emerging patterns and putative mechanisms.

Tropical forests are experiencing large-scale structural changes, the most apparent of which may be the increase in liana (woody vine) abundance and biomass. Lianas permeate most lowland tropical forests, where they can have a huge effect on tree diversity, recruitment, growth and survival, which, in turn, can alter tree community composition, carbon storage and carbon, nutrient and water fluxes. Consequently, increasing liana abundance and biomass have potentially profound ramifications for tropical forest composition and functioning. Currently, eight studies support the pattern of increasing liana abundance and biomass in American tropical and subtropical forests, whereas two studies, both from Africa, do not. The putative mechanisms to explain increasing lianas include increasing evapotranspirative demand, increasing forest disturbance and turnover, changes in land use and fragmentation and elevated atmospheric CO₂. Each of these mechanisms probably contributes to the observed patterns of increasing liana abundance and biomass, and the mechanisms are likely to be interrelated and synergistic. To determine whether liana increases are occurring throughout the tropics and to determine the mechanisms responsible for the observed patterns, a widespread network of large-scale, long-term monitoring plots combined with observational and manipulative studies that more directly investigate the putative mechanisms are essential.

Biomass resilience of Neotropical secondary forests

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha−1), corresponding to a net carbon uptake of 3.05 Mg C ha−1 yr−1, 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha−1) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Ecological differentiation in xylem cavitation resistance is associated with stem and leaf structural traits

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P₅₀) related to dry season leaf water potentials and stem and leaf traits. P₅₀-values ranged from -0.8 to -6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade-tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber-value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade-off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P₅₀, but pioneers and deciduous species had smaller hydraulic safety margins than shade-tolerants and evergreens. A trade-off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species.

Biodiversity of West African Forests: An Ecological Atlas of Woody Plant Species

The rain forests of West Africa have been designated as one of the worlds hotspots of biodiversity. They extend from Ghana to Senegal and are referred to as the Upper Guinean forests. Because of their isolated position, they harbor a large number of rare and endemic animal and plant species.This book focuses on the biodiversity and ecology of these forests. It analyzes the factors that give rise to biodiversity and structure tropical plant communities. It also includes an atlas with ecological profiles of rare plant species and large timber species.

The intermediate disturbance hypothesis applies to tropical forests, but disturbance contributes little to tree diversity

The intermediate disturbance hypothesis (IDH) predicts local species diversity to be maximal at an intermediate level of disturbance. Developed to explain species maintenance and diversity patterns in species-rich ecosystems such as tropical forests, tests of IDH in tropical forest remain scarce, small-scale and contentious. We use an unprecedented large-scale dataset (2504 one-hectare plots and 331,567 trees) to examine whether IDH explains tree diversity variation within wet, moist and dry tropical forests, and we analyse the underlying mechanism by determining responses within functional species groups. We find that disturbance explains more variation in diversity of dry than wet tropical forests. Pioneer species numbers increase with disturbance, shade-tolerant species decrease and intermediate species are indifferent. While diversity indeed peaks at intermediate disturbance levels little variation is explained outside dry forests, and disturbance is less important for species richness patterns in wet tropical rain forests than previously thought.