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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2023.1067552</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Spatiotemporal dynamic of subtropical forest carbon storage and its resistance and resilience to drought in China</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Yan</surname>
<given-names>Mengjie</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Mao</surname>
<given-names>Fangjie</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1918527"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Du</surname>
<given-names>Huaqiang</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/503093"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Li</surname>
<given-names>Xuejian</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Chen</surname>
<given-names>Qi</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ni</surname>
<given-names>Chi</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Huang</surname>
<given-names>Zihao</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Xu</surname>
<given-names>Yanxin</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Gong</surname>
<given-names>Yulin</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Guo</surname>
<given-names>Keruo</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Sun</surname>
<given-names>Jiaqian</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Xu</surname>
<given-names>Cenheng</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>State Key Laboratory of Subtropical Silviculture, Zhejiang Agricultural &amp; Forestry (A &amp; F) University</institution>, <addr-line>Hangzhou</addr-line>, &#xa0;<country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Key Laboratory of Carbon Cycling in Forest Ecosystems and Carbon Sequestration of Zhejiang Province, Zhejiang A &amp; F University</institution>, <addr-line>Hangzhou</addr-line>, &#xa0;<country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>School of Environmental and Resources Science, Zhejiang A &amp; F University</institution>, <addr-line>Hangzhou</addr-line>, &#xa0;<country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Andreia Michelle Smith-Moritz, University of California, Davis, United States</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Tanzeel Ja Farooqi, Yibin University, China; Chengcheng Gang, Institute of Soil and Water Conservation (CAS), China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Fangjie Mao, <email xlink:href="mailto:mfangjie@gmail.com">mfangjie@gmail.com</email>
</p>
</fn>
<fn fn-type="other" id="fn002">
<p>This article was submitted to Plant Biophysics and Modeling, a section of the journal Frontiers in Plant Science</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>17</day>
<month>01</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>14</volume>
<elocation-id>1067552</elocation-id>
<history>
<date date-type="received">
<day>12</day>
<month>10</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>03</day>
<month>01</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Yan, Mao, Du, Li, Chen, Ni, Huang, Xu, Gong, Guo, Sun and Xu</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Yan, Mao, Du, Li, Chen, Ni, Huang, Xu, Gong, Guo, Sun and Xu</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Subtropical forests are rich in vegetation and have high photosynthetic capacity. China is an important area for the distribution of subtropical forests, evergreen broadleaf forests (EBFs) and evergreen needleleaf forests (ENFs) are two typical vegetation types in subtropical China. Forest carbon storage is an important indicator for measuring the basic characteristics of forest ecosystems and is of great significance for maintaining the global carbon balance. Drought can affect forest activity and may even lead to forest death and the stability characteristics of different forest ecosystems varied after drought events. Therefore, this study used meteorological data to simulate the standardized precipitation evapotranspiration index (SPEI) and the Biome-BGC model to simulate two types of forest carbon storage to quantify the resistance and resilience of EBF and ENF to drought in the subtropical region of China. The results show that: 1) from 1952 to 2019, the interannual drought in subtropical China showed an increasing trend, with five extreme droughts recorded, of which 2011 was the most severe one; 2) the simulated average carbon storage of the EBF and ENF during 1985-2019 were 130.58 t&#xb7;hm<sup>-2</sup> and 78.49 t&#xb7;hm<sup>-2</sup>, respectively. The regions with higher carbon storage of EBF were mainly concentrated in central and southeastern subtropics, where those of ENF mainly distributed in the western subtropic; 3) The median of resistance of EBF was three times higher than that of ENF, indicating the EBF have stronger resistance to extreme drought than ENF. Moreover, the resilience of two typical forest to 2011 extreme drought and the continuous drought events during 2009 - 2011 were similar. The results provided a scientific basis for the response of subtropical forests to drought, and indicating that improve stand quality or expand the plantation of EBF may enhance the resistance to drought in subtropical China, which provided certain reference for forest protection and management under the increasing frequency of drought events in the future.</p>
</abstract>
<kwd-group>
<kwd>drought</kwd>
<kwd>carbon storage</kwd>
<kwd>resistance</kwd>
<kwd>resilience</kwd>
<kwd>subtropical China</kwd>
<kwd>forest ecosystem</kwd>
</kwd-group>
<contract-num rid="cn001">No. 31901310, 32171785, 32201553, 31670644, U1809208</contract-num>
<contract-sponsor id="cn001">National Natural Science Foundation of China<named-content content-type="fundref-id">10.13039/501100001809</named-content>
</contract-sponsor>
<counts>
<fig-count count="11"/>
<table-count count="5"/>
<equation-count count="15"/>
<ref-count count="123"/>
<page-count count="18"/>
<word-count count="8017"/>
</counts>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title>1 Introduction</title>
<p>Carbon storage generally refers to the storage of carbon elements in each carbon pool of the forest ecosystem at a certain point in time, and is the result of years of accumulation in the forest ecosystem (<xref ref-type="bibr" rid="B79">Sun and Liu, 2020</xref>). It is not only an important indicator that reflects the basic characteristics of the forest ecological environment (<xref ref-type="bibr" rid="B15">Fang et&#xa0;al., 2014</xref>) but also a theoretical basis for evaluating forest structure, function, and production potential (<xref ref-type="bibr" rid="B4">Bonan, 2008</xref>; <xref ref-type="bibr" rid="B57">Mitchard, 2018</xref>). However, frequent drought induced by climate change greatly affects the carbon sequestration process of forest ecosystem (<xref ref-type="bibr" rid="B71">Schwalm et&#xa0;al., 2012</xref>).</p>
<p>At present, the physical process underlying drought impacts on forests has been well studied. In general, drought slows down forest activities and affects forest stability by reducing forest productivity. Extreme drought directly or indirectly affects forest GPP (<xref ref-type="bibr" rid="B103">Xu et&#xa0;al., 2019</xref>) and terrestrial carbon sinks (<xref ref-type="bibr" rid="B38">Jung et&#xa0;al., 2017</xref>), and may even lead to forest death (<xref ref-type="bibr" rid="B61">Park Williams et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B24">Hartmann et&#xa0;al., 2018</xref>). However, the response of terrestrial ecosystems to drought is one of the largest uncertainties in the carbon cycle (<xref ref-type="bibr" rid="B68">Reichstein et&#xa0;al., 2013</xref>) and is not well represented in current climate-vegetation models (<xref ref-type="bibr" rid="B2">Anderegg et&#xa0;al., 2015</xref>). Therefore, analyzing how forests respond and adapt to drought has become a focal issue in the study of extreme events in the context of climate change. Drought monitoring at the spatial scale is generally evaluated by drought indices (<xref ref-type="bibr" rid="B88">Vicente-Serrano et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B18">Gobena and Gan, 2013</xref>; <xref ref-type="bibr" rid="B6">Center et&#xa0;al., 2017</xref>). Among them, the Standardized Precipitation Evapotranspiration Index (SPEI) characterizes the degree of deviation of a region&#x2019;s dry and wet conditions from the normal year by standardizing the cumulative probability value of the difference between potential evapotranspiration (PET) and precipitation (<xref ref-type="bibr" rid="B101">Xia et&#xa0;al., 2019</xref>), and has been widely used in drought detection research around the world (<xref ref-type="bibr" rid="B90">Vicente-Serrano and Trigo, 2011</xref>; <xref ref-type="bibr" rid="B122">Zhuang et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B42">Lei et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B101">Xia et&#xa0;al., 2019</xref>).</p>
<p>The effects of drought on ecosystem stability can be expressed using ecosystem resistance and resilience (<xref ref-type="bibr" rid="B84">Tilman and Downing, 1994</xref>; <xref ref-type="bibr" rid="B10">De Keersmaecker et&#xa0;al., 2014</xref>), which are two factors that fully consider the immediate and legacy effects of drought on forest ecosystems (<xref ref-type="bibr" rid="B33">Ivits et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B64">Pennekamp et&#xa0;al., 2018</xref>). Resistance expressed as the ability of the ecosystem to maintain its original state under disturbance (<xref ref-type="bibr" rid="B52">Lloret et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B86">Van Ruijven and Berendse, 2010</xref>), and resilience represents the ability of the ecosystem to recover to a normal state from the disturbance (<xref ref-type="bibr" rid="B27">Holling, 1973</xref>). Scholars have previously analyzed the resistance and resilience of species at the biome level based on the perspective of experiments and modeling (<xref ref-type="bibr" rid="B29">Hoover et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B11">De Keersmaecker et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B17">Gazol et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B50">Li et&#xa0;al., 2018</xref>). <xref ref-type="bibr" rid="B32">Isbell et&#xa0;al. (2015)</xref> defined a dimensionless measure of resistance and resilience of grassland ecosystems by measuring the productivity of grassland systems in North America. <xref ref-type="bibr" rid="B30">Huang and Xia (2019)</xref> directly quantified the resistance and resilience of global ecosystems to drought by measuring the ecosystem function change. However, few studies have quantified forest resistance and resilience to drought by measuring changes in forest carbon pool function during and after drought, and the response mechanism of forest carbon pool change to drought has not been clarified.</p>
