Global Ecology and Biogeography

Global patterns and climate drivers of terrestrial ecosystem

water-use efficiency deduced from satellite products and
carbon cycle models

Journal: Global Ecology and Biogeography

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Manuscript ID: GEB-2014-0450

Manuscript Type: Research Papers

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Date Submitted by the Author: 14-Nov-2014

Complete List of Authors: Sun, Yan; Peking University, Sino-French Institute for Earth System
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Science, College of Urban and Environmental Sciences

Piao, Shilong; Peking University, Sino-French Institute for Earth System
Science, College of Urban and Environmental Sciences; Chinese Academy
of Sciences, Institute of Tibetan Plateau Research
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Huang, Mengtian; Peking University, Sino-French Institute for Earth

System Science, College of Urban and Environmental Sciences
Ciais, Philippe; CEA CNRS UVSQ, Laboratoire des Sciences du Climat et de
l’Environnement
Zeng, Zhenzhong; Peking University, Sino-French Institute for Earth
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System Science, College of Urban and Environmental Sciences

Cheng, Lei; CSIRO, Land and Water Flagship
Li, Xiran; Peking University, Sino-French Institute for Earth System
Science, College of Urban and Environmental Sciences
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Zhang, Xinping; Peking University, Sino-French Institute for Earth System

Science, College of Urban and Environmental Sciences
Mao, Jiafu ; Oak Ridge National Laboratory, Climate Change Science
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Institute and Environmental Sciences Division

Poulter, Ben; Montana State University, Institute on Ecosystems and the
Department of Ecology
Shi, Xiaoying; Oak Ridge National Laboratory, Climate Change Science
Institute and Environmental Sciences Division
Wang, Ying-Ping; CSIRO, Ocean and Atmosphere Flagship

Water-use Efficiency, inherent water-use efficiency, transpiration-based

Keywords:
water-use efficiency, remote sensing, process-based model, climate drivers
Page 1 of 36 Global Ecology and Biogeography

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4 1 Global patterns and climate drivers of terrestrial ecosystem water-use efficiency
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2 deduced from satellite products and carbon cycle models
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10 3 Yan Sun1, Shilong Piao1,2*, Mengtian Huang1, Philippe Ciais3, Zhenzhong Zeng1, Lei Cheng4,
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12 4 Xiran Li1, Xinping Zhang1, Jiafu Mao5, Ben Poulter6, Xiaoying Shi5, Ying-Ping Wang7
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16 5 Sino-French Institute for Earth System Science, College of Urban and Environmental
17 6 Sciences, Peking University, Beijing 100871, China
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7 Institute of Tibetan Plateau Research, Chinese Academy of Sciences, Beijing 100085,
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22 8 China.
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9 LSCE, UMR CEA-CNRS, Bat. 709, CE, L'Orme des Merisiers, F-91191 Gif-sur-Yvette,
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10 France
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30 11 CSIRO Land and Water Flagship, GPO Box 1666, Canberra ACT 2601, 8 Australia
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12 Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of
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34 13 Sciences, Beijing 100101, China
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37 14 Climate Change Science Institute and Environmental Sciences Division, Oak Ridge National
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42 16 Montana State University, Institute on Ecosystems and the Department of Ecology,
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CSIRO Ocean and Atmosphere Flagship, PMB #1, Aspendale, Victoria 3195, Australia
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49 19 Running title: Global patterns of Ecosystem WUE
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52 20 Keywords: Water-use Efficiency, inherent water-use efficiency, transpiration-based
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54 21 water-use efficiency, remote sensing, process-based model, climate drivers
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57 22 * Author for correspondence: Shilong Piao (slpiao@pku.edu.cn)
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 Global Ecology and Biogeography Page 2 of 36

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3 23 Abstract
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6 24 Aim Ecosystem water-use efficiency (WUE), defined as the ratio of annual gross primary
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8 25 productivity (GPP) to annual evapotranspiration (ET), quantifies the water costs of ecosystem
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10 26 carbon acquisition through photosynthesis. Here, we investigated how ecosystem WUE varies
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27 spatially under different climate conditions, and how those variations of WUE differ from
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15 28 transpiration-based water-use efficiency (WUEt) and transpiration-based inherent water-use
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17 29 efficiency (IWUEt).
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30 Location Global
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24 31 Methods Two remote-sensing datasets of GPP and ET, and four process-based biosphere
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26 32 model estimates of GPP and ET.
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30 33 Results Global WUE estimated by the two satellite datasets are 1.8 and 1.5 g C m-2 mm-1
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32 34 respectively, which are lower than the simulations from four process-based models (2.0 ± 0.3
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34 35 g C m-2 mm-1). Model-estimated WUE is 16% and 22% higher than that derived from satellite
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36 36 data in tropical and temperate biomes, and even up to 53% higher in the boreal zone.
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37 Precipitation is the main control on the WUE spatial distribution for temperate and tropical
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41 38 regions, but temperature dominates in areas north of 50°N. The values of WUE, produced
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43 39 either from satellite datasets or model simulations, are higher in wetter regions with higher
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45 40 GPP. WUEt exhibits a much lower, near-zero precipitation-sensitivity (SP) than WUE. This
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47 41 finding is contrary to leaf and plant level observations of higher WUEt in dry regions. Thus,
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50 42 we also calculated IWUEt, the product of WUEt and water vapour pressure deficit. IWUEt is
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52 43 found rather conservative with spatially increasing precipitation, in agreement with leaf and
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54 44 plant level measurements.
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Page 3 of 36 Global Ecology and Biogeography

