Evidence for a three-way trade-off between nitrogen and phosphorus competitive abilities and cell size in phytoplankton

Trade-offs among functional traits are essential for explaining community structure and species coexistence. While two-way trade-offs have been investigated in many systems, higher-dimensional trade-offs remain largely hypothetical. Here we demonstrate a three-way trade-off between cell size and competitive abilities for nitrogen and phosphorus in marine and freshwater phytoplankton. At a given cell size, competitive abilities for N and P are negatively correlated, but as cell size increases, competitive ability decreases for both nutrients. The relative importance of the two trade-off axes appears to be environment dependent, suggesting different selective pressures: freshwater phytoplankton separate more along the N vs. P competition axis, and marine phytoplankton separate more along the nutrient competition vs. cell size axis. Our results demonstrate the multidimensional nature of key trade-offs among traits and suggest that such trade-offs may drive species interactions and structure ecological communities.     

Alternative strategies in avian scavengers: how subordinate species foil the despotic distribution

Trade-offs in species’ traits can mediate competition and enable coexistence. A key challenge in ecology is understanding the role of species’ trade-offs in maintaining diversity, and evolutionary trade-offs between the abilities of competing species are best understood by considering how competitive advantages change along an environmental gradient. Previous studies of such trade-offs are generally limited to two-species systems and a single trade-off. In this study, I consider the effect of trade-offs in search efficiency and competitive abilities on habitat use patterns among a diverse avian scavenger guild. I hypothesize that species’ dominance status and search efficiency will both be correlated with patch quality. Using counts of searching birds in areas that vary in habitat quality in terms of both wildlife and human settlement density and observations at experimental carcasses, I assess the competitive ability, search efficiency, and habitat use of seven avian scavenger species in Masai Mara National Reserve, Kenya. Findings support the hypothesis with Bateleurs, a species with high search efficiency, and Ruppell’s, Lappet-faced, and White-backed vultures, species with high individual or social dominance, preferentially exploiting habitats of high quality, while Tawny eagles and Hooded vultures, species with low search efficiency and competitive ability, prefer habitats of low quality. This paper demonstrates the importance of considering multiple strategies for assessing the effect of competition on habitat use within complex communities.     

Species phylogenetic relatedness, priority effects, and ecosystem functioning

Species immigration history can structure ecological communities through priority effects, which are often mediated by competition. As competition tends to be stronger between species with more similar niches, we hypothesize that species phylogenetic relatedness, under niche conservatism, may be a reasonable surrogate of niche similarity between species, and thus influence the strength of priority effects. We tested this hypothesis using a laboratory microcosm experiment in which we established bacterial species pools with different levels of phylogenetic relatedness and manipulated the immigration history of species from each pool into microcosms. Our results showed that strong priority effects, and hence multiple community states, only emerged for the species pool with the greatest phylogenetic relatedness. Community assembly also resulted in a significant positive relationship between bacterial phylogenetic diversity and ecosystem functions. Interestingly, these results emerged despite a lack of phylogenetic conservatism for most of the bacterial functional traits considered. Our results highlight the utility of phylogenetic information for understanding the structure and functioning of ecological communities, even when phylogenetically conserved functional traits are not identified or measured.     

Ecology of hemiepiphytism in fig species is based on evolutionary correlation of hydraulics and carbon economy

