Detrital nutrient content determines growth rate and elemental composition of bromeliad‐dwelling insects

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Growth rate–stoichiometry couplings in diverse biota

Biological stoichiometry provides a mechanistic theory linking cellular and biochemical features of co‐evolving biota with constraints imposed by ecosystem energy and nutrient inputs. Thus, understanding variation in biomass carbon : nitrogen : phosphorus (C : N : P) stoichiometry is a major priority for integrative biology. Among various factors affecting organism stoichiometry, differences in C : P and N : P stoichiometry have been hypothesized to reflect organismal P‐content because of altered allocation to P‐rich ribosomal RNA at different growth rates (the growth rate hypothesis, GRH). We tested the GRH using data for microbes, insects, and crustaceans and we show here that growth, RNA content, and biomass P content are tightly coupled across species, during ontogeny, and under physiological P limitation. We also show, however, that this coupling is relaxed when P is not limiting for growth. The close relationship between P and RNA contents indicates that ribosomes themselves represent a biogeochemically significant repository of P in ecosystems and that allocation of P to ribosome generation is a central process in biological production in ecological systems.     

Interspecific competition alters leaf stoichiometry in 20 grassland species

The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species reflects resource capture dynamics under competition. However, the impact of local plant diversity on species‐specific stoichiometry remains poorly studied. Here, we analyze how spatial and temporal diversity in resource‐acquisition traits affects leaf elemental stoichiometry of plants (i.e. the result of resource capture) and how flexible this stoichiometry is depending on the functional composition of the surrounding community. Therefore, we assessed inter‐ and intraspecific variations of leaf carbon (C), nitrogen (N), and phosphorus (P) (and their ratios) of 20 grassland species in a large trait‐based plant diversity experiment located in Jena (Germany) by measuring leaf elemental concentrations at the species‐level along a gradient in plant trait dissimilarity. Our results show that plants showed large intra‐ and interspecific variation in leaf stoichiometry, which was only partly explained by the functional group identity (grass or herb) of the species. Elemental concentrations (N, P, but not C) decreased with plant species richness, and species tended to become more deviant from their monoculture stoichiometry with increasing trait dissimilarity in the community. These responses differed among species, some consistently increased or decreased in P and N concentrations; for other species, the negative or positive change in P and N concentrations increased with increasing trait difference between the target species and the remaining community. The strength of this relationship was significantly associated to the relative position of the species along trait gradients related to resource acquisition. Trait‐difference and trait‐diversity thus were important predictors of how species’ resource capture changed in competitive neighbourhoods.     

刨花楠不同相对生长速率下林木叶片碳氮磷的适应特征

分别对9年生与13年生刨花楠林木叶片氮磷养分之间关系及林木生物量相对生长速率与叶片碳氮磷化学计量比关系进行分析,探讨不同相对生长速率下的林木叶片N、P养分适应特征,并检验相对生长速率假说理论对刨花楠树种的适应性。结果表明:两种年龄刨花楠林木生物量相对生长速率、叶片C、N、P含量及其计量比值均存在显著差异;同一年龄的林木叶片N、P之间存在显著相关性,二者具有协同相关性;9年生林木叶片P含量及C∶P、N∶P与生物量相对生长速率呈二次曲线相关,而13年生林木叶片N、P含量及C∶N、C∶P、N∶P则与生物量相对生长速率均呈线性相关。研究表明,在能满足植物生长所需养分供给的土壤环境中,叶片N、P含量与林木相对生长速率间呈线性正相关,但当土壤中养分供应满足不了植物高速生长时,植物则会对有限的养分资源进行适应性调整。     

Stoichiometric imbalances between detritus and detritivores are related to shifts in ecosystem functioning

