Mart Vlam

Tropical forests and global change: filling knowledge gaps.

Tropical forests will experience major changes in environmental conditions this century. Understanding their responses to such changes is crucial to predicting global carbon cycling. Important knowledge gaps exist: the causes of recent changes in tropical forest dynamics remain unclear and the responses of entire tropical trees to environmental changes are poorly understood. In this Opinion article, we argue that filling these knowledge gaps requires a new research strategy, one that focuses on trees instead of leaves or communities, on long-term instead of short-term changes, and on understanding mechanisms instead of documenting changes. We propose the use of tree-ring analyses, stable-isotope analyses, manipulative field experiments, and well-validated simulation models to improve predictions of forest responses to global change.

Detecting long‐term growth trends using tree rings: a critical evaluation of methods

Tree-ring analysis is often used to assess long-term trends in tree growth. A variety of growth-trend detection methods (GDMs) exist to disentangle age/size trends in growth from long-term growth changes. However, these detrending methods strongly differ in approach, with possible implications for their output. Here, we critically evaluate the consistency, sensitivity, reliability and accuracy of four most widely used GDMs: conservative detrending (CD) applies mathematical functions to correct for decreasing ring widths with age; basal area correction (BAC) transforms diameter into basal area growth; regional curve standardization (RCS) detrends individual tree-ring series using average age/size trends; and size class isolation (SCI) calculates growth trends within separate size classes. First, we evaluated whether these GDMs produce consistent results applied to an empirical tree-ring data set of Melia azedarach, a tropical tree species from Thailand. Three GDMs yielded similar results - a growth decline over time - but the widely used CD method did not detect any change. Second, we assessed the sensitivity (probability of correct growth-trend detection), reliability (100% minus probability of detecting false trends) and accuracy (whether the strength of imposed trends is correctly detected) of these GDMs, by applying them to simulated growth trajectories with different imposed trends: no trend, strong trends (-6% and +6% change per decade) and weak trends (-2%, +2%). All methods except CD, showed high sensitivity, reliability and accuracy to detect strong imposed trends. However, these were considerably lower in the weak or no-trend scenarios. BAC showed good sensitivity and accuracy, but low reliability, indicating uncertainty of trend detection using this method. Our study reveals that the choice of GDM influences results of growth-trend studies. We recommend applying multiple methods when analysing trends and encourage performing sensitivity and reliability analysis. Finally, we recommend SCI and RCS, as these methods showed highest reliability to detect long-term growth trends.

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate-vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO2 -fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest-stand level, but insights into species-specific growth changes - that ultimately determine community-level responses - are lacking. Here, we analyse species-specific growth changes on a centennial scale, using growth data from tree-ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size-class isolation) growth-trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8-10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large-scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO2 would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.

Tree growth variation in the tropical forest: understanding effects of temperature, rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree-ring study over a 30-year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO2 ) in different combinations to estimate the contribution of each climate factor in explaining the inter-annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter-annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and - to a lesser extent - by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter-annual fluctuations in rainfall. Minimum temperature and atmospheric CO2 concentration did not significantly contribute to explaining the inter-annual variation in stem growth. Our innovative approach - combining a simulation model with historical data on tree-ring growth and climate - allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.

Does biomass growth increase in the largest trees? Flaws, fallacies and alternative analyses

Summary The longstanding view that biomass growth in trees typically follows a rise-and-fall unimodal pattern has been challenged by studies concluding that biomass growth increases with size even among the largest stems in closed forests and in open competition-free environments. We highlight challenges and pitfalls that influence such views and interpretations. The ability to observe and calibrate biomass change in large stems requires adequate data regarding these specific stems. Data checking and control procedures can bias estimates of biomass growth and generate false increases with stem size. It is important to distinguish aggregate and individual-level trends, as these are distinct: a failure to distinguish how biomass growth varies among and within stems results in flawed interpretations. Our assessment of biomass growth in 706 tropical forest stems indicates that biomass growth patterns often plateau for extended periods, with no significant difference in the number of stems indicating positive and negative trends in all but one of the 14 species. Nonetheless, when comparing individual growth during the most recent five years, 13 out of our 14 species indicate that biomass growth increases with size even among the largest sizes. Thus, individual and aggregate patterns of biomass growth with size are distinct. Claims concerning general biomass growth patterns for large trees remain unconvincing. We suggest how future studies can improve our knowledge of growth patterns in and among large trees. This article is protected by copyright. All rights reserved.

