Ecology Letters (2011) 14: 397-406 ABSTRACT: 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.
We compared the leaf traits and plant performance of 53 co-occurring tree species in a semi-evergreen tropical moist forest community. The species differed in all leaf traits analyzed: leaf life span ...varied 11-fold among species, specific leaf area 5-fold, mass-based nitrogen 3-fold, mass-based assimilation rate 13-fold, mass-based respiration rate 15-fold, stomatal conductance 8-fold, and photosynthetic water use efficiency 4-fold. Photosynthetic traits were strongly coordinated, and specific leaf area predicted mass-based rates of assimilation and respiration; leaf life span predicted many other leaf characteristics. Leaf traits were closely associated with growth, survival, and light requirement of the species. Leaf investment strategies varied on a continuum trading off short-term carbon gain against long-term leaf persistence that, in turn, is linked to variation in whole-plant growth and survival. Leaf traits were good predictors of plant performance, both in gaps and in the forest understory. High growth in gaps is promoted by cheap, short-lived, and physiologically active leaves. High survival in the forest understory is enhanced by the formation of long-lived well protected leaves that reduce biomass loss by herbivory, mechanical disturbance, or leaf turnover. Leaf traits underlay this growth--survival trade-off; species with short-lived, physiologically active leaves have high growth but low survival. This continuum in leaf traits, through its effect on plant performance, in turn gives rise to a continuum in species' light requirements.
Understanding the relationship between stand-level tree diversity and productivity has the potential to inform the science and management of forests. History shows that plant diversity-productivity ...relationships are challenging to interpret—and this remains true for the study of forests using non-experimental field data. Here we highlight pitfalls regarding the analyses and interpretation of such studies. We examine three themes: 1) the nature and measurement of ecological productivity and related values; 2) the role of stand history and disturbance in explaining forest characteristics; and 3) the interpretation of any relationship. We show that volume production and true productivity are distinct, and neither is a demonstrated proxy for economic values. Many stand characteristics, including diversity, volume growth and productivity, vary intrinsically with succession and stand history. We should be characterising these relationships rather than ignoring or eliminating them. Failure to do so may lead to misleading conclusions. To illustrate, we examine the study which prompted our concerns —Liang et al. (Science 354:aaf8957, 2016)— which developed a sophisticated global analysis to infer a worldwide positive effect of biodiversity (tree species richness) on “forest productivity” (stand level wood volume production). Existing data should be able to address many of our concerns. Critical evaluations will improve understanding.
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 ($H_{\text{max}}$) 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$H_{\text{max}}$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$H_{\text{max}}$varied strongly from negative to positive, depending on the reference sizes used. Stem height was positively related to$H_{\text{max}}$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$H_{\text{max}}$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$H_{\text{max}}$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$H_{\text{max}}$, 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$H_{\text{max}}$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.
Over half of the world's forests are disturbed, and the rate at which ecosystem processes recover after disturbance is important for the services these forests can provide. We analyze the drivers' ...underlying changes in rates of key ecosystem processes (biomass productivity, litter productivity, actual litter decomposition, and potential litter decomposition) during secondary succession after shifting cultivation in wet tropical forest of Mexico.
We test the importance of three alternative drivers of ecosystem processes: vegetation biomass (vegetation quantity hypothesis), community-weighted trait mean (mass ratio hypothesis), and functional diversity (niche complementarity hypothesis) using structural equation modeling. This allows us to infer the relative importance of different mechanisms underlying ecosystem process recovery.
Ecosystem process rates changed during succession, and the strongest driver was aboveground biomass for each of the processes. Productivity of aboveground stem biomass and leaf litter as well as actual litter decomposition increased with initial standing vegetation biomass, whereas potential litter decomposition decreased with standing biomass. Additionally, biomass productivity was positively affected by community-weighted mean of specific leaf area, and potential decomposition was positively affected by functional divergence, and negatively by community-weighted mean of leaf dry matter content.
Our empirical results show that functional diversity and community-weighted means are of secondary importance for explaining changes in ecosystem process rates during tropical forest succession. Instead, simply, the amount of vegetation in a site is the major driver of changes, perhaps because there is a steep biomass buildup during succession that overrides more subtle effects of community functional properties on ecosystem processes. We recommend future studies in the field of biodiversity and ecosystem functioning to separate the effects of vegetation quality (community-weighted mean trait values and functional diversity) from those of vegetation quantity (biomass) on ecosystem processes and services.
Many studies suggest that biodiversity may be particularly important for ecosystem multifunctionality, because different species with different traits can contribute to different functions. Support, ...however, comes mostly from experimental studies conducted at small spatial scales in low-diversity systems. Here, we test whether different species contribute to different ecosystem functions that are important for carbon cycling in a high-diversity human-modified tropical forest landscape in Southern Mexico. We quantified aboveground standing biomass, primary productivity, litter production, and wood decomposition at the landscape level, and evaluated the extent to which tree species contribute to these ecosystem functions. We used simulations to tease apart the effects of species richness, species dominance and species functional traits on ecosystem functions. We found that dominance was more important than species traits in determining a species' contribution to ecosystem functions. As a consequence of the high dominance in human-modified landscapes, the same small subset of species mattered across different functions. In human-modified landscapes in the tropics, biodiversity may play a limited role for ecosystem multifunctionality due to the potentially large effect of species dominance on biogeochemical functions. However, given the spatial and temporal turnover in species dominance, biodiversity may be critically important for the maintenance and resilience of ecosystem functions.
