Decreased soil cation exchange capacity across ...

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Sep 21, 2017 - 26. • The decreases were detected on both the Tibetan Plateau and the Inner Mongolian. 27. Plateau, among various grassland types. 28.
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Decreased soil cation exchange capacity across northern China’s

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grasslands over the last three decades

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Kai Fang1,2, Dan Kou1,2, Guanqin Wang1,2, Leiyi Chen1, Jinzhi Ding1,2, Fei Li1,2, Guibiao Yang1,2, Shuqi Qin1,2, Li Liu1,2, Qiwen Zhang1,2 and Yuanhe Yang1,2*

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Chinese Academy of Sciences, Beijing 100093, China

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University of Chinese Academy of Sciences, Beijing 100049, China

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Corresponding author: Dr. Yuanhe Yang, tel.: + 86 10-6283 6638, fax: + 86

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10-6283 6632, E-mail: [email protected]

State Key Laboratory of Vegetation and Environmental Change, Institute of Botany,

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Revised manuscript submitted to Journal of Geophysical Research:

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Biogeosciences

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21th- Sep.-2017

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Manuscript information: 31 pages, 1 table, 5 figures and 1 supplementary dataset.

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Key Points:

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 Decreases in topsoil cation exchange capacity were observed across northern

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China’s grasslands over the last three decades.  The decreases were detected on both the Tibetan Plateau and the Inner Mongolian Plateau, among various grassland types.  The decreases were attributed to soil carbon loss, soil desertification, soil acidification, and extreme precipitation.

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Abstract

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Cation exchange capacity (CEC) helps soils hold nutrients and buffer pH, making it

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vital for maintaining basic function of terrestrial ecosystems. However, little is known

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about the temporal dynamics of CEC over broad geographical scales. In this study, we

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used Random Forest method to compare historical CEC data from the 1980s with new

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data from the 2010s across northern China’s grasslands. We found that topsoil CEC in

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the 2010s was significantly lower than in the 1980s, with an overall decline of about

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14%. Topsoil CEC decreased significantly in alpine meadow, alpine steppe, meadow

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steppe, and typical steppe by 11%, 20%, 27% and 9% respectively. Desert steppe was

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the only ecosystem type which experienced no significant change. CEC was

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positively related to soil carbon content, silt content, and mean annual precipitation,

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suggesting that the decline was potentially associated with soil organic carbon loss,

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soil degradation, soil acidification, and extreme precipitation across northern China’s

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grasslands since the 1980s. Overall, our results demonstrate topsoil CEC loss due to

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environmental changes, which may alter the vegetation community composition and

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its productivity and thus trigger grassland dynamics under a changing environment.

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Keywords: environmental change, geochemistry, grassland ecosystems, soil carbon

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loss, soil desertification, soil acidification, extreme precipitation

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1. Introduction

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Cation exchange capacity (CEC) is usually expressed as the number of moles of

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exchangeable cations adsorbed by electrostatic force per unit mass soil. CEC plays an

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important role in holding soil nutrients [Brady and Weil, 2002] and buffering soil pH

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changes [Xu et al., 2012; Luo et al., 2015], which makes it vital for maintaining the

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structure and function of terrestrial ecosystems. CEC can affect ecosystem structure

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and function by controlling the supply of exchangeable cations (i.e., Ca2+, Mg2+, K+,

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Na+, Al3+, and Fe3+) in the soil [Lucas et al., 2011; Mueller et al., 2012]. Given that

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different plant species require different concentrations and ratios of Ca2+, Mg2+, K+

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and Na+, changes in these cations can alter plant biodiversity [Chen et al., 2013].

