中国石松类和蕨类植物多样性研究进展

doi:10.17520/biods.2022381

生物多样性 ›› 2022, Vol. 30 ›› Issue (7): 22381. DOI: 10.17520/biods.2022381

所属专题：物种形成与系统进化

• 纪念第19届国际植物学大会召开5周年专题 • 上一篇下一篇

中国石松类和蕨类植物多样性研究进展

王婷¹, 舒江平¹, 顾钰峰¹, 李艳清¹, 杨拓¹, 徐洲锋³, 向建英², 张宪春⁴^,^*(), 严岳鸿¹^,^*()

1.深圳市兰科植物保护研究中心兰科植物保护与利用国家林业和草原局重点实验室, 广东深圳 518114
2.西南林业大学生物多样性保护学院, 昆明 650224
3.中国科学院华南植物园, 广州 510650
4.中国科学院植物研究所系统与进化植物学国家重点实验室, 北京 100093

收稿日期:2022-07-07 接受日期:2022-07-20 出版日期:2022-07-20 发布日期:2022-07-20
通讯作者: 张宪春,严岳鸿
作者简介:zhangxc@ibcas.ac.cn
E-mail: yhyan@sibs.ac.cn;
第一联系人：
^# 共同第一作者
基金资助:
国家自然科学基金(32170216);广东省基础与应用基础研究基金(2021A1515010920)

Insight into the studies on diversity of lycophytes and ferns in China

Ting Wang¹, Jiangping Shu¹, Yufeng Gu¹, Yanqing Li¹, Tuo Yang¹, Zhoufeng Xu³, Jianying Xiang², Xianchun Zhang⁴^,^*(), Yuehong Yan¹^,^*()

1. Key Laboratory of National Forestry and Grassland Administration for Orchid Conservation and Utilization, The Orchid Conservation and Research Centre of Shenzhen, Shenzhen, Guangdong 518114
2. College of Biodiversity Conservation, Southwest Forestry University, Kunming 650224
3. South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650
4. State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093

Received:2022-07-07 Accepted:2022-07-20 Online:2022-07-20 Published:2022-07-20
Contact: Xianchun Zhang,Yuehong Yan
About author:First author contact:
^# These authors contributed equally to this work

摘要/Abstract

摘要：

石松类和蕨类植物是维管植物的第二大类群, 其起源可追溯到4亿年前。在被子植物出现之前, 石松类和蕨类植物在古地球生态系统中占主导地位, 其重要性一直延续到现在。自20世纪40年代开始, 中国石松类和蕨类植物研究就令世界瞩目, 尤其是2017年第19届国际植物学大会在中国深圳召开后的5年时间里, 中国石松类和蕨类植物研究更是面向世界、走向国际, 研究更为广泛的科学问题, 在物种多样性、保护、系统演化和生态适应性等方面取得了一系列重要研究进展。2017-2022年, 多个中国研究团队利用多组学数据构建了世界石松类和蕨类植物科级水平的生命之树并提出了关键性状孢子囊环带演化的新模式; 解决了石松类和蕨类植物中目级、科级、属级和种级众多关键的系统分类学等问题, 发表了106个新分类群; 开展了大量的植物区系调查和研究, 出版了6部中国石松类和蕨类植物多样性专著和1部世界性专著; 对65种国家重点保护的石松类和蕨类植物进行了迁地保护, 同时实现了桫椤科、水蕨属(Ceratopteris)、观音座莲属(Angiopteris)和鹿角蕨(Platycerium wallichii)等重点保护类群的孢子繁殖; 在系统发育框架下, 研究了石松类和蕨类植物的生态修复功能和生态适应性演化。通过对2017-2022年研究成果的总结和思考, 本文对未来石松类和蕨类植物的发展提出以下建议: (1)提高中国寡型科属以及世界性大科大属的关注力度; (2)加强西藏、四川等薄弱地区石松类和蕨类植物的调查研究, 并结合新技术, 如DNA条形码等以提高区系调查中物种鉴别的效率和准确性; (3)运用多学科交叉的研究方法厘清各科、属、种间系统关系的同时, 还应加强系统和生态适应性演化之间的协同研究; (4)关注石松类和蕨类植物系统位置作为陆生维管植物演化起点的共性科学问题; (5)加强石松类和蕨类植物系统分类学与生态学、植物化学、保护生物学等学科间交叉合作研究。

