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No claim to original US government works New Phytologist © 2018 New Phytologist Trust Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

This is the final version of the article. Available from Wiley via the DOI in this record. ; Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs. ; This paper is a product of the European Union's Seventh Framework Programme AMAZALERT project (282664). The field data used in this study have been generated by the RAINFOR network, which has been supported by a Gordon and Betty Moore Foundation grant, the European Union's Seventh Framework Programme projects 283080, 'GEOCARBON'; and 282664, 'AMAZALERT'; ERC grant 'Tropical Forests in the Changing Earth System'), and Natural Environment Research Council (NERC) Urgency, Consortium and Standard Grants 'AMAZONICA' (NE/F005806/1), 'TROBIT' (NE/D005590/1) and 'Niche Evolution of South American Trees' (NE/I028122/1). Additional data were included from the Tropical Ecology Assessment and Monitoring (TEAM) Network – a collaboration between Conservation International, the Missouri Botanical Garden, the Smithsonian Institution and the Wildlife Conservation Society, and partly funded by these institutions, the Gordon and Betty Moore Foundation, and other donors. Fieldwork was also partially supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico of Brazil (CNPq), project Programa de Pesquisas Ecológicas de Longa Duração (PELD-403725/2012-7). A.R. acknowledges funding from the Helmholtz Alliance 'Remote Sensing and Earth System Dynamics'; L.P., M.P.C. E.A. and M.T. are partially funded by the EU FP7 project 'ROBIN' (283093), with co-funding for E.A. from the Dutch Ministry of Economic Affairs (KB-14-003-030); B.C. [was supported in part by the US DOE (BER) NGEE-Tropics project (subcontract to LANL). O.L.P. is supported by an ERC Advanced Grant and is a Royal Society-Wolfson Research Merit Award holder. P.M. acknowledges support from ARC grant FT110100457 and NERC grants NE/J011002/1, and T.R.B. acknowledges support from a Leverhulme Trust Research Fellowship.

Understanding the processes that determine aboveground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity (woody NPP) and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size-structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influence AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates, and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP, and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs. This article is protected by copyright. All rights reserved. ; This paper is a product of the European Union's Seventh Frame-work Programme AMAZALERT project (282664). The field dataused in this study have been generated by the RAINFOR net-work, which has been supported by a Gordon and Betty MooreFoundation grant, the European Union's Seventh FrameworkProgramme projects 283080, 'GEOCARBON'; and 282664,'AMAZALERT'; ERC grant 'Tropical Forests in the ChangingEarth System'), and Natural Environment Research Council(NERC) Urgency, Consortium and Standard Grants 'AMAZO-NICA' (NE/F005806/1), 'TROBIT' (NE/D005590/1) and 'NicheEvolution of South American Trees' (NE/I028122/1). Additionaldata were included from the Tropical Ecology Assessment andMonitoring (TEAM) Network – a collaboration between Conser-vation International, the Missouri Botanical Garden, the Smith-sonian Institution and the Wildlife Conservation Society, andpartly funded by these institutions, the Gordon and Betty MooreFoundation, and other donors. Fieldwork was also partially sup-ported by Conselho Nacional de Desenvolvimento Cientı´fico eTecnolo´gico of Brazil (CNPq), project Programa de PesquisasEcolo´gicas de Longa Duracßa˜o (PELD-403725/2012-7). A.R.acknowledges funding from the Helmholtz Alliance 'RemoteSensing and Earth System Dynamics'; L.P., M.P.C. E.A. andM.T. are partially funded by the EU FP7 project 'ROBIN'(283093), with co-funding for E.A. from the Dutch Ministry ofEconomic Affairs (KB-14-003-030); B.C. [was supported in partby the US DOE (BER) NGEE-Tropics project (subcontract toLANL). O.L.P. is supported by an ERC Advanced Grant and is aRoyal Society-Wolfson Research Merit Award holder. P.M.acknowledges support from ARC grant FT110100457 and NERCgrants NE/J011002/1, and T.R.B. acknowledges support from aLeverhulme Trust Research Fellowship.