Carbon sink of soil inorganic carbon in arid regions and its contribution to carbon sequestration and emission reduction: A review
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摘要:研究目的
干旱区土壤无机碳作为全球碳循环举足轻重的组成部分，其碳汇效应不容忽视。
研究方法本文查阅了大量国内外干旱区土壤无机碳的相关文献，重点对土壤无机碳汇确认、碳库组成、来源识别，以及碳汇影响因素进行了系统性归纳总结。
研究结果干旱区无机碳汇效应伴随着干旱区负通量研究得到确认，但其碳库组成十分复杂，包括了液相碳库与固相碳库。其中液相储库主要以可溶性碳酸盐形式赋存于干旱区地下水体；固相储库则为以固相碳酸盐矿物的形式赋存在土壤中，依据不同成因来源分为成岩碳酸盐与成土碳酸盐，后者又细分为碳质成土碳酸盐与硅质成土碳酸盐。成土碳酸盐中的硅质成土碳酸盐具备真正长期稳定的碳汇效应。无机碳汇的影响因素复杂，包括了自然的气候、土壤性质与深度、生物作用、成土母质、土壤有机质等因素，以及土地利用与土地覆盖、农业管理措施等人为因素。
结论干旱区土壤无机碳对全球碳循环研究极其重要，当前研究主要聚焦在土壤无机碳来源分辨，碳汇效应强度确认与固碳潜力量化，以及影响因素明确与人为干预的可能性评估等方面。在实现“双碳目标”驱动下，查清干旱—半干旱地区土壤无机碳源汇过程与影响因素必将是未来的研究热点，也是解决“碳失汇”科学难题的突破点，极大地推动全球碳循环研究。
创新点:（1）干旱区土壤无机碳碳库组成与碳汇效应是近几十年来研究热点，作为碳中和路线中人为固碳端的重要组成，缺乏系统和完善的综述文献，本文总结了土壤无机碳碳汇确认过程、碳库组成、来源识别方法，为干旱区土壤无机碳在全球碳循环中的碳汇作用与贡献研究提供参考；（2）本文梳理归纳了干旱区土壤无机碳碳汇作用及影响因素，为土壤无机碳实现人为调控固碳增汇打下理论基础，为调控的方向和强度提供参考。
Abstract:This paper is the result of environmental geological survey engineering.
ObjectiveAs a pivotal component of the global carbon cycle, the role of soil inorganic carbon in arid regions as a carbon sink cannot be ignored.
MethodsThis paper reviewed a large amount of literature related to soil inorganic carbon in arid regions at home and abroad, and focused on the confirmation of soil inorganic carbon carbon sink, carbon pool composition, source identification, and carbon sink influencing factors in a systematic summary.
ResultsThe role of inorganic carbon carbon sinks in arid regions was confirmed along with the study of negative fluxes in arid regions, but the composition of its carbon pool is very complex, including liquid−phase carbon pools and solid−phase carbon pools. The liquid−phase reservoir is mainly in the form of Dissolved Inorganic Carbon in the groundwater of the arid regions; the solid−phase reservoir is the solid−phase Soil Inorganic Carbon in the soil profile, which is divided into Lithogenic Carbonate and Pedogenic Carbonate according to different genetic sources, and the latter is subdivided into carbonaceous soil−forming carbonate and silicic soil−forming carbonate . The SPC in PC has a real long−term stable carbon sink. The factors influencing inorganic carbon sinks are complex, including natural factors: climate, soil properties and depth, biological effects, soil−forming parent material, soil organic matter, etc.; anthropogenic factors: land use and land cover, agricultural management measures (irrigation and fertilization), etc.
ConclusionsSoil inorganic carbon in drylands is extremely important for global carbon sequestration, and current research focuses on the identification of soil inorganic carbon sources, confirmation of carbon sink strength and quantification of carbon sequestration potential, as well as the clarification of influencing factors and assessment of the possibility of human intervention. Driven by the goal of achieving carbon peaking and carbon neutrality goals, the identification of soil inorganic carbon sources and influencing factors will be a research hotspot in the future within the global region, especially in arid and semi−arid regions. It will be a breakthrough point to solve the scientific problem of "Missing carbon sink", which will greatly promote the research of Global Carbon Cycle.
