High-dimensional coexistence of temperate tree species: functional traits, demographic rates, life-history stages, and their physical context

Theoretical models indicate that trade-offs between growth and survival strategies of tree species can lead to coexistence across life history stages (ontogeny) and physical conditions experienced by individuals. There exist predicted physiological mechanisms regulating these trade-offs, such as an investment in leaf characters that may increase survival in stressful environments at the expense of investment in bole or root growth. Confirming these mechanisms, however, requires that potential environmental, ontogenetic, and trait influences are analyzed together. Here, we infer growth and mortality of tree species given size, site, and light characteristics from forest inventory data from Wisconsin to test hypotheses about growth-survival trade-offs given species functional trait values under different ontogenetic and environmental states. A series of regression analyses including traits and rates their interactions with environmental and ontogenetic stages supported the relationships between traits and vital rates expected from the expectations from tree physiology. A combined model including interactions between all variables indicated that relationships between demographic rates and functional traits supports growth-survival trade-offs and their differences across species in high-dimensional niche space. The combined model explained 65% of the variation in tree growth and supports a concept of community coexistence similar to Hutchinson's n-dimensional hypervolume and not a low-dimensional niche model or neutral model.     

Trade-offs between seedling growth and survival in deciduous broadleaved trees in a temperate forest

BACKGROUND AND AIMS: In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest. METHODS: The relationship between seedling survival and seedling relative growth rate in mass (RGR(M)) or height (RGR(H)) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes. KEY RESULTS: Within each species, both RGR(M) and RGR(H) usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGR(M) and RGR(H) in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG. CONCLUSIONS: The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.     

Light environment and tree strategies in a Bolivian tropical moist forest: an evaluation of the light partitioning hypothesis

Light partitioning is thought to contribute to the coexistence of rain forest tree species. This study evaluates the three premises underlying the light partitioning hypothesis; 1) there is a gradient in light availability at the forest floor, 2) tree species show a differential distribution with respect to light, and 3) there is a trade-off in species performance that explains their different positions along the light gradient. To address these premises, we studied the light environment, growth, and survival of saplings of ten non-pioneer tree species in a Bolivian moist forest. Light availability in the understorey was relatively high, with a mean canopy openness of 3.5% and a mean direct site factor of 6.8%. Saplings of two light demanding species occurred at significantly higher light levels than the shade tolerant species. The proportion of saplings in low-light conditions was negatively correlated with the successional position of the species. Light-demanding species were characterised by a low share of their saplings in low-light conditions, a high sapling mortality, a fast height growth and a strong growth response to light. These data show that all three premises for light partitioning are met. There is a clear gradient in shade-tolerance within the group of non-pioneer species leading to a tight packing of species along the small range of light environments found in the understorey. This revised version was published online in August 2006 with corrections to the Cover Date.     

Performance Trade-offs Driven by Morphological Plasticity Contribute to Habitat Specialization of Bornean Tree Species

Growth-survival trade-offs play an important role in niche differentiation of tropical tree species in relation to light-gradient partitioning. However, the mechanisms that determine differential species performance in response to light and soil resource availability are poorly understood. To examine responses to light and soil nutrient availability, we grew seedlings of five tropical tree species for 12 mo at < 2 and 18 percent full sunlight and in two soil types representing natural contrasts in nutrient availability within a lowland dipterocarp forest in North Borneo. We chose two specialists of nutrient-rich and nutrient-poor soils, respectively, and one habitat generalist. Across all species, growth was higher in high than low light and on more nutrient rich soil. Although species differed in growth rates, the ranking of species, in terms of growth, was consistent across the four treatments. Nutrient-rich soils improved seedling survival and increased growth of three species even under low light. Slower-growing species increased root allocation and reduced specific leaf area (SLA) and leaf area ratio (LAR) in response to decreased nutrient supply. All species increased LAR in response to low light. Maximum growth rates were negatively correlated with survival in the most resource-limited environment. Nutrient-poor soil specialists had low maximum growth rates but high survival at low resource availability. Specialists of nutrient-rich soils, plus the habitat generalist, had the opposite suite of traits. Fitness component trade-offs may be driven by both light and belowground resource availability. These trade-offs contribute to differentiation of tropical tree species among habitats defined by edaphic variation.     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

