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Tropical soils contain huge carbon stocks, which climate warming is projected to reduce by stimulating organic matter decomposition, creating a positive feedback that will promote further warming. Models predict that the loss of carbon from warming soils will be mediated by microbial physiology, but no empirical data are available on the response of soil carbon and microbial physiology to warming in tropical forests, which dominate the terrestrial carbon cycle. Here we show that warming caused a considerable loss of soil carbon that was enhanced by associated changes in microbial physiology. By translocating soils across a 3000 m elevation gradient in tropical forest, equivalent to a temperature change of ± 15 °C, we found that soil carbon declined over 5 years by 4% in response to each 1 °C increase in temperature. The total loss of carbon was related to its original quantity and lability, and was enhanced by changes in microbial physiology including increased microbial carbon‐use‐efficiency, shifts in community composition towards microbial taxa associated with warmer temperatures, and increased activity of hydrolytic enzymes. These findings suggest that microbial feedbacks will cause considerable loss of carbon from tropical forest soils in response to predicted climatic warming this century.

The functional traits of organisms within multispecies assemblages regulate biodiversity effects on ecosystem functioning. Yet how traits should assemble to boost multiple ecosystem functions simultaneously (multifunctionality) remains poorly explored. In a multibiome litter experiment covering most of the global variation in leaf trait spectra, we showed that three dimensions of functional diversity (dispersion, rarity, and evenness) explained up to 66% of variations in multifunctionality, although the dominant species and their traits remained an important predictor. While high dispersion impeded multifunctionality, increasing the evenness among functionally dissimilar species was a key dimension to promote higher multifunctionality and to reduce the abundance of plant pathogens. Because too-dissimilar species could have negative effects on ecosystems, our results highlight the need for not only diverse but also functionally even assemblages to promote multifunctionality. The effect of functionally rare species strongly shifted from positive to negative depending on their trait differences with the dominant species. Simultaneously managing the dispersion, evenness, and rarity in multispecies assemblages could be used to design assemblages aimed at maximizing multifunctionality independently of the biome, the identity of dominant species, or the range of trait values considered. Functional evenness and rarity offer promise to improve the management of terrestrial ecosystems and to limit plant disease risks., This work was funded by the British Ecological Society (SR17\1297 grant, PI: P.G.-P.) and by the European Research Council (ERC Grant Agreement #647038, BIODESERT, PI: F.T.M.). Y.L.B.-P. was supported by a Marie Sklodowska-Curie Actions Individual Fellowship within the European Program Horizon 2020 (DRYFUN Project #656035). H.S. was supported by a Juan de la Cierva-Formación grant from the Span ish Ministry of Economy and Competitiveness (FJCI-2015-26782). F.T.M. and S.A. were supported from the Generalitat Valenciana (CIDEGENT/ 2018/041). M.D. was supported by a Formación del Profesorado Universi tario (FPU) fellowship from the Spanish Ministry of Education, Culture and Sports (FPU-15/00392). S.A. was supported by the Spanish MINECO for financial support via the DIGGING_DEEPER project through the 2015 to 2016 BiodivERsA3/FACCE‐JPI joint call for research proposals. B.K.S. re search on biodiversity-ecosystem functions was supported by the Austra lian Research Council (DP170104634 and DP190103714). P.G.-P. was supported by a Ramón y Cajal grant from the Spanish Ministry of Science and Innovation (RYC2018-024766-I). R.M. was supported by MINECO (Grants CGL2014-56567-R and CGL2017-83855-R)

Understanding how evolutionary constraints shape the elevational distributions of tree lineages provides valuable insight into the future of tropical montane forests under global change. With narrow elevational ranges, high taxonomic turnover, frequent habitat specialization, and exceptional levels of endemism, tropical montane forests and trees are predicted to be highly sensitive to environmental change. Using plot census data from a gradient traversing > 3,000 m in elevation on the Amazonian flank of the Peruvian Andes, we employ phylogenetic approaches to assess the influence of evolutionary heritage on distribution trends of trees at the genus‐level. We find that closely related lineages tend to occur at similar mean elevations, with sister genera pairs occurring a mean 254 m in elevation closer to each other than the mean elevational difference between non‐sister genera pairs. We also demonstrate phylogenetic clustering both above and below 1,750 m a.s.l, corresponding roughly to the cloud‐base ecotone. Belying these general trends, some lineages occur across many different elevations. However, these highly plastic lineages are not phylogenetically clustered. Overall, our findings suggest that tropical montane forests are home to unique tree lineage diversity, constrained by their evolutionary heritage and vulnerable to substantial losses under environmental changes, such as rising temperatures or an upward shift of the cloud‐base.

The sizes and shapes of tree crowns are of fundamental importance in ecology, yet understanding the forces that determine them remains elusive. A cardinal question facing ecologists is the degree to which general and non-specific versus ecological and context-dependent processes are responsible for shaping tree crowns. Here, we test this question for the first time across diverse tropical ecosystems. Using trees from 20 plots varying in elevation, precipitation, and ecosystem type (savanna-forest transitions) across the paleo- and neo-tropics, we test the relationship between crown dimensions and tree size. By analyzing these scaling relationships across environmental gradients, biogeographic regions, and phylogenetic distance, we extend Metabolic Scaling Theory (MST) predictions to include how local selective pressures shape variation in crown dimensions. Across all sites, we find strong agreement between mean trends and MST predictions for the scaling of crown size and shape, but large variation around the mean. While MST explained approximately half of the observed variation in tree crown dimensions, we find that local, ecosystem, and phylogenetic predictors account for the half of the residual variation. Crown scaling does not change significantly across regions, but does change across ecosystem types, where savanna tree crowns grow more quickly with tree girth than forest tree crowns. Crowns of legumes were wider and more voluminous than those of other taxa. Thus, while MST can accurately describe the central tendency of tree crown size, local ecological conditions and evolutionary history appear to modify the scaling of crown shape. Importantly, our extension of MST incorporating these differences accounts for the mechanisms driving variation in the scaling of crown dimensions across the tropics. We present allometric equations for the prediction of crown dimensions across tropical ecosystems. These results are critical when scaling the function of individual trees to larger spatial scales or incorporating the size and shape of tree crowns in global biogeochemical models.