How are resource consumption and growth rates of litter‐consuming detritivores affected by imbalances between consumer and litter C:N:P ratios? To address this question, we offered leaf litter as food to three aquatic detritivore species, which represent a gradient of increasing body N:P ratios: a crustacean, a caddisfly and a stonefly. The detritivores were placed in microcosms and submerged in a natural stream. Four contrasting leaf species were offered, both singly and in two‐species mixtures, to obtain different levels of stoichiometric imbalance between the resources and their consumers. The results suggest that detritivore growth was constrained by N rather than C or P, even though 1) the N:P ratios of the consumers’ body tissue was relatively low and 2) microbial leaf conditioning during the experiment reduced the N:P imbalance between detritivores and leaf litter. This surprisingly consistent N limitation may be a consequence of cumulative N‐demand arising from the production of N‐rich chitin in the exoskeletons of all three consumer species, which is lost during regular moults, in addition to N‐demand for silk production by the caddisfly. These N requirements are not commonly quantified in stoichiometric analyses of arthropod consumers. There was no evidence for compensatory feeding, but when offered mixed‐species litter varying in C:N:P ratios, detritivores consumed more of the litter species showing the highest N:P and lowest C:N ratio, accelerating the mass loss of the preferred leaf species in the litter mixture. These results show that imbalances in consumer–resource stoichiometry can have contrasting effects on coupled processes, highlighting a challenge in developing a mechanistic understanding of the role of stoichiometry in regulating ecosystem processes such as leaf litter decomposition.     

滇池流域富磷区不同土壤磷水平下植物叶片的养分化学计量特征

滇池流域是我国典型的富磷区, 分析该区域内不同土壤磷含量下主要植物的化学计量特征, 有助于理解该区域的生态环境特点和生态恢复的特殊性。该研究测定了滇池流域滇中地区75种常见植物叶片的碳(C)、氮(N)、磷(P)及钾(K)含量, 综合分析了该区域不同土壤磷水平(富磷和正常)下不同生活型植物叶片的C、N、P和K的计量特征。结果表明, 研究区域植物叶片C、N和K含量的算术平均数分别是441.42、16.17和13.57 mg·g^-1, P含量的几何平均数为1.92 mg·g^-1, 植物叶片的N、P和K含量之间呈显著的正相关; 富磷区域植物叶片的P和K含量显著高于正常区域, N/P、K/P显著低于正常区域。无论是富磷还是正常区域, 草本植物的N、P和K含量均高于木本植物, 乔木与灌木差异不明显。植物叶片的P含量及N/P与土壤磷水平呈显著相关; 叶片N/P分析结果表明, N是影响滇池流域植物生长和群落恢复的主要限制元素。研究指出, 在滇池流域增加陆地植物群落及生态系统的氮素来源是进行生态修复和面源污染防治的重要切入点。     

云南西双版纳常见寄生植物生态化学计量特征及其与寄主的关系

对云南西双版纳地区常见的半寄生植物澜沧江寄生(Scurrula chingii var. yunnanensis)、五蕊寄生(Dendrophthoe pentandra)和全寄生植物优势种金灯藤(Cuscuta japonica)的生态化学计量特征及其与寄主关系进行研究。结果表明,相较于普通高等陆生植物,两种半寄生植物叶片的C、N含量及N:P较低,P含量较高,全寄生植物金灯藤的N、P含量较高,N:P较低;两种半寄生植物C:N、C:P均显著高于全寄生植物金灯藤。两类寄生植物C、N、P生态化学计量特征季节差异不显著。相较于全寄生植物,半寄生植物与寄主间具有更强的相关性。全寄生植物金灯藤的化学计量内稳性指数显著高于两种半寄生植物,全寄生植物内稳性雨季显著高于旱季。     

Close linkages between leaf functional traits and soil and leaf C:N:P stoichiometry under altered precipitation in a desert steppe in northwestern China