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Besides being essential to plant growth [Likens et al., 1998; McLaughlin and Wimmer,

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1999], the availability of these nutrients affects plant tolerance to drought, frost, and

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pathogens [DeHayes et al., 1999; Demchik and Sharpe, 2000], therefore affecting

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primary productivity and overall structure of terrestrial ecosystems. In contrast,

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increased Al3+ and Fe3+ could limit vegetation productivity because of its toxicity

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[Lieb et al., 2011]. CEC can also maintain the stability of terrestrial ecosystems by

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buffering changes in soil acidity [Bowman et al., 2008], since soil pH decreases could

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alter biological activity and cation supply and have negative impacts on terrestrial

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ecosystems [Kirk et al., 2010]. Overall, in the context of environmental changes, a

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deep understanding of the temporal dynamics of CEC is crucial for predicting the

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responses of ecosystem structure and function, and in guiding policies that maintain

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the stability of terrestrial ecosystems. 3

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Given the importance of CEC, studies about the effects of environmental changes on

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CEC have gradually accumulated in recent decades [Watmough and Dillon, 2003;

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Högberg et al., 2006; Fissore et al., 2007; Ruiz Sinoga et al., 2012; Xu et al., 2012; Lu

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et al., 2015]. It has been reported that CEC exhibited positive correlations with

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2:1-type clay minerals [Xu et al., 2012] and organic matter content [Ruiz Sinoga et al.,

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2012; Xu et al., 2012]. Due to the close relationships between soil organic matter and

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climatic parameters, CEC has also been observed to be positively related with

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precipitation [Ruiz Sinoga et al., 2012] but negatively related with mean annual

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temperature [Fissore et al., 2007]. These studies have generated broad understanding

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of spatial variation and environmental drivers of CEC over broad geographic scales,

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but little is known about temporal CEC dynamics at large spatial scales. Long-term

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site monitoring has demonstrated cation loss from soils due to soil acidification,

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harvesting, and land-use change [Watmough and Dillon, 2003; Högberg et al., 2006;

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Lu et al., 2015]. However, these observations are limited to the site scale and it is

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unknown if they are representative of larger landscape or continental trends.

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Considering that the driving factors for CEC dynamics (e.g., climatic factors, edaphic

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factors, and human activity) exhibit substantial variations in space and time, a

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comprehensive study about the temporal dynamics of CEC along broad environmental

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gradients is needed.

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China’s grasslands are distributed across large geographic scales, making them an 4

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ideal ecosystem to explore the temporal dynamics of CEC for three reasons. First,

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along with the geographic gradient, grassland types are diverse (including both alpine

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and temperate grasslands) and environmental parameters such as precipitation and

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edaphic variables exhibit a wide range [Liao and Jia, 1996]. Second, since the 1980s,

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northern China has experienced significant environmental changes, which mainly

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include the following: (1) Overgrazing and cultivation [Bridges and Oldeman, 1999;

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Yu et al., 2012] have led to soil desertification [Yang et al., 2007; Mu et al., 2013] and

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pasture degradation [Dai et al., 2011], thereby decreasing grassland productivity

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[Babel et al., 2014] and organic carbon storage [Xie et al., 2007; Dai et al., 2011]

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across the study area; (2) Elevated atmospheric acid deposition has induced soil

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acidification across northern China, with a decrease in soil pH of 0.63 units over the

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past two decades [Yang et al., 2012b]; (3) The frequency and intensity of extreme

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precipitation has increased across northern China [Zhai et al., 2005], specifically,

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precipitation in the desert region of China has increased more than 40% since the

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beginning of the 21st century [Xu et al., 2010]. Third, soil physical and chemical

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properties during the 1980s were well documented in historical soil inventory data

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[Office of Soil Census of Inner Mongolia, 1994; Tibet Land Management Bureau,

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1994; Office of Agriculture Resource Layouting of Qinghai, 1997]. Together, these

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geographic conditions, environmental changes, and historical inventory make possible

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an investigation of the temporal dynamics of CEC over the last three decades.

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In this study, we investigated changes in CEC across northern China’s grasslands 5

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from the 1980s to 2010s. We collected surface soil samples from 251 sites on the

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Tibetan Plateau and Inner Mongolian Plateau from 2013-2015, comparing these data

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with 125 soil profiles measured during the 1980s, obtained from China’s Second

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National Soil Inventory. We combined these two datasets of CEC with the related

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environmental parameters, and used Random Forest method to explore the changes of

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CEC between the two periods. We hypothesized that topsoil CEC across northern

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China’s grasslands would show a significant decrease due to the substantial

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environmental changes over the past three decades.