Ultraviolet (UV) radiation is a small fraction of the solar spectrum, which acts as a key environmental modulator of plant function affecting metabolic regulation and growth. Plant species endemic to the Andes are well adapted to the harsh features of high-altitude climate, including high UV radiation. Maca (Lepidium meyenii Walpers) is a member of Brassicaceae family native to the central Andes of Peru, which grows between 3500 and 4500 m of altitude, where only highland grasses and few hardy bushes can survive. Even though maca has been the focus of recent researches, mainly due to its nutraceutical properties, knowledge regarding its adaptation mechanisms to these particular natural environmental conditions is scarce. In this study, we manipulated solar UV radiation by using UV-transmitting (Control) or blocking (UV-block) filters under field conditions (4138 m above the sea level) in order to understand the impact of UV on morphological and physiological parameters of maca crops over a complete growing season. Compared to the UV-blocking filter, under control condition a significant increase of hypocotyl weight was observed during the vegetative phase together with a marked leaf turnover. Although parameters conferring photosynthetic performance were not altered by UV, carbohydrate allocation between above and underground organs was affected. Control condition did not influence the content of secondary metabolites such as glucosinolates and phenolic compounds in hypocotyls, while some differences were observed in the rosettes. These differences were mainly related to leaf turnover and the protection of new young leaves in control plants. Altogether, the data suggest that maca plants respond to strong UV radiation at high altitudes by a coordinated remobilization and relocation of metabolites between source and sink organs via a possible UV signaling pathway.

Foliar trait adaptation to sun and shade has been extensively studied in the context of photosynthetic performance of plants, focusing on nitrogen allocation, light capture and use via chlorophyll pigments and leaf morphology; however, less is known about the potential sun-shade dichotomy of other functionally important foliar traits. In this study, we measured 19 traits in paired sun and shade leaves along a 3,500-m elevation gradient in southern Peru to test whether the traits differ with canopy position, and to assess if relative differences vary with species composition and/or environmental filters. We found significant sun-shade differences in leaf mass per area (LMA), photosynthetic pigments (Chl ab and Car), and δ13C. Sun-shade offsets among these traits remained constant with elevation, soil substrates, and species compositional changes. However, other foliar traits related to structure and chemical defense, and those defining general metabolic processes, did not differ with canopy position. Our results suggest that whole-canopy function is captured in many traits of sun leaves; however, photosynthesis-related traits must be scaled based on canopy light extinction. These findings show that top-of-canopy measurements of foliar chemistry from spectral remote sensing approaches map directly to whole-canopy foliar traits including shaded leaves that cannot be directly observed from above.

Remote sensing of trait-based ecology can improve our understanding of the effect of trait diversity on ecosystem functioning., Spatially continuous data on functional diversity will improve our ability to predict global change impacts on ecosystem properties. We applied methods that combine imaging spectroscopy and foliar traits to estimate remotely sensed functional diversity in tropical forests across an Amazon-to-Andes elevation gradient (215 to 3537 m). We evaluated the scale dependency of community assembly processes and examined whether tropical forest productivity could be predicted by remotely sensed functional diversity. Functional richness of the community decreased with increasing elevation. Scale-dependent signals of trait convergence, consistent with environmental filtering, play an important role in explaining the range of trait variation within each site and along elevation. Single- and multitrait remotely sensed measures of functional diversity were important predictors of variation in rates of net and gross primary productivity. Our findings highlight the potential of remotely sensed functional diversity to inform trait-based ecology and trait diversity-ecosystem function linkages in hyperdiverse tropical forests.

Nutrients constrain the soil carbon cycle in tropical forests, but we lack knowledge on how these constraints vary within the soil microbial community. Here, we used in situ fertilization in a montane tropical forest and in two lowland tropical forests on contrasting soil types to test the principal hypothesis that there are different nutrient constraints to different groups of microorganisms during the decomposition of cellulose. We also tested the hypotheses that decomposers shift from nitrogen to phosphorus constraints from montane to lowland forests, respectively, and are further constrained by potassium and sodium deficiency in the western Amazon. Cellulose and nutrients (nitrogen, phosphorus, potassium, sodium, and combined) were added to soils in situ, and microbial growth on cellulose (phospholipid fatty acids and ergosterol) and respiration were measured. Microbial growth on cellulose after single nutrient additions was highest following nitrogen addition for fungi, suggesting nitrogen as the primary limiting nutrient for cellulose decomposition. This was observed at all sites, with no clear shift in nutrient constraints to decomposition between lowland and montane sites. We also observed positive respiration and fungal growth responses to sodium and potassium addition at one of the lowland sites. However, when phosphorus was added, and especially when added in combination with other nutrients, bacterial growth was highest, suggesting that bacteria out-compete fungi for nitrogen where phosphorus is abundant. In summary, nitrogen constrains fungal growth and cellulose decomposition in both lowland and montane tropical forest soils, but additional nutrients may also be of critical importance in determining the balance between fungal and bacterial decomposition of cellulose.