<p>Forest carbon storage estimation methods commonly include plot survey, remote sensing inversion, and ecosystem model. The sample plot survey seemed as the most accurate method to measure the carbon storage of forest ecosystems, while considerable amount of time costs, manpower, material resources limited restricted its use (<xref ref-type="bibr" rid="B55">Mickler et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B7">Chave et&#xa0;al., 2003</xref>). The remote sensing inversion could obtain vegetation carbon storage in real time and on a large scale. However, due to technical limitations and the lack of description of plant physiology, the estimation of underground carbon storage still remains large uncertainty (<xref ref-type="bibr" rid="B79">Sun and Liu, 2020</xref>). Ecosystem models can mechanistically describe key carbon fixation processes, which is suitable for large-scale research. At present, a large number of carbon storage estimation models have been developed. Among them, the Biome-BGC model is a typical ecosystem process model that can simulate physiological and ecological processes, such as photosynthesis, respiration, and decomposition of ecosystems at different scales, and it is widely used worldwide (<xref ref-type="bibr" rid="B69">Running, 1993</xref>; <xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B99">Wu et&#xa0;al., 2014</xref>).</p>
<p>The subtropical forest ecosystems in the East Asian monsoon region have a net ecosystem productivity of 0.72 Pg C&#xb7;a<sup>-1</sup>, indicating them play a non-negligible role for mitigating global warming (<xref ref-type="bibr" rid="B107">Yu et&#xa0;al., 2014</xref>). China is an important distribution area of subtropical forests in East Asia (<xref ref-type="bibr" rid="B120">Zhou et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B106">Yan et&#xa0;al., 2006</xref>), in which evergreen broadleaf forest (EBF) and evergreen needleleaf forest (ENF) are two most widely distributed forest types with great carbon sink potential (<xref ref-type="bibr" rid="B25">He et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B54">Mao et&#xa0;al., 2022</xref>). However, drought happened recent years significantly affects the carbon balance of EBF and ENF ecosystems, such as the extreme drought in 2003 caused a 55% annual NEP decline in the planted ENF of QianYanZhou (<xref ref-type="bibr" rid="B22">Gu et&#xa0;al., 2008</xref>), and <xref ref-type="bibr" rid="B76">Song et&#xa0;al. (2017)</xref> found the water use efficiency (WUE) greatly increased in the driest year (2009) due to a larger decline in evapotranspiration than gross primary productivity. In addition, due to the complex climatic conditions (<xref ref-type="bibr" rid="B23">Hanson and Weltzin, 2000</xref>; <xref ref-type="bibr" rid="B102">Xie et&#xa0;al., 2015</xref>), different in drought intensities and duration (<xref ref-type="bibr" rid="B59">Niu et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B89">Vicente-Serrano et&#xa0;al., 2014</xref>), and various in vegetation physiological (<xref ref-type="bibr" rid="B49">Liu et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B45">Li et&#xa0;al., 2020</xref>), the impacts of drought on EBF and ENF in subtropical China may have spatial heterogeneous.</p>
<p>This study takes EBF and ENF as the research object, calculated the SPEI in subtropical China, and analyzed the spatiotemporal characteristics of subtropical extreme drought from 1952 to 2019. Then, the driven datasets of Biome-BGC model were collected, and the spatiotemporal evolution trend of vegetation carbon storage of EBF and ENF from 1985 to 2019 were simulated. Finally, extreme droughts and continuous droughts were extracted, and the resistance and resilience of subtropical forests to extreme droughts were analyzed based on the simulated carbon storage. The findings of this study can provide a scientific basis for enhancing the conservation and management of subtropical forests in the context of future global warming, and it has certain enlightenment significance for the forest to resist extreme drought.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<title>2 Materials and methods</title>
<sec id="s2_1">
<title>2.1 Study area</title>
<p>The study area is the entire subtropical region of China, which is located to the south of the Qinling Mountains and Huaihe River, north of Leizhou Peninsula, and east of the Hengduan Mountains (22&#xb0; -34&#xb0;N, 98&#xb0; -122&#xb0;E). The terrain is low in the west and high in the east. China&#x2019;s subtropics belong to the east coast humid monsoon area, and the region is the warmest and hottest compared to the same latitude, except for desert areas. Moreover, the rainfall in the study region is far more abundant than that in the same latitude worldwide (<xref ref-type="bibr" rid="B105">Yang et&#xa0;al., 2006</xref>). The average annual temperature ranges from -1 to 24&#xb0;C, and the average annual precipitation ranges from 450 to 2125mm. China has preserved the best subtropical evergreen forest ecosystem which is the main component of China&#x2019;s subtropical forests (<xref ref-type="bibr" rid="B44">Lin et&#xa0;al., 2020</xref>) and accounts for approximately 25% of China&#x2019;s land area. Dominant families in such ecosystems are the <italic>Cyclobalanopsis Oerst</italic>, <italic>Castanopsis Spach</italic>, and <italic>Lithocarpus</italic> of <italic>Fagaceae</italic>.</p>
</sec>
<sec id="s2_2">
<title>2.2 Data acquisition and processing</title>
<sec id="s2_2_1">
<title>2.2.1 Meteorological data</title>
<p>Meteorological data for the study area from 1952 to 2019, including the daily maximum temperature, minimum temperature, solar radiation, precipitation, relative humidity, and average wind speed, were obtained from the National Meteorological Information Center of the China Meteorological Administration (<uri xlink:href="http://data.cma.cn">http://data.cma.cn</uri>). The processing steps for the meteorological data were as follows: 1.&#xa0;a spatial resolution of 1&#xa0;km was used to interpolate meteorological data from 824 meteorological stations using the inverse distance weight method; 2. temperature was corrected based on altitude, assuming a temperature drop rate of 6.5&#xb0;C&#xb7;km<sup>-1</sup> (<xref ref-type="bibr" rid="B5">Cao et&#xa0;al., 2017</xref>); Then, solar radiation was simulated using the method of <xref ref-type="bibr" rid="B37">Ju et&#xa0;al. (2006)</xref> according to the sunshine duration of each station. 3. meteorological data were obtained for the subtropical region in China through extraction by mask. The average monthly data can be obtained by averaging and summing the corresponding daily scale data (as shown in <xref ref-type="fig" rid="f1">
<bold>Figures&#xa0;1D, E</bold>
</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Extent of Subtropical China and related datasets used in this study <bold>(A)</bold> elevation; <bold>(B)</bold> abundance of EBF; <bold>(C)</bold> ENF abundance of ENF; <bold>(D)</bold> annual average temperature; and <bold>(E)</bold> average precipitation).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g001.tif"/>
</fig>
</sec>
<sec id="s2_2_2">
<title>2.2.2 Elevation data</title>
<p>The elevation data (as shown in <xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1A</bold>
</xref>) used here are from the ASTER Global Digital Elevation Model version 3 (<xref ref-type="bibr" rid="B16">Fujisada et&#xa0;al., 2005</xref>), and the spatial resolution of the data is 1&#xb0;, thus, re-projection of the data must be performed before use. The geographic coordinates of the data were converted to projection coordinates, and the data were resampled to 1&#xa0;km. Finally, elevation data of the subtropical region of China were obtained by mask clipping.</p>
</sec>
<sec id="s2_2_3">
<title>2.2.3 Soil data</title>
<p>Subtropical soil data were obtained by Harmonized World Soil Database ver. 1.2 (<uri xlink:href="http://iiasa.acat/Research/LUC/luc07/External-World-soil-database">http://iiasa.acat/Research/LUC/luc07/External-World-soil-database</uri>) (<xref ref-type="bibr" rid="B96">Wieder et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B100">Xianyong and Hao, 2018</xref>), which has a spatial resolution of 0.05<sup>&#xb0;</sup>. The original data needed to be reprojected and resampled to a resolution of 1&#xa0;km, and then the soil texture data in the subtropical region of China could be obtained by mask cutting. Finally, data on the clay, sand, and silt particles were extracted according to field values to obtain the percentage of data subtropical clay, sand, and silt in the study area (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>The percentage of clay <bold>(A)</bold>, silt <bold>(B)</bold>, and sand <bold>(C)</bold> of subtropical China.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g002.tif"/>
</fig>
</sec>
<sec id="s2_2_4">
<title>2.2.4 Subtropical forest abundance data</title>
<p>In this study, the abundance data of subtropical EBF and ENF were used to simulate carbon storage, and these data were obtained from the Global 30&#xa0;m Fine Land Cover Dynamic Monitoring Product from 1985 to 2020 (<xref ref-type="bibr" rid="B115">Zhang et&#xa0;al., 2021</xref>), which were released by the Space Information Innovation Institute of the Chinese Academy of Sciences. This dataset uses full-time series Landsat satellite data and relies on the Google Earth Engine cloud computing platform to achieve a global annual land cover map containing more than 30 land cover types. Before simulating carbon storage, the data were first trimmed to the subtropical region of China, and then the data for the two forest types were extracted according to the fields values. Finally, the forest distribution data at 30&#xa0;m resolution were linear upscaled to achieve the 1&#xa0;km fraction data using the local average method (<xref ref-type="bibr" rid="B72">Shang et&#xa0;al., 2013</xref>) (<xref ref-type="fig" rid="f1">
<bold>Figures&#xa0;1B, C</bold>
</xref>). Because the time interval for land-use type data was five years, the forest abundance in years that were unclassified at the time of simulation was replaced with stand data from neighboring years.</p>
</sec>
</sec>
<sec id="s2_3">
<title>2.3 Spatial and temporal patterns and analysis of subtropical drought based on the SPEI</title>
<p>In this study, the SPEI were calculated based on soil heat flux, meteorological data, saturated water vapor pressure, sunshine duration, wind speed, and latitude. The calculation process of the SPEI based on this algorithm is as follows (<xref ref-type="bibr" rid="B123">Zotarelli et&#xa0;al., 2010</xref>).</p>
<p>1). The core of the SPEI algorithm is the calculation for PET. The Penman-Monteith (P-M) formula considered heat and aerodynamics factors, and is more consistent with the measured evapotranspiration (<xref ref-type="bibr" rid="B63">Penman, 1948</xref>; <xref ref-type="bibr" rid="B35">Jensen et&#xa0;al., 1990</xref>). The P-M formula is as follows:</p>
<disp-formula>
<label>(1)</label>
