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Aim Palms are an iconic, diverse and often abundant component of tropical ecosystems that provide many ecosystem services. Being monocots, tree palms are evolutionarily, morphologically and physiologically distinct from other trees, and these differences have important consequences for ecosystem services (e.g., carbon sequestration and storage) and in terms of responses to climate change. We quantified global patterns of tree palm relative abundance to help improve understanding of tropical forests and reduce uncertainty about these ecosystems under climate change. Location Tropical and subtropical moist forests. Time period Current. Major taxa studied Palms (Arecaceae). Methods We assembled a pantropical dataset of 2,548 forest plots (covering 1,191ha) and quantified tree palm (i.e., ≥10cm diameter at breast height) abundance relative to co‐occurring non‐palm trees. We compared the relative abundance of tree palms across biogeographical realms and tested for associations with palaeoclimate stability, current climate, edaphic conditions and metrics of forest structure. Results On average, the relative abundance of tree palms was more than five times larger between Neotropical locations and other biogeographical realms. Tree palms were absent in most locations outside the Neotropics but present in >80% of Neotropical locations. The relative abundance of tree palms was more strongly associated with local conditions (e.g., higher mean annual precipitation, lower soil fertility, shallower water table and lower plot mean wood density) than metrics of long‐term climate stability. Life‐form diversity also influenced the patterns; palm assemblages outside the Neotropics comprise many non‐tree (e.g., climbing) palms. Finally, we show that tree palms can influence estimates of above‐ground biomass, but the magnitude and direction of the effect require additional work. Conclusions Tree palms are not only quintessentially tropical, but they are also overwhelmingly Neotropical. Future work to understand the contributions of tree palms to biomass estimates and carbon cycling will be particularly crucial in Neotropical forests.

Why do some forests produce biomass more efficiently than others? Variations in Carbon Use Efficiency (CUE: total Net Primary Production (NPP)/ Gross Primary Production (GPP)) may be due to changes in wood residence time (Biomass/NPPwood), temperature, or soil nutrient status. We tested these hypotheses in 14, one ha plots across Amazonian and Andean forests where we measured most key components of net primary production (NPP: wood, fine roots, and leaves) and autotrophic respiration (Ra; wood, rhizosphere, and leaf respiration). We found that lower fertility sites were less efficient at producing biomass and had higher rhizosphere respiration, indicating increased carbon allocation to belowground components. We then compared wood respiration to wood growth and rhizosphere respiration to fine root growth and found that forests with residence times 40 yrs. A comparison of rhizosphere respiration to fine root growth showed that rhizosphere growth respiration was significantly greater at low fertility sites. Overall, we found that Amazonian forests produce biomass less efficiently in stands with residence times >40 yrs and in stands with lower fertility, but changes to long-term mean annual temperatures do not impact CUE.