Shade tolerance,canopy gaps and mechanisms of coexistence of forest trees

The belief that canopy gaps are important for the maintenance of tree species diversity appears to be widespread, but there have been no formal theoretical models to assess under what conditions gap phase processes allow coexistence. Much of the empirical research on niche differentiation in response to gaps has focused on evidence for an interspecific tradeoff between low light survival and high light growth. The objectives of this study are first to distinguish the possible mechanisms allowing coexistence based on this tradeoff, and second, to explore their limitations. We present a theory of forest dynamics driven by small‐scale disturbances as a special case of the theory of coexistence in variable environments. We demonstrate that temporal and spatial heterogeneity in light conditions that results from canopy gaps can allow stable coexistence as a result of three previously documented general mechanisms: ‘relative non‐linearity’, ‘the successional niche’ and the ‘storage effect’. We find that temporal fluctuations in light availability alone allow the stable coexistence of only two species. Spatial variation in disturbance synchronicity and intensity allows three species to coexist in a narrow parameter space. The rate of extinction is, however, extremely slow and there is transient coexistence of a larger number of species for a long period of time. We conclude that while the low light survival/high light growth tradeoff may be ubiquitous in forest tree species, it is unlikely to function as an important mechanism for the stable coexistence of several tree species.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Performance of trees in forest canopies: explorations with a bottom-up functional-structural plant growth model

Here we present a functional-structural plant model that integrates the growth of metamers into a growing, three-dimensional tree structure, and study the effects of different constraints and strategies on tree performance in different canopies. The tree is a three-dimensional system of connected metamers, and growth is defined by the flush probability of metamers. Tree growth was simulated for different canopy light environments. The result suggest that: the constraints result in an exponential, logistic and decay phase; a mono-layered-leaf crown results from self-shading in a closed canopy; a strong apical control results in slender trees like tall stature species; the interaction between weak apical control and light response results in a crown architecture and performance known from short stature species in closed forest; correlated leaf traits explain interspecific differences in growth, survival and adult stature. The model successfully unravels the interaction effects of different constraints and strategies on tree growth in different canopy light environments.     

Influence of light level on the growth and morphology of saplings in a panamanian forest

Leaf spacing and aboveground growth were monitored in saplings of ten species in a range of light environments in a Panamanian lowland humid forest. One- to 2-m-tall individuals of the chosen species had intermediate to large leaves on stems with few or no branches. Saplings in high light environments grew faster in height and produced more biomass per unit leaf area than shaded saplings for all species. These growth responses involved morphological plasticity with greater extension per unit biomass increment increasing the height growth rate of gap-grown saplings and greater biomass allocation to leaves decreasing the whole plant light compensation point in shade. The relative performance of the species also varied across the light gradient and was related to differences in leaf lifespan and specific leaf mass. Light-demanding species grew as rapidly in shade as shade-tolerant species, but the shorter leaf lifespan of the former necessitates higher production rates to maintain a given leaf area, largely excluding light-demanders from shaded understory locations. Height growth rate was positively correlated with leaf spacing for each species, and differences between species in the height growth rate-internode length relationship were related to interspecific differences in specific leaf mass. Thus, sapling growth histories may be inferred from their morphologies.     

How tree species identity and diversity affect light transmittance to the understory in mature temperate forests

Light is a key resource for plant growth and is of particular importance in forest ecosystems, because of the strong vertical structure leading to successive light interception from canopy to forest floor. Tree species differ in the quantity and heterogeneity of light they transmit. We expect decreases in both the quantity and spatial heterogeneity of light transmittance in mixed stands relative to monocultures, due to complementarity effects and niche filling. We tested the degree to which tree species identity and diversity affected, via differences in tree and shrub cover, the spatiotemporal variation in light availability before, during, and after leaf expansion. Plots with different combinations of three tree species with contrasting light transmittance were selected to obtain a diversity gradient from monocultures to three species mixtures. Light transmittance to the forest floor was measured with hemispherical photography. Increased tree diversity led to increased canopy packing and decreased spatial light heterogeneity at the forest floor in all of the time periods. During leaf expansion, light transmittance did differ between the different tree species and timing of leaf expansion might thus be an important source of variation in light regimes for understory plant species. Although light transmittance at the canopy level after leaf expansion was not measured directly, it most likely differed between tree species and decreased in mixtures due to canopy packing. A complementary shrub layer led, however, to similar light levels at the forest floor in all species combinations in our plots. Synthesis. We find that a complementary shrub layer exploits the higher light availability in particular tree species combinations. Resources at the forest floor are thus ultimately determined by the combined effect of the tree and shrub layer. Mixing species led to less heterogeneity in the amount of light, reducing abiotic niche variability.     