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2. Materials and Methods

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2.1. Study Area

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Our study area mainly contained alpine grasslands on the Tibetan Plateau and

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temperate grasslands on the Inner Mongolian Plateau (Figure 1), whose longitudes

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and latitudes are 80.10°-121.93° E and 28.07°-49.62° N, respectively. During the

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2010s, the mean annual temperature (MAT) varied from -3.06 °C to 8.13 °C, and the

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mean annual precipitation (MAP) varied from 111.51 mm to 719.28 mm. The

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dominant grassland types are alpine meadow (AM), alpine steppe (AS), meadow

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steppe (MS), typical steppe (TS) and desert steppe (DS) [Chinese Academy of

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Sciences, 2001]. The dominant species are Kobresia pygmaea and K. tibetica in the

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AM, Stipa purpurea and Festuca ovina in the AS, S. baicalensis and Leymus

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chinensis in the MS, S. grandis and S. krylovii in the TS, and S. klemenzii, and S.

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breviflora in the DS [Ma et al., 2010]. According to the classification of the World 6

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Reference Base for Soil Resources 2006, the soil types related to the five grassland

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types were defined as Cambisols (in AM and AS), Chernozems (in MS), Kastanozems

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(in TS) and Calcisols (in DS) [IUSS Working Group WRB, 2007].

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2.2. Historical observations during the 1980s

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China’s Second National Soil Inventory during 1979-1989 recorded 74 and 51

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representative grassland soil profiles on the Tibetan Plateau and Inner Mongolian

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Plateau, respectively (Figure 1). From the 1980s data, we obtained CEC and other

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edaphic properties (e.g. SOC content, pH, and silt content) in the A horizon of 125

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typical profiles, with a median depth of 15 cm [Office of Soil Census of Inner

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Mongolia, 1994; Tibet Land Management Bureau, 1994; Office of Agriculture

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Resource Layouting of Qinghai, 1997]. According to the historical records, CEC was

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determined by hydrochloric acid neutralization titration, after extraction with a mixed

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solution consisting of 1 M ammonium acetate and 0.005 M EDTA (adjusted to pH 7.0

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for acidic soil and pH 8.5 for alkaline soil, respectively), and subsequent distillation of

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ammonium [Chang, 1979; Sparks et al., 1996]. SOC content was determined by

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ferrous sulfate titration, after potassium dichromate oxidation [Nelson and Sommers,

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1982]. Soil pH was measured by potentiometry, using a water soil ratio of 2.5:1

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[Koncki et al., 1992]. Silt content (soil texture) was measured by the sieve-hydrometer

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method, based on the speed of particle sedimentation at multiple time steps [Di

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Stefano et al., 2010]. Grassland types were determined by the dominant species

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related to the local soil types. Meanwhile, based on the available detailed 7

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administrative address, longitude and latitude at these sites were determined from a

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digital map developed by the National Administration of Surveying, Mapping, and

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Geoinformation (http://map.tianditu.com/map/index.html). Historical MAT and MAP

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at these sites were extracted using Kriging interpolation based on atmospheric

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temperature and precipitation data from 120 meteorological stations on the Tibetan

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Plateau and the Inner Mongolian Plateau from 1982-1989 (http://data.cma.cn/).

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2.3. Field inventory during the 2010s

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To characterize the current status of grassland soil CEC, we investigated 251 sites

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(173 on the Tibetan Plateau [Ding et al., 2016] and 78 on the Inner Mongolian Plateau)

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throughout northern China’s grasslands during July and August of 2013-2015. The

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sampling sites during the 2010s covered extensive climatic gradients and major

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grassland types (Figure 1). Specifically, latitude and longitude were acquired using

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GPS (M-GIS T20, China), and grassland types were determined by the dominant

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species present. At each site, we set up five 1 m×1 m quadrats at every corner and in

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the center of a 10 m×10 m square. We further selected three quadrats along a

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diagonal line of the square to take topsoil (0-10 cm) samples. Soil samples were

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collected in plastic bags and sent to the laboratory.