<mml:math display="block" id="M1">
<mml:mrow>
<mml:mi>P</mml:mi>
<mml:mi>E</mml:mi>
<mml:mi>T</mml:mi>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mn>0.408</mml:mn>
<mml:mo stretchy="false">(</mml:mo>
<mml:msub>
<mml:mi>R</mml:mi>
<mml:mi>s</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mi>G</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>+</mml:mo>
<mml:mi>&#x3b3;</mml:mi>
<mml:mfrac>
<mml:mrow>
<mml:mn>900</mml:mn>
</mml:mrow>
<mml:mrow>
<mml:mi>T</mml:mi>
<mml:mo>+</mml:mo>
<mml:mn>273</mml:mn>
</mml:mrow>
</mml:mfrac>
<mml:msub>
<mml:mi>u</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mo stretchy="false">(</mml:mo>
<mml:msub>
<mml:mi>e</mml:mi>
<mml:mi>s</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub>
<mml:mi>e</mml:mi>
<mml:mi>a</mml:mi>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mrow>
<mml:mi>&#x394;</mml:mi>
<mml:mo>+</mml:mo>
<mml:mi>&#x3b3;</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>+</mml:mo>
<mml:mn>0.34</mml:mn>
<mml:msub>
<mml:mi>u</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where R<sub>s</sub> is the daily average net radiation flux of the plant surface (MJ&#xb7;m<sup>-2</sup>&#xb7;day<sup>-1</sup>); G is the soil heat flux (MJ&#xb7;m<sup>-2</sup>&#xb7;day<sup>-1</sup>); &#x3b3; is the humidity constant (kPa&#xb7;&#xb0;C<sup>-1</sup>); T is the average temperature(&#xb0;C); U<sub>2</sub> is the average wind speed at 2&#xa0;m per day (m&#xb7;day<sup>-1</sup>); and e<sub>s</sub>, e<sub>a</sub> and &#x394; are the saturated water vapor pressure (kPa), actual water vapor pressure (kPa) and slope of the saturated water vapor pressure-temperature curve, respectively. R<sub>s</sub>, G, &#x3b3;, and other parameters were calculated using the FAO-56 P-M updating equation recommended by the FAO (<xref ref-type="bibr" rid="B1">Allen et&#xa0;al., 1998</xref>).</p>
<p>2). Calculate the measurement of deficit water balance (D<sub>i</sub>):</p>
<disp-formula>
<label>(2)</label>
<mml:math display="block" id="M2">
<mml:mrow>
<mml:msub>
<mml:mi>D</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>P</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mi>P</mml:mi>
<mml:mi>E</mml:mi>
<mml:msub>
<mml:mi>T</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</disp-formula>
<p>3). Create a series of cumulative water deficits at different time scales <inline-formula>
<mml:math display="inline" id="im1">
<mml:mrow>
<mml:msubsup>
<mml:mtext>D</mml:mtext>
<mml:mrow>
<mml:mtext>n</mml:mtext>
</mml:mrow>
<mml:mrow>
<mml:mtext>k</mml:mtext>
</mml:mrow>
</mml:msubsup>
</mml:mrow>
</mml:math>
</inline-formula>:</p>
<disp-formula>
<label>(3)</label>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:msubsup>
<mml:mi>D</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>k</mml:mi>
</mml:msubsup>
<mml:mo>=</mml:mo>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>0</mml:mn>
</mml:mrow>
<mml:mrow>
<mml:mi>k</mml:mi>
<mml:mo>-</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:munderover>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:msub>
<mml:mi>P</mml:mi>
<mml:mrow>
<mml:mi>n</mml:mi>
<mml:mo>-</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mi>P</mml:mi>
<mml:mi>E</mml:mi>
<mml:msub>
<mml:mi>T</mml:mi>
<mml:mrow>
<mml:mi>n</mml:mi>
<mml:mo>-</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mstyle>
<mml:mtext>&#x2009;</mml:mtext>
<mml:mi>n</mml:mi>
<mml:mo>&#x2265;</mml:mo>
<mml:mi>k</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where k is the time scale (generally on month) and n is the number of calculations.</p>
<p>4). Choose the log-logistic probability distribution (negative values can be interpreted and modeled with different shapes for the frequencies of the D series at different timescales) to normalize the D series:</p>
<disp-formula>
<label>(4)</label>
<mml:math display="block" id="M4">
<mml:mrow>
<mml:mi>F</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>x</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mrow>
<mml:mo>[</mml:mo>
<mml:mrow>
<mml:mn>1</mml:mn>
<mml:mo>+</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:mrow>
<mml:mfrac>
<mml:mi>&#x3b1;</mml:mi>
<mml:mrow>
<mml:mi>x</mml:mi>
<mml:mo>-</mml:mo>
<mml:mi>&#x3b3;</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo>)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mi>&#x3b2;</mml:mi>
</mml:msup>
</mml:mrow>
<mml:mo>]</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mrow>
<mml:mo>-</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:msup>
</mml:mrow>
</mml:math>
</disp-formula>
<p>In formula (4) the following parameters are defined:</p>
<disp-formula>
<label>(5)</label>
<mml:math display="block" id="M5">
<mml:mrow>
<mml:mi>&#x3b1;</mml:mi>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mn>2</mml:mn>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
<mml:mi>&#x3b2;</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>&#x393;</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>+</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>&#x3b2;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mi>&#x393;</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>-</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>&#x3b2;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(6)</label>
<mml:math display="block" id="M6">
<mml:mrow>
<mml:mi>&#x3b2;</mml:mi>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mn>2</mml:mn>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
</mml:mrow>
<mml:mrow>
<mml:mn>6</mml:mn>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mn>6</mml:mn>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(7)</label>
<mml:math display="block" id="M7">
<mml:mrow>
<mml:mi>&#x3b3;</mml:mi>
<mml:mo>=</mml:mo>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mi>&#x3b1;</mml:mi>
<mml:mi>&#x393;</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>+</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>&#x3b2;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mi>&#x393;</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>-</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>&#x3b2;</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>&#x393;</italic> is a factorial function and w<sub>0</sub>, w<sub>1</sub>, and w<sub>2</sub> are the probability-weighted moments of D<sub>i</sub>.</p>
<disp-formula>
<label>(8)</label>
<mml:math display="block" id="M8">
<mml:mrow>
<mml:msub>
<mml:mi>w</mml:mi>
<mml:mi>s</mml:mi>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mn>1</mml:mn>
<mml:mi>N</mml:mi>
</mml:mfrac>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>N</mml:mi>
</mml:munderover>
<mml:mrow>
<mml:msup>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>-</mml:mo>
<mml:msub>
<mml:mi>F</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mi>s</mml:mi>
</mml:msup>
<mml:msub>
<mml:mi>D</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(9)</label>
<mml:math display="block" id="M9">
<mml:mrow>
<mml:msub>
<mml:mi>F</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>-</mml:mo>
<mml:mn>0.35</mml:mn>
</mml:mrow>
<mml:mi>N</mml:mi>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where N is the number of months involved in the calculation.</p>
<p>5). Standardize the cumulative probability density to obtain the SPEI (the standard value of SPEI is 0, and the standard deviation is 1):</p>
<p>First, calculate the size of P:</p>
<disp-formula>
<label>(10)</label>
<mml:math display="block" id="M10">
<mml:mrow>
<mml:mi>P</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>-</mml:mo>
<mml:mi>F</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>x</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<p>&#x2460;</p>
<p>
<inline-formula>
<mml:math display="inline" id="im2">
<mml:mrow>
<mml:mi>W</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
<mml:mtext>&#xa0;</mml:mtext>
<mml:mi>P</mml:mi>
<mml:mo>&#x2264;</mml:mo>
<mml:mn>0.5</mml:mn>
<mml:mo>,</mml:mo>
<mml:mi>w</mml:mi>
<mml:mo>=</mml:mo>
<mml:msqrt>
<mml:mrow>
<mml:mo>-</mml:mo>
<mml:mn>2</mml:mn>
<mml:mi>l</mml:mi>
<mml:mi>n</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>P</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:msqrt>
</mml:mrow>
</mml:math>
</inline-formula>,</p>
<disp-formula>
<label>(11)</label>
<mml:math display="block" id="M11">
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:mi>P</mml:mi>
<mml:mi>E</mml:mi>
<mml:mi>I</mml:mi>
<mml:mo>=</mml:mo>
<mml:mi>w</mml:mi>
<mml:mo>-</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mi>w</mml:mi>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:msup>
<mml:mi>w</mml:mi>
<mml:mrow>
<mml:mn>2</mml:mn>
</mml:mrow>
</mml:msup>
</mml:mrow>
<mml:mrow>
<mml:mn>1</mml:mn>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>d</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mi>w</mml:mi>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>d</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:msup>
<mml:mi>w</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>d</mml:mi>
<mml:mn>3</mml:mn>
</mml:msub>
<mml:msup>
<mml:mi>w</mml:mi>
<mml:mrow>
<mml:mn>3</mml:mn>
</mml:mrow>
</mml:msup>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<p>&#x2461;</p>
<p>
<inline-formula>
<mml:math display="inline" id="im3">
<mml:mrow>
<mml:mi>W</mml:mi>
<mml:mi>h</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>P</mml:mi>
<mml:mo>&gt;</mml:mo>
<mml:mn>0.5</mml:mn>
<mml:mo>,</mml:mo>
<mml:mi>P</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>P</mml:mi>
<mml:mo>,</mml:mo>
<mml:mi>w</mml:mi>
<mml:mo>=</mml:mo>
<mml:msqrt>
<mml:mrow>
<mml:mo>-</mml:mo>
<mml:mn>2</mml:mn>
<mml:mi>l</mml:mi>
<mml:mi>n</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>P</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:msqrt>
</mml:mrow>
</mml:math>
</inline-formula>,</p>
<disp-formula>
<label>(12)</label>
<mml:math display="block" id="M12">
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:mi>P</mml:mi>
<mml:mi>E</mml:mi>
<mml:mi>I</mml:mi>
<mml:mo>=</mml:mo>
<mml:mo>-</mml:mo>
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:mrow>
<mml:mi>w</mml:mi>
<mml:mo>-</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mn>0</mml:mn>
</mml:msub>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mi>w</mml:mi>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>c</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:msup>
<mml:mi>w</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
<mml:mrow>
<mml:mn>1</mml:mn>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>d</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mi>w</mml:mi>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>d</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:msup>
<mml:mi>w</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo>+</mml:mo>
<mml:msub>
<mml:mi>d</mml:mi>
<mml:mn>3</mml:mn>
</mml:msub>
<mml:msup>
<mml:mi>w</mml:mi>
<mml:mn>3</mml:mn>
</mml:msup>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo>)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>In the above equations, c<sub>0 =</sub> 2.515517, c<sub>1 =</sub> 0.802853, c<sub>2 =</sub> 0.010328, d<sub>1 =</sub> 1.432788, d<sub>2 =</sub> 0.189269, and d<sub>3 =</sub> 0.001308.</p>