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3 45 Main conclusions For arid ecosystems water losses are uncorrelated with vegetation
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5 46 productivity leading to lower WUE, whereas their control of canopy conductance is
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7 47 intrinsically efficient to maintain higher IWUEt. Water-use efficiency as defined in three
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10 48 different ways produce different behavior, suggesting that separate responses of each water
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12 49 loss composition should be paid attention in comparing WUE trends.
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 Global Ecology and Biogeography Page 4 of 36

1
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3 51 INTRODUCTION
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6 52 Photosynthesis is the largest component of the terrestrial carbon balance (Houghton, 2007). In
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9 53 this process CO2 diffuses through leaf stomata and mesophyll cell walls and is assimilated
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11 54 into carbohydrate (Sage, 1994). At the same time as CO2 is diffusing through leaf stomata,
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13 55 water vapour inside the leaf diffuses out as transpiration. Transpiration and photosynthesis are
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15 56 thus tightly coupled through leaf stomatal behaviour.
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19 57 The coupling between carbon and water exchange at the leaf level can be quantified by ratio
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21 58 of its photosynthesis to transpiration, or water use efficiency at leaf scale. But at ecosystem
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23 59 scale, water-use efficiency (WUE) is generally defined as the ratio of annual gross primary
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60 production (GPP) to annual evapotranspiration (ET). ET includes: transpiration, water loss
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28 61 from canopy interception and bare soil evaporation. Ecosystem WUE is thus regulated by
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30 62 canopy conductance that is estimated by integrating leaf stomatal conductance over canopy
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32 63 leaf area index. Aerodynamic conductance and vegetation coverage also affect ecosystem
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34 64 WUE through determining the extent to which water and energy fluxes interact with the
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65 atmosphere (Jarvis & McNaughton, 1986). Water loss associated with ecosystem carbon
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39 66 assimilation can be divided into two components: (1) ‘physiologically productive water’
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41 67 channeled through transpiration during the growing season; and (2) ‘non-productive’ water
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43 68 use: canopy interception and bare soil evaporation (Ponton et al., 2006; Hu et al., 2008). The
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69 partitioning of total ecosystem water loss between these productive and non-productive water
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48 70 uses can affect the coupling of water and carbon cycle at regional and global scales (Jackson
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50 71 et al., 2005).
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53 72 At leaf or canopy scale, optimal theories have been developed and used to explain quite
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56 73 successfully the responses of WUE to various environmental factors such as increased CO2
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58 74 and available soil water (Manzoni et al., 2011; Medlyn et al., 2011). Leaf WUE has been
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Page 5 of 36 Global Ecology and Biogeography