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We analyzed soil CEC and other edaphic properties in the laboratory. First, composite

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soil samples (equally mixed subsamples from three samples from each site) were

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air-dried and sieved (< 2 mm), and fine roots were manually removed. To avoid the 8

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potential uncertainties caused by measuring method, soil CEC, SOC content, pH, and

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silt content were all measured by the same methods as in the 1980s. MAT and MAP at

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these sites were extracted in the same method but from meteorological data of

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2010-2015.

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2.4. Data Analyses

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To examine the dynamics of soil CEC from 1980s to 2010s, it is necessary to have

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time-paired data for each sampling site. This method has been used to examine soil

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organic [Yu et al., 2009; Yang et al., 2014] and inorganic carbon [Yang et al., 2012a]

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dynamics, as well as soil pH changes [Yang et al., 2012b]. Because the site locations

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in the two sampling periods were inconsistent, we used the Random Forest model to

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estimate CEC values for the missing points from each sampling period. Random

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Forest extends Classification and Regression Tree analyses to improve prediction

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accuracy without excessive computation [Breiman, 2001a, 2001b; Wiesmeier et al.,

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2011; Ziegler and Konig, 2014; Were et al., 2015]. To reduce the uncertainties caused

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by the uneven distribution of sampling sites during the two periods, we selected a

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grassland transect (including 91 among the 125 sites in the 1980s and 189 among the

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251 sites in the 2010s) from the central-southern Tibetan Plateau to the eastern Inner

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Mongolian Plateau (Figure 1) to examine the CEC dynamics. Briefly, the dataset of

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measured CEC was split into two parts: a training dataset used to construct the

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Random Forest model (90% of the data points) and a test dataset used to assess the

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performance of the model (10% of the data points). Based on the historical soil 9

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inventory training dataset from the 1980s and related environmental variables (i.e.,

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longitude, latitude, grassland type, MAT, MAP, SOC, pH, and silt content), we

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constructed a Random Forest model to predict CEC for those sites without

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measurement in the 1980s (189 sites that surveyed during the 2010s). By doing so, we

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obtained a dataset consisting of 189 data points including measured soil CEC from the

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2010s and corresponding estimated data for the 1980s (detailed procedure in Figure 2).

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Similarly, using the training dataset from the current survey during the 2010s and the

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corresponding environmental data, we created another Random Forest model to

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simulate CEC during the 2010s for those sites without measurements from the 2010s

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(91 sites measured in the 1980s). Thereafter, we obtained another dataset consisting of

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91 data points including measured CEC from 1980s and estimated data from the

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2010s (Figure 2). By combining these data sets, we finally generated a time-paired

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dataset across the 280 sites with either measured or estimated data for the two

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sampling periods for each sampling site. We used Pearson correlation coefficient (r)

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and root-mean-squared error (RMSE) to test the accuracy of the model predictions.

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We repeated the Random Forest training 1000 times, and the averages were treated as

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the predicted CEC values. All the Random Forest Model computations were

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conducted using R software [R Development Core Team, 2012]. Based on the

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time-paired dataset obtained through the Random Forest method, we used a paired

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t-test to explore the temporal dynamics of CEC. We also examined the relationships

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between CEC dynamics and environmental variables using linear regression model.

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All the statistical analyses were performed using SPSS 20.0 (IBM SPSS, Chicago, 10

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Illinois, USA), with a significance level of 0.05.

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3. Results

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3.1. Reliable Estimation by the Random Forest Model

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To test the accuracy of the model predictions, we examined the correlation between

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the predicted and measured CEC values. Our results showed that during the 1980s, r

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and RMSE between the testing data set and the measured values were 0.90 and 5.41

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cmol kg-1, respectively (Figure 3a). Similarly, r and RMSE during the 2010s were

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0.93 and 3.48 cmol kg-1, respectively (Figure 3b). These results showed high

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correlation and low estimation error, indicating a reliable prediction ability for the

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Random Forest model.

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3.2. Temporal Dynamics of CEC across Northern China’s Grasslands

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Topsoil CEC across northern China’s grasslands during the 2010s was significantly

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lower than the 1980s (P < 0.001; Figure 4a). According to the results of 280 sampling

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sites within the transect, topsoil CEC significantly declined by about 14% (Table 1).