Leaf functional traits are important for characterizing plant nutrient strategies. The C:N:P stoichiometric balance in soils and plants, which could indicate types of nutrient limitation, is altered under changing precipitation patterns. However, whether such alterations affect leaf functional traits remains unclear. We conducted a three-year simulated precipitation experiment in a desert steppe in northwestern China to determine changes in leaf photosynthetic traits and nutrient conservation traits in five plant species and tested the relationships of these traits with soil and leaf C:N:P stoichiometry. The five species showed few changes in their leaf traits under drought conditions, but they adjusted these traits (especially P traits) under extremely wet conditions (50% increase in precipitation). Improved leaf photosynthetic N and P use, lowered leaf P uptake, and enhanced leaf N resorption might help Lespedeza potaninii to rely less on soil nutrients in extremely wet environments than other species do. Leaf photosynthetic traits were regulated primarily by soil and leaf C:N:P stoichiometry. Leaf nutrient conservation traits were controlled by both leaf C:N:P stoichiometry and soil properties (i.e., enzyme activity and microbial biomass), a condition especially true for P traits. The results suggest that precipitation-induced alteration in the C:N:P stoichiometric balance might have important influences on plant nutrient use strategies and even on the nutrient cycling of desert steppes.

     

C:N:P Stoichiometry and Leaf Traits of Halophytes in an Arid Saline Environment,Northwest China

Salinization is an important and increasingly prevalent issue which has broad and profound effects on plant survival and distribution pattern. To understand the patterns and potential drivers of leaf traits in saline environments, we determined the soil properties, leaf morphological traits (specific leaf area, SLA, and leaf dry matter content, LDMC), leaf chemical traits (leaf carbon, C, nitrogen, N, and phosphorus, P, stoichiometry) based on 142 observations collected from 23 sites in an arid saline environment, which is a vulnerable ecosystem in northwest China. We also explored the relationships among leaf traits, the responses of leaf traits, and plant functional groups (herb, woody, and succulent woody) to various saline environments. The arid desert halophytes were characterized by lower leaf C and SLA levels, higher N, but stable P and N:P. The leaf morphological traits were correlated significantly with the C, N, and P contents across all observations, but they differed within each functional group. Succulent woody plants had the lowest leaf C and highest leaf N levels among the three functional groups. The growth of halophytes might be more limited by N rather than P in the study area. GLM analysis demonstrated that the soil available nutrients and plant functional groups, but not salinity, were potential drivers of leaf C:N:P stoichiometry in halophytes, whereas species differences accounted for the largest contributions to leaf morphological variations. Our study provides baseline information to facilitate the management and restoration of arid saline desert ecosystem.     

Zooplankton eat what they need: copepod selective feeding and potential consequences for marine systems

Herbivores are generally faced with a plethora of resources which differ in quality. Therefore, they should be able to select foods which most closely match their metabolic needs. Here, we tested the hypothesis that copepods of the species Acartia tonsa select prey cells based on quality differences within prey species. We assessed age‐specific variation in feeding behaviour and evaluated the potential consequences of such variation for nutrient cycles. Nauplii (young) stages characterized by a low nitrogen to phosphorus (N:P) ratio in their body tissue selected for phosphorus‐rich food, while older copepodite stages with higher body N:P selected for nitrogen‐rich food. Further, the analysis of a 35‐year data set in the southern North Sea revealed a positive correlation between the abundance of nauplii and the ratio of dissolved inorganic N:P, thus suggesting that P‐availability for primary producers declines with the population densities of nauplii. Our findings demonstrate that a combination of stage‐specific selective feeding and body stoichiometry has the potential to affect cycling of limiting nutrients when consumer populations change in composition.     

Microhabitat specialization and similarity of scale-insect assemblages on different fruit trees and in different countries

Abstract. 1. Assemblages of scale-insects on deciduous fruit trees were compared in a wide geographic range of Palaearctic subregions.
2. The parts of all host plant species in the whole geographic region were unsaturated with scale species. There was no clear niche expansion, nor any niche segregation between scale-insect species.
3. These results show that even plants such as deciduous trees, rich in coccoid species, are not fully utilized by scale insects.
4. The similarity of the scale-insect faunas on different plant species could be explained partly by the distribution patterns of the insects, and partly by phylogenetic connection between host plants.     