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3 75 observed or predicted to increase under mild water stress, which theoretically results from
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5 76 decreased stomatal conductance (Manzoni et al., 2011; Niu et al., 2011). However,
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7 77 non-stomatal limitations, including reduced Rubisco activity and electron transport capacity,
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10 78 are also found to play a significant role in regulating leaf photosynthetic rate (Zhou et al.,
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12 79 2013). As water stress becomes more severe, those internal controls will dominate, and reduce
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14 80 leaf WUE (Manzoni et al., 2011). Moreover, the response of leaf WUE to water stress varies
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16 81 among species owing to different water use strategies (Zhou et al., 2013).
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20 82 At ecosystem scale, interactions between leaf physiology and canopy leaf area dynamics,
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22 83 carbon allocation and so on result in complex responses to changes in external environment
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24 84 (Niu et al., 2008). Water loss by soil evaporation may respond to environmental change quite
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26 85 differently from transpiration at ecosystem scale. Furthermore, differences in ecological states
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86 related to stand age, plant type, structure and composition of ecosystems (Niu et al., 2011)
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31 87 may lead to different water-use strategies by different ecosystems or under different climate
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33 88 conditions.
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89 Although ecosystem-scale WUE has been measured and compared in many studies, no
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39 90 consistent conclusions have emerged about the variability of mean WUE across climate
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41 91 gradients. For example, some studies found that ecosystem WUE is higher in drier regions
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43 92 (Yu et al., 2008; Lu & Zhuang, 2010; Ito & Inatomi, 2012), a finding which is consistent with
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93 that for leaf-scale responses, but contrary to the results based on eddy-covariance
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48 94 measurements of CO2 and water fluxes by Hu et al. (2008), who found that ecosystem WUE
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50 95 increases increased with precipitation. By defining WUE as the ratio of aboveground net
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52 96 primary productivity (NPP) to ET, Ponce-Campos et al. (2013) found WUE a rather
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54 97 conservative mean quantity across different biomes. They also found that the response of
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57 98 ecosystem WUE to changes in precipitation varied over time, or WUE increased to a
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3 99 maximum limit during drought, and decreased to a common value in wet years. Moreover,
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5 100 simulations by Tian et al. (2010) suggest that the relationship between WUE (calculated as
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7 101 the NPP to ET ratio) and annual mean precipitation is ecosystem-specific. For example, WUE
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10 102 decreased for wetland and cropland, but increased for forest and grassland ecosystems in
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12 103 response to an increase in precipitation.
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15 104 Some of those diverging responses may be explained by the different definition of WUE. It
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105 brings difficulties to draw firm conclusions about the causes of spatial gradients in WUE at
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20 106 regional to global scale based on the results of previous studies. Furthermore responses of
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22 107 ecosystem WUE to environmental changes can vary in space and time (Ponce-Campos et al.,
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24 108 2013). Contradictions between local studies can be partly due to their lack of spatial
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26 109 representativeness. In addition, many previous studies have empirically correlated WUE with
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110 precipitation gradients but ignored the confounding effects of temperature, radiation and
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31 111 nutrient availability (Hu et al., 2008; Yu et al., 2008; Ponce-Campos et al., 2013). Therefore,
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33 112 those empirical relationships do allow us to gain further insight about the key controls of
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35 113 ecosystem WUE at different temporal and spatial scales.
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39 114 In this study we investigated the global spatial patterns of ecosystem WUE. Spatial drivers
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41 115 and limiting climatic and ecological factors controlling WUE spatial variations were analyzed
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43 116 using a spatial regression analysis of WUE upon temperature, precipitation and solar radiation.
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117 Because temperature, rainfall and radiation co-vary spatially, multi-regression analysis
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48 118 between WUE and these three climate variables were used to identify the dominant global
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50 119 driver for regions under different climates. In addition, two other definitions (water uses
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52 120 transpiration-based water-use efficiency, i.e., WUEt, and inherent water-use efficiency, i.e.,
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54 121 IWUEt) discriminating the productive water uses from non-productive water consumption
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3 122 were used to investigate how different water uses affect the coupling of water and carbon
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5 123 cycles.
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9 124 Material and methods
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12 125 Data-driven GPP products
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16 126 The MODIS and JUNG sets of gridded GPP data products are derived from climate and
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18 127 satellite data, and from satellite, climate and flux-tower data, respectively. MODIS GPP is
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20 128 estimated at 1 km spatial resolution from a light-use efficiency (LUE) model, as part of the
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22 129 operational MODIS algorithms (Running et al., 2004) using meteorological data from NASA
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130 Data Assimilation Office and detailed vegetation information (land cover and FPAR) derived
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27 131 from the Moderate Resolution Imaging Spectroradiometer (MODIS) satellite from 2000 to
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29 132 present (Running et al., 2004; Zhao et al., 2005; Turner et al., 2006). JUNG GPP is another
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31 133 data-oriented product derived by extrapolating the flux-tower observations from 178 sites in
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33 134 space and time using climate data and remotely sensed fAPAR data from 1982 to 2008 (Jung