Highlights:(1) The composition of soil inorganic carbon pool and the role of carbon sink in arid regions is an international research hotspot in recent decades. As an important component of the anthropogenic carbon sequestration end of the carbon neutral route, there is a lack of systematic and complete review literature about this. This paper summarizes the carbon sink confirmation process, carbon pool composition, source identification methods, to provide a reference for studying the role and contribution of soil inorganic carbon in the global carbon cycle in drylands. (2) This paper composes and summarizes the carbon carbon sink role and the influencing factors of soil inorganic carbon in arid regions, and lays a theoretical foundation for soil inorganic carbon to achieve anthropogenic regulation of carbon sequestration and sink enhancement, and provides reference for the direction and intensity of regulation.
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1. 研究目的（Objective）
湘中坳陷作为南方复杂构造区页岩气勘探的热点地区之一，也是中国油气勘探久攻未克的地区。前期在湘中地区北部的涟源凹陷泥盆系和石炭系获得了页岩气突破和发现，证实了湘中地区上古生界页岩气资源丰富。但对湘中地区南部的邵阳凹陷调查程度较为薄弱，针对邵阳凹陷二叠系仅开展了少量基础地质调查工作，页岩气资源潜力评价方面的工作尤为欠缺。本次研究依托邵阳湘邵地1井（XSD1井）钻探工程建立了邵阳凹陷二叠系地层层序序列，揭示了主要含气页岩层系的分布特征，获取了含气性评价参数，对湘中地区二叠系页岩气勘探开发和重新评价湘中坳陷页岩气资源潜力具有重要的现实意义。
2. 研究方法（Methods）
中国地质调查局武汉地质调查中心在收集分析区域地质相关资料的基础上，结合邵阳凹陷短陂桥向斜的煤田浅钻、非震物探等资料开展页岩气地质综合评价，采用页岩埋深500~4500 m，页岩有机碳含量≥1.0%，页岩厚度≥15 m，页岩有机质热演化程度1.0%~3.5%的评价参数在短陂桥向斜区优选页岩气远景区，论证部署了1口小口径页岩气地质调查井—XSD1井，湖南煤田地质勘查有限公司组织实施钻探（图 1a）。该井采样全井段取心钻井工艺，测井选取PSJ-2数字测井系统，录井采用SK-2000G气测录井，钻获二叠系大隆组156.05 m（暗色硅质页岩、钙质泥岩94.48 m），龙潭组349.95 m（暗色泥岩216.93 m，粉砂质泥岩36.9 m），对这两套层系共采集暗色泥岩样品33件，进行解析气含量测定分析，落实了含气性评价参数。
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图 1 湘邵地1井构造位置图（a）、主要含气层段岩性和含气性参数柱状图（b）、以及富集模式图（c）Figure 1. Structural location of Xiangshaodi 1 well (a), lithology and gas-bearing parameters of main gas bearing zones (b), and enrichment mode (c)3. 结果（Results）
本次样品分析工作由武汉地质调查中心古生物与生命-环境协同演化重点实验室完成，采用YSQ-IIIA岩石解析气测定仪（燃烧法）对含气段岩心共计33件样品进行分析。该井钻获二叠系大隆组厚度156.05 m，为一套硅质岩、硅质页岩、炭质钙质泥岩地层。其中在井深842~930.2 m硅质页岩、钙质泥岩段，气测全烃值从1.06%上升至16.54%，甲烷值从1.01%上升至14.04%，13件大隆组硅质页岩现场解析总含气量为1.29~9.97 m3/t，平均4.85 m3/t。实现了湘中坳陷二叠系页岩气新发现，有效拓展了华南地区大隆组勘探范围。
钻获龙潭组厚度349.95 m，上段为一套细砂岩、粉砂岩夹泥岩潮坪相沉积地层，下段为一套炭质泥岩、粉砂质泥岩夹薄层细砂岩泻湖相沉积地层。在井深1013.4~1048 m泥岩与粉砂岩互层段气测全烃值最高可达19.87%，甲烷值最高为16.94%，7件泥岩与粉砂岩样品现场解析总含气量0.57~3.42 m3/t，平均1.78 m3/t；井深1088.10~1199.75 m泥岩夹泥质粉砂岩含气层111.6 m，气测全烃值最高可达28.2%，甲烷值最高为23.6%，13件泥岩、粉砂质泥岩样品现场解析总含气量0.90~4.55 m3/t，平均2.01 m3/t（图 1b），首次查明了湘中坳陷二叠系龙潭组非常规油气分布特点。
通过区域地质背景分析，并结合煤田区域地质资料，本研究认为滑脱断裂（F9）上下盘具有不同的页岩气聚集条件。滑脱断裂之上由一系列的同向逆断层形成的逆冲推覆体，地层变形强烈，且裂缝发育，导致页岩气保存条件变差。滑脱断裂下盘是页岩气主要富集区，地层平缓，不发育次级通天断裂，与下盘地层形成反向遮挡，易形成封闭，保存条件良好（图 1c）。
4. 结论（Conclusions）
