1. Natural populations deliver a wide range of products that provide income for millions of peopl... more 1. Natural populations deliver a wide range of products that provide income for millions of people and need to be exploited sustainably. Large heterogeneity in individual performance within these exploited populations has the potential to improve population recovery after exploitation and thus help sustain yields over time. 2. We explored the potential of using individual heterogeneity to design smarter harvest schemes, by sparing individuals that contribute most to future productivity and population growth, using the understorey palm Chamaedorea elegans as a model system. Leaves of this palm are an important non-timber forest product and longterm inter-individual growth variability can be evaluated from internode lengths. 3. We studied a population of 830 individuals, half of which was subjected to a 67% defoliation treatment for 3 years. We measured effects of defoliation on vital rates and leaf size-a trait that determines marketability. We constructed integral projection models in which vital rates depended on stem length, past growth rate and defoliation and evaluated transient population dynamics to quantify population development and leaf yield. We then simulated scenarios in which we spared individuals that were either most important for population growth or had leaves smaller than marketable size. 4. Individuals varying in size or past growth rate responded similar to leaf harvesting in terms of growth and reproduction. By contrast, a reduction in survival chance was smaller in large individuals than in small ones. Simulations showed that harvestinduced population decline was greatly reduced when sparing individuals from size and past growth classes that contributed most to population growth. Under this scenario, cumulative leaf harvest over 20 years was somewhat reduced, but longterm leaf production was sustained. A threefold increase in leaf yield was generated when individuals with small leaves were spared. This study demonstrates the potential to create smarter systems of palm leaf harvest by accounting for individual heterogeneity within exploited populations. Sparing individuals that contribute most to population growth ensured sustained leaf production over time. The concepts and methods presented This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
Among tropical forests, lianas are predicted to have a growth advantage over trees during seasona... more Among tropical forests, lianas are predicted to have a growth advantage over trees during seasonal drought, with substantial implications for tree and forest dynamics. We tested the hypotheses that lianas maintain higher water status than trees during seasonal drought and that lianas maximize leaf cover to match high, dry-season light conditions, while trees are more limited by moisture availability during the dry season. We monitored the seasonal dynamics of predawn and midday leaf water potentials and leaf phenology for branches of 16 liana and 16 tree species in the canopies of two lowland tropical forests with contrasting rainfall regimes in Panama. In a wet, weakly seasonal forest, lianas maintained higher water balance than trees and maximized their leaf cover during dry-season conditions, when light availability was high, while trees experienced drought stress. In a drier, strongly seasonal forest, lianas and trees displayed similar dry season reductions in leaf cover following strong decreases in soil water availability. Greater soil moisture availability and a higher capacity to maintain water status allow lianas to maintain the turgor potentials that are critical for plant growth in a wet and weakly seasonal forest but not in a dry and strongly seasonal forest.
Shifting cultivation is the main land-use system transforming landscapes in riverine Amazonia. In... more Shifting cultivation is the main land-use system transforming landscapes in riverine Amazonia. Increased concentration of the human population around villages and increasing market integration during the last decades may be causing agricultural intensification. Studies have shown that agricultural intensification, i.e. higher number of swidden-fallow cycles and shorter fallow periods, reduces crop productivity of swiddens and the regrowth capacity of fallows, undermining the resilience of the shifting cultivation system as a whole. We investigated the temporal and spatial dynamics of shifting cultivation in Brazilian Amazonia to test the hypotheses that (i) agriculture has become more intensive over time, and (ii) patterns of land-use intensity are related to land accessibility and human population density. We applied a breakpoint-detection algorithm to Landsat time-series spanning three decades and retrieved the temporal dynamics of shifting cultivation fields, which go through alternating phases of crop production (swidden) and secondary forest regrowth (fallow). We found that fallow-period length has decreased from 6.4 to 5.1 years on average, and that expansion over old-growth forest has slowed down over time. Shorter fallow periods and higher frequency of slash and burn cycles are practiced closer to residences and around larger villages. Our results indicate that shifting cultivation in riverine Amazonia has gone through a process of agricultural intensification in the past three decades. The resulting landscape is predominantly covered by young secondary forests ( 12 yrs old), and 20% of it have gone through intensive use. Reversing this trend and avoiding the negative consequences of agricultural intensification requires land use planning that accounts for the constraints of land use in riverine areas.