<p>The SPEI usually has a variety of time scales, such as monthly, seasonal, and annual. Compared to the SPEI with shorter time scales for targeting meteorological and agricultural drought, the SPEI with a longer time scale is more sensitive to hydrological drought (<xref ref-type="bibr" rid="B34">Ivits et&#xa0;al., 2014</xref>), moreover, vegetation germination in biological communities is mainly affected by accumulated precipitation in the previous 12 months (<xref ref-type="bibr" rid="B87">Vicente-Serrano, 2006</xref>). Therefore, in this study, the SPEI value at the 12-month scale (hereafter referred to as SPEI<sub>12</sub>) was used to characterize the interannual drought in subtropical regions, and the linear regression equation was used to evaluate the trend and characteristics of SPEI<sub>12</sub> over time based on the following formula:</p>
<disp-formula>
<label>(13)</label>
<mml:math display="block" id="M13">
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>l</mml:mi>
<mml:mi>o</mml:mi>
<mml:mi>p</mml:mi>
<mml:mi>e</mml:mi>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mi>y</mml:mi>
<mml:mo>&#xd7;</mml:mo>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>y</mml:mi>
</mml:munderover>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>&#xd7;</mml:mo>
<mml:mi>S</mml:mi>
<mml:mi>N</mml:mi>
<mml:msub>
<mml:mi>P</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
</mml:mrow>
</mml:mstyle>
<mml:mo>-</mml:mo>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>y</mml:mi>
</mml:munderover>
<mml:mi>i</mml:mi>
</mml:mstyle>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>y</mml:mi>
</mml:munderover>
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:mi>N</mml:mi>
<mml:msub>
<mml:mi>P</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
<mml:mrow>
<mml:mi>y</mml:mi>
<mml:mo>&#xd7;</mml:mo>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>y</mml:mi>
</mml:munderover>
<mml:mrow>
<mml:msup>
<mml:mi>i</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
</mml:mstyle>
<mml:mo>-</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:mrow>
<mml:mstyle displaystyle="true">
<mml:munderover>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>y</mml:mi>
</mml:munderover>
<mml:mi>i</mml:mi>
</mml:mstyle>
</mml:mrow>
<mml:mo>)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>slope</italic> is the changing trend (when <italic>slope &gt;</italic>0, it indicates that SPEI<sub>12</sub> is increasing; when <italic>slope</italic>&lt;0, then the SPEI<sub>12</sub> is decreasing), y is the number of drought years, <italic>i</italic> is the number of years (i=1, 2&#x2026; n), and SNP<sub>i</sub> is the SPEI<sub>12</sub> value of the i<sub>th</sub> year.</p>
<p>On the spatial scale, the drought classification criteria (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>) (<xref ref-type="bibr" rid="B6">Center et&#xa0;al., 2017</xref>) were used to define the drought conditions among different regions in subtropical China. For the time series, the overall mean value could not meet the criteria for drought classification due to the spatial variation of SPEI<sub>12</sub>. Therefore, in this study, the threshold value of the SPEI<sub>12</sub> percentile was used to define the annual drought conditions with reference to the precipitation percentile threshold method (<xref ref-type="bibr" rid="B111">Zhai and Pan, 2003</xref>), as shown in <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Drought grade of standardized precipitation evapotranspiration index.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">SPEI</th>
<th valign="top" align="center">Drought levels</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">-0.5&lt;SPEI</td>
<td valign="top" align="left">No Drought</td>
</tr>
<tr>
<td valign="top" align="left">-1.0&lt;SPEI&#x2264;-0.5</td>
<td valign="top" align="left">Mild Drought</td>
</tr>
<tr>
<td valign="top" align="left">-0.5&lt;SPEI&#x2264;-1</td>
<td valign="top" align="left">Moderate Drought</td>
</tr>
<tr>
<td valign="top" align="left">-1&lt;SPEI&#x2264;-1.5</td>
<td valign="top" align="left">Severe Drought</td>
</tr>
<tr>
<td valign="top" align="left">-1.5&lt;SPEI&#x2264;-2</td>
<td valign="top" align="left">Extreme Drought</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>SPEI percentile threshold for determining water balance conditions.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">SPEI Percentile</th>
<th valign="top" align="center">Conditions</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">&#x2265;90%</td>
<td valign="top" align="left">Extreme Wet</td>
</tr>
<tr>
<td valign="top" align="left">75%-90%</td>
<td valign="top" align="left">Moderate wet</td>
</tr>
<tr>
<td valign="top" align="left">25%-75%</td>
<td valign="top" align="left">Normal</td>
</tr>
<tr>
<td valign="top" align="left">10%-25%</td>
<td valign="top" align="left">Moderate Drought</td>
</tr>
<tr>
<td valign="top" align="left">&#x2264;10%</td>
<td valign="top" align="left">Extreme Drought</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s2_4">
<title>2.4 Spatiotemporal simulation and trend analysis of subtropical forest carbon storage</title>
<p>In this study, the carbon storage of vegetation in two types of forests in subtropical China from 1985 to 2019 was simulated with a spatial resolution of 1km and temporal resolution on a daily time steps.</p>
<p>The Biome-BGC model simulates the physiological and ecological processes of vegetation that control the material cycle and energy flow of the ecosystem, including canopy radiation, photosynthesis, stomatal conductance, autotrophic respiration, heterotrophic respiration, phenological dynamics, and evapotranspiration. The model can simulate the energy and carbon-nitrogen water cycles between the atmosphere, vegetation, and soil of the terrestrial ecosystem in daily steps to estimate the storage and flux fluxes among carbon, nitrogen, and water pools (<xref ref-type="bibr" rid="B69">Running, 1993</xref>).</p>
<p>The input data to the Biome-BGC model included vegetation abundance data, topographic data, soil data, and meteorological data, which has been described in Section 2.2.</p>
<p>This study used 67 physiological and ecological parameters to run the model. At present, few studies have focused on the physiology and ecology parameters of vegetation. In this study, the proportion of nitrogen in the Rubisco enzyme, litter coefficient at the leaf replacement period, and the carbon-nitrogen distribution ratio of each part were obtained by an iterative method (<xref ref-type="bibr" rid="B53">Lu et&#xa0;al., 2016</xref>). In addition, the default ENF or EBF parameters provided by White (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>) were adopted if the parameters could not be determined through the literature. Some parameter values are shown in <xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>.</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Main parameters input to the Biome-BGC model.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Parameter</th>
<th valign="top" align="center">EBF</th>
<th valign="top" align="center">ENF</th>
<th valign="top" align="center">Unit</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Froot turnover</td>
<td valign="top" align="center">0.7</td>
<td valign="top" align="left">0.7 (<xref ref-type="bibr" rid="B53">Lu et&#xa0;al., 2016</xref>)</td>
<td valign="top" align="left">a<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Specific leaf area</td>
<td valign="top" align="center">10</td>
<td valign="top" align="left">12 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">m&#xb7;Kg C<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">The proportion of unstable substances in fine roots</td>
<td valign="top" align="center">34</td>
<td valign="top" align="left">34 (<xref ref-type="bibr" rid="B47">Liu and Fei, 2013</xref>)</td>
<td valign="top" align="left">%</td>
</tr>
<tr>
<td valign="top" align="left">Fine root cellulose ratio</td>
<td valign="top" align="center">44</td>
<td valign="top" align="left">44 (<xref ref-type="bibr" rid="B47">Liu and Fei, 2013</xref>)</td>
<td valign="top" align="left">%</td>
</tr>
<tr>
<td valign="top" align="left">Fine root lignin ratio</td>
<td valign="top" align="center">24</td>
<td valign="top" align="left">22 (<xref ref-type="bibr" rid="B28">Holling et&#xa0;al., 1995</xref>)</td>
<td valign="top" align="left">%</td>
</tr>
<tr>
<td valign="top" align="left">The ratio of nitrogen to Rubisco enzyme</td>
<td valign="top" align="center">0.08</td>
<td valign="top" align="left">0.07 (<xref ref-type="bibr" rid="B53">Lu et&#xa0;al., 2016</xref>)</td>
<td valign="top" align="left">kg N Rub&#xb7;kg<break/>N leaf<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Leaf carbon and nitrogen ratio</td>
<td valign="top" align="center">42</td>
<td valign="top" align="left">42 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">kg C&#xb7;kg N<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Litter carbon-nitrogen ratio</td>
<td valign="top" align="center">49</td>
<td valign="top" align="left">93 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">kg C&#xb7;kg N<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Fine roots carbon-nitrogen ratio</td>
<td valign="top" align="center">58</td>
<td valign="top" align="left">58 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">kg C&#xb7;kg N<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Living wood carbon-nitrogen ratio</td>
<td valign="top" align="center">50</td>
<td valign="top" align="left">58 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">kg C&#xb7;kg N<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Deadwood carbon-nitrogen ratio</td>
<td valign="top" align="center">550</td>
<td valign="top" align="left">730 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">kg C&#xb7;kg N<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">The biggest stomatal conductance</td>
<td valign="top" align="center">0.006</td>
<td valign="top" align="left">0.006 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">m&#xb7;s<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">The surface conductance</td>
<td valign="top" align="center">0.00006</td>
<td valign="top" align="left">0.00006 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">m&#xb7;s<sup>-1</sup>
</td>
</tr>
<tr>
<td valign="top" align="left">Boundary layer conductance</td>
<td valign="top" align="center">0.09</td>
<td valign="top" align="left">0.01 (<xref ref-type="bibr" rid="B95">White et&#xa0;al., 2000</xref>)</td>
<td valign="top" align="left">m&#xb7;s<sup>-1</sup>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>First, the model was spun-up, meaning that the carbon, nitrogen, and water storage of the ecosystem when the annual change in the soil carbon pool was less than 0.0005&#xa0;kg C&#xb7;m<sup>-2</sup> (<xref ref-type="bibr" rid="B83">Thornton, 2010</xref>) was used as the initial condition of the simulation, and then the vegetation carbon storage of the EBF and ENF in subtropical China from 1985 to 2019 was simulated.</p>