The carbon isotopic composition of plant leaf wax biomarkers is commonly used to reconstruct paleoenvironmental conditions. Adding to the limited calibration information available for modern tropical forests, we analyzed plant leaf and leaf wax carbon isotopic compositions in forest canopy trees across a highly biodiverse, 3.3 km elevation gradient on the eastern flank of the Andes Mountains. We sampled the dominant tree species and assessed their relative abundance in each tree community. In total, 405 sunlit canopy leaves were sampled across 129 species and nine forest plots along the elevation profile for bulk leaf and leaf wax n-alkane (C27–C33) concentration and carbon isotopic analyses (δ13C); a subset (76 individuals, 29 species, five forest plots) were additionally analyzed for n-alkanoic acid (C22–C32) concentrations and δ13C. δ13C values display trends of +0.87 ± 0.16‰ km−1 (95% CI, r2 = 0.96, p

Funding: SF was supported by the São Paulo Research Foundation (FAPESP 2014/13322-9), SF and MG were supported by NERC (AMAZONICA NE/F005806/1, ECOFOR NE/K01944X/1, BIO-RED NE/N012542/1) and ERC (GEM-TRAIT 321121). NF was supported by a Marie Curie Intra-European Fellowship within the EU FP7. BC was partially supported by the EU FP7 project AMAZALERT. Data collection following RAINFOR protocols was supported by several grants from NERC, the EU (T-FORCES 103919, PAN-AMAZONIA 74254, GEOCARBON 283080), and the Moore Foundation (1656, 3000). Climate, species composition, and soils are thought to control carbon cycling and forest structure in Amazonian forests. Here, we add a demographics scheme (tree recruitment, growth, and mortality) to a recently developed non-demographic model—the Trait-based Forest Simulator (TFS)—to explore the roles of climate and plant traits in controlling forest productivity and structure. We compared two sites with differing climates (seasonal vs. aseasonal precipitation) and plant traits. Through an initial validation simulation, we assessed whether the model converges on observed forest properties (productivity, demographic and structural variables) using datasets of functional traits, structure, and climate to model the carbon cycle at the two sites. In a second set of simulations, we tested the relative importance of climate and plant traits for forest properties within the TFS framework using the climate from the two sites with hypothetical trait distributions representing two axes of functional variation (“fast” vs. “slow” leaf traits, and high vs. low wood density). The adapted model with demographics reproduced observed variation in gross (GPP) and net (NPP) primary production, and respiration. However, NPP and respiration at the level of plant organs (leaf, stem, and root) were poorly simulated. Mortality and recruitment rates were underestimated. The equilibrium forest structure differed from observations of stem numbers suggesting either that the forests are not currently at equilibrium or that mechanisms are missing from the model. Findings from the second set of simulations demonstrated that differences in productivity were driven by climate, rather than plant traits. Contrary to expectation, varying leaf traits had no influence on GPP. Drivers of simulated forest structure were complex, with a key role for wood density mediated by its link to tree mortality. Modeled mortality and recruitment rates were linked to plant traits alone, drought-related mortality was not accounted for. In future, model development should focus on improving allocation, mortality, organ respiration, simulation of understory trees and adding hydraulic traits. This type of model that incorporates diverse tree strategies, detailed forest structure and realistic physiology is necessary if we are to be able to simulate tropical forest responses to global change scenarios. Publisher PDF

Wet organic-rich mineral and peat soils in the tropical Andes represent a potentially significant, but little studied, source of methane to the atmosphere. Here we report the results of field and laboratory measurements of soil&ndash, atmosphere methane exchange and associated environmental variables from freely draining upland and inundation prone wetland soils in a humid puna ecosystem in the Southeastern Andes of Peru. Between seasons and across the landscape soil&ndash, atmosphere exchange varied between uptake and emission. Notable hotspots of methane emission, peaking during the wet season, were observed from both upland and wetland soils with particularly strong emissions from moss-accumulating topographic lows. This variability was best explained by the influence of oxygen concentration on methane production in superficial soil horizons.

Understanding variation in key functional traits across gradients in high diversity systems and the ecology of community changes along gradients in these systems is crucial in light of conservation and climate change. We examined inter- and intraspecific variation in leaf mass per area (LMA) of sun and shade leaves along a 3330-m elevation gradient in Peru, and in sun leaves across a forest–savanna vegetation gradient in Brazil. We also compared LMA variance ratios (T-statistics metrics) to null models to explore internal (i.e., abiotic) and environmental filtering on community structure along the gradients. Community-weighted LMA increased with decreasing forest cover in Brazil, likely due to increased light availability and water stress, and increased with elevation in Peru, consistent with the leaf economic spectrum strategy expected in colder, less productive environments. A very high species turnover was observed along both environmental gradients, and consequently, the first source of variation in LMA was species turnover. Variation in LMA at the genus or family levels was greater in Peru than in Brazil. Using dominant trees to examine possible filters on community assembly, we found that in Brazil, internal filtering was strongest in the forest, while environmental filtering was observed in the dry savanna. In Peru, internal filtering was observed along 80% of the gradient, perhaps due to variation in taxa or interspecific competition. Environmental filtering was observed at cloud zone edges and in lowlands, possibly due to water and nutrient availability, respectively. These results related to variation in LMA indicate that biodiversity in species rich tropical assemblages may be structured by differential niche-based processes. In the future, specific mechanisms generating these patterns of variation in leaf functional traits across tropical environmental gradients should be explored. Fil: Neyret, Margot. Ecole Normale Supérieure; Francia Fil: Bentley, Lisa Patrick. University of Oxford; Reino Unido. Sonoma State University; Estados Unidos Fil: Oliveras, Imma. University of Oxford; Reino Unido. Wageningen University; Alemania Fil: Marimon, Beatriz S.. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Marimon-Junior, Ben Hur. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Almeida de Oliveira, Edmar. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Barbosa Passos, Fabio. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Castro Ccoscco, Rosa. Universidad San Antonio Abad de Cusco; Perú Fil: Santos, Josias dos. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Matias Reis, Simone. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Morandi, Paulo S.. Universidade Federal do Mato Grosso do Sul; Brasil Fil: Rayme Paucar, Gloria. Universidad San Antonio Abad de Cusco; Perú Fil: Robles Caceres, Arturo. Universidad San Antonio Abad de Cusco; Perú Fil: Valdez Tejeira, Yolvi. Universidad San Antonio Abad de Cusco; Perú Fil: Yllanes Choque, Yovana. Universidad San Antonio Abad de Cusco; Perú Fil: Salinas, Norma Beatriz. University of Oxford; Reino Unido. Pontificia Universidad Católica de Perú; Perú Fil: Shenkin, Alexander. University of Oxford; Reino Unido Fil: Asner, Gregory P.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Enquist, Brian J.. Arizona State University; Estados Unidos Fil: Malhi, Yadvinder. University of Oxford; Reino Unido