The need for a canopy perspective to understand the importance of phenotypic plasticity for promoting species coexistence and light-use complementarity in forest ecosystems

Because of their overwhelming size over other organisms, trees define the structural and energetic properties of forest ecosystems. From grasslands to forests, leaf area index, which determines the amount of light energy intercepted for photosynthesis, increases with increasing canopy height across the various terrestrial ecosystems of the world. In vertically well-developed forests, niche differentiation along the vertical gradient of light availability may promote species coexistence. In addition, spatial and temporal differentiation of photosynthetic traits among the coexisting tree species (functional diversity) may promote complementary use of light energy, resulting in higher biomass and productivity in multi-species forests. Trees have evolved retaining high phenotypic plasticity because the spatial/temporal distribution of resources in forest ecosystems is highly heterogeneous and trees modify their own environment as they increase nearly 1,000 times in size through ontogeny. High phenotypic plasticity may enable coexistence of tree species through divergence in resource-rich environments, as well as through convergence in resource-limited environments. We propose that the breadth of individual-level phenotypic plasticity, expressed at the metamer level (leaves and shoots), is an important factor that promotes species coexistence and resource-use complementarity in forest ecosystems. A cross-biome comparison of the link between plasticity of photosynthesis-related traits and stand productivity will provide a functional explanation for the relationship between species assemblages and productivity of forest ecosystems.     

Wood mechanics, allometry, and life-history variation in a tropical rain forest tree community

Wood density plays a central role in the life-history variation of trees, and has important consequences for mechanical properties of wood, stem and branches, and tree architecture. Wood density, modulus of rupture, modulus of elasticity, and safety factors for buckling and bending were determined for saplings of 30 Bolivian rain forest tree species, and related to two important life-history axes: juvenile light demand and maximum adult stature. Wood density was strongly positively related to wood strength and stiffness. Species safety factor for buckling was positively related to wood density and stiffness, but tree architecture (height : diameter ratio) was the strongest determinant of mechanical safety. Shade-tolerant species had dense and tough wood to enhance survival in the understorey, whereas pioneer species had low-density wood and low safety margins to enhance growth in gaps. Pioneer and shade-tolerant species showed opposite relationships between species traits and adult stature. Light demand and adult stature affect wood properties, tree architecture and plant performance in different ways, contributing to the coexistence of rain forest species.     

The significance of ratios of detritus types and micro-organism productivity to competitive interactions between aquatic insect detritivores

Investigations of competitive interactions emphasize non-detrital resources, even though detritus is a major component of most food webs. Studies of competing species focus usually on single resource types, although consumers in nature are likely to encounter mixtures of resource types that may affect whether competition results in exclusion or coexistence. The invasive mosquito Aedes albopictus is capable of excluding the native mosquito Ochlerotatus triseriatus in competition for single detritus types in laboratory and field microcosms. In this study, we used nine ratios of two detritus types (animal and leaf) common in natural containers to test whether detritus ratios affect the outcome of competition. Under intraspecific and interspecific competition, A. albopictus attained higher survival and estimated population growth rate than did O. triseriatus. Unlike past studies, both species had positive growth and high adult survival, with little evidence of competitive effects, under one resource ratio (10:1 ratio of leaf : animal detritus) regardless of mosquito densities, suggesting potential coexistence. Path analysis showed that densities of larvae had negative effects on population growth for O. triseriatus but not for A. albopictus, indicating competitive superiority of A. albopictus. Population growth of both species was affected strongly by the direct paths from animal (positive) and leaf (negative) detritus, and the indirect effect of leaf detritus via bacterial production (positive). Field sampling established that detritus entered real tree holes in ratios similar to those in our experiment, suggesting that natural variation in detritus ratios may influence local coexistence of these species. Seasonal variation in ratios of plant and animal detritus indicated that temporal as well as spatial variation in inputs may be important for potential coexistence.     

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.     

Light and competition gradients fail to explain the coexistence of shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings

Background and Aims

The coexistence of forest tree species has often been linked to differences among species in terms of their response to light availability during the regeneration stage. From this perspective, species coexistence results from growth–growth or mortality–growth trade-offs along spatial light gradients. Experimental evidence of growth–growth trade-offs in natural conditions is sparse due to various confounding factors that potentially hinder the relationship. This study examined growth hierarchies along light gradients between two tree species with contrasting shade tolerance by controlling potential confounding factors such as seedling size, seedling status, seedling density and species composition.

Methods

Natural regenerated shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings were used, and growth rankings over a 4-year period were compared in 8- to 10-year-old tree seedlings.

Key results

No rank reversal occurs between the two species along the light gradient, or along the density, mixture or seedling size gradients. The shade-tolerant species was always the more competitive of the two. Pronounced effects of initial size on seedling growth were observed, whereas the effects of light and competition by neighbours were of secondary importance. The paramount effect of size, which results from the asymmetric nature of interseedling competition, gives a strong advantage to tall seedlings over the long term.

Conclusions

This study extends previous efforts to identify potential drivers of rank reversals in young tree mixtures. It does not support the classical assumption that spatial heterogeneity in canopy opening explains the coexistence of the two species studied. It suggests that spatial variation in local size hierarchies among seedlings that may be caused by seedling emergence time or seedling initial performance is the main driver of the dynamics of these mixed stands.     