<p>Based on the simulation results, the linear regression analysis was used to calculate the trend of simulated carbon storage. The equation is similar to Eq. 13, where <italic>slope</italic> is the changing trend (when <italic>slope &gt;</italic>0, carbon storage is increasing, and when <italic>slope</italic>&lt;0, carbon storage is decreasing), y is the number of simulated years, i is the number of years (<italic>i</italic>=1, 2,&#x2026;, n), and SNP<sub>i</sub> is the value of carbon storage in the <italic>i</italic>
<sub>th</sub> year.</p>
</sec>
<sec id="s2_5">
<title>2.5 Evaluation of forest resistance and resilience</title>
<p>Forest ecosystem resistance (Rt) represents the ability of a forest ecosystem to maintain its original state under drought disturbance, whereas forest ecosystem resilience (Rs) describes the ability of a forest to recover from drought disturbance to a normal state (<xref ref-type="bibr" rid="B86">Van Ruijven and Berendse, 2010</xref>; <xref ref-type="bibr" rid="B10">De Keersmaecker et&#xa0;al., 2014</xref>). These parameters represent functions of the stability of an ecosystem (<xref ref-type="bibr" rid="B10">De Keersmaecker et&#xa0;al., 2014</xref>). In this study, formulas 14 and 15 were used to quantitatively explain the Rt and Rs of subtropical forests to drought, where the values of Rt and Rs are unitless to facilitate the comparison of the stability of forest ecosystems with two different levels of productivity.</p>
<disp-formula>
<label>(14)</label>
<mml:math display="block" id="M14">
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mi>t</mml:mi>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mover accent="true">
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mi>n</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="true">&#xaf;</mml:mo>
</mml:mover>
</mml:mrow>
<mml:mrow>
<mml:mrow>
<mml:mo>|</mml:mo>
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mi>e</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mover accent="true">
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mi>n</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="true">&#xaf;</mml:mo>
</mml:mover>
</mml:mrow>
<mml:mo>|</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(15)</label>
<mml:math display="block" id="M15">
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mi>s</mml:mi>
<mml:mo>=</mml:mo>
<mml:mrow>
<mml:mo>|</mml:mo>
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mi>e</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mover accent="true">
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mi>n</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="true">&#xaf;</mml:mo>
</mml:mover>
</mml:mrow>
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mrow>
<mml:mi>e</mml:mi>
<mml:mo>+</mml:mo>
<mml:mi>i</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:mover accent="true">
<mml:mrow>
<mml:msub>
<mml:mi>Y</mml:mi>
<mml:mi>n</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="true">&#xaf;</mml:mo>
</mml:mover>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo>|</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where Y<italic>
<sub>n</sub>
</italic> is the carbon storage in the normal year from 1985 to 2019, Y<italic>
<sub>e</sub>
</italic> is the carbon storage in the extreme drought year, and Y<italic>
<sub>e</sub>
</italic>
<sub>+</sub>
<italic>
<sub>i</sub>
</italic> is the forest carbon storage in year <italic>i</italic> after the event. Since the effects of drought on forest growth may last for several years and lead to legacy effects (<xref ref-type="bibr" rid="B2">Anderegg et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B70">Schwalm et&#xa0;al., 2017</xref>), <italic>i</italic> =1, 2, and 4 was used in this study to quantitatively analyze the resilience of carbon storage in the first, second, and fourth years after drought (in addition, the water balance of the forest in the four years after drought should also be considered). A higher Rt value indicates that the forest is more resistant to drought and a higher Rs value means stronger forest resilience (<xref ref-type="bibr" rid="B30">Huang and Xia, 2019</xref>).</p>
<p>In this study, we first analyzed the spatiotemporal distribution trend of the SPEI<sub>12</sub> in the subtropical region of China from 1952 to 2019. Extreme drought years were determined based on the drought time series. Combined with the simulation results of carbon storage in subtropical forests, the resistance and resilience of forests affected by extreme drought were analyzed. As the frequency of drought changes, the acclimation of forests to drought may also change accordingly (<xref ref-type="bibr" rid="B2">Anderegg et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B32">Isbell et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B3">Anderegg et&#xa0;al., 2020</xref>). Therefore, whether continuous drought occurred before and after the drought year was determined in this study, and whether continuous drought affects forest resistance and resilience is also a significant step in this research and discussion.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<title>3 Result</title>
<sec id="s3_1">
<title>3.1 Spatiotemporal characteristics of SPEI and drought trend</title>
<p>The time series of SPEI<sub>12</sub> in the subtropical region from 1952 to 2019 are shown in <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>. During the 68 years, subtropical forests in China experienced five extreme drought events, which occurred every 13.6 years. According to the spatial variation trends in drought (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>), SPEI<sub>12</sub> showed a decreasing trend in 62.13% of the region, with an overall decrease of 0.036(10&#xa0;a)<sup>-1</sup>. <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref> showed the trends and characteristics of SPEI<sub>12</sub> with time using the linear regression equation (Eq. 13). The regions with a serious downward trend were mainly distributed in most of the western subtropical region, especially in parts of the Sichuan and Yunnan provinces, with a downward trend of more than 0.15 (10a)<sup>-1</sup>.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Spatiotemporal trend of SPEI<sub>12</sub>: <bold>(A)</bold> Time series and percentiles of drought conditions of SPEI<sub>12</sub> from 1952 to 2019 (the solid red lines at the 10th and 90th percentiles represent thresholds for extreme drought and extreme wetness, respectively, and the dashed red lines at the 25th and 75th percentiles represent thresholds for moderate drought and moderate wetness, respectively and different regions represent different drought conditions); and <bold>(B)</bold> spatial variation trend of subtropical drought from 1952 to 2019.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g003.tif"/>
</fig>
<p>According to <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>, two extreme drought events occurred after 1985 in 2009 and 2011. Because the SPEI<sub>12</sub> value was lower in 2011, 2011 was defined as the extreme drought year in this study. Therefore, this study focused on the resistance of subtropical forests in 2011 to drought and their resilience at the first to fourth year after the 2011 extreme drought, which covers the period from 2011 to 2015.</p>
<p>To compare the different drought conditions of the two forest types, we calculated the pixel proportions of the two forest types under different drought conditions (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). According in <xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>, the areas of extreme and severe drought were concentrated in the central to southwestern region of subtropical China, where the forest cover rate was high. The results for EBF showed that 19.3% suffered from extreme drought (SPEI<sub>12</sub>&#x2264;-2) and 26.6% suffered from severe drought (-2&lt; SPEI<sub>12</sub>&#x2264;-1.5), while those for ENF showed that 31.9% suffered from extreme drought (SPEI<sub>12</sub>&#x2264;-2) and 21.2% suffered from severe drought condition (-2&lt; SPEI<sub>12</sub>&#x2264;-1.5). The results showed that most EBF and ENF in subtropical regions of China suffered from severe to extreme drought in 2011.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Spatial distribution of different drought levels of two typical subtropical forests in 2011 <bold>(A)</bold>, 2012 <bold>(B)</bold>, 2013 <bold>(C)</bold>, and 2015 <bold>(D)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g004.tif"/>
</fig>
<p>The value of SPEI12 in 2012 is -0.028, and the water balance condition is normal as shown in <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>. As shown in <xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>, only 8.84% of EBF pixels and 1.7% of ENF pixels are under severe to extreme drought conditions in 2012, and the majority of them are under normal water balance condition.</p>
<p>The SPEI<sub>12</sub> value in 2013 was -0.526, which represents a moderate drought according to the time series. However, according to the spatial distribution (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4C</bold>
</xref>), only a few forest pixels in the subtropical region of China were affected by extreme to severe drought in 2013, and the subtropical forests as a whole had normal moisture status.</p>
<p>The SPEI<sub>12</sub> value was 0.587 in 2015, indicating the absence of drought. Further analysis showed that the vast majority of forest pixels in the EBF (95.34%) and ENF (79.56%) were in the normal to wet state without drought in 2015 (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4D</bold>
</xref>).</p>
</sec>
<sec id="s3_2">
<title>3.2 Spatial and temporal variation trends in carbon storage in subtropical forests</title>
<p>The spatial distribution of average vegetation carbon storage of EBF and ENF during 1985-2019 were shown in <xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>. As shown in the figure, the average vegetation carbon storage of EBF and ENF ranged in 124.03 - 143.45&#xa0;t hm<sup>-2</sup> (130.58 &#xb1; 10.02&#xa0;t hm<sup>-2</sup>) and 77.21 - 82.59&#xa0;t hm<sup>-2</sup> (78.49 &#xb1; 8.49&#xa0;t hm<sup>-2</sup>), respectively. In terms of spatial distribution, the regions with higher carbon storage of EBF were mainly concentrated in central and southeastern subtropics, where those of ENF mainly distributed in the western subtropic, such as Tibet, Yunnan and Sichuan province.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Spatial distribution of average carbon storage for two types of forests during 1985-2019 <bold>(A)</bold> EBF and <bold>(B)</bold> ENF).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g005.tif"/>
</fig>