Made available in DSpace on 2019-10-04T12:31:04Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-11-19 UK Natural Environment Research Council European Research Council Advanced Investigator Award (GEM-TRAIT) Nature Conservancy-Oxford Martin School Climate Partnership NERC Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Gordon and Betty Moore Foundation Sime Darby Foundation Project USA-NAS/PEER Project ReFlor FAPEMAT Empresa Brasileira de Pesquisa Agropecuaria - Embrapa European Research Council (H2020-MSCA-RISE-2015) UK government Darwin Initiative Nature Conservancy UK Natural Environment Research Council (NERC) Jackson Foundation Meteorological extreme events such as El Nino events are expected to affect tropical forest net primary production (NPP) and woody growth, but there has been no large-scale empirical validation of this expectation. We collected a large high-temporal resolution dataset (for 1-13 years depending upon location) of more than 172 000 stem growth measurements using dendrometer bands from across 14 regions spanning Amazonia, Africa and Borneo in order to test how much month-to-month variation in stand-level woody growth of adult tree stems (NPPstem) can be explained by seasonal variation and interannual meteorological anomalies. A key finding is that woody growth responds differently to meteorological variation between tropical forests with a dry season (where monthly rainfall is less than 100 mm), and aseasonal wet forests lacking a consistent dry season. In seasonal tropical forests, a high degree of variation in woody growth can be predicted from seasonal variation in temperature, vapour pressure deficit, in addition to anomalies of soil water deficit and shortwave radiation. The variation of aseasonal wet forest woody growth is best predicted by the anomalies of vapour pressure deficit, water deficit and shortwave radiation. In total, we predict the total live woody production of the global tropical forest biome to be 2.16 Pg C yr(-1), with an interannual range 1.96-2.26 Pg C yr(-1) between 1996-2016, and with the sharpest declines during the strong El Nino events of 1997/8 and 2015/6. There is high geographical variation in hotspots of El Nino-associated impacts, with weak impacts in Africa, and strongly negative impacts in parts of Southeast Asia and extensive regions across central and eastern Amazonia. Overall, there is high correlation (r = -0.75) between the annual anomaly of tropical forest woody growth and the annual mean of the El Nino 3.4 index, driven mainly by strong correlations with anomalies of soil water deficit, vapour pressure deficit and shortwave radiation. This article is part of the discussion meeting issue 'The impact of the 2015/2016 El Nino on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'. Univ Oxford, Sch Geog & Environm, Environm Change Inst, South Parks Rd, Oxford OX1 3QY, England IIAP, Iquitos, Peru No Arizona Univ, Sch Informat Comp & Cyber Syst, Flagstaff, AZ 86011 USA Univ Stirling, Fac Nat Sci, Stirling FK9 4LA, Scotland CENAREST, Inst Rech Ecol Trop, BP 842, Libreville, Gabon ANPN, BP 20379, Libreville, Gabon Univ Exeter, Coll Life & Environm Sci, Geog, Amory Bldg, Exeter EX4 4RJ, Devon, England Univ Autonoma Gabriel Rene Moreno, Museo Hist Nat Noel Kempff Mercado, Ave Irala 565,Casilla Postal 2489, Santa Cruz, CA, Bolivia Forestry Res Inst Ghana, Kumasi, Ghana Woods Hole Res Ctr, Falmouth, MA USA Univ Nacl San Antonio Abad Cusco, Cuzco, Peru Univ Estado Mato Grosso, Programa Posgrad Ecol & Convervacao, BR-78690000 Nova Xavantina, MT, Brazil Ctr Euromediterraneo Cambiamente Climat, Lecce, Italy Minist Foret & Environm, BP199, Libreville, Gabon Amazon Environm Res Inst IPAM, Canarana, MG, Brazil Univ Estadual Paulista, Dept Ecol, BR-13506900 Rio Claro, SP, Brazil Embrapa Amazonia Oriental, Trav Dr Eneas Pinheiro S-N,CP 48, BR-66095100 Belem, PA, Brazil Univ La Serena, Dept Biol, La Serena, Chile Univ Lancaster, Lancaster Environm Ctr, Lancaster LA1 4YQ, England Univ Fed Para, Inst Geosciencias, Belem, Para, Brazil Australian Natl Univ, Res Sch Biol, Canberra, ACT 2601, Australia Univ Edinburgh, Sch Geosci, Edinburgh EH93FF, Midlothian, Scotland Lund Univ, Phys Geog & Ecosyst Sci, Lund, Sweden Univ Leeds, Sch Geog, Leeds, W Yorkshire, England Univ Estadual Paulista, Dept Ecol, BR-13506900 Rio Claro, SP, Brazil UK Natural Environment Research Council: NE/P001092/1 European Research Council Advanced Investigator Award (GEM-TRAIT): 321131 NERC: NE/I014705/1 NERC: NE/K016369/1 NERC: NE/F005776/1 NERC: NE/K016385/1 NERC: NE/J011002/1 CNPq: 457914/2013-0/MCTI/ CNPq/FNDCT/LBA/ESECAFLOR CNPq: 403725/ 2012-7 CNPq: 441244/2016-5 CNPq: 457602/2012-0 Project USA-NAS/PEER: PGA-2000005316 Project ReFlor FAPEMAT: 0589267/2016 CNPq: 574008/2008-0 Empresa Brasileira de Pesquisa Agropecuaria - Embrapa: SEG: 02.08.06.005.00 European Research Council (H2020-MSCA-RISE-2015): 691053-ODYSSEA UK government Darwin Initiative: 17-023 UK Natural Environment Research Council (NERC): NE/F01614X/1 UK Natural Environment Research Council (NERC): NE/G000816/1 UK Natural Environment Research Council (NERC): NE/K016431/1 UK Natural Environment Research Council (NERC): NE/P004512/1 : PQ-2