We examined whether variations in photosynthetic capacity are linked to variations in the environment and/or associated leaf traits for tropical moist forests (TMFs) in the Andes/western Amazon regions of Peru. We compared photosynthetic capacity (maximal rate of carboxylation of Rubisco (Vcmax), and the maximum rate of electron transport (Jmax)), leaf mass, nitrogen (N) and phosphorus (P) per unit leaf area (Ma, Na and Pa, respectively), and chlorophyll from 210 species at 18 field sites along a 3300-m elevation gradient. Western blots were used to quantify the abundance of the CO2-fixing enzyme Rubisco. Area- and N-based rates of photosynthetic capacity at 25°C were higher in upland than lowland TMFs, underpinned by greater investment of N in photosynthesis in high-elevation trees. Soil [P] and leaf Pa were key explanatory factors for models of area-based Vcmax and Jmax but did not account for variations in photosynthetic N-use efficiency. At any given Na and Pa, the fraction of N allocated to photosynthesis was higher in upland than lowland species. For a small subset of lowland TMF trees examined, a substantial fraction of Rubisco was inactive. These results highlight the importance of soil- and leaf-P in defining the photosynthetic capacity of TMFs, with variations in N allocation and Rubisco activation state further influencing photosynthetic rates and N-use efficiency of these critically important forests. © 2016 The Authors. New Phytologist © 2016 New Phytologist Trust

Average responses of forest foliar traits to elevation are well understood, but far less is known about trait distributional responses to elevation at multiple ecological scales. This limits our understanding of the ecological scales at which trait variation occurs in response to environmental drivers and change. We analyzed and compared multiple canopy foliar trait distributions using field sampling and airborne imaging spectroscopy along an Andes-to-Amazon elevation gradient. Field-estimated traits were generated from three community-weighting methods, and remotely sensed estimates of traits were made at three scales defined by sampling grain size and ecological extent. Field and remote sensing approaches revealed increases in average leaf mass per unit area (LMA), water, nonstructural carbohydrates (NSCs) and polyphenols with increasing elevation. Foliar nutrients and photosynthetic pigments displayed little to no elevation trend. Sample weighting approaches had little impact on field-estimated trait responses to elevation. Plot representativeness of trait distributions at landscape scales decreased with increasing elevation. Remote sensing indicated elevation-dependent increases in trait variance and distributional skew. Multiscale invariance of LMA, leaf water and NSC mark these traits as candidates for tracking forest responses to changing climate. Trait-based ecological studies can be greatly enhanced with multiscale studies made possible by imaging spectroscopy. Fil: Asner, Gregory P.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Martin, Roberta E.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Anderson, Christopher Brian. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Kryston, Katherine. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Vaughn, Nicholas. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Knapp, David E.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Bentley, Lisa Patrick. University of Oxford; Reino Unido Fil: Shenkin, Alexander. University of Oxford; Reino Unido Fil: Salinas, Norma. University of Oxford; Reino Unido. Pontificia Universidad Católica de Perú; Perú Fil: Sinca, Felipe. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Tupayachi, Raul. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Quispe Huaypar, Katherine. Universidad Nacional de San Antonio Abad del Cusco; Perú Fil: Montoya Pillco, Milenka. Universidad Nacional de San Antonio Abad del Cusco; Perú Fil: Ccori Álvarez, Flor Delis. Universidad Nacional de San Antonio Abad del Cusco; Perú Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Enquist, Brian J.. Arizona State University; Estados Unidos Fil: Malhi, Yadvinder. University of Oxford; Reino Unido

More than 200 years ago, Alexander von Humboldt reported that tropical plant species richness decreased with increasing elevation and decreasing temperature. Surprisingly, coordinated patterns in plant, bacterial, and fungal diversity on tropical mountains have not yet been observed, despite the central role of soil microorganisms in terrestrial biogeochemistry and ecology. We studied an Andean transect traversing 3.5 km in elevation to test whether the species diversity and composition of tropical forest plants, soil bacteria, and fungi follow similar biogeographical patterns with shared environmental drivers. We found coordinated changes with elevation in all three groups: species richness declined as elevation increased, and the compositional dissimilarity among communities increased with increased separation in elevation, although changes in plant diversity were larger than in bacteria and fungi. Temperature was the dominant driver of these diversity gradients, with weak influences of edaphic properties, including soil pH. The gradients in microbial diversity were strongly correlated with the activities of enzymes involved in organic matter cycling, and were accompanied by a transition in microbial traits towards slower‐growing, oligotrophic taxa at higher elevations. We provide the first evidence of coordinated temperature‐driven patterns in the diversity and distribution of three major biotic groups in tropical ecosystems: soil bacteria, fungi, and plants. These findings suggest that interrelated and fundamental patterns of plant and microbial communities with shared environmental drivers occur across landscape scales. These patterns are revealed where soil pH is relatively constant, and have implications for tropical forest communities under future climate change.

The temperature sensitivity of soil organic matter (SOM) decomposition in tropical forests will influence future climate. Studies of a 3.5-kilometer elevation gradient in the Peruvian Andes, including short-term translocation experiments and the examination of the long-term adaptation of biota to local thermal and edaphic conditions, have revealed several factors that may regulate this sensitivity. Collectively this work suggests that, in the absence of a moisture constraint, the temperature sensitivity of decomposition is regulated by the chemical composition of plant debris (litter) and both the physical and chemical composition of preexisting SOM: higher temperature sensitivities are found in litter or SOM that is more chemically complex and in SOM that is less occluded within aggregates. In addition, the temperature sensitivity of SOM in tropical montane forests may be larger than previously recognized because of the presence of “cold-adapted” and nitrogen-limited microbial decomposers and the possible future alterations in plant and microbial communities associated with warming. Studies along elevation transects, such as those reviewed here, can reveal factors that will regulate the temperature sensitivity of SOM. They can also complement and guide in situ soil-warming experiments, which will be needed to understand how this vulnerability to temperature may be mediated by altered plant productivity under future climatic change.

