Pieter A. Zuidema

Relating tree growth to rainfall in Bolivian rain forests: a test for six species using tree ring analysis

Many tropical regions show one distinct dry season. Often, this seasonality induces cambial dormancy of trees, particularly if these belong to deciduous species. This will often lead to the formation of annual rings. The aim of this study was to determine whether tree species in the Bolivian Amazon region form annual rings and to study the influence of the total amount and seasonal distribution of rainfall on diameter growth. Ring widths were measured on stem discs of a total of 154 trees belonging to six rain forest species. By correlating ring width and monthly rainfall data we proved the annual character of the tree rings for four of our study species. For two other species the annual character was proved by counting rings on trees of known age and by radiocarbon dating. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall in certain periods of the year, indicating that rainfall plays a major role in tree growth. Three species showed a strong relationship with rainfall at the beginning of the rainy season, while one species is most sensitive to the rainfall at the end of the previous growing season. These results clearly demonstrate that tree ring analysis can be successfully applied in the tropics and that it is a promising method for various research disciplines.

Demography of the Brazil nut tree ( Bertholletia excelsa ) in the Bolivian Amazon: impact of seed extraction on recruitment and population dynamics

A demographic study was carried out on Bertholletia excelsa, the Brazil nut tree, in two primary forest sites in Northern Bolivia where Brazil nuts have been harvested for several decades. In spite of the large proportion (93%) of seeds that are harvested, reasonable densities of recently emerged seedlings were found. Seeds of Bertholletia are contained in woody fruits that are primarily opened by agoutis. Most fruits are left untouched on the forest floor for 1-2 y before they are opened, possibly due to high energetic costs of fruit opening just after fruit fall. However, the proportion of viable seeds is strongly reduced in older fruits. Growth in diameter at breast height (dbh) was low for pole-sized trees ( 100 cm dbh) and peaked for intermediate-sized trees (30-60 cm). These trees often attained a growth rate of > 1.5 cm y −1 , which is high compared with other non-pioneer tropical trees. This, and the strong growth response to increased light availability found for seedlings and saplings, suggest that Bertholletia excelsa can be classified as a gap-dependent species. Matrix popula- tion models were constructed for both study populations. Population growth rates (λ) were close to one, and were most sensitive to persistence in one size category. Age estimates revealed that age at first reproduction (at dbh > 60 cm) amounts to over 120 y, and age in the last category (dbh > 160 cm) to almost 300 y. Given the continuous rejuvenation of the population, the stable population size, the high age at maturity and the long reproductive period, it is concluded that current levels of Brazil nut extraction may be sustained at least for several decades and perhaps for even longer periods.

Dendroecology in the tropics: a review

Over the last decade the field of tropical dendroecology has developed rapidly and major achievements have been made. We reviewed the advances in three main themes within the field. First, long chronologies for tropical tree species were constructed which allowed climate reconstructions, revealed sources of climatic variation and clarified climate–growth relations. Other studies combined tree-ring data and stable isotope (13C and 18O) measurements to evaluate the response of tropical trees to climatic variation and changes. A second set of studies assessed long-term growth patterns of individual trees throughout their life. These studies enhanced the understanding of growth trajectories to the canopy, quantified autocorrelated tree growth and yielded new estimates of tree ages. Such studies were also used to reconstruct the disturbance history of tropical forests. The last set of studies applied tree-ring data to growth models. Tree-ring data can replace diameter measurements from research plots, provide additional information to construct population models, improve timber yield models and validate model output. Based on our review, we propose two main directions for future research. (1) An evaluation of the causes and consequences of growth variation within and among trees and their relation to environmental variation. Studies evaluating this directly contribute to improved understanding of tropical tree ecology. (2) The simultaneous measurement of widths and stable isotope fractions in tree rings offers the potential to study responses of trees to climatic change. Given the major role of tropical forests in the global carbon cycle, knowing these responses is of high priority.