Stoichiometric traits (N:P) of understory plants contribute to reductions in plant diversity following long‐term nitrogen addition in subtropical forest

Nitrogen enrichment is pervasive in forest ecosystems, but its influence on understory plant communities and their stoichiometric characteristics is poorly understood. We hypothesize that when forest is enriched with nitrogen (N), the stoichiometric characteristics of plant species explain changes in understory plant diversity. A 13‐year field experiment was conducted to explore the effects of N addition on foliar carbon (C): N: phosphorus (P) stoichiometry, understory plant species richness, and intrinsic water use efficiency (iWUE) in a subtropical Chinese fir forest. Four levels of N addition were applied: 0, 6, 12, and 24 g m⁻² year⁻¹. Individual plant species were categorized into resistant plants, intermediate resistant plants, and sensitive plants based on their response to nitrogen addition. Results showed that N addition significantly decreased the number of species, genera, and families of herbaceous plants. Foliar N:P ratios were greater in sensitive plants than resistant or intermediate resistant plants, while iWUE showed an opposite trend. However, no relationship was detected between soil available N and foliar N, and soil N:P and foliar N:P ratios. Our results indicated that long‐term N addition decreased the diversity of understory plants in a subtropical forest. Through regulating water use efficiency with N addition, sensitive plants change their N:P stoichiometry and have a higher risk of mortality, while resistant plants maintain a stable N:P stoichiometry, which contributes to their survival. These findings suggest that plant N:P stoichiometry plays an important role in understory plant performance in response to environmental change of N.     

UV‐B impact on aphid performance mediated by plant quality and plant changes induced by aphids

Plants face various abiotic and biotic environmental factors and therefore need to adjust their phenotypic traits on several levels. UV‐B radiation is believed to impact herbivorous insects via host plant changes. Plant responses to abiotic challenges (UV‐B radiation) and their interaction with two aphid species were explored in a multifactor approach. Broccoli plants [Brassica oleracea L. convar. botrytis (L.), Brassicaceae] were grown in two differently covered greenhouses, transmitting either 80% (high UV‐B) or 4% (low UV‐B) of ambient UV‐B. Three‐week‐old plants were infested with either specialist cabbage aphids [Brevicoryne brassicae (L.), Sternorrhyncha, Aphididae] or generalist green peach aphids [Myzus persicae (Sulzer), Sternorrhyncha, Aphididae]. Plants grown under high‐UV‐B intensities were smaller and had higher flavonoid concentrations. Furthermore, these plants had reduced cuticular wax coverage, whereas amino acid concentrations of the phloem sap were little influenced by different UV‐B intensities. Cabbage aphids reproduced less on plants grown under high UV‐B than on plants grown under low UV‐B, whereas reproduction of green peach aphids in both plant light sources was equally poor. These results are likely related to the different specialisation‐dependent sensitivities of the two species. The aphids also affected plant chemistry. High numbers of cabbage aphid progeny on low‐UV‐B plants led to decreased indolyl glucosinolate concentrations. The induced change in these glucosinolates may depend on an infestation threshold. UV‐B radiation considerably impacts plant traits and subsequently affects specialist phloem‐feeding aphids, whereas aphid growth forces broccoli to generate specific defence responses.     

Does host‐plant diversity explain species richness in insects? A test using Coccidae (Hemiptera)

1. The megadiverse herbivores and their host plants are a major component of biodiversity, and their interactions have been hypothesised to drive the diversification of both. 2. If plant diversity influences the diversity of insects, there is an expectation that insect species richness will be strongly correlated with host‐plant species richness. This should be observable at two levels (i) more diverse host‐plant groups should harbour more species of insects, and (ii) the species richness of a group of insects should correlate with the richness of the host groups it uses. However, such a correlation is also consistent with a hypothesis of random host use, in which insects encounter and use hosts in proportion to the diversity of host plants. Neither of these expectations has been widely tested. 3. These expectations were tested using data from a species‐rich group of insects – the Coccidae (Hemiptera). 4. Significant positive correlations were found between the species richness of coccid clades (genera) and the species richness of the host‐plant family or families upon which the clades occur. On a global scale, more closely related plant families have more similar communities of coccid genera but the correlation is weak. 5. Random host use could not be rejected for many coccids but randomisation tests and similarity of coccid communities on closely related plant families show that there is non‐random host use in some taxa. Overall, our results support the idea that plant diversity is a driver of species richness of herbivorous insects, probably via escape‐and‐radiate or oscillation‐type processes.     