Growth responses of 15 rain-forest tree species to a light gradient: the relative importance of morphological and physiological traits

1. Growth of seedlings of 15 rain-forest tree species was compared under controlled conditions, at six different light levels (3, 6, 12, 25, 50 and 100% daylight).
2. Most plant variables showed strong ontogenetic changes; they were highly dependent on the biomass of the plant.
3. Growth rate was highest at intermediate light levels (25–50%) above which it declined. Most plant variables showed a curvilinear response to irradiance, with the largest changes at the lowest light levels.
4. There was a consistent ranking in growth between species; species that were fast growing in a low-light environment were also fast growing in a high-light environment.
5. At low light, interspecific variation in relative growth rate was determined mainly by differences in a morphological trait, the leaf area ratio (LAR), whereas at high light it was determined mainly by differences in a physiological trait, the net assimilation rate (NAR).
6. NAR became a stronger determinant of growth than LAR in more than 10–15% daylight. As light availability in the forest is generally much lower than this threshold level, it follows that interspecific variation in growth in a forest environment is mainly owing to variation in morphology.     

Close relationship between leaf life span and seedling relative growth rate in temperate hardwood species

The life span of resource-acquiring organs (leaves, shoots, fine roots) is closely associated with species successional position and environmental resource availability. We examined to what extent leaf life span is related to inter- and intraspecific variation in seedling relative growth rate (RGR). We examined relationships between relative growth rate in mass (RGR_M) or height (RGR_H) and leaf life span, together with classical RGR_M components [net assimilation rate (NAR), specific leaf area (SLA), leaf weight ratio (LWR), and leaf area ratio (LAR)] for seedlings of five hardwood species of different successional position across a wide range of environmental resource availability, including the presence or absence of leaf litter in shaded forest understory, small canopy gaps, and large canopy gaps. Both SLA and LAR were negatively correlated with RGR_M along the environmental gradient for all species. However, positive correlations were observed among species within microsites, indicating that these two components cannot consistently explain the variation in RGR_M. Both NAR and LWR affect interspecific, but not intraspecific, variation in RGR_M. Leaf life span was negatively correlated with either RGR_M or RGR_H in both inter- and intraspecific comparisons. Species with short-lived, physiologically active leaves have high growth rates, particularly in resource-rich environments. Consequently, leaf life span is a good predictor of seedling RGR. Leaf life span affects plant performance and has a strong and consistent effect on tree seedling growth, even among contrasting environments.     

Changes in Leaf Phenology are Dependent on Tree Height inAcer mono,a Deciduous Broad-leaved Tree

Ontogenetic changes in leaf phenology of a hardwood tree,Acer mono, were investigated in individuals in different size classes in a temperate forest. Leaf emergence was earliest in current-year seedlings, and was increasingly delayed with increasing height of the individual. The shorter the tree, the longer the duration of leaf emergence. Timing of leaf emergence of the dominant heterospecific canopy trees was almost identical to that of conspecific adults; understorey light then gradually decreased with expansion of canopy leaves. These traits indicate that smaller individuals that receive the least light in summer can acquire favourable light for a longer period in spring than taller plants even in a forest understorey, but the advantage decreases with increasing plant height. Changes in the duration of leaf emergence and leaf longevity in response to environmental light regime [sun (forest edge)vs. shade (forest understorey)], were greatest for current year seedlings but decreased with increasing plant height. These results suggest that the plastic response of leaf phenology in juvenile stages may reduce the risk of losing an entire cohort in spatially heterogenous environments in the understorey of temperate forests.     

Light‐based Regeneration Niches: Evidence from 21 Dipterocarp Species using Size‐specific RGRs

A continuing challenge in tropical ecology is to explain the coexistence of large numbers of rain forest tree species. One possible coexistence mechanism is partitioning of the highly variable and dynamic forest light environment, in which species that grow better in one light treatment grow worse in another. To test whether species respond differently to the light environment, we estimated growth rates of 21 Dipterocarpaceae species from Malaysian Borneo grown in shade houses for 2 yr in three light treatments (0.3%, 3%, and 18% full sunlight). We made regular measurements of height, diameter, and aboveground biomass, enabling us to calculate growth rates for each response. We estimated size‐specific growth rates using nonlinear mixed‐effects models, as average relative growth rate was strongly size dependent. For all species, the greatest diameter growth rate was achieved in 18 percent full sunlight, whereas for five of the twenty‐one species, the greatest height growth rate was achieved in three percent full sunlight. We investigated correlations among growth rates in different light treatments, but no negative correlations were found, indicating that species growing well in one light treatment did not grow poorly in the others. There were substantial crossovers, however, in species ranks among the three light treatments, indicating that there was no single growth rate hierarchy common to all light treatments. The lack of a single consistent growth hierarchy across light treatments indicates that heterogeneity in the forest light environment could contribute to the maintenance of the diversity of Dipterocarpaceae found in lowland Bornean rain forests via light‐based regeneration niches.