<p>According to the analysis of spatiotemporal characteristics of subtropical drought, follows we mainly focus on the vegetation carbon storage of two types of forest during 2011-2015 for better understanding the resistance and resilience of subtropical forests to drought. The monthly time series and spatial trends of carbon storage of two subtropical forest are shown in <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>. As shown in <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6A</bold>
</xref>, the carbon storage of EBF fluctuates within a certain range, whereas the ENF fluctuates relatively little and continues to increase each year. From 2011 to 2015, the mean values of the overall annual trend of carbon storage for the two forest species in subtropical China were -0.042 (EBF) and -0.013 (ENF). The overall carbon storage of subtropical evergreen forests showed a slightly decreasing trend, and the annual variation ranged mainly between -0.25 and 0.25. In subtropical China, 60.5% of the EBF and 51.18% of the ENF pixels showed a downward trend from 2011 to 2015. <xref ref-type="fig" rid="f6">
<bold>Figures&#xa0;6B, C</bold>
</xref> shows the trends and characteristics of carbon storage changes over time evaluated using the linear regression equation (Eq. 13). The spatial distribution of the changing trend in carbon storage in the two forests (<xref ref-type="fig" rid="f6">
<bold>Figures&#xa0;6B, C</bold>
</xref>) was relatively complex: the two forests showed an overall downward trend; the pixels in the middle and southeast of the EBF showed an overall upward trend, and the pixels from northwest to southeast of the ENF showed an upward trend.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Time serials and spatial trends of carbon storage from 2011 to 2015 <bold>(A)</bold>. Monthly carbon storage; <bold>(B)</bold> trends of EBF; and <bold>(C)</bold>. trends of ENF).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g006.tif"/>
</fig>
</sec>
<sec id="s3_3">
<title>3.3 Stability of the EBF and ENF in subtropical China during drought in 2011</title>
<p>To evaluate the stability of forests in subtropical China to drought, we first explored the resistance of the two forest types to drought in 2011 and the resilience of the forests at 1, 2, and 4 years after 2011.</p>
<sec id="s3_3_1">
<title>3.3.1 Comparison of resistance of two forest types to drought in 2011</title>
<p>The resistance levels of the EBF pixels and ENF pixels for 2011 are shown in <xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7A</bold>
</xref>. According to the calculation results, the ranges of Rt<sub>EBF</sub> and Rt<sub>ENF</sub> were different to some extent and had values of 0.15 - 78.40 and 0.01 - 26.47, respectively. The median resistance value of the EBF calculated based on simulated carbon storage was 12.21, and that of the ENF was 3.85. These results clearly show that the resistance of the EBF to the 2011 drought, which was based on carbon storage, was significantly higher than that of the ENF. The spatial distributions of the Rt<sub>EBF</sub> and Rt<sub>ENF</sub> are shown in <xref ref-type="fig" rid="f8">
<bold>Figures&#xa0;8A, B</bold>
</xref>. In the western region of the subtropical zone, the EBF exhibits weak resistance to drought, while in the central to eastern region, the EBF exhibits strong resistance to drought, gradually increasing from west to east. The overall spatial distribution of ENF resistance was relatively average, with the western subtropical region being slightly stronger than the central and eastern regions. There are some differences in the spatial distributions of the EBF and ENF resistance. The results showed that the two forest types responded differently to the 2011 drought.</p>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>Resistance of the EBF and ENF to drought in 2011 <bold>(A)</bold>, and their resilience to the 2011 extreme drought after the first <bold>(B)</bold>, second <bold>(C)</bold>, and fourth <bold>(D)</bold> year.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g007.tif"/>
</fig>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>Spatial distribution of resistance to 2011 drought <bold>(A)</bold>. EBF and <bold>(B)</bold> ENF).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g008.tif"/>
</fig>
</sec>
<sec id="s3_3_2">
<title>3.3.2 Comparison of EBF and ENF resilience</title>
<p>
<xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref> shows the spatial distribution of resilience of the EBF and ENF at 1, 2, and 4 years after the 2011 drought. As for the resilience of the two forest types one year after the drought, the median resilience of the EBF and ENF were 1.00 and 0.947, respectively (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7B</bold>
</xref>), which means that the resilience levels of the two forest types in the first year after drought were similar, with the resilience of the EBF being slightly stronger than that of the ENF. The spatial distribution of resilience of the EBF and ENF showed certain differences (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9A, B</bold>
</xref>). Overall, the spatial distribution of EBF resilience was relatively average, whereas the resilience of ENF was weak in the northwestern part of the subtropical region and strong in the central to eastern part.</p>
<fig id="f9" position="float">
<label>Figure&#xa0;9</label>
<caption>
<p>Spatial distribution of resilience of EBF and ENF to 2011 extreme drought after the first <bold>(A, B)</bold>, second <bold>(C, D)</bold>, and fourth <bold>(E, F)</bold> year.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g009.tif"/>
</fig>
<p>In the second year after the 2011 drought (2013), the median resilience of the two forest types was 1.00 and 0.87, respectively (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7C</bold>
</xref>). The resilience level of the EBF was similar to that one year later, while the resilience level of the ENF slightly decreased. The spatial distribution of EBF resilience in the two years after the drought was similar to that in the year after the drought, which was relatively average overall (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9C, D</bold>
</xref>). However, the resilience level of the ENF in the southern subtropical region showed a clearly decreasing trend in the two years after the drought, which was clearly different from that in the northern region, which may be related to the uncertainty of carbon storage simulation.</p>
<p>In the fourth year (2015) after the 2011 drought, the resilience levels of the two forests were generally similar (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7D</bold>
</xref>), with median resilience values of 1.26 and 1.33, respectively, indicating that the resilience of both forests had increased. The spatial distribution of resilience at the fourth years after the drought is shown in <xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9 E, F</bold>
</xref>, the resilience levels of both forests in the central subtropical region were significantly higher than those in the western and southeastern subtropical regions, and higher levels of resilience areas were mainly distributed in Yunnan, Sichuan, Chongqing, Guizhou and other regions of the junction, which also represent areas that suffered more severe drought mentioned in Section 3.1. From the perspective of spatial distribution, the resilience of the two types of forests increased in the central subtropical region but decreased in the western subtropical region decreased. The inconsistency of resilience in different regions may be caused by the spatial heterogeneity of drought levels and the different spatial distributions of forests. Thus, the water-heat balance conditions of different pixels and the relationship between forest abundance data and the resilience level must be further compared. The resilience values indicate that both forest types will return to normal conditions within four years.</p>
</sec>
</sec>
<sec id="s3_4">
<title>3.4 Effects of continuous drought on forest resilience</title>
<p>As mentioned above, changes in drought frequency affect forest resistance and resilience; therefore, this section examines the effects of continuous drought on forest resistance and resilience.</p>
<p>In this study, the year of extreme drought was 2011. Pixels that were or were not affected by drought in 2009, 2010, and 2011 were divided into two categories, and pixels that presented the two different situations in the three years were combined. The forest pixels that experienced three consecutive drought years (2009-2011), two consecutive drought years (2010-2011), and one drought year (2011) were screened and divided into three climatic combinations. <xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref> shows the resistance and resilience of the two forests under the three drought combinations.</p>
<fig id="f10" position="float">
<label>Figure&#xa0;10</label>
<caption>
<p>Statistical of Rt and Rs of two typical subtropical forests during and after the 2011 extreme drought. Each row represents a different combination of droughts. &#x201c;N&#x201d; represents normal, &#x201c;D&#x201d; represents drought, and &#x201c;NDD&#x201d; represents pixels that were normal in 2009 but drought in 2010 and 2011, i.e., pixels that suffered drought for two consecutive years. The first column shows the Rt in 2011, and columns two to four show the resilience of the first, second, and fourth years after the drought.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g010.tif"/>
</fig>
<p>
<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref> shows that the EBF generally has higher resistance under three different continuous drought conditions, that is, subtropical EBF in China can quickly adapt to drought. In terms of resilience, under the condition of a different combination of drought, two kinds of resilience showed similar levels of forest, and in the fourth year after the drought, two kinds of resilience of forest showed very similar levels, suggesting that the two kinds of forest will recover to normal levels after four years of drought. These results support the idea that forests need at least one year to recover from interannual droughts (<xref ref-type="bibr" rid="B70">Schwalm et&#xa0;al., 2017</xref>).</p>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<title>4 Discussion</title>
<sec id="s4_1">
<title>4.1 Uncertainty analysis of SPEI calculation</title>