Soils store about four times as much carbon as plant biomass1, and soil microbial respiration releases about 60 petagrams of carbon per year to the atmosphere as carbon dioxide2. Short-term experiments have shown that soil microbial respiration increases exponentially with temperature3. This information has been incorporated into soil carbon and Earth-system models, which suggest that warming-induced increases in carbon dioxide release from soils represent an important positive feedback loop that could influence twenty-first-century climate change4. The magnitude of this feedback remains uncertain, however, not least because the response of soil microbial communities to changing temperatures has the potential to either decrease5,6,7 or increase8,9 warming-induced carbon losses substantially. Here we collect soils from different ecosystems along a climate gradient from the Arctic to the Amazon and investigate how microbial community-level responses control the temperature sensitivity of soil respiration. We find that the microbial community-level response more often enhances than reduces the mid- to long-term (90 days) temperature sensitivity of respiration. Furthermore, the strongest enhancing responses were observed in soils with high carbon-to-nitrogen ratios and in soils from cold climatic regions. After 90 days, microbial community responses increased the temperature sensitivity of respiration in high-latitude soils by a factor of 1.4 compared to the instantaneous temperature response. This suggests that the substantial carbon stores in Arctic and boreal soils could be more vulnerable to climate warming than currently predicted. This work was carried out with Natural Environment Research Council (NERC) funding (grant number NE/H022333/1). K.K. was supported by an Academy of Finland post-doctoral research grant while finalizing this manuscript. P.M. was supported by ARC FT110100457 and NERC NE/G018278/1, and B.K.S by the Grain Research and Development Corporation and ARC DP130104841.

Solar irradiance and precipitation are the most likely drivers of the seasonal variation of net primary productivity (NPP) in tropical forests. Since their roles remain poorly understood, we use litter traps, dendrometer bands and census data collected from one hectare permanent plots to quantify the seasonality of above-groundNPPcomponents and weather parameters in 13 sites distributed along a 2800-m altitudinal gradient ranging from lowland Amazonia to the high Andes. We combine canopy leaf area index and litterfall data to describe the seasonality of canopy production. We hypothesize that solar irradiance is the primary driver of canopy phenology in wetter sites, whereas precipitation drives phenology in drier systems. The seasonal rhythm of canopyNPPcomponents is in synchrony with solar irradiance at all altitudes. Leaf litterfall peaks in the late dry season, both in lowland (averaging 0.54 ± 0.08 Mg C ha y−1, n = 5) and montane forests (averaging 0.29 ± 0.04 Mg C ha y−1, n = 8). Peaks in above-ground coarse woodyNPPappears to be triggered by the onset of rainfall in seasonal lowland rain forests (averaging 0.26 ± 0.04 Mg C ha y−1, n = 5, in November), but not in montane cloud forests.

Aim: To examine variation in the phylogenetic diversity (PD) of tree communities across geographical and environmental gradients in Amazonia. Location: Two hundred and eighty-three c. 1 ha forest inventory plots from across Amazonia. Methods: We evaluated PD as the total phylogenetic branch length across species in each plot (PDss), the mean pairwise phylogenetic distance between species (MPD), the mean nearest taxon distance (MNTD) and their equivalents standardized for species richness (ses.PDss, ses.MPD, ses.MNTD). We compared PD of tree communities growing (1) on substrates of varying geological age; and (2) in environments with varying ecophysiological barriers to growth and survival. Results: PDss is strongly positively correlated with species richness (SR), whereas MNTD has a negative correlation. Communities on geologically young- and intermediate-aged substrates (western and central Amazonia respectively) have the highest SR, and therefore the highest PDss and the lowest MNTD. We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses.PDss and ses.MNTD. MPD and ses.MPD are strongly correlated with how evenly taxa are distributed among the three principal angiosperm clades and are both highest in western Amazonia. Meanwhile, seasonally dry tropical forest (SDTF) and forests on white sands have low PD, as evaluated by any metric. Main conclusions: High ses.PDss and ses.MNTD reflect greater lineage diversity in communities. We suggest that high ses.PDss and ses.MNTD in western Amazonia results from its favourable, easy-to-colonize environment, whereas high values in the Brazilian and Guianan Shields may be due to accumulation of lineages over a longer period of time. White-sand forests and SDTF are dominated by close relatives from fewer lineages, perhaps reflecting ecophysiological barriers that are difficult to surmount evolutionarily. Because MPD and ses.MPD do not reflect lineage diversity per se, we suggest that PDss, ses.PDss and ses.MNTD may be the most useful diversity metrics for setting large-scale conservation priorities. © 2015 John Wiley & Sons Ltd.