The Amazon Basin has experienced more variable climate over the last decade, with a severe and widespread drought in 2005 causing large basin-wide losses of biomass. A drought of similar climatological magnitude occurred again in 2010; however, there has been no basin-wide ground-based evaluation of effects on vegetation. We examine to what extent the 2010 drought affected forest dynamics using ground-based observations of mortality and growth from an extensive forest plot network. We find that during the 2010 drought interval, forests did not gain biomass (net change: −0.43 Mg ha−1, confidence interval (CI): −1.11, 0.19, n = 97), regardless of whether forests experienced precipitation deficit anomalies. This contrasted with a long-term biomass sink during the baseline pre-2010 drought period (1998 to pre-2010) of 1.33 Mg ha−1 yr−1 (CI: 0.90, 1.74, p

Changes to the carbon cycle in tropical forests could affect global climate, but predicting such changes has been previously limited by lack of field-based data. Here we show seasonal cycles of the complete carbon cycle for 14, 1ha intensive carbon cycling plots which we separate into three regions: humid lowland, highlands, and dry lowlands. Our data highlight three trends: (1) there is differing seasonality of total net primary productivity (NPP) with the highlands and dry lowlands peaking in the dry season and the humid lowland sites peaking in the wet season, (2) seasonal reductions in wood NPP are not driven by reductions in total NPP but by carbon during the dry season being preferentially allocated toward either roots or canopy NPP, and (3) there is a temporal decoupling between total photosynthesis and total carbon usage (plant carbon expenditure). This decoupling indicates the presence of nonstructural carbohydrates which may allow growth and carbon to be allocated when it is most ecologically beneficial rather than when it is most environmentally available.

Understanding the relationship between photosynthesis, net primary productivity and growth in forest ecosystems is key to understanding how these ecosystems will respond to global anthropogenic change, yet the linkages among these components are rarely explored in detail. We provide the first comprehensive description of the productivity, respiration and carbon allocation of contrasting lowland Amazonian forests spanning gradients in seasonal water deficit and soil fertility. Using the largest data set assembled to date, ten sites in three countries all studied with a standardized methodology, we find that (i) gross primary productivity (GPP) has a simple relationship with seasonal water deficit, but that (ii) site-to-site variations in GPP have little power in explaining site-to-site spatial variations in net primary productivity (NPP) or growth because of concomitant changes in carbon use efficiency (CUE), and conversely, the woody growth rate of a tropical forest is a very poor proxy for its productivity. Moreover, (iii) spatial patterns of biomass are much more driven by patterns of residence times (i.e. tree mortality rates) than by spatial variation in productivity or tree growth. Current theory and models of tropical forest carbon cycling under projected scenarios of global atmospheric change can benefit from advancing beyond a focus on GPP. By improving our understanding of poorly understood processes such as CUE, NPP allocation and biomass turnover times, we can provide more complete and mechanistic approaches to linking climate and tropical forest carbon cycling.

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.