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36 135 et al., 2009). This latter monthly GPP product covers the period 1982-2008 at 0.5° spatial
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38 136 resolution, but many of the flux tower observations only cover much shorter periods.
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41 137 Data-driven ET products
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45 138 Global ET also cannot be measured directly at present, and are estimated using algorithms
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47 139 based on global meteorology data. MODIS ET is estimated using the Penman-Monteith
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49 140 approach with the enhanced vegetation index (EVI) fields from MODIS as input (Mu et al.,
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141 2011; MOD16A2, http://www.ntsg.umt.edu/, accessed 16 September 2013). JUNG ET is
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54 142 another dataset based on model tree ensemble (MTE) regression, which extrapolates flux
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56 143 tower data as described for GPP above, yet with an additional correction for energy balance
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3 144 closure assuming constant Bowen ratios (Jung et al., 2010). The spatial resolution of MODIS
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5 145 ET and JUNG ET are same as their corresponding GPP products.
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9 146 Process-based models
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12 147 Process-based models incorporate empirical knowledge of mechanisms, and mass, water and
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14 148 energy conservation principles to simulate fluxes of CO2 and water. Four models (CLM,
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149 CABLE, LPJ and ORCHIDEE) were used in this study at a global resolution of 0.5°. We use
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21 151 (http://dods.extra.cea.fr/data/p529viov/cruncep/) and annual CO2 concentration over the
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153 physiological processes controlling spatial gradients of WUE are incorporated in each model.
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28 154 Particularly, CLM and CABLE take into account the effect of nitrogen-limitation on
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30 155 ecosystem photosynthesis. Information about each model is presented in Table S1, and more
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32 156 details are given by Mao et al. (2013) for CLM, Wang et al. (2011) and Zhang et al. (2013)
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34 157 for CABLE, Sitch et al. (2003) for LPJ, and Krinner et al. (2005) for ORCHIDEE.
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38 158 Analyses
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41 159 GPP and ET from MODIS were re-gridded to 0.5° resolution using nearest-neighbour pixels.
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160 Monthly GPP and ET from remote-sensing data and from model simulations were summed to
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46 161 give annual totals. Mean annual GPP and ET were then calculated for the period from 2000 to
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48 162 the most recent year available (different time periods covered by each dataset, see Table S1).
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50 163 For both GPP and ET, the 0.5° grid cells with low productivity defined by a mean annual
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52 164 Normalized Difference Vegetation Index (NDVI, from AVHRR NDVI3g data) lower than 0.1
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55 165 were discarded (11% of the land area).
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3 166 Assuming homogeneous vegetation cover within each 0.5° grid cell, the ecosystem WUE was
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5 167 calculated as:
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8 ‫ܲܲܩ‬
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‫ܶܧ‬
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12 169 with GPP in g C m-2 yr-1 and ET in mm yr-1. For the remote-sensing datasets and process
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14 170 models, we used only GPP and ET produced or simulated consistently by each data-driven
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171 algorithm or model. Transpiration-based WUE (WUEt) was defined as the ratio of annual
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ܹܷ‫= ݐܧ‬
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26 174 T is only available from the process model outputs.
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29 175 Previous studies by Beer et al. (2007, 2009) suggested that ‘intrinsic water use efficiency’,
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176 IWUE, that accounts for the effect of VPD on WUE should provide a better measure of
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34 177 ecosystem water use efficiency to other climatic variables:
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36 ‫ܦܸܲ × ܲܲܩ‬
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41 179 where VPD is the diurnal vapour pressure deficit (hPa) averaged over the growing season
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43 180 (defined here as the months when NDVI > 0.1) from Global Monitoring and Assimilation
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45 181 Office (GMAO, http://disc.sci.gsfc.nasa.gov/daac-bin/FTPSubset.pl ). Mean inherent
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47 182 transpiration-based water use efficiency, IWUEt, can then be calculated from the output of
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183 each model as:
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56 185 Annual WUE in each 0.5° grid cell was projected in a two-dimensional space with mean
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58 186 annual temperature (MAT; ranging from -20 to 30°C as the horizontal axis) and mean annual
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3 187 precipitation (MAP; ranging from 0 to 2000 mm as the vertical axis) binned into intervals of
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5 188 0.5°C MAT and 20 mm MAP. For all grid cells within the same MAT and MAP pixel,
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7 189 area-weighted averages of GPP, ET and then WUE were calculated.
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11 190 For the multiple regression analysis, spatial climate gradients were calculated using a moving
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13 191 window of 4.5°×4.5°, i.e., with the 81 pixels surrounding each 0.5° grid cell. The effects of
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15 192 precipitation, temperature, and radiation on WUE were estimated separately by controlling for
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193 the effects of other variables.
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21 194 Results
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195 Global WUE
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28 196 Using two sets of satellite data products, we obtain an estimate of global WUE of 1.8 g C m-2
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30 197 mm-1 and 1.5 g C m-2 mm-1, from MODIS and JUNG data respectively (Fig. 1). Across
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32 198 different regions, estimates of WUE from MODIS are higher than those from JUNG in
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35 199 tropical (30°S-30°N) and temperate (30°N-50°N and 30°S-50°S) zones, but slightly lower in
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40 201 Process-based model simulations produce on average higher estimates (by ≈ 21%) of global
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202 WUE than those derived from remote sensing, particularly in boreal zone, where the mean
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45 203 estimate of WUE from process-based models is 54% higher than the estimate from the