关键词: 维管植物, 新分类群, 国家重点保护植物, 系统发育

Abstract

Background: Lycophytes and ferns are the second largest group of vascular plants, dating back as far as 400 million years ago. Before the domination of angiosperms, lycophytes and ferns were the key component of ancient terrestrial ecosystems, and are still important to modern ecosystems today. Since the 1940s, the study of lycophytes and ferns in China has attracted worldwide attention. In the past five years since the 19th International Botanical Congress (IBC) held in Shenzhen in 2017, research on lycophytes and ferns in China has expanded internationally and focuses on a wider range of scientific issues. A series of important advances have been made on plant diversity, conservation, phylogeny, and ecological adaptation.
Results: From 2017 to 2022, the tree of life for lycophytes and ferns in the world has been reconstructed by the family level, and a new evolutionary model for the sporangium was proposed. Some of the phylogenetic and taxonomic problems of order, family, genus and species of lycophytes and ferns were solved, meanwhile 106 new taxa were published. A large number of floristic collections and studies have been conducted, and six Chinese plus one worldwide monographs on the diversity of lycophytes and ferns have been published. In addition, 65 species of lycophytes and ferns under state priority conservation have been ex situ protected, and spore reproduction has been achieved in priority conservation groups such as Cyatheaceae, Ceratopteris, Angiopteris and Platycerium wallichiia. The ecological restoration and adaptive evolution of lycophytes and ferns were studied under the phylogeny.
Perspectives: After literature survey on the research of lycophytes and ferns in China from 2017 to 2022, we make the following suggestions for future studies: (1) Focus future research on Chinese oligotypic families/genera and the worldwide mega-diverse families/genera. (2) Prioritize research in under-studied areas, such as Tibet and Sichuan, and utilize new technologies (e.g., DNA barcoding) to improve the efficiency and accuracy of species identification. (3) When clarifying the systematic relationship between families, genera, and species by using multi-disciplinary approaches, it is necessary to conduct research on plant evolutionary history and adaptation mechanisms. (4) Research efforts should focus on the common scientific conjecture of the systematic position of lycophytes and fern as the origin for the evolution of terrestrial vascular plants. (5) Pay more attention to the cooperation among the phylogenetics, ecology, phytochemistry, conservation biology and other subjects of lycophytes and ferns.

Key words: vascular plants, new taxa, wild plants under state priority conservation, phylogeny

王婷, 舒江平, 顾钰峰, 李艳清, 杨拓, 徐洲锋, 向建英, 张宪春, 严岳鸿 (2022) 中国石松类和蕨类植物多样性研究进展. 生物多样性, 30, 22381. DOI: 10.17520/biods.2022381.

Ting Wang, Jiangping Shu, Yufeng Gu, Yanqing Li, Tuo Yang, Zhoufeng Xu, Jianying Xiang, Xianchun Zhang, Yuehong Yan (2022) Insight into the studies on diversity of lycophytes and ferns in China. Biodiversity Science, 30, 22381. DOI: 10.17520/biods.2022381.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2022381