Woody hemiepiphytic species (Hs) are important components of tropical rain forests, and they have been hypothesized to differ from non-hemiepiphytic tree species (NHs) in adaptations relating to water relations and carbon economy; but few studies have been conducted comparing ecophysiological traits between the two growth forms especially in an evolutionary context. Using common-garden plants of the genus Ficus, functional traits related to plant hydraulics and carbon economy were compared for seven NHs and seven Hs in their adult terrestrial "tree-like" growth phase. We used phylogenetically independent contrasts to test the hypothesis that differences in water availability selected for contrasting suites of traits in Hs and NHs, driving evolutionary correlations among functional traits including hydraulic conductivity and photosynthetic traits. Species of the two growth forms differed in functional traits; Hs had substantially lower xylem hydraulic conductivity and stomatal conductance, and higher instantaneous photosynthetic water use efficiency. Leaf morphological and structural traits also differed strikingly between the two growth forms. The Hs had significantly smaller leaves, higher leaf mass per area (LMA), and smaller xylem vessel lumen diameters. Across all the species, hydraulic conductivity was positively correlated with leaf gas exchange indicating high degrees of hydraulic-photosynthetic coordination. More importantly, these correlations were supported by correlations implemented on phylogenetic independent contrasts, suggesting that most trait correlations arose through repeated convergent evolution rather than as a result of chance events in the deep nodes of the lineage. Vatiation in xylem hydraulic conductivity was also centrally associated with a suite of other functional traits related to carbon economy and growth, such as LMA, water use efficiency, leaf nutrient concentration, and photosynthetic nutrient use efficiency, indicating important physiological constraints or trade-offs among functional traits. Shifts in this trait cluster apparently related to the adaptation to drought-prone canopy growth during the early life cycle of Hs and clearly affected ecophysiology of the later terrestrial stage of these species. Evolutionary flexibility in hydraulics and associated traits might be one basis for the hyper-diversification of Ficus species in tropical rain forests.     

Effects of plant species traits on ecosystem processes: experiments in the Patagonian steppe

Several experiments have shown that aboveground net primary productivity increases with plant species richness. The main mechanism proposed to explain this relationship is niche complementarity, which is determined by differences in plant traits that affect resource use. We combined field and laboratory experiments using the most abundant species of the Patagonian steppe to identify which are the traits that determine niche complementarity in this ecosystem. We estimated traits that affect carbon, water, microclimate, and nitrogen dynamics. The most important traits distinguishing among species, from the standpoint of their effects on ecosystem functioning, were potential soil nitrification, rooting depth, and soil thermal amplitude. Additionally, we explored the relationship between trait diversity and aboveground net primary production (ANPP) using a manipulative field experiment. ANPP and the fraction of ANPP accounted for by trait diversity increased with number of traits. The effect of trait diversity decreased as the number of traits increased. Here, the use of traits gave us a mechanistic understanding of niche complementarity in the Patagonian steppe.     

Food resource partitioning in a Mediterranean demersal fish assemblage: the effect of body size and niche width

We investigated the effects of body size, feeding strategy and depth distribution on the trophic resource partitioning among the 26 dominant fish consumers in a fish assemblage on the central Mediterranean shelf-break. The fish assemblage was structured in two major trophic guilds: epibenthic and zooplanktonic feeders, according to the position of each predator along the benthos–plankton gradient. Within each main guild, the species were segregated along a prey-size or fish-size gradient into five further guilds. Fish size and prey size were strongly correlated, indicating that the prey-size niche can be well explained by predator size. Fish consumers showed a significant negative correlation between the similarity in prey type and the similarity in depth distribution; most species with similar trophic preferences segregated along the depth dimension. The only predators overlapping in both food and depth preferences were those with a more specialist trophic behavior. These results suggest that fish body size and depth preferences are the two main niche dimensions, explaining a large part of the coexistence between the Mediterranean shelf-break fish consumers.     

The probabilistic niche model reveals substantial variation in the niche structure of empirical food webs

The structure of food webs, complex networks of interspecies feeding interactions, plays a crucial role in ecosystem resilience and function, and understanding food web structure remains a central problem in ecology. Previous studies have shown that key features of empirical food webs can be reproduced by low-dimensional "niche" models. Here we examine the form and variability of food web niche structure by fitting a probabilistic niche model to 37 empirical food webs, a much larger number of food webs than used in previous studies. The model relaxes previous assumptions about parameter distributions and hierarchy and returns parameter estimates for each species in each web. The model significantly outperforms previous niche model variants and also performs well for several webs where a body-size-based niche model performs poorly, implying that traits other than body size are important in structuring these webs' niche space. Parameter estimates frequently violate previous models' assumptions: in 19 of 37 webs, parameter values are not significantly hierarchical, 32 of 37 webs have nonuniform niche value distributions, and 15 of 37 webs lack a correlation between niche width and niche position. Extending the model to a two-dimensional niche space yields networks with a mixture of one- and two-dimensional niches and provides a significantly better fit for webs with a large number of species and links. These results confirm that food webs are strongly niche-structured but reveal substantial variation in the form of the niche structuring, a result with fundamental implications for ecosystem resilience and function.     