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47 204 remote-sensing data products. The mean global WUE from different process-based models is
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49 205 estimated at 2.0 g C m-2 mm-1, varying from 1.7 g C m-2 mm-1 by CLM to 2.3 g C m-2 mm-1 by
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55 207 Global spatial patterns of WUE
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3 208 The two estimates of WUE derived from remote-sensing products both show large spatial
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5 209 variations (Figs 2a-b), with significant differences between different datasets, especially in
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7 210 dryland areas (MAP < 250 mm). WUE over northern Europe (ranging from 2.5 to 3.0 g C m-2
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10 211 mm-1), northern Eurasia (≈ 50°N), tropical rainforest areas and southeastern North America
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12 212 (temperate and sub-tropical) are consistently higher than other regions, while estimates of the
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14 213 WUE < 1.0 g C m-2 mm-1 are found in the Qinghai Tibet Plateau, India and northwestern
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16 214 North America.
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20 215 Similar spatial patterns of WUE are simulated by the process-based models, although the
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22 216 estimates of WUE values from the results of four process-based models are much higher than
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24 217 estimates from remote-sensing products, especially in mid- and high latitude regions. For
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26 218 regions north of 50°N, the average WUE values over four model simulations are generally
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219 larger than 2.0 g C m-2 mm-1 and higher than data-driven ones by up to 40% (Figs 2g-h).
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31 220 However, WUE values over high latitudes vary greatly among different simulations, which
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33 221 ranging from < 0.5 g C m-2 mm-1 for CLM to > 3.5 g C m-2 mm-1 for LPJ (Figs 2c-f).
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222 Variation of the sensitivities of WUE to MAP and MAT
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40 223 Spatial variations of WUE with MAT and MAP are quite similar between the estimates from
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42 224 satellite data and process-based models (see Fig. 3). Globally, estimates of WUE using both
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225 methods are highest in regions with optimal hydro-thermal conditions (i.e., the certain ratios
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47 226 of MAP to MAT). However, estimate of WUE from remote-sensing products is maximal at a
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49 227 MAP of ≈ 1300 mm and MAT between 5 to 10°C (Fig. 3a), while maximal WUE estimates
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51 228 from the process-based models are found in regions with MAP in the range of 600-800 mm
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53 229 and MAT between 2 and 6°C (Fig. 3b).
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3 230 However, variations of WUE with MAP or MAT, measured by precipitation-sensitivity (SP)
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5 231 and temperature-sensitivity (ST), from two different approaches are quite different (insets of
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7 232 Fig. 3). SP is positive for the results from both remote-sensing products and process-based
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10 233 models, even in the cold regions with MAT lower than 0°C. But, SP estimated from
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12 234 remote-sensing products decreases with MAT when MAT < 10°C, then increases with MAT
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14 235 when 10°C < MAT < 15°C, and decreases with MAT when MAT > 15°C (Fig. 3a). In
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16 236 contrast, SP estimated from process-based models increases with MAT when -10°C < MAT
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237 <10°C, then remains almost constant when MAT > 10°C (Fig. 3b). In terms of ST,
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21 238 remote-sensing products present positive values in arid regions with MAP less than 200 mm
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241 Geographical spatial sensitivity of WUE to climate variables
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32 242 In climate space, multiple regression analysis shows that WUE is significantly (p < 0.05)
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34 243 positively related to precipitation for ≈ 40% of the vegetated land both in the remote-sensing
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244 products and in the models (Fig. 4a, 4b). Positive correlation mainly occurs south of 50°N,
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39 245 with lower SP (< 1‰ mm-1; p < 0.05) found in humid areas and higher values (> 4‰ mm-1; p
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41 246 < 0.05) in drier regions such as the western United States, central Asia and dry areas of
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43 247 northeast Asia. At high latitudes (60°N - 90°N), WUE over ≈ 40% of the pixels (p < 0.05) is
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248 negatively associated with increasing precipitation. The biggest differences in the WUE-MAP
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48 249 spatial correlation between remote-sensing data and models are found for tropical drylands. In
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50 250 these regions, model-simulated WUE is significantly positive correlated with MAP, while
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52 251 remote-sensing estimated WUE responds negatively to MAP (Fig. S2). Moreover, for tropical
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54 252 rainforests, modelled WUE is positively correlated with precipitation; but estimates from the
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3 253 remote-sensing products show spatially varying responses to changing precipitation, with
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5 254 much weaker sensitivity than that from model simulations.
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9 255 Both remote-sensing and model results show broadly consistent variation of WUE with MAT
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11 256 (Figs 4c-d). Generally, WUE is positively correlated with temperature at high latitudes (about
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13 257 50% pixels north of 50°N ; p < 0.05), with ST ranging from 5% °C-1 to 20% °C-1 from
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15 258 remote-sensing products, but smaller ST of less than 10% °C-1 from model results. On the
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259 contrary, WUE is negatively correlated with MAT over ≈ 30% (p < 0.05) of the pixels south
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20 260 of 50°N. For tropical rainforest ecosystems, the remote-sensing products and model results
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22 261 consistently produce a weak negative sensitivity of WUE to temperature (ST > -10% °C-1; p <
r
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24 262 0.05).
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28 263 In terms of the impact of mean annual solar downward radiation (RS), both remote-sensing
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30 264 and model WUE results show positive correlations between WUE and RS over 20% to 30% (p
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32 265 < 0.05) pixels north of 50°N, with radiation-sensitivity (SR) higher than 6% per W m-2 (Figs