https://www.biodiversity-science.net/CN/Y2022/V30/I7/22381

图/表 10

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科名 Famliy			世界物种数 No. of species in the world (a)	世界新增物种数 No. of new species in the world (b)		中国物种数No. of species in China			中国新增物种数 No. of new species in China		世界新增物种数/世界物种数 b/a	中国期望新增物种数 Expected values of new species in China	实际值与期望值的比较 Comparison of actual and expected values
石松科	Lycopodiaceae		388	42		69			3		0.11	7.47	0
卷柏科	Selaginellaceae		700	32		73			8		0.05	3.34	1
水韭科	Isoetaceae		250	22		5			4		0.09	0.44	1
木贼科	Equisetaceae		15	2		10			0		0.13	1.33	0
松叶蕨科	Psilotaceae		17	0		1			0		0.00	0.00	0
瓶尔小草科	Ophioglossaceae		112	13		22			0		0.12	2.55	0
合囊蕨科	Marattiaceae		111	19		30			1		0.17	5.14	0
紫萁科	Osmundaceae		18	5		8			0		0.28	2.22	0
膜蕨科	Hymenophyllaceae		434	44		51			2		0.10	5.17	0
马通蕨科	Matoniaceae		4	0		NA			NA		0.00	NA	NA
双扇蕨科	Dipteridaceae		11	1		5			1		0.09	0.45	1
里白科	Gleicheniaceae		157	7		16			1		0.04	0.71	1
海金沙科	Lygodiaceae		40	0		9			0		0.00	0.00	0
莎草蕨科	Schizaeaceae		35	6		2			0		0.17	0.34	0
双穗蕨科	Anemiaceae		115	0		NA			NA		0.00	NA	NA
槐叶蘋科	Salviniaceae		21	2		5			0		0.10	0.48	0
蘋科	Marsileaceae		61	2		3			0		0.03	0.10	0
伞序蕨科	Thyrsopteridaceae		1	0		NA			NA		0.00	NA	NA
柱囊蕨科	Loxsomataceae		2	0		NA			NA		0.00	NA	NA
垫囊蕨科	Culcitaceae		2	0		NA			NA		0.00	NA	NA
瘤足蕨科	Plagiogyriaceae		15	0		8			0		0.00	0.00	0
金毛狗科	Cibotiaceae		9	0		2			0		0.00	0.00	0
丝囊蕨科	Metaxyaceae		6	0		NA			NA		0.00	NA	NA
蚌壳蕨科	Dicksoniaceae		35	8		NA			NA		0.23	NA	NA
桫椤科	Cyatheaceae		643	127		14			2		0.20	2.77	0
袋囊蕨科	Saccolomataceae		18	0		NA			NA		0.00	NA	NA
花楸蕨科	Cystodiaceae		1	0		NA			NA		0.00	NA	NA
番茄蕨科	Lonchitidaceae		2	0		NA			NA		0.00	NA	NA
鳞始蕨科	Lindsaeaceae		234	0		17			0		0.00	0.00	0
凤尾蕨科	Pteridaceae		1,211	579		235			10		0.48	112.36	0
碗蕨科	Dennstaedtiaceae		265	58		53			0		0.22	11.60	0
冷蕨科	Cystopteridaceae		37	0		20			0		0.00	0.00	0
轴果蕨科	Rhachidosoraceae		8	0		5			0		0.00	0.00	0
肠蕨科	Diplaziopsidaceae		4	0		3			0		0.00	0.00	0
链脉蕨科	Desmophlebiaceae		2	0		NA			NA		0.00	NA	NA
半网蕨科	Hemidictyaceae		1	0		NA			NA		0.00	NA	NA
铁角蕨科	Aspleniaceae		730	153		108			13		0.21	22.64	0
岩蕨科	Woodsiaceae		39	86		24			1		2.21	52.92	0
球子蕨科	Onocleaceae		5	0		4			0		0.00	0.00	0
乌毛蕨科	Blechnaceae		265	40		14			0		0.15	2.11	0
蹄盖蕨科	Athyriaceae		650	7		282			5		0.01	3.04	1
金星蕨科	Thelypteridaceae		1,034	644		199			2		0.62	123.94	0
翼盖蕨科	Didymochlaenaceae		1	0		1			0		0.00	0.00	0
肿足蕨科	Hypodematiaceae		22	7		13			7		0.32	4.14	1
鳞毛蕨科	Dryopteridaceae		2,115	161		496			34		0.08	37.76	0
肾蕨科	Nephrolepidaceae		19	0		5			0		0.00	0.00	0
藤蕨科	Lomariopsidaceae	69			59		4	2		0.86		3.42		0
三叉蕨科	Tectariaceae	250			1		42	1		0.00		0.17		1
蓧蕨科	Oleandraceae	15			4		5	0		0.27		1.33		0
骨碎补科	Davalliaceae	65			10		17	0		0.15		2.62		0
水龙骨科	Polypodiaceae	1,652			594		267	5		0.36		96.00		0