One of the major challenges in ecology is to understand how ecosystems respond to changes inenvironmental conditions, and how taxonomic and functional diversity mediate these changes. Inthis study, we use a trait-spectra and individual-based model, to analyse variation in forest primaryproductivity along a 3.3 km elevation gradient in the Amazon-Andes. The model accuratelypredicted the magnitude and trends in forest productivity with elevation, with solar radiation andplant functional traits (leaf dry mass per area, leaf nitrogen and phosphorus concentration, andwood density) collectively accounting for productivity variation. Remarkably, explicit representationof temperature variation with elevation was not required to achieve accurate predictions offorest productivity, as trait variation driven by species turnover appears to capture the effect oftemperature. Our semi-mechanistic model suggests that spatial variation in traits can potentiallybe used to estimate spatial variation in productivity at the landscape scale. Fil: Fyllas, Nikolaos M.. University of Oxford; Reino Unido Fil: Patrick Bentley, Lisa. University of Oxford; Reino Unido Fil: Shenkin, Alexander. University of Oxford; Reino Unido Fil: Asner, Gregory P.. Carnegie Institution for Science. Department of Global Ecology; Reino Unido Fil: Atkin, Owen K.. The Australian National University. ARC Centre of Excellence in Plant Energy Biology. Research School of Biology; Australia Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Enquist, Brian J.. Arizona State University; Estados Unidos Fil: Farfan Rios, William. University Wake Forest; Estados Unidos Fil: Gloor, Emanuel. University of Leeds; Reino Unido Fil: Guerrieri, Rossella. Universitat Autònoma de Barcelona; España. University of Edinburgh; Reino Unido Fil: Huaraca Huasco, Walter. Universidad Nacional de San Antonio Abad del Cusco; Perú Fil: Ishida, Yoko. James Cook University; Australia Fil: Martin, Roberta E.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Meir, Patrick. University of Edinburgh; Reino Unido. The Australian National University. Research School of Biology. Division of Plant Sciences; Australia Fil: Phillips, Oliver. University of Leeds; Reino Unido Fil: Salinas, Norma. University of Oxford; Reino Unido. Pontificia Universidad Católica de Perú; Perú Fil: Silman, Miles. University Wake Forest; Estados Unidos Fil: Weerasinghe, Lasantha K.. The Australian National University. Research School of Biology. Division of Plant Sciences; Australia Fil: Zaragoza Castells, Joana. The Australian National University. Research School of Biology. Division of Plant Sciences; Australia. University of Exeter; Reino Unido Fil: Malhi, Yadvinder. University of Oxford; Reino Unido

The net primary productivity, carbon (C) stocks and turnover rates (i.e. C dynamics) of tropical forests are an important aspect of the global C cycle. These variables have been investigated in lowland tropical forests, but they have rarely been studied in tropical montane forests (TMFs). This study examines spatial patterns of above- and belowground C dynamics along a transect ranging from lowland Amazonia to the high Andes in SE Peru. Fine root biomass values increased from 1.50 Mg C ha−1 at 194 m to 4.95 ± 0.62 Mg C ha−1 at 3020 m, reaching a maximum of 6.83 ± 1.13 Mg C ha−1 at the 2020 m elevation site. Aboveground biomass values decreased from 123.50 Mg C ha−1 at 194 m to 47.03 Mg C ha−1 at 3020 m. Mean annual belowground productivity was highest in the most fertile lowland plots (7.40 ± 1.00 Mg C ha−1 yr−1) and ranged between 3.43 ± 0.73 and 1.48 ± 0.40 Mg C ha−1 yr−1 in the premontane and montane plots. Mean annual aboveground productivity was estimated to vary between 9.50 ± 1.08 Mg C ha−1 yr−1 (210 m) and 2.59 ± 0.40 Mg C ha−1 yr−1 (2020 m), with consistently lower values observed in the cloud immersion zone of the montane forest. Fine root C residence time increased from 0.31 years in lowland Amazonia to 3.78 ± 0.81 years at 3020 m and stem C residence time remained constant along the elevational transect, with a mean of 54 ± 4 years. The ratio of fine root biomass to stem biomass increased significantly with increasing elevation, whereas the allocation of net primary productivity above- and belowground remained approximately constant at all elevations. Although net primary productivity declined in the TMF, the partitioning of productivity between the ecosystem subcomponents remained the same in lowland, premontane and montane forests.

Aboveground primary productivity is widely considered to be limited by phosphorus (P) availability in lowland tropical forests and by nitrogen (N) availability in montane tropical forests. However, the extent to which this paradigm applies to belowground processes remains unresolved. We measured indices of soil microbial nutrient status in lowland, sub-montane and montane tropical forests along a natural gradient spanning 3400 m in elevation in the Peruvian Andes. With increasing elevation there were marked increases in soil concentrations of total N, total P, and readily exchangeable P, but a decrease in N mineralization determined by in situ resin bags. Microbial carbon (C) and N increased with increasing elevation, but microbial C : N : P ratios were relatively constant, suggesting homeostasis. The activity of hydrolytic enzymes, which are rich in N, decreased with increasing elevation, while the ratio of enzymes involved in the acquisition of N and P increased with increasing elevation, further indicating an increase in the relative demand for N compared to P with increasing elevation. We conclude that soil microorganisms shift investment in nutrient acquisition from P to N between lowland and montane tropical forests, suggesting that different nutrients regulate soil microbial metabolism and the soil carbon balance in these ecosystems.

Climate change is affecting the amount and complexity of plant inputs to tropical forest soils. This is likely to influence the carbon (C) balance of these ecosystems by altering decomposition processes e.g., “positive priming effects” that accelerate soil organic matter mineralization. However, the mechanisms determining the magnitude of priming effects are poorly understood. We investigated potential mechanisms by adding 13C labeled substrates, as surrogates of plant inputs, to soils from an elevation gradient of tropical lowland and montane forests. We hypothesized that priming effects would increase with elevation due to increasing microbial nitrogen limitation, and that microbial community composition would strongly influence the magnitude of priming effects. Quantifying the sources of respired C (substrate or soil organic matter) in response to substrate addition revealed no consistent patterns in priming effects with elevation. Instead we found that substrate quality (complexity and nitrogen content) was the dominant factor controlling priming effects. For example a nitrogenous substrate induced a large increase in soil organic matter mineralization whilst a complex C substrate caused negligible change. Differences in the functional capacity of specific microbial groups, rather than microbial community composition per se, were responsible for these substrate-driven differences in priming effects. Our findings suggest that the microbial pathways by which plant inputs and soil organic matter are mineralized are determined primarily by the quality of plant inputs and the functional capacity of microbial taxa, rather than the abiotic properties of the soil. Changes in the complexity and stoichiometry of plant inputs to soil in response to climate change may therefore be important in regulating soil C dynamics in tropical forest soils. Refereed/Peer-reviewed

Background: Tropical montane cloud forests (TMCF) are unique ecosystems with high biodiversity and large carbon reservoirs. To date there have been limited descriptions of the carbon cycle of TMCF. Aims: We present results on the production, allocation and cycling of carbon for two mid-elevation (1500–1750 m) tropical montane cloud forest plots in San Pedro, Kosnipata Valley, Peru. Methods: We repeatedly recorded the components of net primary productivity (NPP) using biometric measurements, and autotrophic (R a) and heterotrophic (R h) respiration, using gas exchange measurements. From these we estimated gross primary productivity (GPP) and carbon use efficiency (CUE) at the plot level. Results: The plot at 1500 m was found very productive, with our results comparable with the most productive lowland Amazonian forests. The plot at 1750 m had significantly lower productivity, possibly because of greater cloud immersion. Both plots had similar patterns of NPP allocation, a substantial seasonality in NPP com...