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However, substantial variations existed among the various grassland types (i.e., AM,

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AS, MS, TS and DS). Topsoil CEC did not exhibit a significant change in DS perhaps

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because of its lower initial values, but decreased by 11%, 20%, 27% and 9% in AM,

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AS, MS and TS, respectively (Figure 4b; Table 1).

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4. Discussion

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There has been significant loss of topsoil CEC across northern China’s grasslands 11

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over the last three decades. A decline in base cations over a longer time period was

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also detected in North America [Watmough and Dillon, 2003], Europe [Högberg et al.,

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2006], Oceania [Hartemink, 1998; Curtin et al., 2015], Africa [Jaiyeoba, 2003] and

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Asia [Ali et al., 1997; Lu et al., 2015], indicating that this phenomenon is widespread.

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The decrease of CEC observed in this study is within the range of previous studies,

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varying from 8% to 50% [Ali et al., 1997; Hartemink, 1998; Jaiyeoba, 2003; Curtin et

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al., 2015]. Such a pattern could be due to the following four phenomena. First, a

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reduction in SOC content can lead to a decrease in CEC. Overgrazing and cultivation

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in past decades [Bridges and Oldeman, 1999; Yang et al., 2007; Yu et al., 2012; Mu et

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al., 2013] has increasingly degraded grasslands [Dai et al., 2011] and causing declines

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in grassland productivity [Babel et al., 2014]. Consequently, the SOC stock of China’s

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grasslands decreased by 3.56 Pg C (1 Pg = 1015 g), dropping from approximately

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37.71 to 34.15 Pg C from the 1980s to the 2000s, with particularly marked changes in

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Tibet, Qinghai, and Inner Mongolia [Xie et al., 2007]. The decrease in SOC has a

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profound influence on CEC for two reasons. One is that the dissociation of some

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functional groups (e.g., carboxyl and phenolic hydroxyl) decreases with the decrease

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in SOC. As a result, the amount of negative charge possessed by humus is reduced,

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weakening the complexation of the humus with exchangeable base cations [Brady and

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Weil, 2002; Chapin et al., 2011]. The other is that cation-bridging (e.g., Ca2+, Fe3+,

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Al3+) between clay minerals and humus can decline due to a reduction in SOC,

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making humus-clay mineral complexes unstable [Brady and Weil, 2002; von Lutzow

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et al., 2006; Mueller et al., 2012]. Consequently, the protection for SOM against 12

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decomposition is weakened and causes the decrease in CEC. Consistent with this

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inference, our results showed that CEC was positively correlated with SOC, and its

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variations were mostly explained by SOC in both the 1980s (TP: r2 = 0.72, P < 0.001;

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IM: r2 = 0.71, P < 0.001; Figure 5a) and 2010s (TP: r2 = 0.89, P < 0.001; IM: r2 =

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0.84, P < 0.001; Figure 5e). Many studies also have found a positive relationship

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between CEC and SOC [Tůma et al., 2011; Lu et al., 2014; Gruba and Mulder, 2015].

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Second, the CEC reduction can also be caused by soil desertification. It has been

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reported that soils in arid and semi-arid regions were more likely to be degraded

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[Bridges and Oldeman, 1999]. Both the Tibetan Plateau and Inner Mongolian Plateau,

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which are located in arid and semi-arid regions, have been suffering the effects of

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climate warming, human disturbance, overgrazing, cultivation, and pikas, all of which

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can result in soil degradation and desertification [Yang et al., 2007; Xue et al., 2009;

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Xue et al., 2011; Yu et al., 2012; Qin et al., 2015]. It has been reported that lots of

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Inner Mongolian grasslands have undergone degradation or desertification from 1985

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to 2000 [Mu et al., 2013]. Once soil desertification occurs, wind erosion alters soil

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texture, blowing away fine particles (e.g. clay and silt) and leaving coarse and barren

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sand [Chepil, 1957; Hoffmann et al., 2008; Amundson et al., 2015]. Moreover, with

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the decrease in particle contents, the surface areas provided for exchangeable cations

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become smaller [Brady and Weil, 2002]. Furthermore, soil desertification causes a

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decrease in soil productivity and SOC content [Larney et al., 1998; Yan et al., 2005].