科名 Famliy			世界物种数 No. of species in the world (a)	世界新增物种数 No. of new species in the world (b)		中国物种数No. of species in China			中国新增物种数 No. of new species in China		世界新增物种数/世界物种数 b/a	中国期望新增物种数 Expected values of new species in China	实际值与期望值的比较 Comparison of actual and expected values
石松科	Lycopodiaceae		388	42		69			3		0.11	7.47	0
卷柏科	Selaginellaceae		700	32		73			8		0.05	3.34	1
水韭科	Isoetaceae		250	22		5			4		0.09	0.44	1
木贼科	Equisetaceae		15	2		10			0		0.13	1.33	0
松叶蕨科	Psilotaceae		17	0		1			0		0.00	0.00	0
瓶尔小草科	Ophioglossaceae		112	13		22			0		0.12	2.55	0
合囊蕨科	Marattiaceae		111	19		30			1		0.17	5.14	0
紫萁科	Osmundaceae		18	5		8			0		0.28	2.22	0
膜蕨科	Hymenophyllaceae		434	44		51			2		0.10	5.17	0
马通蕨科	Matoniaceae		4	0		NA			NA		0.00	NA	NA
双扇蕨科	Dipteridaceae		11	1		5			1		0.09	0.45	1
里白科	Gleicheniaceae		157	7		16			1		0.04	0.71	1
海金沙科	Lygodiaceae		40	0		9			0		0.00	0.00	0
莎草蕨科	Schizaeaceae		35	6		2			0		0.17	0.34	0
双穗蕨科	Anemiaceae		115	0		NA			NA		0.00	NA	NA
槐叶蘋科	Salviniaceae		21	2		5			0		0.10	0.48	0
蘋科	Marsileaceae		61	2		3			0		0.03	0.10	0
伞序蕨科	Thyrsopteridaceae		1	0		NA			NA		0.00	NA	NA
柱囊蕨科	Loxsomataceae		2	0		NA			NA		0.00	NA	NA
垫囊蕨科	Culcitaceae		2	0		NA			NA		0.00	NA	NA
瘤足蕨科	Plagiogyriaceae		15	0		8			0		0.00	0.00	0
金毛狗科	Cibotiaceae		9	0		2			0		0.00	0.00	0
丝囊蕨科	Metaxyaceae		6	0		NA			NA		0.00	NA	NA
蚌壳蕨科	Dicksoniaceae		35	8		NA			NA		0.23	NA	NA
桫椤科	Cyatheaceae		643	127		14			2		0.20	2.77	0
袋囊蕨科	Saccolomataceae		18	0		NA			NA		0.00	NA	NA
花楸蕨科	Cystodiaceae		1	0		NA			NA		0.00	NA	NA
番茄蕨科	Lonchitidaceae		2	0		NA			NA		0.00	NA	NA
鳞始蕨科	Lindsaeaceae		234	0		17			0		0.00	0.00	0
凤尾蕨科	Pteridaceae		1,211	579		235			10		0.48	112.36	0
碗蕨科	Dennstaedtiaceae		265	58		53			0		0.22	11.60	0
冷蕨科	Cystopteridaceae		37	0		20			0		0.00	0.00	0
轴果蕨科	Rhachidosoraceae		8	0		5			0		0.00	0.00	0
肠蕨科	Diplaziopsidaceae		4	0		3			0		0.00	0.00	0
链脉蕨科	Desmophlebiaceae		2	0		NA			NA		0.00	NA	NA
半网蕨科	Hemidictyaceae		1	0		NA			NA		0.00	NA	NA
铁角蕨科	Aspleniaceae		730	153		108			13		0.21	22.64	0
岩蕨科	Woodsiaceae		39	86		24			1		2.21	52.92	0
球子蕨科	Onocleaceae		5	0		4			0		0.00	0.00	0
乌毛蕨科	Blechnaceae		265	40		14			0		0.15	2.11	0
蹄盖蕨科	Athyriaceae		650	7		282			5		0.01	3.04	1
金星蕨科	Thelypteridaceae		1,034	644		199			2		0.62	123.94	0
翼盖蕨科	Didymochlaenaceae		1	0		1			0		0.00	0.00	0
肿足蕨科	Hypodematiaceae		22	7		13			7		0.32	4.14	1
鳞毛蕨科	Dryopteridaceae		2,115	161		496			34		0.08	37.76	0
肾蕨科	Nephrolepidaceae		19	0		5			0		0.00	0.00	0
藤蕨科	Lomariopsidaceae	69			59		4	2		0.86		3.42		0
三叉蕨科	Tectariaceae	250			1		42	1		0.00		0.17		1
蓧蕨科	Oleandraceae	15			4		5	0		0.27		1.33		0
骨碎补科	Davalliaceae	65			10		17	0		0.15		2.62		0
水龙骨科	Polypodiaceae	1,652			594		267	5		0.36		96.00		0