Background: The forests of north-west Amazonia are characterised by the highest levels of tree diversity in the world, high rainfall and relatively fertile soils. Aims: Here we present a comprehensive description of the carbon cycle of two 1 ha forests plots in Allpahuayo, near Iquitos, Peru, one on an occasionally inundated alisol/gleysol landscape, the other on an arenosol (sandy soil). Methods: Data on the components of net primary productivity (NPP) and autotrophic respiration were collected over the period 2009–2011, and summed to estimate gross primary productivity (GPP) and carbon use efficiency (CUE). Results: Overall, these forests showed high values of GPP (39.05 ± 4.59 and 41.88 ± 4.60 Mg C ha−1 year−1). Despite the lack of a dry season, the forests showed distinct seasonality in tree growth, litterfall, flowering and fine root productivity. This showed that tropical forests with little seasonality in water supply can still exhibit distinct seasonality in NPP and carbon use, apparently synchron...

Aim Large trees (d.b.h. 70 cm) store large amounts of biomass. Several studies suggest that large trees may be vulnerable to changing climate, poten- tially leading to declining forest biomass storage. Here we determine the importance of large trees for tropical forest biomass storage and explore which intrinsic (species trait) and extrinsic (environment) variables are associated with the density of large trees and forest biomass at continental and pan-tropical scales. Location Pan-tropical.

The key role of tropical forest belowground carbon stocks and fluxes is well recognised as one of the main components of the terrestrial ecosystem carbon cycle. This study presents the first detailed investigation of spatial and temporal patterns of fine root stocks and fluxes in tropical forests along an elevational gradient, ranging from the Peruvian Andes (3020 m) to lowland Amazonia (194 m), with mean annual temperatures of 11.8 degrees C to 26.4 degrees C and annual rainfall values of 1900 to 1560 mm yr(-1), respectively. Specifically, we analyse abiotic parameters controlling fine root dynamics, fine root growth characteristics, and seasonality of net primary productivity along the elevation gradient. Root and soil carbon stocks were measured by means of soil cores, and fine root productivity was recorded using rhizotron chambers and ingrowth cores. We find that mean annual fine root below ground net primary productivity in the montane forests (0-30 cm depth) ranged between 4.27 +/- 0.56 Mg C ha(-1) yr(-1) (1855 m) and 1.72 +/- 0.87 Mg C ha(-1) yr(-1) (3020 m). These values include a correction for finest roots (< 0.6 mm diameter), which we suspect are under sampled, resulting in an underestimation of fine roots by up to 31% in current ingrowth core counting methods. We investigate the spatial and seasonal variation of fine root dynamics using soil depth profiles and an analysis of seasonal amplitude along the elevation gradient. We report a stronger seasonality of NPPFineRoot within the cloud immersion zone, most likely synchronised to seasonality of solar radiation. Finally, we provide the first insights into root growth characteristics along a tropical elevation transect: fine root area and fine root length increase significantly in the montane cloud forest. These insights into belowground carbon dynamics of tropical lowland and montane forests have significant implications for our understanding of the global tropical forest carbon cycle. (Less)

The seasonality of solar irradiance and precipitation may regulate seasonal variations in tropical forests carbon cycling. Controversy remains over their importance as drivers of seasonal dynamics of net primary productivity in tropical forests. We use ground data from nine lowland Amazonian forest plots collected over 3 years to quantify the monthly primary productivity (NPP) of leaves, reproductive material, woody material, and fine roots over an annual cycle. We distinguish between forests that do not experience substantial seasonal moisture stress (“humid sites”) and forests that experience a stronger dry season (“dry sites”). We find that forests from both precipitation regimes maximize leaf NPP over the drier season, with a peak in production in August at both humid (mean 0.39 ± 0.03 Mg C ha−1 month−1 in July, n = 4) and dry sites (mean 0.49 ± 0.03 Mg C ha−1 month−1 in September, n = 8). We identify two distinct seasonal carbon allocation patterns (the allocation of NPP to a specific organ such as wood leaves or fine roots divided by total NPP). The forests monitored in the present study show evidence of either (i) constant allocation to roots and a seasonal trade-off between leaf and woody material or (ii) constant allocation to wood and a seasonal trade-off between roots and leaves. Finally, we find strong evidence of synchronized flowering at the end of the dry season in both precipitation regimes. Flower production reaches a maximum of 0.047 ± 0.013 and 0.031 ± 0.004 Mg C ha−1 month−1 in November, in humid and dry sites, respectively. Fruitfall production was staggered throughout the year, probably reflecting the high variation in varying times to development and loss of fruit among species. (Less)