Background: The forests of western Amazonia are known to be more dynamic that the better-studied forests of eastern Amazonia, but there has been no comprehensive description of the carbon cycle of a western Amazonian forest. Aims: We present the carbon budget of two forest plots in Tambopata in south-eastern Peru, western Amazonia. In particular, we present, for the first time, the seasonal variation in the detailed carbon budget of a tropical forest. Methods: We measured the major components of net primary production (NPP) and total autotrophic respiration over 3–6 years. Results: The NPP for the two plots was 15.1 ± 0.8 and 14.2 ± 1.0 Mg C ha−1 year−1, the gross primary productivity (GPP) was 35.5 ± 3.6 and 34.5 ± 3.5 Mg C ha−1 year−1, and the carbon use efficiency (CUE) was 0.42 ± 0.05 and 0.41 ± 0.05. NPP and CUE showed a large degree of seasonality. Conclusions: The two plots were similar in carbon cycling characteristics despite the different soils, the most notable difference being high allocation ...

The functional role of herbivores in tropical rainforests remains poorly understood. We quantified the magnitude of, and underlying controls on, carbon, nitrogen and phosphorus cycled by invertebrate herbivory along a 2800 m elevational gradient in the tropical Andes spanning 12°C mean annual temperature. We find, firstly, that leaf area loss is greater at warmer sites with lower foliar phosphorus, and secondly, that the estimated herbivore-mediated flux of foliar nitrogen and phosphorus from plants to soil via leaf area loss is similar to, or greater than, other major sources of these nutrients in tropical forests. Finally, we estimate that herbivores consume a significant portion of plant carbon, potentially causing major shifts in the pattern of plant and soil carbon cycling. We conclude that future shifts in herbivore abundance and activity as a result of environmental change could have major impacts on soil fertility and ecosystem carbon sequestration in tropical forests.

Background: In tropical montane cloud forests (TMCFs) fires can be a frequent source of disturbance near the treeline. Aims: To identify how forest structure and tree species composition change in response to fire and to identify fire-tolerant species, and determine which traits or characteristics are associated with fire tolerance. Methods: Shifts in forest structure and diversity were assessed on 15 paired plots in burnt and unburnt (control) forests, along a fire chronosequence. Species were classified as fire-sensitive, fire survivors or fire thrivers, using a fire tolerance index. Regression and tree analyses were used to determine which traits contributed to the fire tolerance of species. Results: There were more small- and medium-sized trees in the burnt plots independent of time since the last fire. There were shifts in family importance value and in species diversity among the plots. Of the 73 species analysed there were 39 fire-sensitive species, 19 fire survivors and 14 fire thrivers. Sprouting ability and biomass showed a positive relationship with the fire tolerance of species. Conclusions: There were immediate as well as long-term (up to 15 years) effects of fire on forest structure and species composition, suggesting that TMCFs take more than 15 years to regenerate from a fire. Long-term studies are needed to fully understand regeneration patterns of TMCFs after fires. Keywords: bark thickness; fire sensitivity; Peruvian Andes; species diversity; sprouting, tree size; wood density

1. The Andes are predicted to warm by 3–5 °C this century with the potential to alter the processes regulating carbon (C) cycling in these tropical forest soils. This rapid warming is expected to stimulate soil microbial respiration and change plant species distributions, thereby affecting the quantity and quality of C inputs to the soil and influencing the quantity of soil-derived CO2 released to the atmosphere. 2. We studied tropical lowland, premontane and montane forest soils taken from along a 3200-m elevation gradient located in south-east Andean Peru. We determined how soil microbial communities and abiotic soil properties differed with elevation. We then examined how these differences in microbial composition and soil abiotic properties affected soil C-cycling processes, by amending soils with C substrates varying in complexity and measuring soil heterotrophic respiration (RH). 3. Our results show that there were consistent patterns of change in soil biotic and abiotic properties with elevation. Microbial biomass and the abundance of fungi relative to bacteria increased significantly with elevation, and these differences in microbial community composition were strongly correlated with greater soil C content and C:N (nitrogen) ratios. We also found that RH increased with added C substrate quality and quantity and was positively related to microbial biomass and fungal abundance. 4. Statistical modelling revealed that RH responses to changing C inputs were best predicted by soil pH and microbial community composition, with the abundance of fungi relative to bacteria, and abundance of gram-positive relative to gram-negative bacteria explaining much of the model variance. 5. Synthesis. Our results show that the relative abundance of microbial functional groups is an important determinant of RH responses to changing C inputs along an extensive tropical elevation gradient in Andean Peru. Although we do not make an experimental test of the effects of climate change on soil, these results challenge the assumption that different soil microbial communities will be ‘functionally equivalent’ as climate change progresses, and they emphasize the need for better ecological metrics of soil microbial communities to help predict C cycle responses to climate change in tropical biomes.