Forest fragmentation and biodiversity: the case for intermediate-sized conservation areas

Understanding the effects of forest fragmentation on biodiversity is essential for successful and efficient forest conservation. Four factors may cause loss of biodiversity in forest fragments: the effect of non-random sampling of the original forest, reduced forest size, isolation and edge effects. A review of 58 papers on effects of forest fragmentation reveals that general conclusions from fragmentation research are biased due to a focus on birds, on size-effects rather than isolation, and on species presence rather than population sizes. Perhaps the most important finding is that current knowledge on fragmentation effects is based mainly on studies in small fragments (10 000 - 100 000 ha) and should focus on the biological and human-induced processes which determine species presistence.

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.

Fast–slow continuum and reproductive strategies structure plant life-history variation worldwide

Significance Schedules of survival, growth, and reproduction define life-history strategies across species. Understanding how life-history strategies are structured is fundamental to our understanding of the evolution, abundance, and distribution of species. We found that life-history strategies of 418 plant species worldwide are explained by an axis representing the pace of life and another representing the wide range of reproductive strategies. This framework predicts responses to perturbations and long-term population performance, showing great promise as a predictive tool for plant population responses to environmental change. The identification of patterns in life-history strategies across the tree of life is essential to our prediction of population persistence, extinction, and diversification. Plants exhibit a wide range of patterns of longevity, growth, and reproduction, but the general determinants of this enormous variation in life history are poorly understood. We use demographic data from 418 plant species in the wild, from annual herbs to supercentennial trees, to examine how growth form, habitat, and phylogenetic relationships structure plant life histories and to develop a framework to predict population performance. We show that 55% of the variation in plant life-history strategies is adequately characterized using two independent axes: the fast–slow continuum, including fast-growing, short-lived plant species at one end and slow-growing, long-lived species at the other, and a reproductive strategy axis, with highly reproductive, iteroparous species at one extreme and poorly reproductive, semelparous plants with frequent shrinkage at the other. Our findings remain consistent across major habitats and are minimally affected by plant growth form and phylogenetic ancestry, suggesting that the relative independence of the fast–slow and reproduction strategy axes is general in the plant kingdom. Our findings have similarities with how life-history strategies are structured in mammals, birds, and reptiles. The position of plant species populations in the 2D space produced by both axes predicts their rate of recovery from disturbances and population growth rate. This life-history framework may complement trait-based frameworks on leaf and wood economics; together these frameworks may allow prediction of responses of plants to anthropogenic disturbances and changing environments.

Do Persistently Fast‐Growing Juveniles Contribute Disproportionately to Population Growth? A New Analysis Tool for Matrix Models and Its Application to Rainforest Trees

Plants and animals often exhibit strong and persistent growth variation among individuals within a species. Persistently fast‐growing individuals have a higher chance of reaching reproductive size, do so at a younger age, and therefore contribute disproportionately to population growth (λ). Here we introduce a new approach to quantify this “fast‐growth effect.” We propose using age‐size‐structured matrix models in which persistently fast and slow growers are distinguished as they occur in relatively young and old age classes for a given size category. Life‐cycle pathways involving fast growth can then be identified, and their contribution to λ is quantified through loop analysis. We applied this approach to an example species, the tropical rainforest tree Cedrela odorata, that shows persistent growth variation among individuals. Loop analysis showed that juvenile trees reaching the 10‐cm diameter class at below‐median age contributed twice as much to λ as slow juvenile growers. Fast growth to larger‐diameter categories also contributed disproportionately to λ. The results were robust to changes in parameter values and life‐history trade‐offs. These results show that the fast‐growth effect can be strong in long‐lived species. Persistent growth differences among individuals should therefore be accommodated for in demographic models and life‐history studies.

Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.

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.

Incorporating persistent tree growth differences increases estimates of tropical timber yield

Sustainable tropical forest management using selective logging, like any natural resource management system, requires reliable estimates of future yields. Commonly, timber yield predictions are calculated using the average of diameter growth rates in the population and thus ignore variation among trees. Using tree-ring analysis for three Amazonian timber species, we show that strong and persistent growth differences exist among individual trees. These differences substantially affect predictions of future timber yield. After incorporating realistic growth variation, we calculated timber yields that were 36–50% higher than those calculated using average growth rates. Even so, the regrowth of timber volume in a 20-year interval between logging events is low (18–24%) due to the old age of trees that are logged at first harvest. Improved yield estimates are important to evaluate the economic viability of sustainable forest management, a hotly debated issue in conservation circles.