What enables coexistence in plant communities? Weak versus strong species traits and the role of local processes

Explaining the coexistence of species that basically depend on the same resources has been a brainteaser for generations of ecologists. Different mechanisms have been proposed to facilitate coexistence in plant communities, where space is an important resource. Using a stochastic cellular automaton simulation model we analyze - separately and in combination - the influence of different species traits and processes which alter local competition on the coexistence of plant species over a fixed time horizon. We show that different species traits operate on different time scales in competition. We therefore suggest the concept of weak versus strong traits according to short- or long-term exclusion of species differing in these traits. As a consequence, highly non-linear trade-offs between weak and strong traits can result in communities. Furthermore, we found that trade-offs based on physiological species traits such as plant lifetime, dispersal range and plant growth, did not support broad and long-term coexistence—further processes such as density-dependent mortality and light-dependent colonization were necessary. This suggests that coexistence in plant communities requires (stabilizing) local processes to support the (equalizing) trade-offs in species traits.     

Consequences of niche overlap for ecosystem functioning: an experimental test with pond grazers

While the number of studies investigating the effects of species diversity on ecosystem properties continues to expand, few have explicitly examined how ecosystem functioning depends quantitatively on the degree of niche complementarity among species. We report the results of a microcosm experiment where similarity in habitat use among aquatic snail species was evaluated as a predictor of changes in community and ecosystem properties due to increasing species richness. Replicate microcosms with all possible one- and two-species combinations of a guild of six snail species were stocked with identical initial snail biomass. Microcosms with two species of snails had greater final snail biomass, lower attached algae biomass, and less total organic matter than monocultures. Snail species differed in their use of five distinct habitat types in the microcosms. Similarity in habitat use between a species pair was negatively related to the magnitude of change (e.g., deltaEF [change in ecosystem function]) in dissolved oxygen. periphyton biomass, and accrual of organic matter with a change in diversity. However, using the most stringent criterion for complementarity effects (e.g., Dmax [proportional deviation of the total polyculture yield from the highest yielding monoculture]), a relationship between species' niche similarity and changes in function with increasing species richness was only observed for dissolved oxygen. The identity of snail species present in the microcosms had strong effects on total organic matter, snail biomass, dissolved oxygen, periphyton biomass, and sedimentation rate. In this study, herbivore identity, sampling effects, and niche complementarity all appear to contribute to species richness effects on pond ecosystem properties and community structure. The analytical approach employed here may profitably be used in other systems to quantify the role of niche complementarity in species richness-ecosystem function relationships.     

Differences in sensory ecology contribute to resource partitioning in the bats Myotis bechsteinii and Myotis nattereri (Chiroptera: Vespertilionidae)

Coexisting animal species frequently differ in resource use in at least one niche dimension and thus avoid competition. While a range of morphological differences that lead to differentiation in animals' mechanical access to food have been identified, the role of sensory differences in within-guild niche differentiation has received less attention. We tested the hypothesis that differences in sensory access to prey contribute to resource partitioning between potentially competing species using two sympatric, similar-sized, congeneric bat species as a model system. Nursery colonies of Natterer's bat (Myotis nattereri) and Bechstein's bat (Myotis bechsteinii) roost in bat boxes in the same orchard and forage in forests and orchards nearby. In observations and behavioural experiments with freshly captured M. bechsteinii, we showed that individuals are able to hunt using prey-generated sound alone. In contrast, M. nattereri rarely uses prey-generated sound, but instead is able to find prey by echolocation very close to vegetation. In accordance with these behavioural data, we showed that M. bechsteinii has significantly larger ears than M. nattereri, providing it with superior detection and localization abilities for relatively low-frequency prey rustling sounds. We hypothesized that these differences in sensory ecology of the two syntopic, congeneric species would contribute to resource partitioning, so that M. bechsteinii would find more noisy prey taxa, possibly hidden in vegetation, by listening for prey sounds, while M. nattereri would have better access to still prey using echolocation or associative learning. Analysis of faecal samples collected on the same nights from bat boxes occupied by each species corroborated this prediction. The diets of the two species differed significantly, reflecting their different prey perception techniques and thereby supporting the hypothesis that differences in sensory ecology contribute to niche differentiation. Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users. B.M. Siemers and S.M. Swift contributed equally to this work.     