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All of these processes will reduce CEC, along with soil desertification. Our data 13

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support the above conclusions, with CEC positively correlated with silt content

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(1980s TP: r2 = 0.12, P < 0.01; IM: r2 = 0.71, P < 0.001; Figure 5b; 2010s TP: r2 =

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0.25, P < 0.001; IM: r2 = 0.68, P < 0.001; Figure 5f), but negatively associated with

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sand content (1980s TP: r2 = 0.12, P < 0.01; IM: r2 = 0.62, P < 0.001; Figure 5c;

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2010s TP: r2 = 0.14, P < 0.001; IM: r2 = 0.67, P < 0.001; Figure 5g).

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Third, the CEC reduction can still be induced by soil acidification. It has been

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reported that enhanced atmospheric nitrogen deposition [Liu et al., 2013] and sulfur

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deposition [Zhao et al., 2009] resulted in soil acidification across China’s grasslands

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[Yang et al., 2012b]. Sustained acidification leads to a lower soil pH, which can be

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buffered by carbonate and base cations. It has been hypothesized that when the soil

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pH is over 7.5, the change is mainly buffered by carbonate [Bowman et al., 2008;

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Yang et al., 2012a, b], while the exchangeable base cations play a lesser role. When

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the soil pH is in the range of 4.5-7.5, it is primarily the exchangeable cations that

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buffer pH change [Bowman et al., 2008]. Exchangeable ions including Ca2+, Mg2+, K+

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and Na+ are adsorbed on the surface of the soil colloid, making CEC value higher.

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With the increases in acidity, H+ gradually occupies the exchangeable sites. At the

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same time, base cations disassociate and are more likely to be leached by rainfall,

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resulting in decreased CEC [Watmough and Dillon, 2003; Lapenis et al., 2004; Lieb et

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al., 2011; Chen et al., 2013; Lu et al., 2014]. This classic hypothesis is supported by

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experimental studies on long-term dynamics of CEC and base cations at the local

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scale [Watmough and Dillon, 2003; Högberg et al., 2006; Lu et al., 2015]. To be 14

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specific, results from a manipulative experiment also showed that soil pH and base

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saturation significantly increased when acid addition stopped [Högberg et al., 2006].

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Therefore, we argue that acidification mainly affects CEC when the pH was low (e.g.

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in AM and MS).

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Fourth, the decrease in CEC is possibly related to extreme precipitation. In recent

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decades, extreme precipitation has been observed more frequently [IPCC, 2007;

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Moldan et al., 2012]. Recent studies indicated that the frequency and intensity of

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extreme precipitation were gradually increasing across western China and the Tibetan

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Plateau [Zhai et al., 2005; Yang et al., 2008; Xu et al., 2010], making soil base cations

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leached. Normally, precipitation is positively correlated with SOC [Ruiz Sinoga et al.,

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2012], which is strongly associated with CEC [Gruba and Mulder, 2015]. Normally,

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precipitation is positively correlated with SOC [Ruiz Sinoga et al., 2012], which is

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strongly associated with CEC [Gruba and Mulder, 2015]. In both periods, our results

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showed that CEC increased along the precipitation gradient (1980s TP: r2 = 0.12, P
Mg2+ > K+ > Na+, and so, if CEC is reduced, base cations

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will be lost in different degrees. As a result, species that rely on a certain cation may

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decrease or even become extinct on long time scales. In addition, in alpine meadow

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and meadow steppe environments with a low pH, soil acidification caused by nitrogen

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and sulfur deposition leads to the decrease in CEC, eventually releasing Al3+ and Fe3+

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into the soil [Bowman et al., 2008]. Consequently, species that are sensitive to Al3+ or

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Fe3+ may decrease. Second, ecosystem function across northern China’s grasslands

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may be weakened when CEC is reduced. The decrease in CEC and the increase in

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Al3+ and Fe3+ weaken the abilities of the nutrient (i.e., Ca2+, Mg2+, K+ and Na+) supply

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and plant tolerance to drought, frost or pathogen attacks [DeHayes et al., 1999;

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Demchik and Sharpe, 2000]. Thus, to some degree, it may potentially decrease

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vegetation productivity [Likens et al., 1998; McLaughlin and Wimmer, 1999]. To

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avoid the ecological consequences mentioned above, it is vital to reduce the

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interference of human activities in these ecosystems and strictly control the intensity 16

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of grazing and cultivation [Mu et al., 2013]. Reducing the emissions of SO2 and NOx

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is also essential to relieve soil acidification and environment aggravation caused by

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continuous CEC loss [Lucas et al., 2011].