15N in tree rings as a bio-indicator of changing nitrogen cycling in tropical forests: an evaluation at three sites using two sampling methods

Anthropogenic nitrogen deposition is currently causing a more than twofold increase of reactive nitrogen input over large areas in the tropics. Elevated 15N abundance (δ15N) in the growth rings of some tropical trees has been hypothesized to reflect an increased leaching of 15N-depleted nitrate from the soil, following anthropogenic nitrogen deposition over the last decades. To find further evidence for altered nitrogen cycling in tropical forests, we measured long-term δ15N values in trees from Bolivia, Cameroon, and Thailand. We used two different sampling methods. In the first, wood samples were taken in a conventional way: from the pith to the bark across the stem of 28 large trees (the “radial” method). In the second, δ15N values were compared across a fixed diameter (the “fixed-diameter” method). We sampled 400 trees that differed widely in size, but measured δ15N in the stem around the same diameter (20 cm dbh) in all trees. As a result, the growth rings formed around this diameter differed in age and allowed a comparison of δ15N values over time with an explicit control for potential size-effects on δ15N values. We found a significant increase of tree-ring δ15N across the stem radius of large trees from Bolivia and Cameroon, but no change in tree-ring δ15N values over time was found in any of the study sites when controlling for tree size. This suggests that radial trends of δ15N values within trees reflect tree ontogeny (size development). However, for the trees from Cameroon and Thailand, a low statistical power in the fixed-diameter method prevents to conclude this with high certainty. For the trees from Bolivia, statistical power in the fixed-diameter method was high, showing that the temporal trend in tree-ring δ15N values in the radial method is primarily caused by tree ontogeny and unlikely by a change in nitrogen cycling. We therefore stress to account for tree size before tree-ring δ15N values can be properly interpreted.

Tree age distributions reveal large-scale disturbance-recovery cycles in three tropical forests

Over the past few decades there has been a growing realization that a large share of apparently ‘virgin’ or ‘old-growth’ tropical forests carries a legacy of past natural or anthropogenic disturbances that have a substantial effect on present-day forest composition, structure and dynamics. Yet, direct evidence of such disturbances is scarce and comparisons of disturbance dynamics across regions even more so. Here we present a tree-ring based reconstruction of disturbance histories from three tropical forest sites in Bolivia, Cameroon, and Thailand. We studied temporal patterns in tree regeneration of shade-intolerant tree species, because establishment of these trees is indicative for canopy disturbance. In three large areas (140–300 ha), stem disks and increment cores were collected for a total of 1154 trees (>5 cm diameter) from 12 tree species to estimate the age of every tree. Using these age estimates we produced population age distributions, which were analyzed for evidence of past disturbance. Our approach allowed us to reconstruct patterns of tree establishment over a period of around 250 years. In Bolivia, we found continuous regeneration rates of three species and a peaked age distribution of a long-lived pioneer species. In both Cameroon and Thailand we found irregular age distributions, indicating strongly reduced regeneration rates over a period of 10–60 years. Past fires, windthrow events or anthropogenic disturbances all provide plausible explanations for the reported variation in tree age across the three sites. Our results support the recent idea that the long-term dynamics of tropical forests are impacted by large-scale disturbance-recovery cycles, similar to those driving temperate forest dynamics.

Sapwood allocation in tropical trees: a test of hypotheses

Carbon allocation to sapwood in tropical canopy trees is a key process determining forest carbon sequestration, and is at the heart of tree growth and dynamic global vegetation models (DGVM). Several allocation hypotheses exist including those applying assumptions on fixed allocation, pipe model, and hierarchical allocation between plant organs. We use a tree growth model (IBTREE) to evaluate these hypotheses by comparing simulated sapwood growth with 30 year tree ring records of the tropical long-lived tree Toona ciliata M. Roem. in Thailand. Simulated annual variation in wood production varied among hypotheses. Observed and simulated growth patterns matched most closely (r2=0.70) when hierarchical allocation was implemented, with low priority for sapwood. This allocation method showed realistic results with respect to reserve dynamics, partitioning and productivity and was the only one able to capture the large annual variation in tree ring width. Consequently, this method might also explain the large temporal variation in diameter growth and the occurrence of missing rings often encountered in other tropical tree species. Overall, our results show that sapwood growth is highly sensitive to allocation principles, and that allocation assumptions may greatly influence estimated carbon sequestration of tropical forests under climatic change.