Multidimensional trade-offs in species responses to disturbance: implications for diversity in a subtropical forest

Species employ diverse strategies to cope with natural disturbance, but the importance of these strategies for maintaining tree species diversity in forests has been debated. Mechanisms that have the potential to promote tree species coexistence in the context of repeated disturbance include life history trade-offs in colonization and competitive ability or in species' ability to survive at low resource conditions and exploit the temporary resource-rich conditions often generated in the wake of disturbance (successional niche). Quantifying these trade-offs requires long-term forest monitoring and modeling. We developed a hierarchical Bayes model to investigate the strategies tree species employ to withstand and recover from hurricane disturbance and the life history trade-offs that may facilitate species coexistence in forests subject to repeated hurricane disturbance. Unlike previous approaches, our model accommodates temporal variation in process error and observations from multiple sources. We parameterized the model using growth and mortality data from four censuses of a 16-ha plot taken every five years (1990-2005), together with damage data collected after two hurricanes and annual seed production data (1992-2005). Species' susceptibilities to hurricane damage as reflected by changes in diameter growth and fecundity immediately following a storm were weak, highly variable, and unpredictable using traditional life history groupings. The lower crowding conditions (e.g., high light) generated in the wake of storms, however, led to greater gains in growth and fecundity for pioneer and secondary-forest species than for shade-tolerant species, in accordance with expectation of life history. We found moderate trade-offs between survival in high crowding conditions, a metric of competitive ability, and long-distance colonization. We also uncovered a strong trade-off between mean species fecundity in low crowding conditions, a metric of recovery potential, and competitive ability. Trade-offs in competitive and colonization ability, in addition to successional niche processes, are likely to contribute to species persistence in these hurricane-impacted forests. The strategies species employ to cope with hurricane damage depend on the degree to which species rely on sprouting, repair of adult damage, changes in demographic rates in response to enhanced resource availability after storms, or long-distance dispersal as recovery mechanisms.     

Behavioral responses to echolocation calls from sympatric heterospecific bats: implications for interspecific competition

Mutual recognition is the product of species coexistence, and has direct effects on survival and reproduction of animals. Bats are able to discriminate between sympatric different heterospecifics based on their echolocation calls, which has been shown both in free-flying and captive bats. To date, however, the factors that may determine the behavioral responses of bats to echolocation calls from sympatric heterospecifics have rarely been tested, especially under well-controlled conditions in captive bats. Hence, we aimed at tackling this question by performing playback experiments (habituation–dishabituation) with three horseshoe bat species within the constant-frequency bat guild, which included big-eared horseshoe bats (Rhinolophus macrotis), Blyth’s horseshoe bats (Rhinolophus lepidus), and Chinese horseshoe bats (Rhinolophus sinicus). We studied the behavioral responses of these three species to echolocation calls of conspecifics, to other two species, and to another heterospecifics bat, Stoliczka’s trident bat (Asellisus stoliczkanus), which also belongs to this guild. We found that the three rhinolophid species displayed a series of distinct behaviors to heterospecific echolocation but few to conspecific calls after habituation, suggesting that they may have been able to discriminate sympatric heterospecific echolocation calls from those of conspecifics. Interestingly, the behavioral responses to heterospecific calls were positively correlated with the interspecific overlap index in trophic niche, whereas call design had only a minor effect. This implies that the behavioral responses of these bats to heterospecific echolocation calls may be related to the degree of interspecific food competition.     