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34 266 4e-f). However, negative correlations dominate in the low and mid-latitude regions with
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267 considerable variation of SR ranging from near-zero to values < -6% per W m-2. SR in
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39 268 mid-latitudes (-1% per W m-2 ~ -3% per W m-2) is larger in magnitude than in tropical forest
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41 269 regions (> -1% per W m-2). In addition, data-driven results show positive SR over most of
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43 270 Australia, part of North Africa and southern North America, which contradict with model
44
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271 simulations.
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49 272 DISCUSSION
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52 273 Major differences between data-driven and modelled WUE
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56 274 The two WUE estimations based on remote-sensing datasets differ from each other in both
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58 275 global mean values and spatial patterns, especially for regions with low ET, such as tropical
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3 276 drylands. Differences of WUE between MODIS and JUNG are primarily attributable to their
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5 277 systematic differences in estimated ET. ET produced by MODIS is considerably lower in dry
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7 278 areas but generally higher in humid regions than JUNG ET. These differences in ET lead to
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10 279 great differences in the magnitude of WUE, as well as spatial gradients of WUE between
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12 280 MODIS- and JUNG-based estimates. Uncertainties in ET estimated from remote-sensing can
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14 281 have a number of causes such as: ignoring terrestrial water storage terms, the lack of energy
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16 282 balance closure in flux-tower datasets, and landscape-level heterogeneity (Zeng et al., 2012).
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283 In addition, biases in GPP from satellite products also lead to errors in WUE estimates.
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21 284 MODIS and JUNG all use satellite fAPAR to drive empirical models for GPP, but this
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23 285 methodology is considered to be uncertain because cloudiness can cause unspecified biases in
24
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25 286 fAPAR (Baret et al., 2007). Moreover, Lee et al. (2013) found that enhanced vegetation index
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27 287 (EVI) in Amazon remains high, whereas sun-induced chlorophyll fluorescence (SIF) from
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30 288 GOSAT SIF suggests a decrease in GPP during the dry season. Thus GPP estimates made
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32 289 using current fAPAR can be positively biased in tropical regions. For GPP derived from
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34 290 JUNG, uncertainty increases for boreal and tropical regions, where few flux sites are located.
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291 Compared with remote-sensing based WUE, model estimates are generally higher (Fig. 1, Fig.
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40 292 2), indicating that some of the processes involved are poor simulations of the actual
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42 293 physiological and ecological mechanisms. As shown by Fig. 5, averaged GPP across four
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44 294 models is 28% higher than remote-sensing derived GPP, whereas mean modelled ET is not
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46 295 significantly different from the data-driven result. Moreover, differences in modelled
47
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49 296 processes such as carbon-nitrogen interactions, soil depth maps, root profiles, and soil water
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51 297 holding capacity parameters, as well as different function of water stress (Sulman et al., 2012)
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53 298 are all responsible for the different coupling between GPP and ET in each model, and thus
54
55 299 lead to larger uncertainties in WUE estimates.
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3 300 Nutrient resource availability also regulates plant productivity (Cleveland et al., 2013) as well
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5 301 as other ecosystem processes, and eventually impacts WUE. Fisher et al. (2012) even inverted
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7 302 regions with nutrient limitation from NDVI deviations from a maximum NDVI-ET slope,
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10 303 which is close to a maximum envelope for WUE. Previous studies on the temporal change of
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12 304 productivity concluded that nitrogen limitation reduces the response of GPP to elevated
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14 305 atmospheric CO2 (Thornton et al., 2007; Zaehle et al., 2014). Spatially, the insufficient
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16 306 availability of nitrogen is widespread in terrestrial vegetation (Wang et al., 2010) and nitrogen
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307 limitation increases from the tropics to high latitudes (Wang et al., 2010). Wang et al. (2010)
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21 308 estimated that net primary productivity (NPP) is reduced by 35% - 40% at high latitudes by
22
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23 309 nitrogen limitation and reported that simulations without considering the nitrogen limitation
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25 310 overestimate NPP by about 29%. For tropical ecosystems, the impact of nitrogen limitation on
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27 311 plant growth is weaker, because in tropical ecosystems ~ 18% of NPP is contributed by
28
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30 312 external nitrogen inputs (Cleveland et al., 2013). Some studies have also suggested that
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32 313 phosphorus rather than nitrogen limitation constrains NPP across much of the tropics where
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34 314 nitrogen is plentiful (Wang et al., 2010; Cleveland et al., 2013); this is because new
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36 315 phosphorus inputs are very small relative to plant demand in the tropics (Cleveland et al.,
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316 2013).
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42 317 Failure to include human disturbance in the models may also leads to biases in vegetation
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44 318 productivity. None of the models used for this study take land-use change into account; the
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46 319 models may thus contain much too great an area of forest and hence higher simulated GPP
47
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49 320 (Bondeau et al., 2007). Neglecting air pollution also causes a reduction in modelled GPP by
50
51 321 deteriorating physiological and ecological processes. Although many studies have reported
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53 322 that nitrogen limitation significantly suppresses ET through reducing leaf area and stomatal
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55 323 conductance under photosynthesis down-regulation (Lee et al., 2013), our comparison
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3 324 between remote-sensing based and modelled ET shows no evident high-bias in the model
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5 325 simulations (Fig. 5b).