<p>The Thornthwaite (TW) (<xref ref-type="bibr" rid="B82">Thornthwaite, 1948</xref>) and P-M (<xref ref-type="bibr" rid="B58">Monteith, 1965</xref>) formulas were commonly used to calculate the PET in the SPEI estimation. The SPEI calculated based on TW formula was easily to implement, while it may indicate excessive dry conditions due to the influence of temperature, under the significantly increased temperatures in recent years (<xref ref-type="bibr" rid="B116">Zhang et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B60">Nouri and Homaee, 2020</xref>; <xref ref-type="bibr" rid="B19">Guo et&#xa0;al., 2022</xref>). Compared to the TW formula, which only considers temperature, the P-M formula considered variety of meteorological factors, which are more complex, and is generally more consistent with actual evapotranspiration (<xref ref-type="bibr" rid="B35">Jensen et&#xa0;al., 1990</xref>), specially it enable to describe the regions influenced by aerodynamic factors (<xref ref-type="bibr" rid="B48">Liu and Jiang, 2015</xref>). Consider, these two methods were widely used in the drought detection researches, further validation is required to determine which algorithm is better characterized in subtropical China.</p>
<p>The SPEI results were limited by meteorological stations, whereas some areas had few meteorological stations. Therefore, the accuracy of the SPEI in characterizing drought conditions in the study area needs to be confirmed by further comparative analysis with more observational data. The results of the temporal analysis indicate that the SPEI<sub>12</sub> shows a significant downward trend from the 1950s to the 2010s, the interannual drought in the subtropical region of China shows an increasing trend from the 1950s to the 2010s, which is consistent with previous studies (<xref ref-type="bibr" rid="B48">Liu and Jiang, 2015</xref>). Spatial analysis showed a clear trend of drought in the subtropical southwest, especially in the Sichuan Basin, which is consistent with previous research findings (<xref ref-type="bibr" rid="B51">Li et&#xa0;al., 2012</xref>). A comparison of historical drought events showed that in 1982, Ningbo, Jinhua, Wenzhou, Jiujiang, and Nanchang (<xref ref-type="bibr" rid="B114">Zhang and Liu, 1993</xref>; <xref ref-type="bibr" rid="B113">Zhang et&#xa0;al., 2003</xref>); in 1986, Nanchang, Shangrao, Ji &#x2018;an, and Ganzhou (<xref ref-type="bibr" rid="B114">Zhang and Liu, 1993</xref>; <xref ref-type="bibr" rid="B113">Zhang et&#xa0;al., 2003</xref>); in 1991, south Guangdong and Guangxi (<xref ref-type="bibr" rid="B92">Wang and Chen, 2012</xref>); in 1994, Suzhou and Shanghai (<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al., 2003</xref>); and in 2000, moderate to severe drought in Yangzhou, Nanjing, Hefei and Anqing (<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al., 2003</xref>). The simulation results of SPEI in this study were consistent with typical historical drought events (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>).</p>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>Latitude and longitude of cities where historical drought events occurred.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Time</th>
<th valign="top" align="center">City</th>
<th valign="top" align="center">Latitude and longitude</th>
<th valign="top" align="center">Literature</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">1982</td>
<td valign="top" align="left">Ningbo</td>
<td valign="top" align="left">28&#xb0;51&#x2019;-30&#xb0;33&#x2019;N,120&#xb0;55&#x2019;-122&#xb0;16&#x2019;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1982</td>
<td valign="top" align="left">Jinhua</td>
<td valign="bottom" align="left">28&#xb0;32&#x2019;-29&#xb0;41&#x2019;N,119&#xb0;14&#x2019;-120&#xb0;46&#x2019;30&#x201d;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1982</td>
<td valign="top" align="left">Wenzhou</td>
<td valign="bottom" align="left">27&#xb0;03&#x2019;-28&#xb0;36&#x2019;N, 119&#xb0;37&#x2019;-121&#xb0;18&#x2019;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B114">Zhang and Liu (1993)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1982</td>
<td valign="bottom" align="left">Jiujiang</td>
<td valign="bottom" align="left">28&#xb0;47&#x2019;-30&#xb0;06&#x2019;N,113&#xb0;57&#x2019;-116&#xb0;53&#x2019;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B114">Zhang and Liu (1993)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1982</td>
<td valign="bottom" align="left">Nanchang</td>
<td valign="bottom" align="left">28&#xb0;10&#x2019;-29&#xb0;11&#x2019;N,115&#xb0;27&#x2019;-116&#xb0;35&#x2019;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B114">Zhang and Liu (1993)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1986</td>
<td valign="bottom" align="left">Nanchang</td>
<td valign="bottom" align="left">28&#xb0;10&#x2019;-29&#xb0;11&#x2019;N,115&#xb0;27&#x2019;-116&#xb0;36&#x2019;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B114">Zhang and Liu (1993)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1986</td>
<td valign="bottom" align="left">Shangrao</td>
<td valign="bottom" align="left">27&#xb0;48&#xb4;-29&#xb0;42&#xb4;N,116&#xb0;13&#xb4;-118&#xb0;29&#xb4;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B114">Zhang and Liu (1993)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1986</td>
<td valign="bottom" align="left">Ji&#x2019;an</td>
<td valign="bottom" align="left">25&#xb0;58&#x2032;-27&#xb0;57&#x2032;N, 113&#xb0;46&#x2019;-115&#xb0;56&#x2019;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1986</td>
<td valign="bottom" align="left">Ganzhou</td>
<td valign="bottom" align="left">24&#xb0;29&#x2032;-27&#xb0;09&#x2032;N,113&#xb0;54&#x2032;-116&#xb0;38&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1994</td>
<td valign="bottom" align="left">Suzhou</td>
<td valign="bottom" align="left">30&#xb0;47&#x2032;-32&#xb0;02&#x2032;N,119&#xb0;55&#x2032;-121&#xb0;20&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1994</td>
<td valign="bottom" align="left">Shanghai</td>
<td valign="bottom" align="left">30&#xb0;40&#x2032;-31&#xb0;53&#x2032;N,120&#xb0;52&#x2032;-122&#xb0;12&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000</td>
<td valign="bottom" align="left">Yangzhou</td>
<td valign="bottom" align="left">32&#xb0;15&#x2032;-33&#xb0;25&#x2032;N,119&#xb0;01&#x2032;-119&#xb0;54&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000</td>
<td valign="bottom" align="left">Nanjing</td>
<td valign="bottom" align="left">31&#xb0;14&#x2032;-32&#xb0;37&#x2032;N, 118&#xb0;22&#x2032;-119&#xb0;14&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000</td>
<td valign="bottom" align="left">Hefei</td>
<td valign="bottom" align="left">30&#xb0;56&#x2032;-32&#xb0;33&#x2032;N,116&#xb0;40&#x2032;-117&#xb0;58&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000</td>
<td valign="bottom" align="left">Anqing</td>
<td valign="bottom" align="left">29&#xb0;47&#x2032;-31&#xb0;16&#x2032;N,115&#xb0;45&#x2032;-117&#xb0;44&#x2032;E</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B113">Zhang et&#xa0;al. (2003)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s4_2">
<title>4.2 Simulation uncertainty of carbon storage in subtropical forests</title>
<p>The uncertainty of the Biome-BGC model may be divided into the uncertainty of the model structure, input variables, and uncertainty of model parameters (<xref ref-type="bibr" rid="B46">Li and Sun, 2018</xref>). First, the uncertainty of the model structure may be due to inadequate simulation of carbon, nitrogen, and water cycles in the ecosystem, which may lead to the difference between the simulation and observations (<xref ref-type="bibr" rid="B8">Churkina et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B26">Hidy et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B75">Smith and Dukes, 2013</xref>). Impacts like human activities (<xref ref-type="bibr" rid="B12">Du et&#xa0;al., 2021</xref>) (e.g., management practices and forest wildfires) on the carbon and nitrogen water cycle were not considered in this simulation, which may create uncertainty. Second, the uncertainty of the input variables may be due to errors or inadequacies in the collection and statistics of input data (<xref ref-type="bibr" rid="B39">Jung et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B13">Eastaugh et&#xa0;al., 2011</xref>). Although the forest abundance data used in this simulation are from fine-scale classification products with a resolution of 30&#xa0;m, resampling to 1&#xa0;km does not avoid pixel mixing and thus errors (<xref ref-type="bibr" rid="B80">Su et&#xa0;al., 2022</xref>). Third, the uncertainty of the model parameters may be due to their different effects of the model parameters on the output results under different conditions (<xref ref-type="bibr" rid="B81">Tatarinov and Cienciala, 2006</xref>; <xref ref-type="bibr" rid="B40">Kang, 2016</xref>; <xref ref-type="bibr" rid="B67">Raj et&#xa0;al., 2018</xref>). For some difficult-to-obtain parameters, this study was obtained by reviewing the literature and directly using the model defaults, which may cause uncertainty in the results.</p>
<p>However, by comparing the simulated values with the observed values in the Zhejiang forest inventory sample plots, and using evaluation indicators such as correlation coefficient and root mean square error to analyze the simulation accuracy, we found that the simulated carbon storage values of both forest types in this study were correlated with the observed values, as shown in <xref ref-type="fig" rid="f11">
<bold>Figure&#xa0;11</bold>
</xref>, and the mean values of carbon storage of both forest types were within the range of the values reported in previous studies (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>).</p>
<fig id="f11" position="float">
<label>Figure&#xa0;11</label>
<caption>
<p>The comparison between simulated and observed vegetation carbon storage of two typical subtropical forests in Zhejiang Province (<bold>A</bold>. EBF and <bold>B</bold>. ENF).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-14-1067552-g011.tif"/>
</fig>
<table-wrap id="T5" position="float">
<label>Table&#xa0;5</label>
<caption>
<p>Comparison between the mean values of carbon storage of two forest types simulated in this study and previous studies.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">The time range</th>
<th valign="top" align="center">EBF(t&#xb7;hm<sup>-2</sup>)</th>
<th valign="top" align="center">ENF(t&#xb7;hm<sup>-2</sup>)</th>
<th valign="top" align="center">Literature</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">1985-2019</td>
<td valign="top" align="center">130.58</td>
<td valign="top" align="center">78.49</td>
<td valign="top" align="left">This study</td>
</tr>
<tr>
<td valign="top" align="left">1994</td>
<td valign="top" align="center">/</td>
<td valign="top" align="center">40</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B21">Gu et&#xa0;al. (2010)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1996</td>
<td valign="top" align="center">66.1</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B56">Minghong et&#xa0;al. (1996)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1999</td>
<td valign="top" align="center">26.3</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B91">Wang (1999)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000</td>
<td valign="top" align="center">100.73</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B121">Zhou et&#xa0;al. (2000)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000</td>
<td valign="top" align="center">/</td>