（1）二叠系大隆组岩性以硅质岩、硅质页岩为主，夹少量灰岩。主要含气段存在于上段硅质页岩段，厚88.2 m，含气量平均为4.85 m3/t，含气性优越，资源潜力大。
（2）二叠系龙潭组上段以致密砂岩气为主，含气量平均为1.78 m3/t；下段以页岩气为主，泥岩厚达177.47 m，含气量平均为2.01 m3/t，具有泥岩厚度大，含气性好等特征。
（3）保存条件是页岩气富集关键，构造改造弱的封闭演化环境有利于页岩气保存，研究区滑脱断裂下盘是页岩气主要富集区，易形成封闭，保存条件良好。
（4）湘邵地1井在二叠系大隆组和龙潭组获得良好的页岩气显示，证实了湘中地区二叠系具有良好的页岩气资源潜力，对湘中地区页岩气资源潜力评价具有重要意义。
5. 基金项目（Fund support）
本文为中国地质调查局项目“中扬子地区油气页岩气调查评价”（DD20221659）资助的成果。
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图 1 全球碳库与碳失汇示意图（据Lal，2019；Naorem et al., 2022修改）
Pg=1015g；人类通过化石燃料排放CO2进入大气的量约5.4 Pg C a−1，破坏森林排放进入大气的量约1.6 Pg C a−1，海洋吸收大气CO2的量约 2 Pg C a−1，大气CO2增加的量约3.3 Pg C a−1，这之间存在着−1.7 Pg C a−1的差值，即5.4 Pg +1.6 Pg −2 Pg −3.3 Pg =1.7 Pg，这部分CO2去向不明，故被称为碳失汇（据方精云和郭兆迪，2007）
Figure 1. Schematic diagram of global carbon pool and missing carbon sink (modified from Lal, 2019; Naorem et al., 2022)
Pg = 1015g; there is a difference of −1.7 Pg a−1 between human CO2 emissions into the atmosphere through fossil fuels of about 5.4 Pg a−1, forest destruction emissions into the atmosphere of about 1.6 Pg a−1, ocean absorption of atmospheric CO2 of about 2 Pg a−1, and increase in atmospheric CO2 of about 3.3 Pg a−1, which is 5.4 Pg + 1.6 Pg − 2 Pg −3.3 Pg =1.7 Pg, and this part of CO2 goes to an unknown destination, which is called missing carbon sink (modified from Fang Jingyun and Guo Zhaodi, 2007)
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图 2 全球土壤无机碳密度分布图及其与年平均降水量低值区的相关性（据Zamanian et al., 2016修改）
Figure 2. Global soil inorganic carbon carbon density distribution map and its correlation with the low annual mean precipitation area (modified from Zamanian et al., 2016)
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图 3 荒漠土壤CO2负通量的对比实验验证（据Ma et al., 2013修改）
Figure 3. Comparative experimental validation of negative CO2 fluxes in desert soils (modified from Ma et al., 2013)
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图 4 土壤无机碳固相、液相储库与转移转化过程示意图
Figure 4. Schematic diagram of soil inorganic carbon solid phase and liquid phase reservoirs and transfer transformation processes
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图 5 土壤无机碳库组成、来源辨析与碳汇作用示意图
Figure 5. Schematic diagram of soil inorganic carbon pool composition, source identification and carbon sink role
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图 6 塔里木盆地土壤无机碳液相储库DIC淋溶传输（据Li et al., 2015）
a—DIC淋溶传输示意图；b—DIC14C测年年龄等值线图；c—塔里木盆地边缘至中心的地下水DIC年龄
Figure 6. DIC leaching transport of soil inorganic carbon liquid phase reservoirs in the Tarim Basin (after Li et al., 2015)
a−Schematic diagram of DIC leaching transport; b−DIC14C dating age contour map; c−DIC age of groundwater from the edge to the center of the Tarim Basin