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Acknowledgments

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Associated data are available in supporting information. We are grateful to the

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members of the IBCAS Sampling Campaign Teams for their assistance in field

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investigations. This work was supported by the National Natural Science Foundation

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of China (31670482 and 41371213), Key Research Program of Frontier Sciences,

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Chinese Academy of Sciences (QYZDB-SSW-SMC049), Chinese Academy of

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Sciences-Peking University Pioneer Cooperation Team, and the Thousand Young

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Talents Program.

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Table 1. Means of CEC and its changes among five grassland types within a grassland

609

transect across northern China from the 1980s-2010s. The values in

610

parenthesis show the 95% confidence interval calculated using the Random

611

Forest Model. Notes: n, sample size; AM, alpine meadow; AS, alpine steppe;

612

MS, meadow steppe; TS, typical steppe; DS, desert steppe. CEC during the 1980s

CEC during the 2010s

Change in CEC

(cmol kg-1)

(cmol kg-1)

(cmol kg-1)

Grassland type

n

Total

280

17.06 [16.04, 18.08] 14.69 [13.76, 15.61]

-2.37 [-2.97, -1.78]

AM

97

22.57 [20.92, 24.23] 20.13 [18.35, 21.91]

-2.45 [-3.71, -1.18]

AS

90

13.84 [12.30, 15.39]

11.07 [9.85, 12.29]

-2.77 [-3.60, -1.94]

MS

16

26.41 [22.14, 30.67] 19.39 [17.24, 21.55]

-7.01 [-10.32, -3.70]

TS

52

13.44 [12.19, 14.69] 12.22 [11.26, 13.18]

-1.22 [-2.17, -0.26]

DS

25

8.83 [7.99, 9.66]

8.72 [7.26, 10.18]

-0.11 [-1.54, 1.32]

613

29

614

Figure Legends

615

Figure 1. The distribution of sampling sites during the 1980s and the 2010s. The

616

background colors show grassland extent and type across northern China

617

[Chinese Academy of Sciences, 2001]. The rectangle represents the

618

grassland transect selected for exploring the temporal dynamics of CEC.

619

Blue points represent the sites during the 1980s and green points represent

620

the 2010s.

621 622 623

Figure 2. Procedure of constructing Random Forest (RF) models to predict data for those sampling sites during the 2000s (a) and during the 1980s (b).

624 625 626

Figure 3. Comparison between predicted and measured CEC for the period of (a) the 1980s and (b) the 2010s.

627 628

Figure 4. Changes in CEC (a) along a grassland transect across northern China’s

629

grasslands and (b) among various grassland types between the 1980s and

630

the 2010s. The inset in panel (a) shows the comparison of CEC between

631

the two periods, with a 1:1 line for reference. Different letters above

632

box-and-whisker plots express significant differences between the two

633

periods. The gap and square, lower and upper edges and bars in the boxes

634

represent median and mean values, 25th and 75th percentiles, and standard

635

deviations of all data, respectively. AM: alpine meadow, AS: alpine steppe, 30

636

MS: meadow steppe, TS: typical steppe, DS: desert steppe.

637 638

Figure 5. Relationships of CEC with (a, e) SOC, (b, f) silt content, (c, g) sand content,

639

and (d, h) MAP. Panels (a-d) and (e-h) illustrate the relationships between

640

CEC and environmental factors during the 1980s and the 2010s,

641

respectively. Data and trends on the Tibetan Plateau are represented by red

642

points and lines, while those on the Inner Mongolian Plateau are shown by

643

blue points and lines.

31

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.