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7
8
9 326 Spatial pattern of WUE
10
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12 327 With the exception of the patterns estimated from MODIS, the spatial distributions of WUE
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14 328 estimated from different datasets consistently show higher values in wetter environments (Fig.
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16
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329 2). For mid- and low latitude ecosystems with seasonal water-limitation (Huxman et al.,
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19 330 2004), WUE spatial gradients are controlled by precipitation rather than by temperature and
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21 331 radiation (Fig. S2). The positive sensitivity of WUE to precipitation gradually declines from
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23 332 desert, through mesic, to humid regions as soil water stress decreases. Semi-arid and mesic
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333 areas, limited by soil water availibility, can utilize higher precipitation to sustain GPP more
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28 334 efficiently than wetter regions (Huxman et al., 2004). The enhancement of ET is less than that
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30 335 of GPP per unit of precipitation increase, which results in a higher WUE for wetter areas
31
32 336 when going from arid to mesic climates. For humid regions such as northern Europe and
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34 337 tropical rainforest, where ecosystems have a high WUE, any increase in precipitation in
35
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338 excess of plant demand becomes runoff, which may even deplete nutrients available for plant
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39 339 growth through leaching (Huxman et al., 2004; Posada & Schuur, 2011). More cloudiness
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41 340 during precipitation also decreases RS, which consequently limits GPP (Scanlon & Albertson,
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43 341 2004). In the context of long-term adaptation in areas with abundant precipitation, plants with
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342 higher resources use efficiency, especially for nutrients rather than water, have a competitive
46
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48 343 advantage in long-term selection (Huxman et al., 2004). Ecological studies of NPP also point
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50 344 out that nitrogen mineralization, water uptake by plants and productivity (NPP) are optimally
51
52 345 adjusted to each other (Roy et al., 2001). As a result, WUE tends to be controlled by the
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54 346 strongest limiting factor. The existence of other limiting factors for WUE results in spatially
55
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57 347 inconsistent responses of the sensitivity of WUE to precipitation between models and
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3 348 remote-sensing results (Figs S2a-b). Without considering nutrient limitation, models predict
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5 349 that WUE is enhanced as precipitation increases; therefore WUE can be positively correlated
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7 350 with precipitation even for rainforest (Figs 4a-b). GPP of ecosystems at high latitudes often is
8
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10 351 co-limited by temperature and radiation instead of precipitation (Figs 4c-f). For those cold
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12 352 regions, air temperature and incoming solar radiation limit GPP, they are positively correlated
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14 353 with ecosystem productivity (Piao et al., 2007) and hence lead to higher WUE. However,
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16 354 observations of the carbon cycle from GOSAT (Frankenberg et al.,2011) suggested that
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355 MODIS and JUNG underestimate GPP in the tropics for savannas where no flux tower
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21 356 observations are available at present, and overestimate GPP in arctic regions. So it is possible
22
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23 357 that estimates of GPP from MODIS and JUNG are too high in tropical forest and boreal forest,
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25 358 and too low in savannas. This would change the spatial pattern of WUE.
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359 Most current models and observations indicate that transpiration dominates the terrestrial
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31 360 water flux emitted to the atmosphere (Lawrence, 2007; Jasechko et al., 2013). Based on this,
32
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33 361 previous studies used ET, expecting it to be close to T, to calculate ecosystem WUE.
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35 362 However, few observation sites are located in arid regions with annual precipitation less than
36
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363 200 mm, at such sites bare soil evaporation contributes most to ET, and acts to decrease WUE.
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40 364 Our results (Fig. 3) show that mean WUE increases with precipitation along climate gradients,
41
42 365 which contradicts evidence from WUE observations at leaf and plant scale that water stress is
43
44 366 found to reduce T per unit of GPP.
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46
47
48 367 To explore whether the inclusion of bare soil evaporation in ET results in this discrepancy, we
49
50 368 used model simulations to calculate the transpiration-based water use efficiency (WUEt)
51
52 369 given by equation (2). WUEt in climate space still shows higher values with larger
53
54 370 precipitation (Fig. 6a), except for cold regions with MAT < 0°C, indicating that the lower
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57 371 WUE in dry regions is not only determined by the increased soil evaporation. This finding is
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3 372 consistent with canopy-scale studies (Baldocchi, 1994; Hu et al., 2008), which pointed out
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5 373 that WUEt at canopy scale is mainly controlled by photosynthetic capacity (i.e., GPP) instead
6
7 374 of T. The denser canopies of vegetation growing in humid conditions have both larger
8
9
10 375 light-bearing surfaces and higher light use efficiency (Hu et al., 2008). Thus vegetation will
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12 376 benefit from the increased diffuse radiation (Knohl & Baldocchi, 2008), and hence increase
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14 377 GPP per unit of water loss. However, we also found that, in climate space, modelled values of
15
16 378 SP for WUEt are generally positive but very close to zero, except for cold regions with MAT <
17
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379 0°C, indicating that arid ecosystems take a number of strategies to reduce transpiration to
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21 380 maintain normal physiological activities during dry seasons (Monasterio & Sarmiento, 1976).
22
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24 381 Increased water vapour pressure deficit at the leaf surface has been found to reduce leaf WUE
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26 382 (Linderson et al., 2012). Similar responses have also been observed at canopy or ecosystem
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383 scale (Ponton et al., 2006; Niu et al., 2011; Linderson et al., 2012). In order to remove the
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31 384 effects of both bare soil evaporation and VPD, here, we now use transpiration-based inherent
32
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33 385 water-use efficiency (IWUEt, equation 4) to investigate variations of modelled IWUEt with