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图 7 景观与剖面尺度中原生碳酸盐与次生碳酸盐赋存形态区别
a—成岩碳酸盐景观与剖面照片，包含灰岩地层原位风化的LC以及冲积搬运的LC碎屑（Monger et al.，2015）；b、e—剖面上的PC钙质菌丝体与胶膜（Zamanian et al.，2016；达佳伟，2020）；c、f—剖面上的PC钙结核（Zamanian et al.，2016；达佳伟，2020）；d—剖面中的PC层状钙板（达佳伟，2020）；g—黄土中的PC钙质根石（Zamanian et al.，2016）
Figure 7. Difference between primary and secondary carbonate fugacity patterns in landscape and profile scale
a−Photos of mafic carbonate landscapes and profiles containing in situ weathered LC from tuff strata and alluvially transported LC debris (Monger et al., 2015); b, e−PC calcareous mycelium with colloidal film in profiles (Zamanian et al., 2016; Da Jiawei, 2020); c, f−PC calcium nodules in profiles (Zamanian et al., 2016; Da Jiawei, 2020); d−PC laminated calcium plates in the profile (Da Jiawei, 2020); g−PC calcareous rhizoliths in the loess (Zamanian et al., 2016)
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图 8 微观尺度中原生碳酸盐与次生碳酸盐赋存形态区别
a—钙结核土壤基质中混入的LC颗粒（刘恋等，2010）；b—含海洋化石的LC颗粒（Monger et al.，2015）；c—钙结核内部PC，包括分布于土壤孔隙中的亮晶方解石以及土壤基质中的泥晶方解石（达佳伟，2020）；d—土壤孔隙璧上的PC钙膜（Zamanian et al.，2016）；e—钙结核中围绕原生石英及长石颗粒生长的PC（达佳伟，2020）；f—假菌丝体中的针状方解石（Monger et al.，2015）；g—镜下PC结核以及碎屑表面PC钙膜（Zamanian et al.，2005）；h—钙化根，可见细胞结构（Zamanian et al.，2016）；i—蚯蚓球晶，正交偏光下（Zamanian et al.，2016）
Figure 8. Difference between primary and secondary carbonate fugitive morphology in microscale
a−LC particles mixed in the soil matrix of calcium nodules (Liu Lian et al., 2010); b−LC particles containing marine fossils (Monger et al., 2015); c−PC inside calcium nodules, including leucocrystalline calcite distributed in soil pores and mud−crystalline calcite in the soil matrix (Da Jiawei, 2020); d−PC calcium film on soil pore jams (Zamanian et al., 2016); e−PC growing around primary quartz and feldspar grains in calcium nodules (Da Jiawei, 2020); f−Nneedle−like calcite in pseudomycorrhiza (Monger et al., 2015); g−Microscopic PC nodules as well as PC calcium films on the surface of debris (Zamanian et al., 2005); h−Calcified roots with visible cellular structures (Zamanian et al., 2016); i− Earthworm spherical crystals, under orthogonal polarization (Zamanian et al., 2016)
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图 9 不同来源成土碳酸盐风化与固碳过程示意图（据Monger et al.，2015修改）
Figure 9. Schematic diagram of carbonate weathering and carbon sequestration processes in different sources of soil−forming carbonates (modified from Monger et al., 2015)
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图 10 碳酸盐世代分类示意图（据Monger et al.，2015修改）
Figure 10. Schematic diagram of carbonate generation classification (modified from Monger et al., 2015)
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图 11 不同pH下溶液中3种碳酸的比例变化曲线