Understanding functional trait-environment relationships (TERs) may improve predictions of community assembly. However, many empirical TERs have been weak or lacking conceptual foundation. TERs based on leaf venation networks may better link individuals and communities via hydraulic constraints. We report measurements of vein density, vein radius, and leaf thickness for more than 100 dominant species occurring in ten forest communities spanning a 3,300 m Andes-Amazon elevation gradient in Peru. We use these data to measure the strength of TERs at community scale and to determine whether observed TERs are similar to those predicted by physiological theory. We found strong support for TERs between all traits and temperature, as well weaker support for a predicted TER between maximum abundance-weighted leaf transpiration rate and maximum potential evapotranspiration. These results provide one approach for developing a more mechanistic trait-based community assembly theory. Fil: Blonder, Benjamin. University of Oxford; Reino Unido Fil: Salinas, Norma. University of Oxford; Reino Unido. Pontificia Universidad Católica de Perú; Perú Fil: Patrick Bentley, Lisa. University of Oxford; Reino Unido Fil: Shenkin, Alexander. University of Oxford; Reino Unido Fil: Chambi Porroa, Percy O.. Universidad Nacional San Antonio Abad del Cusco; Perú Fil: Valdez Tejeira, Yolvi. Universidad Nacional San Antonio Abad del Cusco; Perú Fil: Violle, Cyrille. Université de Montpellier. Centre d’Ecologie Fonctionnelle et Evolutive; Francia Fil: Fyllas, Nikolaos M.. University of Oxford; Reino Unido Fil: Goldsmith, Gregory R.. Paul Scherrer Institute. Ecosystem Fluxes Group. Laboratory for Atmospheric Chemistry; Suiza Fil: Martin, Robert E.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Asner, Gregory P.. Carnegie Institution for Science. Department of Global Ecology; Estados Unidos Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Enquist, Brian J.. Arizona State University; Estados Unidos Fil: Malhi, Yadvinder. University of Oxford; Reino Unido

Soil enzymes are catalysts of organic matter depolymerisation, which is of critical importance for ecosystem carbon (C) cycling. Better understanding of the sensitivity of enzymes to temperature will enable improved predictions of climate change impacts on soil C stocks. These impacts may be especially large in tropical montane forests, which contain large amounts of soil C. We determined the temperature sensitivity (Q 10) of a range of hydrolytic and oxidative enzymes involved in organic matter cycling from soils along a 1900 m elevation gradient (a 10 °C mean annual temperature gradient) of tropical montane forest in the Peruvian Andes. We investigated whether the activity (V max) of selected enzymes: (i) exhibited a Q 10 that varied with elevation and/or soil properties; and (ii) varied among enzymes and according to the complexity of the target substrate for C-degrading enzymes. The Q 10 of V max for β-glucosidase and β-xylanase increased with increasing elevation and declining mean annual temperature. For all other enzymes, including cellobiohydrolase, N-acetyl β-glucosaminidase and phosphomonoesterase, the Q 10 of V max did not vary linearly with elevation. Hydrolytic enzymes that degrade more complex C compounds had a greater Q 10 of V max, but this pattern did not apply to oxidative enzymes because phenol oxidase had the lowest Q 10 value of all enzymes studied here. Our findings suggest that regional differences in the temperature sensitivities of different enzyme classes may influence the terrestrial C cycle under future climate warming.

Background: The slopes of the eastern Andes harbour some of the highest biodiversity on Earth and a high proportion of endemic species. However, there have been only a few and limited descriptions of carbon budgets in tropical montane forest regions. Aims: We present the first comprehensive data on the production, allocation and cycling of carbon for two high elevation (ca. 3000 m) tropical montane cloud forest plots in the Kosñipata Valley, Peruvian Andes. Methods: We measured the main components and seasonal variation of net primary productivity (NPP), autotrophic (Ra) and heterotrophic (Rh) respiration to estimate gross primary productivity (GPP) and carbon use efficiency (CUE) in two 1-ha plots. Results:NPP for the two plots was estimated to be 7.05 ± 0.39 and 8.04 ± 0.47 Mg C ha−1 year−1, GPP to be 22.33 ± 2.23 and 26.82 ± 2.97 Mg C ha−1 year−1 and CUE was 0.32 ± 0.04 and 0.30 ± 0.04. Conclusions: We found strong seasonality in NPP and moderate seasonality of Ra, suggesting that forest NPP is driven by changes in photosynthesis and highlighting the importance of variation in solar radiation. Our findings imply that trees invest more in biomass production in the cooler season with lower solar radiation and more in maintenance during the warmer and high solar radiation period.

Owen K. Atkin [et al.].- Received: 8 July 2014, Accepted: 29 November 2014, Leaf dark respiration (Rdark) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits., Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark., Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8–28°C). By contrast, Rdark at a standard T (25°C, Rdark25) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark25 at a given photosynthetic capacity (Vcmax25) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark25 values at any given Vcmax25 or [N] were higher in herbs than in woody plants., The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs).

Background: The slopes of the eastern Andes harbour some of the highest biodiversity on Earth and a high proportion of endemic species. However, there have been only a few and limited descriptions of carbon budgets in tropical montane forest regions. Aims: We present the first comprehensive data on the production, allocation and cycling of carbon for two high elevation (ca. 3000 m) tropical montane cloud forest plots in the Kosñipata Valley, Peruvian Andes. Methods: We measured the main components and seasonal variation of net primary productivity (NPP), autotrophic (Ra) and heterotrophic (Rh) respiration to estimate gross primary productivity (GPP) and carbon use efficiency (CUE) in two 1-ha plots. Results:NPP for the two plots was estimated to be 7.05 ± 0.39 and 8.04 ± 0.47 Mg C ha−1 year−1, GPP to be 22.33 ± 2.23 and 26.82 ± 2.97 Mg C ha−1 year−1 and CUE was 0.32 ± 0.04 and 0.30 ± 0.04. Conclusions: We found strong seasonality in NPP and moderate seasonality of Ra, suggesting that forest NPP is driven by changes in photosynthesis and highlighting the importance of variation in solar radiation. Our findings imply that trees invest more in biomass production in the cooler season with lower solar radiation and more in maintenance during the warmer and high solar radiation period.