Forest restoration increases global forest area and ecosystem services such as primary productivity and carbon storage. How tree species functional composition impacts the provisioning of these ...services as forests develop is sparsely studied. We used 10-year data from 478 plots with 191,200 trees in a forest biodiversity experiment in subtropical China to assess the relationship between community productivity and community-weighted mean (CWM) or functional diversity (FD) values of 38 functional traits. We found that effects of FD values on productivity became larger than effects of CWM values after 7 years of forest development and that the FD values also became more reliable predictors of productivity than the CWM values. In contrast to CWM, FD values consistently increased productivity across ten different species-pool subsets. Our results imply that to promote productivity in the long term it is imperative for forest restoration projects to plant multispecies communities with large functional diversity.
•We evaluated the performance of 24 tree species in the wet tropics in Mexico.•We relate performance to 13 functional leaf, reproductive and architectural traits.•Screening after 1y may suffice to ...select the best performing species after 3.5y.•Species with small seeds and large crowns are good candidates for pasture plantings.
Functional traits may predict tree growth rates and survival in plantings aimed to accelerate natural succession in pastures. We evaluate the growth and survival of 24 tree species used for forest restoration in pastures in the wet tropics in Mexico for 42months. We relate their performance to 13 underlying functional traits that are important for leaf display, tree architecture and reproduction. Pioneers realized both faster growth rates and higher survival than non-pioneer species. No consistent ontogenetic cross-overs in species growth and survival were found during 42months, which implies that species performance early after planting (12months) predicted performance well after 42months, probably because environmental conditions changed little over time in these early-successional pastures. Nine out of 12 functional traits were good predictors of growth rates and/or survival measures across all 24 species or for pioneer or non-pioneer species separately. Growth rates and survival were positively related to crown size and negatively to seed mass, but they were – surprisingly – not related to the leaf traits we measured. Multivariate plant trait axes explained species growth rates and survival better than individual traits only for the non-pioneer species group. Selection of additional traits may be needed for better predictions of overall performance of restoration plantings. Combining the basic science of functional ecology with the applied goals of forest management and restoration allow us to select the best mid-term performing tree species for planting in pastures using a short-term screening of growth and survival rates. Pioneers and non-pioneer species with small seeds and large crown length are good candidates for plantings in pastures since they have the best mid-term performance.
ABSTRACT
Secondary forests are increasingly important components of human‐modified landscapes in the tropics. Successional pathways, however, can vary enormously across and within landscapes, with ...divergent regrowth rates, vegetation structure and species composition. While climatic and edaphic conditions drive variations across regions, land‐use history plays a central role in driving alternative successional pathways within human‐modified landscapes. How land use affects succession depends on its intensity, spatial extent, frequency, duration and management practices, and is mediated by a complex combination of mechanisms acting on different ecosystem components and at different spatial and temporal scales. We review the literature aiming to provide a comprehensive understanding of the mechanisms underlying the long‐lasting effects of land use on tropical forest succession and to discuss its implications for forest restoration. We organize it following a framework based on the hierarchical model of succession and ecological filtering theory. This review shows that our knowledge is mostly derived from studies in Neotropical forests regenerating after abandonment of shifting cultivation or pasture systems. Vegetation is the ecological component assessed most often. Little is known regarding how the recovery of belowground processes and microbiota communities is affected by previous land‐use history. In published studies, land‐use history has been mostly characterized by type, without discrimination of intensity, extent, duration or frequency. We compile and discuss the metrics used to describe land‐use history, aiming to facilitate future studies. The literature shows that (i) species availability to succession is affected by transformations in the landscape that affect dispersal, and by management practices and seed predation, which affect the composition and diversity of propagules on site. Once a species successfully reaches an abandoned field, its establishment and performance are dependent on resistance to management practices, tolerance to (modified) soil conditions, herbivory, competition with weeds and invasive species, and facilitation by remnant trees. (ii) Structural and compositional divergences at early stages of succession remain for decades, suggesting that early communities play an important role in governing further ecosystem functioning and processes during succession. Management interventions at early stages could help enhance recovery rates and manipulate successional pathways. (iii) The combination of local and landscape conditions defines the limitations to succession and therefore the potential for natural regeneration to restore ecosystem properties effectively. The knowledge summarized here could enable the identification of conditions in which natural regeneration could efficiently promote forest restoration, and where specific management practices are required to foster succession. Finally, characterization of the landscape context and previous land‐use history is essential to understand the limitations to succession and therefore to define cost‐effective restoration strategies. Advancing knowledge on these two aspects is key for finding generalizable relations that will increase the predictability of succession and the efficiency of forest restoration under different landscape contexts.
ABSTRACT
Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified ...tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.
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