Figure 11. Variation curve of the ratio of three carbonic acids in solution of different pH
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图 12 土壤无机碳影响因素关系示意图
Figure 12. Schematic diagram of the factors influencing soil inorganic carbon
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图 13 土壤无机碳溶解损失与源汇效应示意图
Figure 13. Diagram of SIC dissolution loss and source−sink effect
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图 14 植物根系促进周围PC沉淀以及钙质根石（Rhizoliths）形成过程示意图（据Zamanian et al., 2016修改）
Figure 14. Schematic diagram of the process of PC precipitation surrounding plant roots and formation of Rhizoliths (modified from Zamanian et al., 2016)
表 1 全球干旱荒漠区CO2负通量报告统计（据Schlesinger，2017修改）
Table 1 Global negative CO2 flux reporting statistics for arid desert regions (modified from Schlesinger, 2017)
研究地点 年平均
降水量/
mm负通量/
(g C m−2 a−1)文献来源 墨西哥，下加利福尼亚州 174 39~52 Hastings et al., 2005 美国，莫哈韦沙漠 150 102~127 Jasoni et al., 2005；Wohlfahrt et al., 2008 中国，古尔班通古特沙漠 160 62~622 Xie et al., 2009 173 49 Liu et al., 2012 164 25 Ma et al., 2014 中国，宁夏 287 77 Jia et al., 2014 275 28 Liu et al., 2015a 中国，塔里木盆地 21 Li et al., 2015 
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表 2 基于δ13C值估算的不同地区土壤无机碳PC与LC占比
Table 2 Estimated percentage of soil inorganic carbon PC and LC in different regions based on δ13C values
地点 PC占比/
%PC含量/(g/kg) 文献来源 塔里木盆地阿拉尔垦区 1.33~35.7 1.34~56.36 李杨梅等，2018 内蒙古乌兰察布 17.4~83.6 42~177 张林等，2010 俄罗斯 20~50 / Morgun et al., 2008 俄罗斯 66.8~73.8 26.9~60.1 Ryskov et al., 2008 美国德克萨斯州 2~11，
9~20，
60~70，
17~100/ Nordt et al., 1998 美国德克萨斯州 40~90 / Rabenhorst et al., 1983 以色列 30~60 / Magaritz and Amiel, 1980
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表 3 基于87Sr/86Sr值估算的不同地区土壤无机碳SPC与CPC占比
Table 3 Soil inorganic carbon SPC and CPC shares in different regions estimated based on 87Sr/86Sr values
地点 PC钙源/SPC占比 成土母质 文献来源 定
性摩洛哥阿特拉斯 来自降水以及基岩风化 玄武岩 Hamidi et al., 2001 智利阿塔卡马沙漠 沿海地区主要来自海洋气溶胶输入（>50%)，
大陆内部主要来自基岩风化岩浆岩+灰岩+碎屑岩 Rech et al., 2003 印度西高止山脉 来自非钙质基岩风化以及风沙 片麻岩+花岗岩+绿岩 Durand et al., 2006 定
量西班牙两座火山岛 极少 岩浆岩+砂岩+风积物 Huerta et al., 2015 美国新墨西哥州 <2% 非钙质冲积物 Capo and Chadwick，1999 西班牙中部 2.7% ~ 7.8% 花岗岩 Chiquet et al., 1999 澳大利亚中南部 ~10% 花岗岩、玄武岩、角闪岩、绿岩、页岩 Dart et al., 2007 夏威夷岛 33% 玄武岩 Whipkey et al., 2000 美国西南部 39~58% 岩浆岩+灰岩 Naiman et al., 2000 印度南部 24%~82% 片麻岩+超镁质岩 Violette et al., 2010 喀麦隆远北地区 >50% 花岗岩+绿岩 Dietrich et al., 2017 注：定性指研究中基于87Sr/86Sr值推断PC的主要钙源，并未得到明确SPC占比的；定量指研究中基于87Sr/86Sr值得出明确SPC占比或来自硅酸岩风化钙源贡献比例的。
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表 4 不同深度土壤中土壤无机碳占比
Table 4 Percentage of soil inorganic carbon in soils at different depths