Environmental variability and allocation trade-offs maintain species diversity in a process-based model of succulent plant communities

Ecological theory suggests that environmental variability can promote coexistence, provided that species occupy differential niches. In this study, we focus on two questions: (1) Do allocation trade-offs provide a sufficient basis for niche differentiation in succulent plant communities? (2) What is the relative importance of different forms of environmental variability on species diversity and community composition? We approach these questions with a generic, individual-based simulation model. In our model, plants compete for water in a spatially explicit environment. Species differ in their size at maturity and in the allocation of carbon to roots, leaves and storage tissue. The model was fully specified with independent literature data. Model output was compared to characteristics of a species-rich community in the semi-arid Richtersveld (South Africa). The model reproduced the coexistence of plants with different sizes at maturity, the dominance of succulent shrubs, and the level of vegetation cover. We analyzed the effects of three forms of environmental variability: (a) temporal fluctuations in precipitation (rain and fog), (b) spatial heterogeneity of water supply due to run-on and run-off processes and (c) ‘rock pockets’ that limit root competition in space. The three types of variability had differential effects on diversity: diversity exhibited a strong hump-shaped response to temporal variation. Spatial variability increased diversity, with the strongest increase occurring at intermediate levels of temporal variability. Finally, rock pockets had the weakest effect, but contributed to diversity by providing refuges for small species, particularly at low temporal variability. The model thus shows that spatio-temporal variation of resource supply can maintain diversity over long time scales even in small systems, as is the case in the Richtersveld succulent communities. Trade-offs in allocation provide the basis for necessary niche differentiation. By describing resource competition between individual plants, our model provides a mechanistic basis for the link from species traits to community composition at given environmental conditions. It thereby contributes to an understanding of the forces shaping plant communities. Such an understanding is critical to reduce the threats environmental change poses to biodiversity and ecosystem services.     

Epiphyte metapopulation dynamics are explained by species traits, connectivity, and patch dynamics

The colonization-extinction dynamics of many species are affected by the dynamics of their patches. For increasing our understanding of the metapopulation dynamics of sessile species confined to dynamic patches, we fitted a Bayesian incidence function model extended for dynamic landscapes to snapshot data on five epiphytic lichens among 2083 mapped oaks (dynamic patches). We estimate the age at which trees become suitable patches for different species, which defines their niche breadth (number of suitable trees). We show that the colonization rates were generally low, but increased with increasing connectivity in accordance with metapopulation theory. The rates were related to species traits, and we show, for the first time, that they are higher for species with wide niches and small dispersal propagules than for species with narrow niches or large propagules. We also show frequent long-distance dispersal in epiphytes by quantifying the relative importance of local dispersal and background deposition of dispersal propagules. Local stochastic extinctions from intact trees were negligible in all study species, and thus, the extinction rate is set by the rate of patch destruction (tree fall). These findings mean that epiphyte metapopulations may have slow colonization-extinction dynamics that are explained by connectivity, species traits, and patch dynamics.     

Coexistence of the niche and neutral perspectives in community ecology

The neutral theory for community structure and biodiversity is dependent on the assumption that species are equivalent to each other in all important ecological respects. We explore what this concept of equivalence means in ecological communities, how such species may arise evolutionarily, and how the possibility of ecological equivalents relates to previous ideas about niche differentiation. We also show that the co-occurrence of ecologically similar or equivalent species is not incompatible with niche theory as has been supposed, because niche relations can sometimes favor coexistence of similar species. We argue that both evolutionary and ecological processes operate to promote the introduction and to sustain the persistence of ecologically similar and in many cases nearly equivalent species embedded in highly structured food webs. Future work should focus on synthesizing niche and neutral perspectives rather than dichotomously debating whether neutral or niche models provide better explanations for community structure and biodiversity.     

Male mating effort in a polygynous ungulate

Survival and mating success are key fitness components of mammalian males. Because energy is limited, life history theory predicts trade-offs between current and future reproduction. While many studies have examined fitness trade-off in females, we know little about trade-offs faced by males. In polygynous species, male mating success is largely dependent upon intrasexual competition. Consequently, males have greater uncertainty over the benefits of a given allocation than over its costs, and the correlation between mating effort and success is likely much weaker in males than in females. We analyzed 14 years of data on the mating effort and survival of marked bighorn rams to investigate fitness trade-offs. Dominant rams defended single estrous ewes (“tending”) while subordinates attempted to copulate after separating the tending pair (“coursing”). We estimated the participation in tending and coursing for each ram and the effort in searching for breeding opportunities by each ram each year. We compared these three behavioral indices of male mating effort to demographic parameters, individual characteristics, and both yearly and long-term survival. Mating effort during the rut was unrelated to ram overwinter survival, but longevity was positively correlated with mating effort between 2 and 5 years of age. Persistent variation among rams is likely to explain this pattern, suggesting that in natural populations a few high quality males enjoy both high mating success and high survival.An erratum to this article can be found at     