Amazon forests, which store ∼ 50% of tropical forest carbon and play a vital role in global water, energy, and carbon cycling, are predicted to experience both longer and more intense dry seasons by the end of the 21st century. However, the climate sensitivity of this ecosystem remains uncertain: several studies have predicted large-scale die-back of the Amazon, whereas several more recent studies predict that the biome will remain largely intact. Combining remote-sensing and ground-based observations with a size- and age-structured terrestrial ecosystem model, we explore the sensitivity and ecological resilience of these forests to changes in climate. We demonstrate that water stress operating at the scale of individual plants, combined with spatial variation in soil texture, explains observed patterns of variation in ecosystem biomass, composition, and dynamics across the region, and strongly influences the ecosystem's resilience to changes in dry season length. Specifically, our analysis suggests that in contrast to existing predictions of either stability or catastrophic biomass loss, the Amazon forest's response to a drying regional climate is likely to be an immediate, graded, heterogeneous transition from high-biomass moist forests to transitional dry forests and woody savannah-like states. Fire, logging, and other anthropogenic disturbances may, however, exacerbate these climate change-induced ecosystem transitions.

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.

Atmospheric carbon dioxide records indicate that the land surface has acted as a strong global carbon sink over recent decades1, 2, with a substantial fraction of this sink probably located in the tropics3, particularly in the Amazon4. Nevertheless, it is unclear how the terrestrial carbon sink will evolve as climate and atmospheric composition continue to change. Here we analyse the historical evolution of the biomass dynamics of the Amazon rainforest over three decades using a distributed network of 321 plots. While this analysis confirms that Amazon forests have acted as a long-term net biomass sink, we find a long-term decreasing trend of carbon accumulation. Rates of net increase in above-ground biomass declined by one-third during the past decade compared to the 1990s. This is a consequence of growth rate increases levelling off recently, while biomass mortality persistently increased throughout, leading to a shortening of carbon residence times. Potential drivers for the mortality increase include greater climate variability, and feedbacks of faster growth on mortality, resulting in shortened tree longevity5. The observed decline of the Amazon sink diverges markedly from the recent increase in terrestrial carbon uptake at the global scale1, 2, and is contrary to expectations based on models.

What determines the seasonal and interannual variation of growth rates in trees in a tropical forest? We explore this question with a novel four-year high-temporal-resolution data set of carbon allocation from two forest plots in the Bolivian Amazon. The forests show strong seasonal variation in tree wood growth rates, which are largely explained by shifts in carbon allocation, and not by shifts in total productivity. At the deeper soil plot, there was a clear seasonal trade-off between wood and canopy NPP, while the shallower soils plot showed a contrasting seasonal trade-off between wood and fine roots. Although a strong 2010 drought reduced photosynthesis, NPP remained constant and increased in the six-month period following the drought, which indicates usage of significant nonstructural carbohydrate stores. Following the drought, carbon allocation increased initially towards the canopy, and then in the following year, allocation increased towards fine-root production. Had we only measured woody growth at these sites and inferred total NPP, we would have misinterpreted both the seasonal and interannual responses. In many tropical forest ecosystems, we propose that changing tree growth rates are more likely to reflect shifts in allocation rather than changes in overall productivity. Only a whole NPP allocation perspective can correctly interpret the relationship between changes in growth and changes in productivity.