植物生态化学计量特征及其主要假说

植物生态化学计量学是生态化学计量学的重要分支, 主要研究植物器官元素含量的计量特征, 以及它们与环境因子、生态系统功能之间的关系。19世纪, 化学家们通过室内实验, 分析了植物器官的元素含量, 开始了对植物化学元素之间关系的探索。如今, 生态学家通过野外采样和控制实验, 探索植物化学元素计量特征的变化规律、对全球变化的响应以及与植物功能属性之间的关系, 促进了植物生态化学计量学的快速发展。该文在概述植物生态化学计量学发展简史的基础上, 综述了19世纪以来该领域的研究进展。首先, 该文将植物生态化学计量学的发展历程概括为思想萌芽期、假说奠基期和理论构建期3个时期, 对各个时期的主要研究进行了简要回顾和梳理。第二, 概述了植物主要器官的化学计量特征, 尤其是陆生植物叶片氮(N)和磷(P)的计量特征。总体上, 全球陆生植物叶片N、P含量和N:P (质量比)的几何平均值分别为18.74 mg∙g^-1、1.21 mg∙g^-1和15.55 (与16:1的Redfield比一致); 在物种或群落水平上, 叶片N和P含量一般呈现随温度升高、降水增加而降低的趋势。不同生活型植物叶片N和P计量特征差异明显, 尤其是草本植物叶片N和P含量高于木本植物, 落叶阔叶木本植物叶片N和P含量高于常绿木本植物。与叶片相比, 细根和其他器官化学计量特征研究较少。第三, 总结了养分添加实验对植物化学元素计量特征的影响。总体上, N添加一般会提高土壤N的可利用性, 使植物器官中N含量和N:P升高, 在一定程度上提高植物生产力; P添加可能会缓解过量N输入导致的N-P失衡问题, 提高植物器官P含量。但是, 长期过量施肥会打破植物器官原有的元素间计量关系, 导致元素计量关系失衡和生产力下降。第四, 梳理总结了植物生态化学计量学的重要理论、观点和假说, 主要包括刻画化学计量特征与植物生长功能关系的功能关联假说、刻画化学计量特征与环境因子关系的环境关联假说或理论以及刻画化学计量特征与植物进化历史关系的进化关联假说。最后, 指出了植物生态化学计量学研究中存在的问题, 展望了10个未来需要重点关注的研究方向。     

Resolving the ‘nitrogen paradox’ of arbuscular mycorrhizas: fertilization with organic matter brings considerable benefits for plant nutrition and growth

Arbuscular mycorrhizal fungi (AMF) can transfer nitrogen (N) to host plants, but the ecological relevance is debated, as total plant N and biomass do not generally increase. The extent to which the symbiosis is mutually beneficial is thought to rely on the stoichiometry of N, phosphorus (P) and carbon (C) availability. While inorganic N fertilization has been shown to elicit strong mutualism, characterized by improved plant and fungal growth and mineral nutrition, similar responses following organic N addition are lacking. Using a compartmented microcosm experiment, we determined the significance to a mycorrhizal plant of placing a ¹⁵N‐labelled, nitrogen‐rich patch of organic matter in a compartment to which only AMF hyphae had access. Control microcosms denied AMF hyphal access to the patch compartment. When permitted access to the patch compartment, the fungus proliferated extensively in the patch and transferred substantial quantities of N to the plant. Moreover, our data demonstrate that allowing hyphal access to an organic matter patch enhanced total plant N and P contents, with a simultaneous and substantial increase in plant biomass. Furthermore, we demonstrate that organic matter fertilization of arbuscular mycorrhizal plants can foster a mutually beneficial symbiosis based on nitrogen transfer, a phenomenon previously thought irrelevant.     