The effect of natural canopy gaps on the performance of naturally occurring tree seedling populat... more The effect of natural canopy gaps on the performance of naturally occurring tree seedling populations was studied in a tropical rain forest in French Guiana. This was done at two levels of scale. Firstly, on a 20 m × 250 m forest transect intersecting four canopy gaps, it was investigated how patterns of recruitment, growth and survival of seedlings of Cecropia obtusa, Dicorynia guianensis and Pourouma bicolor differed between canopy gaps and closed forest. Secondly, for one large natural canopy gap, performance of seedling cohorts established before ('pre-gap' cohorts) and after ('post-gap' cohorts) gap formation was studied in relation to environmental heterogeneity. The direct site factor (DSF) was used as an indicator of light availability in the gap zone. Cecropia specialised in large gaps, and also in specific sites within the large gap: seedling performance was increased by light, and by dead wood. Pourouma mainly germinated under closed forest conditions, where it can survive for a long period until a canopy gap is formed nearby. Pourouma seedlings adjusted well to the new gap environment but they did not specialise in specific places within the gap. Seedling location of Pourouma seedlings was mainly determined by accidental positioning before gap formation. Dicorynia was able to germinate both in gaps and under closed forest conditions. Although the number of gaps studied in this study was low, the results show that seedling establishment, early growth and survival of the three tree species were affected by canopy gaps and by the environmental heterogeneity within a large gap.
Defoliation is a ubiquitous stressor that can strongly limit plant performance. Tolerance to defo... more Defoliation is a ubiquitous stressor that can strongly limit plant performance. Tolerance to defoliation is often associated with compensatory growth. Genetic variation in tolerance and compensatory growth responses, in turn, play an important role in the evolutionary adaptation of plants to changing disturbance regimes but this issue has been poorly investigated for long-lived woody species. We quantified genetic variation in plant growth and growth parameters, tolerance to defoliation and compensatory responses to defoliation for a population of the understorey palm Chamaedorea elegans. In addition, we evaluated genetic correlations between growth and tolerance to defoliation. We performed a greenhouse experiment with 731 seedlings from 47 families with twelve or more individuals of C. elegans. Seeds were collected in southeast Mexico within a 0.7 ha natural forest area. A two-third defoliation treatment (repeated every two months) was applied to half of the individuals to simulate leaf loss. Compensatory responses in specific leaf area, biomass allocation to leaves and growth per unit leaf area were quantified. We found that growth rate was highly heritable and that plants compensated strongly for leaf loss. However, genetic variation in tolerance, compensation, and the individual compensatory responses was low. We found strong correlations between family mean growth rates in control and defoliation treatments. We did not find indications for growth-tolerance trade-offs: genetic correlation between tolerance and growth rate were not significant. The low genetic variation in tolerance and compensatory responses observed here suggests a low potential for evolutionary adaptation to changes in damage or herbivory, but high ability to adapt to changes in environment that require different growth rates. The strong correlations between family mean growth rates in control and defoliation treatments suggest that performance differences among families are also maintained under stress of disturbance. .
Secondary forests are increasingly important components of human-modified landscapes in the tropi... more 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
Large lianas and trees in the forest canopy are challenged by hydraulic and mechanical failures a... more Large lianas and trees in the forest canopy are challenged by hydraulic and mechanical failures and need to balance hydraulic conductivity, hydraulic safety and mechanical safety. Our study integrates these functions in canopy branches to understand the performance of canopy trees and lianas, and their difference. We sampled and measured branches from 22 species at a canopy crane in the tropical forest at Xishuangbanna, SW China. We quantified the hydraulic conductivity from the xylem-specific hydraulic conductivity (K S ), hydraulic safety from the cavitation resistance (P 50 ) and mechanical safety from the modulus of rupture (MOR) to evaluate trade-offs and differences between lianas and trees. We also measured a number of anatomical features that may influence these three functional traits. Our results suggest the following: trade-offs between hydraulic conductivity, hydraulic safety and mechanical safety are weak or absent; liana branches better resist external mechanical forces (higher MOR) than tree branches; and liana and tree branches were similar in hydraulic performance (K S and P 50 ). The anatomical features underlying K S , P 50 and MOR may differ between lianas and trees. We conclude that canopy branches of lianas and trees diverged in mechanical design due to fundamental differences in wood formation, but converged in hydraulic design.