Background: Dead biomass, including woody debris (WD), is an important component of the carbon cycle in tropical forests. Aims: This study analyses WD (>2 cm) and other above-ground fluxes in mature tropical forest plots along an elevational gradient (210–3025 m above sea level) in southern Peru. Methods: This work was based on inventories of fine and coarse WD (FWD and CWD, respectively), above-ground biomass, and field-based and experimental respiration measurements. Results: Total WD stocks ranged from 6.26 Mg C ha−1 at 3025 m to 11.48 Mg C ha−1 at 2720 m. WD respiration was significantly correlated with moisture content (P < 0.001; R 2 = 0.25), temperature (P < 0.001; R 2 = 0.12) and wood density (P < 0.001; R 2 = 0.16). Controlled experiments showed that both water content and temperature increased respiration rates of individual WD samples. The full breadth of the temperature sensitivity coefficient, or Q 10, estimates, ranging from 1.14–2.13, was low compared to other studies. In addition, temperat...

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 ...

Severe drought in a tropical forest ecosystem suppresses photosynthetic carbon uptake and plant maintenance respiration, but growth is maintained, suggesting that, overall, less carbon is available for tree tissue maintenance and defence, which may cause the subsequent observed increase in tree mortality. The underlying mechanisms that determine the response of tropical forest ecosystems to drought remain poorly understood. Based on observations from a network of intensively measured forest plots in the Amazon basin, this study shows that severe drought suppresses photosynthetic carbon uptake and plant maintenance respiration. Plant growth is maintained however, suggesting that less carbon is available for tree tissue maintenance and defence — which may explain the observed increase in tree mortality that follows a drought. In 2005 and 2010 the Amazon basin experienced two strong droughts1, driven by shifts in the tropical hydrological regime2 possibly associated with global climate change3, as predicted by some global models3. Tree mortality increased after the 2005 drought4, and regional atmospheric inversion modelling showed basin-wide decreases in CO2 uptake in 2010 compared with 2011 (ref. 5). But the response of tropical forest carbon cycling to these droughts is not fully understood and there has been no detailed multi-site investigation in situ. Here we use several years of data from a network of thirteen 1-ha forest plots spread throughout South America, where each component of net primary production (NPP), autotrophic respiration and heterotrophic respiration is measured separately, to develop a better mechanistic understanding of the impact of the 2010 drought on the Amazon forest. We find that total NPP remained constant throughout the drought. However, towards the end of the drought, autotrophic respiration, especially in roots and stems, declined significantly compared with measurements in 2009 made in the absence of drought, with extended decreases in autotrophic respiration in the three driest plots. In the year after the drought, total NPP remained constant but the allocation of carbon shifted towards canopy NPP and away from fine-root NPP. Both leaf-level and plot-level measurements indicate that severe drought suppresses photosynthesis. Scaling these measurements to the entire Amazon basin with rainfall data, we estimate that drought suppressed Amazon-wide photosynthesis in 2010 by 0.38 petagrams of carbon (0.23–0.53 petagrams of carbon). Overall, we find that during this drought, instead of reducing total NPP, trees prioritized growth by reducing autotrophic respiration that was unrelated to growth. This suggests that trees decrease investment in tissue maintenance and defence, in line with eco-evolutionary theories that trees are competitively disadvantaged in the absence of growth6. We propose that weakened maintenance and defence investment may, in turn, cause the increase in post-drought tree mortality observed at our plots.

Understanding climatic controls on tropical forest productivity is key to developing more reliable models for predicting how tropical biomes may respond to climate change. Currently there is no consensus on which factors control seasonal changes in tropical forest tree growth. This study reports the first comprehensive plot-level description of the seasonality of growth in a Peruvian tropical forest. We test whether seasonal and interannual variations in climate are correlated with changes in biomass increment, and whether such relationships differ among trees with different functional traits. We found that biomass increments, measured every 3 months on the two plots, were reduced by between 40 and 55 % in the peak dry season (July–September) relative to peak wet season (January–March). The seasonal patterns of biomass accumulation are significantly (p