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35 386 MAP and MAT. A quite different modelled IWUEt-MAP spatial relationship is observed to
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387 that found for WUE and shows that IWUEt displays negative, but very close to zero, SP for
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40 388 warm regions with MAT > 10°C (Fig. 6b). This finding basically agrees with the leaf WUE
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42 389 conclusion of higher WUEt being found in dry regions, where drought-tolerant species with
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44 390 physiological advantages in resisting drought and holding water dominate the ecosystem
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46 391 (Zhou et al., 2013). For arid regions, more water vapour flux is allocated to bare soil
47
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49 392 evaporation (Hu et al., 2008) and intensive transpiration driven by high VPD. This high water
50
51 393 cost, uncorrelated with vegetation productivity, leads to relatively lower WUE in dry areas
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53 394 than in wetter areas, though modelled plants in these regions conserve a higher IWUEt than in
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55 395 mesic and humid climates. In contrast, for wet regions with high vegetation coverage and high
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396 air humidity, most of the water consumption is closely bound up with vegetation productivity
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1
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3 397 and hence leads to a higher WUE. Moreover, long-term natural selection, i.e., ecosystem
4
5 398 adaptation to local climate and nutrient status (Huxman et al., 2004), shaped the optimal
6
7 399 vegetation composition (Still et al., 2003) and essentially determined the spatial pattern of
8
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10 400 WUE.
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13 401 Considering that poor representation of eco-physiological mechanisms in the models may
14
15 402 lead to unrealistic conclusions, we calculated satellite product IWUE (equation 3) to test the
16
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403 validity of the model results. A lower SP was also estimated by data-driven IWUE than WUE,
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20 404 despite negative values found with MAT > 20°C owing to an extremely low MODIS-ET
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22 405 product (Fig. S2a). These results suggest that the ‘intrinsic changes’ of water and carbon
r
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24 406 correlation actually exist in ecosystems. IWUE derived from models (Fig. S2b) shows a
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26 407 generally similar climatic pattern, except for dry areas with MAP < 200 mm but higher SP
27
28
408 across the whole MAT range with the satellite result. This response to precipitation is
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31 409 different from that estimated from modelling IWUEt (Fig. 6b) with negative but near-zero SP
32
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33 410 across the whole MAT range, indicating the important influence of evaporation on WUE
34
35 411 behaviour. Consistent IWUE-MAP relationships derived from satellite products and models
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412 demonstrates that the conclusions from our calculation of water-use efficiency based on
39
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40 413 transpiration (WUEt, IWUEt) are broadly credible in explaining the spatial patterns of WUE.
41
42
43 414 ACKNOWLEDGEMENT
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47 415 This study was supported by the National Natural Science Foundation of China (41125004),
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49 416 National Basic Research Program of China (2013CB956303), Chinese Ministry of
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51 417 Environmental Protection Grant (201209031), the 111 Project (B14001), and National Youth
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53 418 Top-notch Talent Support Program in China. Jiafu Mao and Xiaoying Shi’s time is supported
54
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56 419 by the US Department of Energy (DOE), Office of Science, Biological and Environmental
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58 420 Research. Oak Ridge National Laboratory is managed by UT-BATTELLE for DOE under
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3 421 contract DE-AC05-00OR22725. The simulation of CLM is supported by the US Department
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5 422 of Energy (DOE), Office of Science, Biological and Environmental Research. Oak Ridge
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7 423 National Laboratory is managed by UT-BATTELLE for DOE under contract
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10 424 DE-AC05-00OR22725.
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13 425
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1
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33 611 BIOSKECH
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36 612 Yan Sun is a PhD candidate at Peking University, Beijing, China. She has broad research
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38 613 interests particularly in vegetation dynamics and the carbon balance of terrestrial ecosystems.
39 614 She also maintains interests in nitrogen and phosphorus cycles.
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3 616 Figure legends
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6 617 Figure 1 Total water-use efficiency (WUE) estimations for global, boreal (50°N-90°N),
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9 618 tropical (30°S-30°N) area and temperate zone of the Northern Hemisphere (30°N-50°N) and
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11 619 Southern Hemisphere (30°S-50°S) from two sets of observed evapotranspiration (ET) and
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13 620 gross primary productivity (GPP) products and four global models. Error bars indicate
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15 621 interannual standard deviations.
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21 623 dataset and (c-f) process-based models, averaged from 2000 to the most recent year available
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625 mean WUE across four models are also shown. Values of mean annual Normalized
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40 631 MAP under same MAT interval) along the MAP gradient. Red lines represent the relative
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633 same small range of MAT) of WUE. Grey scale indicates uncertainties among two
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47 634 data-driven results and four models results, in which the darkest zones represent largest
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52 636 Figure 4 Spatial sensitivity of WUE to: (a, b) precipitation, (c, d) temperature and (e, f) solar
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55 637 radiation, based on remote-sensing data (a, c, e) and process models (b, d, f). Only statistically
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3 639 Figure 5 Comparisons between products from data-driven and outputs simulated by process
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5 640 models: (a) mean GPP and (b) mean ET, the colour bar shows the point intensity.
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9 641 Figure 6 Distribution of (a) mean transpiration-based water-use efficiency (WUEt) and (b)
10 642 transpiration-based inherent water-use efficiency (IWUEt) in MAT-MAP domain from
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12 643 process-based models. The bottom insets show the MAP-sensitivity (red lines) across the
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14 644 MAT range and the grey scale indicates uncertainties among four model results, in which the
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