Ecology and Conservation of the Endangered American Burying Beetle (Nicrophorus americanus)

Field studies were conducted on the western population of the endangered American burying beetle (Nicrophorus americanus) in Oklahoma and Arkansas to determine its habitat affinities. A common cause of declining populations is some specialized adaptation that makes it difficult to respond to a rapidly changing habitat. We evaluated the hypothesis that N. americanus is a habitat specialist in its search for food, preferring mature forests with deep, humic soils. This hypothesis was rejected. Based on comparisons of niche breadth among syntopic congeners and niche overlap, N. americanus is relatively generalized in its use of a range of habitats when searching for food. It is likely that the generalist nature and the endangered status of N. americanus both derive from the fact that it is the largest member of its guild. In comparison to smaller species, N. americanus breeds on larger carcasses, which are more unpredictable in space and time. It is likely, therefore, that N. americanus must search over a larger area and greater diversity of habitats than its smaller congeners.     

Are functional traits good predictors of demographic rates? Evidence from five neotropical forests

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.     

Spatial hyperdynamism in a post-disturbance simulated forest

Competition–colonization models can address the population dynamics of remnants following habitat destruction. Spatially explicit versions have produced qualifications of the extinction debt issue and limited hyperdynamism in populations following habitat destruction. Although spatially explicit, these efforts examined few indicators of the spatial structure of the landscape. An existing model is modified here to represent a difference in niche adaptations as well as the competition–colonization tradeoff. Several landscape metrics are calculated at each iteration. Although the addition of niche differentiation did not change the qualitative outcome of the model, the spatial metrics show that some aspects of landscape structure, i.e., average patch area and proximity, become hyperdynamic and remain so. Small fluctuations in species populations are magnified in their spatial expression because the landscape is simplified.     

Architecture of 54 moist-forest tree species: traits, trade-offs, and functional groups

Tree architecture is an important determinant of the height extension, light capture, and mechanical stability of trees, and it allows species to exploit the vertical height gradient in the forest canopy and horizontal light gradients at the forest floor. Tropical tree species partition these gradients through variation in adult stature (Hmax) and light demand. In this study we compare 22 architectural traits for 54 Bolivian moist-forest tree species. We evaluate how architectural traits related to Hmax vary with tree size, and we present a conceptual scheme in which we combine the two axes into four different functional groups. Interspecific correlations between architecture and Hmax varied strongly from negative to positive, depending on the reference sizes used. Stem height was positively related to Hmax at larger reference diameters (14-80 cm). Species height vs. diameter curves often flattened toward their upper ends in association with reproductive maturity for species of all sizes. Thus, adult understory trees were typically shorter than similar-diameter juveniles of larger species. Crown area was negatively correlated with Hmax at small reference heights and positively correlated at larger reference heights (15-34 m). Wide crowns allow the small understory species to intercept light over a large area at the expense of a reduced height growth. Crown length was negatively correlated with Hmax at intermediate reference heights (4-14 m). A long crown enables small understory species to maximize light interception in a light-limited environment. Light-demanding species were characterized by orthotropic stems and branches, large leaves, and a monolayer leaf arrangement. They realized an efficient height growth through the formation of narrow and shallow crowns. Light demand turned out to be a much stronger predictor of tree architecture than Hmax, probably because of the relatively low, open, and semi-evergreen canopy at the research site. The existence of four functional groups (shade-tolerant, partial-shade-tolerant, and long- and short-lived pioneer) was confirmed by the principal component and discriminant analysis. Both light demand and Hmax capture the major variation in functional traits found among tropical rain forest tree species, and the two-way classification scheme provides a straightforward model to understand niche differentiation in tropical forests.