Intraspecific and interspecific architectural patterns were studied for eight tree species of a B... more Intraspecific and interspecific architectural patterns were studied for eight tree species of a Bornean rain forest. Trees 5-19 m tall in two 4-ha permanent sample plots in primary forest were selected, and three light descriptors and seven architectural traits for each tree were measured. Two general predictions were made: (1) Slow growing individuals (or short ones) encounter lower light, and have flatter crowns, fewer leaf layers, and thinner stems, than do fast growing individuals (or tall ones). ( ) Species with higher shade-tolerance receive less light and have flatter crowns, fewer leaf layers, and thinner stems, than do species with lower shade-tolerance. Shade-tolerance is assumed to decrease with maximum growth rate, mortality rate, and adult stature of a species. Two light descriptors (crown position index and available space, but not canopy height) indicated higher light conditions for trees with more rapid growth, and for trees of greater height. Light levels were higher for species with high maximum growth rates and with greater adult stature. Most intraspecific architectural patterns conformed to the predictions: total leaf area and the number of leaf layers increased with increasing height and higher growth rates, and crown length/tree height ratio and stem slenderness respectively increased and decreased with growth rate. Yet, crown width/tree height ratio and relative crown length did not change with tree height, nor did they with previous growth. Slow growing (and short) trees may not have the reserves to invest in further horizontal crown growth, and to avoid leaf self-shading sufficiently within their relatively narrow crowns. Predictions on interspecific architectural patterns were not supported by the data. Species that were expected to be more shade-tolerant (lower maximum growth, lower mortality, and shorter stature) had deeper crowns, greater leaf areas, and more leaf layers, than did less shade-tolerant species. These patterns may be explained by lower loss rates of branches and leaves of the more shade-tolerant species. These species avoid leaf self-shading by distributing their leaves at the crown periphery. The role of lateral light appears to be more important than hitherto realized. The crown width/height ratio and height/dbh ratio were negatively correlated, both intraspecifically and interspecifically. It is suggested that trees co-ordinate their crown and stem growth so that they maintain their stability at small safety margins in the forest understorey.
Water storage in the stems of woody plants contributes to their responses to short-term water sho... more Water storage in the stems of woody plants contributes to their responses to short-term water shortages. To estimate the contribution of water storage to the daily water budget of trees, time lags of sap flow between different positions of trunk are used as a proxy of stem water storage. In lianas, another large group of woody species, it has rarely been studied whether stored water functions in their daily water use, despite their increasing roles in the carbon and water dynamics of tropical forests caused by their increasing abundance. We hypothesized that lianas would exhibit large time lags due to their extremely long stems, wide vessels and large volume of parenchyma in the stem. We examined time lags in sap flow, diel changes of stem volumetric water content (VWC) and biophysical properties of sapwood of 19 lianas and 26 co-occurring trees from 27 species in 4 forests (karst, tropical seasonal, flood plain and savanna) during a wet season. The plants varied in height/length from <5 to >60 m. The results showed that lianas had significantly higher saturated water content (SWC) and much lower wood density than trees. Seven of 19 liana individuals had no time lags; in contrast, only 3 of 26 tree individuals had no time lags. In general, lianas had shorter time lags than trees in our data set, but this difference was not significant for our most conservative analyses. Across trees and lianas, time lag duration increased with diurnal maximum changeable VWC but was independent of the body size, path length, wood density and SWC. The results suggest that in most lianas, internal stem water storage contributes little to daily water budget, while trees may rely more on stored water in the stem.
Understanding the relationship between stand-level tree diversity and productivity has the potent... more 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.