For decades, the productivity of tropical montane cloud forests (TMCF) has been assumed to be lower than in tropical lowland forests due to nutrient limitation, lower temperatures, and frequent cloud immersion, although actual estimates of gross primary productivity (GPP) are very scarce. Here, we present the results of a process-based modeling estimate of GPP, using a soil–plant–atmosphere model, of a high elevation Peruvian TMCF. The model was parameterized with field-measured physiological and structural vegetation variables, and driven with meteorological data from the site. Modeled transpiration corroborated well with measured sap flow, and simulated GPP added up to 16.2 ± SE 1.6 Mg C ha−1 y−1. Dry season GPP was significantly lower than wet season GPP, although this difference was 17% and not caused by drought stress. The strongest environmental controls on simulated GPP were variation of photosynthetic active radiation and air temperature (T air). Their relative importance likely varies with elevation and the local prevalence of cloud cover. Photosynthetic parameters (V cmax and J max) and leaf area index were the most important non-environmental controls on GPP. We additionally compared the modeled results with a recent estimate of GPP of the same Peruvian TMCF derived by the summing of ecosystem respiration and net productivity terms, which added up to 26 Mg C ha−1 y−1. Despite the uncertainties in modeling GPP we conclude that at this altitude GPP is, conservatively estimated, 30–40% lower than in lowland rainforest and this difference is driven mostly by cooler temperatures than changes in other parameters.

Background: The dry transitional forests of the southern Amazonia have received little attention from a carbon cycling and ecosystem function perspective, yet they represent ecosystems that may be impacted by global climate change in the future. Aims: To compare the full carbon cycle for two 1-ha forest plots that straddle the ecotone between humid forest and dry forest in Amazonia, ca. 100 km from Santa Cruz, Bolivia. Methods: 2.5 years of measurements of the components of net primary production (NPP) and autotrophic respiration were collected. Results: Total NPP was 15.5 ± 0.89 Mg C ha−1 year−1 at the humid site and 11.27 ± 0.68 Mg C ha−1 year−1 at the dry site; a total Gross Primary Production (GPP) of 34.14 ± 2.92 Mg C ha−1 year−1 and 26.88 ± 2.70 Mg C ha−1 year−1 at the two sites. Carbon use efficiency for both sites was higher than reported for other Amazonian forests (0.45 ± 0.05 and 0.42 ± 0.05). Conclusions: Drier soil conditions selected for the dry deciduous tree species which had higher leaf photosynthesis and total GPP. NPP allocation patterns were similar at the two sites, suggesting that in terms of carbon allocation, the dry forests of the southern Amazonia behave as a scaled-down version of wetter humid forests.

We report results from a large-scale nutrient fertilization experiment along a "megadiverse" (154 unique species were included in the study) 3,000-m elevation transect in the Peruvian Andes and adjacent lowland Amazonia. Our objectives were to test if nitrogen (N) and phosphorus (P) limitation shift along this elevation gradient, and to determine how an alleviation of nutrient limitation would manifest in ecosystem changes. Tree height decreased with increasing elevation, but leaf area index (LAI) and diameter at breast height (DBH) did not vary with elevation. Leaf N:P decreased with increasing elevation (from 24 at 200 m to 11 at 3,000 m), suggesting increased N limitation and decreased P limitation with increasing elevation. After 4 years of fertilization (N, P, N + P), plots at the lowland site (200 m) fertilized with N + P showed greater relative growth rates in DBH than did the control plots; no significant differences were evident at the 1,000 m site, and plots fertilized with N at the highest elevation sites (1,500, 3,000 m) showed greater relative growth rates in DBH than did the control plots, again suggesting increased N constraint with elevation. Across elevations in general N fertilization led to an increase in microbial respiration, while P and N + P addition led to an increase in root respiration and corresponding decrease in hyphal respiration. There was no significant canopy response (LAI, leaf nutrients) to fertilization, suggesting that photosynthetic capacity was not N or P limited in these ecosystems. In sum, our study significantly advances ecological understanding of nutrient cycling and ecosystem response in a region where our collective knowledge and data are sparse: we demonstrate N limitation in high elevation tropical montane forests, N and P co-limitation in lowland Amazonia, and a nutrient limitation response manifested not in canopy changes, but rather in stem and belowground changes. © 2012 Springer-Verlag Berlin Heidelberg.