<td valign="top" align="center">80.798</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B108">Yun-Ting and Jiang-Ming (2002)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1977-2008</td>
<td valign="top" align="center">40.7</td>
<td valign="top" align="center">33.9</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B104">Yang et&#xa0;al. (2022)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">1979-2012</td>
<td valign="top" align="center">/</td>
<td valign="top" align="center">21.37</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B43">Li et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2000-2014</td>
<td valign="top" align="center">74.2</td>
<td valign="top" align="center">62</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B119">Zhang et&#xa0;al. (2019b)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2004-2014</td>
<td valign="top" align="center">70.91</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B62">Peng et&#xa0;al. (2016)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2007</td>
<td valign="top" align="center">89.2</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B112">Zhang et&#xa0;al. (2007)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2008</td>
<td valign="top" align="center">53.62</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B93">Wang et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2009</td>
<td valign="top" align="center">/</td>
<td valign="top" align="center">12.72</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B36">Jin et&#xa0;al. (2019)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2009-2018</td>
<td valign="top" align="center">43.7</td>
<td valign="top" align="center">37.5</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B104">Yang et&#xa0;al. (2022)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2010</td>
<td valign="top" align="center">129.34</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B109">Zeng et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2010.5-6</td>
<td valign="top" align="center">134.9</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B110">Zeng et&#xa0;al. (2013)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2011</td>
<td valign="top" align="center">38.92</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B31">Hu et&#xa0;al. (2011)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2011</td>
<td valign="top" align="center">126</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B118">Zhang et&#xa0;al. (2020)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2012</td>
<td valign="top" align="center">97.49</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B78">Sun and Guan (2014)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2012-2013</td>
<td valign="top" align="center">97.30</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B117">Zhang et&#xa0;al. (2019a)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">2010-2050</td>
<td valign="top" align="center">94.71</td>
<td valign="top" align="center">91.33</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B25">He et&#xa0;al. (2017)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s4_3">
<title>4.3 Differences in ecosystem stability between EBF and ENF</title>
<p>This study shows the spatial differences between drought resistance and resilience of the EBF and ENF in the Chinese subtropics (<xref ref-type="fig" rid="f8">
<bold>Figures&#xa0;8</bold>
</xref>, <xref ref-type="fig" rid="f9">
<bold>9</bold>
</xref>). As shown in the figure, the resistance value of the EBF to drought was significantly higher than that of the ENF, indicating that EBF has a stronger drought resistance ability in the subtropical region of China, which is consistent with previous studies (<xref ref-type="bibr" rid="B30">Huang and Xia, 2019</xref>; <xref ref-type="bibr" rid="B73">Shao et&#xa0;al., 2022</xref>). The reason may belong to the higher photosynthesis efficiency of EBF than that of ENF during drought (<xref ref-type="bibr" rid="B98">Wu and Wang, 2022</xref>). Previous studies found that EBF could accelerate the loss of old leaves and maintain the growth of young leaves to maintain the light use efficiency, and increase the carbon sequestration capability (<xref ref-type="bibr" rid="B97">Wu et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B94">Wei et&#xa0;al., 2017</xref>). Although there is significant difference in the resistance of the two forest types, no significant difference in resilience between the two forests 1-4 years after the drought. Previous studies found ecosystem resilience at a large scale can be expressed by their respective WUE levels (<xref ref-type="bibr" rid="B66">Ponce-Campos et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B77">Stan et&#xa0;al., 2021</xref>). Therefore, due to the similar change trend of WUE in the case of changes in hydroclimatic conditions of EBF and ENF (<xref ref-type="bibr" rid="B74">Sharma and Goyal, 2018</xref>; <xref ref-type="bibr" rid="B30">Huang and Xia, 2019</xref>), resulting the similar resilience of EBF and ENF to drought in subtropical China. However, the varies in climate and geological conditions cased spatial difference of WUE in two forests, which may be the reason of spatial heterogeneity in resilience throughout subtropical China (<xref ref-type="bibr" rid="B20">Guo et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B73">Shao et&#xa0;al., 2022</xref>).</p>
<p>EBF and ENF are the predominant vegetation types in subtropical regions of China, especially EBF, which accounts for approximately 60% of global photosynthetic carbon uptake (<xref ref-type="bibr" rid="B57">Mitchard, 2018</xref>). Their stability to drought plays a key role in maintaining the stability of subtropical forest ecosystems. According to the analysis results, there were certain differences in the ecosystem stability of the two types of forests, indicating that the stability of different biomes is different (<xref ref-type="bibr" rid="B65">Pennington and Lavin, 2016</xref>; <xref ref-type="bibr" rid="B17">Gazol et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B3">Anderegg et&#xa0;al., 2020</xref>). Current forest management and protection strategies (e.g. Natural forest resources protection, Returning farmland to forest, etc.) have made great improvement on carbon sequestration capability (<xref ref-type="bibr" rid="B41">Kong et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B85">Tong et&#xa0;al., 2020</xref>), and the increasing forest stand quality further enhanced the LUE and WUE of forest ecosystems (<xref ref-type="bibr" rid="B66">Ponce-Campos et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B20">Guo et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B30">Huang and Xia, 2019</xref>; <xref ref-type="bibr" rid="B73">Shao et&#xa0;al., 2022</xref>), leading to strong resistance and resilience to drought. There is no doubt that these strategies should continue to be implemented and widely promoted to provide strong support for subtropical forests to respond and adapt to climate change. However, whether the stability of evergreen forests could remain at current stage under the continuous increasing severe drought events should be further investigated in the future (<xref ref-type="bibr" rid="B14">Easterling et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B9">Dai, 2013</xref>; <xref ref-type="bibr" rid="B68">Reichstein et&#xa0;al., 2013</xref>).</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<title>5 Conclusion and recommendation</title>
<p>In this study, the FAO-PM algorithm was used to calculate SPEI data, and the Biome-BGC model was used to simulate carbon storage data. The spatiotemporal distribution characteristics of drought in the subtropical regions of China from 1952 to 2019 were analyzed, and the resistance and resilience of two types of forests to drought in the subtropical regions of China were quantified. The following conclusions were drawn:</p>
<list list-type="order">
<list-item>
<p>From 1952 to 2019, China&#x2019;s subtropical forests experienced five extreme drought events, with approximately one every 13.6 years. Two large-scale extreme drought events occurred after 1985 in 2009 and 2011, with 2011 being the year with the most severe and widespread drought. In the EBF, 19.3% suffered from extreme drought and 26.6% suffered from severe drought. In the ENF, 31.9% suffered from extreme drought and 21.2% suffered from severe drought.</p>
</list-item>
<list-item>
<p>From 1985 to 2019, the average carbon storage of vegetation in EBF and ENF in the subtropical region of China was 130.58 t&#xb7;hm<sup>-2</sup> and 78.49 t&#xb7;hm<sup>-2</sup>, respectively. From 2011 to 2015, the mean values of the overall change trend of carbon storage of the EBF and ENF in the subtropical region of China were -0.042 a<sup>-1</sup> and -0.013 a<sup>-1</sup>, respectively. The carbon storage of vegetation in both forests showed a slight downward trend, and the spatial distribution of changes in carbon storage was complex.</p>
</list-item>
<list-item>
<p>There were significant differences in the resistance of the two forest types to extreme drought, with EBF being significantly more resistant to drought than ENF in subtropical China, and EBF and ENF were broadly similar in resilience levels after drought. Therefore, the EBF is better adapted to drought in the subtropical region of China, and its high stability is mainly due to its high resistance to drought. The results indicating that better management level or extend the EBF plantation to increase the proportion of EBF in subtropical forest may enhance the resistance and resilience of the region to severe drought.</p>
</list-item>
</list>
</sec>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>MY: Data curation, Formal analysis, Investigation, Methodology, Validation, Preparation of the first draft. FM: Conceptualization, Methodology, Data Curation, Formal Analysis, Funding Acquisition, Review and editing. HD: Funding Acquisition, Supervision. XL: Methodology, Formal analysis, Data curation. QC: Formal analysis, Investigation. CN:Formal analysis, Investigation. ZH: Formal analysis, Investigation. YX: Formal analysis, Investigation. YG:Formal analysis, Investigation. KG:Formal analysis, Investigation. JS:Formal analysis, Investigation. CX:Formal analysis, Investigation. All authors contributed to the article and approved the submitted version.</p>
</sec>
</body>
<back>
<ack>
<title>Acknowledgments</title>
<p>The authors gratefully acknowledge the support of National Natural Science Foundation of China (No. 31901310, 32171785, 32201553), Leading Goose Project of Science Technology Department of Zhejiang Province (2023C02035), Scientific Research Project of Baishanzu National Park (2022JBGS02), the State Key Laboratory of Subtropical Silviculture Foundation (No. zy20180201).</p>
</ack>
<sec id="s8" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s9" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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