类别 Classification	总数 Total	极危 Critically Endangered	濒危Endangered	易危Vulnerable	近危 Near Threatened	无危 Least Concern	数据缺乏 Data Deficient
中国石松类和蕨类植物 Lycophytes and ferns in China	2,244	43	68	71	66	1,124	872
国家重点保护野生石松类和蕨类植物 The wild lycophytes and ferns under state priority conservation	128	25	8	63	8	5	17
国家重点保护野生石松类和蕨类植物中的迁地保护植物 Ex situ conservation of wild lycophytes and ferns under state priority conservation	62	14	4	27	5	2	7

类别 Classification	总数 Total	极危 Critically Endangered	濒危Endangered	易危Vulnerable	近危 Near Threatened	无危 Least Concern	数据缺乏 Data Deficient
中国石松类和蕨类植物 Lycophytes and ferns in China	2,244	43	68	71	66	1,124	872
国家重点保护野生石松类和蕨类植物 The wild lycophytes and ferns under state priority conservation	128	25	8	63	8	5	17
国家重点保护野生石松类和蕨类植物中的迁地保护植物 Ex situ conservation of wild lycophytes and ferns under state priority conservation	62	14	4	27	5	2	7

科 Family	种 Species	文献 References
石松科 Lycopodiaceae	蛇足石杉 Huperzia serrata	焦瑜等, 2014
石松科 Lycopodiaceae	粗糙马尾杉 Phlegmariurus squarrosus	唐军等, 2015
水韭科 Isoёtaceae	中华水韭 Isoëtes sinensis	路靖等, 2013; 邢建娇等, 2013
水韭科 Isoёtaceae	云贵水韭 Isoëtes yunguiensis	焦瑜等, 2014; 骆强等, 2016
瓶尔小草科 Ophioglossaceae	七指蕨 Helminthostachys zeylanica	宋莉英等, 2015
合囊蕨科 Marattiaceae	福建观音座莲 Angiopteris fokiensis	曾汉元和丁炳扬, 2003; 何俊等, 2021
	秦氏观音座莲 Angiopteris chingii	焦瑜等, 2014
	相马氏观音座莲 Angiopteris somae	焦瑜等, 2014
	圆基观音座莲 Angiopteris subrotundata	焦瑜等, 2014
	河口观音座莲 Angiopteris hokouensis	焦瑜等, 2014
	披针观音座莲 Angiopteris caudatiformis	焦瑜等, 2014
	王氏观音座莲 Angiopteris wangii	焦瑜等, 2014
	天星蕨 Christensenia assamica	焦瑜等, 2014
金毛狗科 Cibotiaceae	金毛狗 Cibotium barometz	张祖荣等, 2010; 邢建娇等, 2012; 王益和, 2012
桫椤科 Cyatheaceae	桫椤 Alsophila spinulosa	王金娟等, 2007; 王辉等, 2013
	中华桫椤 Alsophila costularis	王金娟等, 2007; 刘保东等, 2020
	粗齿桫椤 Alsophila denticulata	曾汉元和丁炳扬, 2003
	笔筒树 Sphaeropteris lepifera	马洪娜等, 2010
	白桫椤 Sphaeropteris brunoniana	蒋胜军等, 2002; 王金娟等, 2007; 陈贵菊等, 2008
	海南白桫椤 Sphaeropteris hainanensis	蒋胜军等, 2002
	滇南桫椤 Alsophila austroyunnanensis	焦瑜等, 2014
	黑桫椤 Alsophila podophylla	张祖荣等, 2010; 焦瑜等, 2014
	大叶黑桫椤 Alsophila gigantea	焦瑜等, 2014; 郎月婷等, 2021
	阴生桫椤 Alsophila latebrosa	焦瑜等, 2014; 郎月婷等, 2021
凤尾蕨科 Pteridaceae	荷叶铁线蕨 Adiantum reniforme	黄芳, 2008; 李洪林等, 2008
	粗梗水蕨 Ceratopteris pteridoides	孙锐等, 2008; 王文明等, 2018
	水蕨 Ceratopteris thalictroides	戴锡玲等, 2005; 陈蔚辉等, 2008; 陈雨虹等, 2010; 朱晓凤等, 2016
乌毛蕨科 Blechnaceae	对开蕨 Asplenium komarovii	刘保东等, 1993; 焦瑜等, 2014; 曾汉元和丁炳扬, 2003; 王蕾等, 2009; 韩汶雨等, 2020
乌毛蕨科 Blechnaceae	苏铁蕨 Brainea insignis	王玥等, 2007; 郭建瑞等, 2008; 夏漪, 2014^①
鹿角蕨科 Platyceriaceae	鹿角蕨 Platycerium wallichii	王卫清等, 2011; 郭捡等, 2012, 2013; 吴建设等, 2020; 叶秀仙等, 2020