研究地点 不同深度SIC分布结论 文献来源 中国 1~3 m储量占比54.9%~88.5% Li et al., 2007 内蒙古 1~3 m储量占比>50% Wang et al., 2010 兰州 1~2 m储量占比50% Zhang et al., 2015 西班牙东南部 1~2 m储量占比51% Dı́az−Hernández et al., 2003 新疆 1 m以下储量占比>80%，3 m以下储量占比>50% Wang et al., 2013a 加拿大萨斯喀彻温省 深层土壤（C层）储量占比几乎100% Landi et al., 2003 内蒙古 0~30 cm、30~100 cm储量占比15%、85% Wang et al., 2013b 黄土高原 0~20 cm、20~50 cm、50~100 cm储量均值2.39、2.92、4.89 Pg Tan et al., 2014 新疆 0~30 cm、30~100 cm密度均值：耕地11.0、30.9 kg C m−2，灌木林地9.8、27.0 kg C m−2 Wang et al., 2015c 青藏高原 0~30 cm、0~50 cm、0~100 cm密度均值5.70、9.10、13.46 kg C m−2 Yang et al., 2010 黄河三角洲 0~20 cm、80~100 cm含量均值为10.48 g∙kg−1、12.72 g∙kg−1 Guo et al., 2016 伊朗西北部 表土平均含量（A层）：7.9%；深层平均含量（C层）：21.2% Raheb et al., 2017
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表 5 不同地区土壤无机碳与土壤有机碳相关性汇总
Table 5 Summary of correlation between soil inorganic carbon and soil organic carbon in different regions
研究地点 SIC与SOC
相关性结论文献来源 甘肃省河西走廊中部 正相关 Su et al., 2010 以色列贝特谢安 Tamir et al., 2012 内蒙古、西藏 Shi et al., 2012 新疆焉耆盆地 Wang et al., 2014 新疆焉耆盆地 Wang et al., 2015b 新疆焉耆盆地 Wang et al., 2015c 黄河三角洲 Guo et al., 2016 中国河北衢州
黄土高原Bughio et al., 2016
Tong et al., 2020室内实验 负相关 Demoling et al., 2007 宁夏 Liu et al., 2014 兰州 Zhang et al., 2015 云南 Li et al., 2016 黄土高原 Zhao et al., 2016 黑河流域 Yang et al., 2018 青藏高原 Du and Gao, 2020 内蒙古、西藏 不相关 Shi et al., 2012 华北平原 Lu et al., 2020 西班牙巴塞罗那 PC与SOC碳
同位素线性相关，
具备成因联系Rovira and Vallejo，2008 美国华盛顿州 Stevenson et al., 2005
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表 6 土地利用类型变化与不同土地覆盖对SIC的影响
Table 6 Effect of land use type change and different land cover on SIC
研究地点 研究结论 文献来源 黄土高原 耕地−草地：SIC减少 Liu et al., 2014 美国南部阿根廷潘帕斯草原 自然土地−灌溉耕地，SIC减少 Kim et al., 2020 俄罗斯库尔斯克 自然土地−灌溉耕地：SIC增加 Mikhailova and Post, 2006 新疆焉耆盆地 Wang et al., 2015b 新疆焉耆盆地 Wang et al., 2015c 美国西北部蛇河平原 Entry et al., 2004 中国 Wu et al., 2009 美国西南部 Nyachoti et al., 2019 新疆塔里木盆地 干旱盐碱地−灌溉耕地：SIC增加 Li et al., 2015 甘肃省河西走廊中部 沙地−灌木、林地、耕地：SIC增加 Su et al., 2010 华北平原 普通耕地−集约化种植耕地：SIC增加 Lu et al., 2020 综述 草地−耕地：SIC增加；沙地−林地：SIC增加；草地、耕地−林地：SIC减少 An et al., 2019 黄土高原 SIC密度：农田=草地>森林 Tan et al., 2014 中国 SIC密度：沙漠、草原>灌木丛、耕地>沼泽、森林、草甸 Mi et al., 2008 内蒙古 SIC密度：沙漠>灌木沙漠>灌木−草原>森林、草原 Wang et al., 2010 中国西北部黑河流域 SIC密度：温带草原>高山草甸 Yang et al., 2018
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表 7 灌溉对土壤SIC的影响及可能原因总结
Table 7 Summary of the effect of irrigation in soil SIC and possible causes
相关性 可能原因 相关文献 研究地点 正相关 灌溉提供溶液环境，加强生物活动，
输入钙镁离子以及DIC，促进PC形成Wu et al., 2009 中国 Zhang et al., 2015 兰州 Bughio et al., 2016 河北衢州 Entry et al., 2004 美国西北部蛇河平原 Li et al., 2015 新疆塔里木盆地 Wang et al., 2016 美国新墨西哥州 Nyachoti et al., 2019 美国西南部 负相关 低EC（碳酸钙不饱和）的大量灌溉
水易造成SIC溶解流失Wu et al., 2008 加利福尼亚 Schindlbacher et al., 2019 德国拜罗伊特市 Kim et al., 2020 美国南部大平原，
阿根廷潘帕斯草原
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