Tracking sucking herbivory with nitrogen isotope labelling: Lessons from an individual trait-based approach

Response and effect traits help to understand how changes in ecological communities (e.g. in response to land use) relate to changes in ecosystem functioning. In grasslands, plants and insect herbivores are involved in many ecosystem processes such as herbivory and plant biomass production. Simultaneous changes in the trait composition of both plants and herbivores should affect herbivory rates, with consequences for plant growth and potentially biomass production. The mechanisms underlying these links are little understood for grasses and sucking insects, which build a major part of grassland communities. In a mesocosm experiment, we manipulated the composition of grasses and sucking herbivores (Hemiptera) to study the role of plant traits, herbivore traits and their interaction on herbivory and plant growth. Because sucking herbivory is generally difficult to quantify, we developed a novel experimental setting, in which we labelled plants with ¹⁵N isotope. This allowed to quantify ¹⁵N uptake and thus sucking rates of individuals. We found that herbivory and simultaneous plant growth reduction are most strongly linked to herbivore species identity. Unexpectedly, herbivory did not increase with herbivore size, but was highest for small species and for thin-bodied Heteroptera. Additionally, herbivory and plant growth reduction depended on the interacting herbivore and plant species, indicating trait matching, which could, however, not be explained with commonly used traits. This indicates that mechanisms linking ecological communities and ecosystem processes are highly context-specific. To understand how global change affects ecosystem functioning, studies need to cover all functionally relevant groups, including plant sap suckers.     

Leaf nitrogen and phosphorus concentrations of woody plants differ in responses to climate,soil and plant growth form

Leaf chemistry is important in predicting the functioning and dynamics of ecosystems. As two key traits, leaf nitrogen (N) and phosphorus (P) concentrations set the limits for plant growth, and leaf N:P ratios indicate the shift between N‐ and P‐limitation. To understand the responses of leaf chemistry to their potential drivers, we measured leaf N and P concentrations of 386 woody species at 14 forest sites across eastern China, and explored the effects of climate, soil, and plant growth form on leaf N, P and N:P ratios. In general, leaf N and P were both negatively related to mean annual temperature and precipitation, and positively related to soil N and P concentrations. Leaf N:P ratios showed opposite trends. General linear models showed that variation in leaf N was mainly determined by a shift in plant growth form (from evergreen broadleaved to deciduous broadleaved to conifer species) along the latitudinal gradient, while variations in leaf P and N:P were driven by climate, plant growth form, and their interaction. These differences may reflect differences in nutrient cycling and physiological regulations of P and N. Our results should help understand the ecological patterns of leaf chemical traits and modeling ecosystem nutrient cycling.     

Indirect food web interactions increase growth of an algivorous stream fish

1. We tested the hypothesis that indirect food web interactions between some common, invertivorous fishes and their prey would positively affect growth of an algivorous fish species. Specifically, we predicted that orangethroat darter (Etheostoma spectabile) would increase periphyton biomass via a top‐down pathway, indirectly enhancing growth of the algivorous central stoneroller minnow (Campostoma anomalum). Moreover, we predicted that sand shiner (Notropis stramineus) would increase periphyton biomass via a bottom‐up pathway and indirectly enhance growth of the stoneroller minnow. 2. In an 83‐day experiment in large, outdoor, stream mesocosms, we stocked two fish species per mesocosm (stoneroller and either darter or shiner), estimated the effects of the invertivorous and grazing fishes on periphyton biomass and estimated growth of the algivorous fish. 3. The darter consumed grazing invertebrates, indirectly increasing periphyton biomass. The shiner consumed terrestrial insects as predicted, but it did not affect periphyton biomass. 4. In support of our hypothesis, the darter indirectly enhanced stoneroller growth. As predicted, stonerollers consumed the increased periphyton in streams with darters, resulting in greater growth, condition and gut fullness compared to streams without darters. No indirect interaction was observed between stonerollers and shiners. 5. Our study suggests that some invertivorous fish species can positively affect growth of algivorous fishes through indirect food web interactions. Thus, in stream communities, it is possible that the loss of a single, invertivorous fish taxon could have negative consequences on algivorous fish populations via the removal of positive indirect food web interactions.     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.