Lianas, woody climbing plants, are increasing in many tropical forests, with cascading effects su... more Lianas, woody climbing plants, are increasing in many tropical forests, with cascading effects such as decreased forest productivity, carbon sequestration, and resilience. Possible causes are increasing forest fragmentation, CO 2 fertilization, and drought. Determining the primary changing species and their underlying vital rates help explain the liana trends. We monitored over 17,000 liana stems for 13 yr in 20 ha of old-growth forest in the Congo Basin, and here we report changes and vital rates for the community and for the 87 most abundant species. The total liana abundance declined from 15,007 lianas in 1994 to 11,090 in 2001 to 9,978 in 2007. Over half (52%) of the evaluated species have significantly declining populations, showing that the community response is not the result of changes in a few dominant species only. Species density change (i.e., the change in number of individuals per hectare) decreased with mortality rate, tended to increase with recruitment rate, but was independent of growth rate. Species change was independent of functional characteristics important for plant responses to fragmentation, CO 2 , and drought, such as lifetime light requirements, climbing and dispersal mechanism, and leaf size. These results indicate that in Congo lianas do not show the reputed global liana increase, but rather a decline, and that elements of the reputed drivers underlying global liana change do not apply to this DR Congo forest. We suggest warfare in the Congo Basin to have decimated the elephant population, leading to less disturbance, forest closure, and declining liana numbers. Our results imply that, in this tropical forest, local causes (i.e., disturbance) override more global causes of liana change resulting in liana decline, which sharply contrasts with the liana increase observed elsewhere.
Tropical forests are experiencing large-scale structural changes, the most apparent of which may ... more 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.
The paradigm that tropical trees with farther seed dispersal experience lower offspring mortality... more The paradigm that tropical trees with farther seed dispersal experience lower offspring mortality is currently based on within-species studies documenting higher survival of offspring located farther from conspecific adults and/or closer to light gaps. We determined whether the paradigm also holds among species by comparing spatial patterns of offspring mortality among three sympatric Neotropical rainforest tree species with the same seed dispersers but with different dispersal abilities. First, we assessed spatially non-random mortality for each species by measuring spatial shifts of the population recruitment curve (PRC) with respect to conspecific adults and light gaps across three early life stages: dispersed seeds, young seedlings and old seedlings. Then, we determined whether PRC shifts were greater for the species with short dispersal distances than for the species with greater dispersal distances. We found that the PRC shifted away from conspecific adults consistently across life stages, but we found no consistent PRC shifts towards gaps, suggesting that mortality was related more to the proximity of conspecifics than to absence of light gaps. PRC shifts away from adults were greatest in the species with the lowest dispersal ability, supporting the paradigm. Differential PRC shifts caused the spatial distribution of offspring to become almost independent with respect to adult trees and gaps in all three species, despite large differences in seed dispersal distance among these species. Our results provide direct empirical support for the paradigm that among tropical trees, species with farther seed dispersal are less impacted by spatially non-random mortality than are similar species with shorter dispersal distances. Thus, greater dispersal effectiveness merits extra investments of trees in seed dispersal ability, even at the cost of fecundity.
Lianas are an important component of most tropical forests, where they vary in abundance from hig... more Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to Wx carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had signiWcantly higher CO 2 assimilation per unit mass (A mass ), nitrogen concentration (N mass ), and 13 C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use eYciency (WUE), but lower leaf structural investments. Seasonal variation in CO 2 assimilation per unit area (A area ), phosphorus concentration per unit mass (P mass ), and photosynthetic N-use eYciency (PNUE), however, was signiWcantly lower in lianas than in trees. For instance, mean tree A area decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana 13 C increased four times more than tree 13 C, with no reduction in PNUE, whereas trees had a signiWcant reduction in PNUE. Lianas had higher A mass than trees throughout the year, regardless of season. Collectively, our Wndings indicate that lianas Wx more carbon and use water and nitrogen more eYciently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.
This paper is NOT THE PUBLISHED VERSION; but the author's final, peer-reviewed manuscript. The pu... more This paper is NOT THE PUBLISHED VERSION; but the author's final, peer-reviewed manuscript. The published version may be accessed by following the link in the citation below.
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