中国石松类和蕨类植物多样性研究进展

Insight into the studies on diversity of lycophytes and ferns in China

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年份 Year	新科 New family	新属 New genus	新种 New Species	新种下分类群 New infraspecific taxon	新组合 New combination	新名称 New name	新记录 New record	新异名 New synonyms	新模式标定 New typification	新订正 New correction
2017	-	-	16	-	2	1	-	1	1	-
2018	1	-	19	-	15	2	2	6	4	-
2019	-	-	18	-	27	-	5	7	12	-
2020	-	3	7	1	9	1	1	1	1	-
2021	-	-	26	-	33	-	1	2	2	1
2022	-	-	15	-	-	-	-	-	-	-
汇总 Sum	1	3	101	1	86	4	9	17	20	1

省级行政区划 Provincial administrative area			2017-2022年					2000-2022年
省级行政区划 Provincial administrative area			新种 New species	新种下分类群 New infraspecific taxon	新记录 New record	采集密度 Sampling density		新种 New species	新种下分类群 New infraspecific taxon	新记录 New record	采集密度 Sampling density
安徽	Anhui		-	-	2	264		2	-	3	267
澳门	Aomen		-	-	-	-		-	-	-	-
北京	Beijing		-	-	-	31		-	1	-	52
重庆	Chongqing		1	-	-	117		6	1	-	142
福建	Fujian		1	-	-	109		3	1	1	174
甘肃	Gansu		-	-	-	101		-	-	1	138
广东	Guangdong		9	-	-	1,943		16	1	-	3,764
广西	Guangxi		16	-	-	124		29	-	3	646
贵州	Guizhou		20	-	-	786		59	7	2	912
海南	Hainan		7	-	-	108		16	-	2	535
河北	Hebei		-	-	-	14		-	-	-	21
黑龙江	Heilongjiang		1	-	-	-		1	-	1	-
河南	Henan		-	-	-	26		-	-	-	28
香港	Hong Kong		1	-	-	-		2	2	-	-
湖北	Hubei		-	-	-	2,407		4	3	-	3,218
湖南	Hunan	4		-	-		102	8	-	1		719
江苏	Jiangsu	-		-	-		69	-	-	-		104
江西	Jiangxi	1		-	-		564	3	1	1		1,329
吉林	Jilin	-		-	-		2	-	-	-		8
辽宁	Liaoning	-		-	-		-	-	-	-		-
内蒙古	Nei Mongol	-		-	-		-	1	-	1		-
宁夏	Ningxia	-		-	-		14	-	-	-		-
青海	Qinghai	-		-	-		9	-	-	-		11
陕西	Shaanxi	-		-	-		92	3	-	1		95
山东	Shandong	3		-	-		56	5	-	-		77
上海	Shanghai	-		-	-		1	-	-			8
山西	Shanxi	-		-	-		37	1	-	1		-
四川	Sichuan	5		-	-		175	16	2	3		837
台湾	Taiwan	13		1	6		2	25	5	10		7
天津	Tianjin	-		-	-		1	-	-	-		-
新疆	Xinjiang	-		-	-		15	-	-	2		44
西藏	Xizang	2		-	-		211	31	5	3		999
云南	Yunnan	33		-	1		2,424	71	17	11		6,826
浙江	Zhejiang	1		-	-		685	5	-	-		724

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