The production of aboveground soft tissue represents an important share of total net primary production in tropical rain forests. Here we draw from a large number of published and unpublished datasets (n=81 sites) to assess the determinants of litterfall variation across South American tropical forests. We show that across old-growth tropical rainforests, litterfall averages 8.61±1.91 Mg ha−1 yr−1 (mean ± standard deviation, in dry mass units). Secondary forests have a lower annual litterfall than old-growth tropical forests with a mean of 8.01±3.41 Mg ha−1 yr−1. Annual litterfall shows no significant variation with total annual rainfall, either globally or within forest types. It does not vary consistently with soil type, except in the poorest soils (white sand soils), where litterfall is significantly lower than in other soil types (5.42±1.91 Mg ha−1 yr−1). We also study the determinants of litterfall seasonality, and find that it does not depend on annual rainfall or on soil type. However, litterfall seasonality is significantly positively correlated with rainfall seasonality. Finally, we assess how much carbon is stored in reproductive organs relative to photosynthetic organs. Mean leaf fall is 5.74±1.83 Mg ha−1 yr−1 (71% of total litterfall). Mean allocation into reproductive organs is 0.69±0.40 Mg ha−1 yr−1 (9% of total litterfall). The investment into reproductive organs divided by leaf litterfall increases with soil fertility, suggesting that on poor soils, the allocation to photosynthetic organs is prioritized over that to reproduction. Finally, we discuss the ecological and biogeochemical implications of these results.

The net primary productivity (NPP) of tropical forests is one of the most important and least quantified components of the global carbon cycle. Most relevant studies have focused particularly on the quantification of the above-ground coarse wood productivity, and little is known about the carbon fluxes involved in other elements of the NPP, the partitioning of total NPP between its above- and below-ground components and the main environmental drivers of these patterns. In this study we quantify the above- and below-ground NPP of ten Amazonian forests to address two questions: (1) How do Amazonian forests allocate productivity among its above- and below-ground components? (2) How do soil and leaf nutrient status and soil texture affect the productivity of Amazonian forests? Using a standardized methodology to measure the major elements of productivity, we show that NPP varies between 9.3±1.3 Mg C ha−1 yr−1 (mean±standard error), at a white sand plot, and 17.0±1.4 Mg C ha−1 yr−1 at a very fertile Terra Preta site, with an overall average of 12.8±0.9 Mg C ha−1 yr−1. The studied forests allocate on average 64±3% and 36±3% of the total NPP to the above- and below-ground components, respectively. The ratio of above-ground and below-ground NPP is almost invariant with total NPP. Litterfall and fine root production both increase with total NPP, while stem production shows no overall trend. Total NPP tends to increase with soil phosphorus and leaf nitrogen status. However, allocation of NPP to below-ground shows no relationship to soil fertility, but appears to decrease with the increase of soil clay content.

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- and Weibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (≤40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8 Mg ha−1 (range 6.6 to 112.4) to 8.0 Mg ha−1 (−2.5 to 23.0). For all plots, aboveground live biomass was −52.2 Mg ha−1 (−82.0 to −20.3 bootstrapped 95% CI), or 13%, lower when including H estimates, with the greatest relative reductions in estimated biomass in forests of the Brazilian Shield, east Africa, and Australia, and relatively little change in the Guiana Shield, central Africa and southeast Asia. Appreciably different stand structure was observed among regions across the tropical continents, with some storing significantly more biomass in small diameter stems, which affects selection of the best height models to reduce uncertainty and biomass reductions due to H. After accounting for variation in H, total biomass per hectare is greatest in Australia, the Guiana Shield, Asia, central and east Africa, and lowest in east-central Amazonia, W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if tropical forests span 1668 million km2 and store 285 Pg C (estimate including H), then applying our regional relationships implies that carbon storage is overestimated by 35 Pg C (31–39 bootstrapped 95% CI) if H is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree H is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of tropical carbon stocks and emissions due to deforestation.