濒危植物百山祖冷杉和资源冷杉的物种划分及其遗传资源的保护

doi:10.17521/cjpe.2022.0295

植物生态学报 ›› 2023, Vol. 47 ›› Issue (12): 1629-1645.DOI: 10.17521/cjpe.2022.0295

濒危植物百山祖冷杉和资源冷杉的物种划分及其遗传资源的保护

杨玲¹^,²^,³^,^*, 梁思琪¹^,²^,³^,^*, 潘佳明¹^,²^,³, 韦金鑫¹^,²^,³, 丁涛⁴, 蒋日红⁵, 邵毅贞⁶, 张宪春¹^,², 刘勇波⁷^,^**(), 向巧萍¹^,²^,^**()

¹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
²国家植物园, 北京 100093
³中国科学院大学生命科学学院, 北京 100049
⁴广西壮族自治区中国科学院广西植物研究所, 广西桂林 541006
⁵广西壮族自治区林业科学研究院, 南宁 530002
⁶河南农业大学生命科学学院, 郑州 450002
⁷中国环境科学研究院, 国家环境保护区域生态过程与功能评估重点实验室, 北京 100012

收稿日期:2022-07-18 接受日期:2023-06-05 出版日期:2023-12-20 发布日期:2023-06-08
通讯作者: **(Liu YB, liuyb@craes.org.cn; Xiang QP, qpxiang@ibcas.ac.cn)
作者简介:^*同等贡献
基金资助:
生态环境部生物多样性调查与评估项目(2019HJ2096001006)

Species delimitation and genetic conservation of the endangered firs Abies beshanzuensis and A. ziyuanensis

YANG Ling¹^,²^,³^,^*, LIANG Si-Qi¹^,²^,³^,^*, PAN Jia-Ming¹^,²^,³, WEI Jin-Xin¹^,²^,³, DING Tao⁴, JIANG Ri-Hong⁵, SHAO Yi-Zhen⁶, ZHANG Xian-Chun¹^,², LIU Yong-Bo⁷^,^**(), XIANG Qiao-Ping¹^,²^,^**()

¹State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
²China National Botanical Garden, Beijing 100093, China
³College of Life Sciences, University of Chinese Academy of Sciences, Beijing 100049, China
⁴Guangxi Institute of Botany, Chinese Academy of Sciences, Guilin, Guangxi 541006, China
⁵Guangxi Forestry Research Institute, Nanning 530002, China
⁶College of Life Sciences, Henan Agricultural University, Zhengzhou 450002, China
⁷State Environment Protection Key Laboratory of Regional Ecological Process and Functional Assessment, Chinese Research Academy of Environmental Sciences, Beijing 100012, China

Received:2022-07-18 Accepted:2023-06-05 Online:2023-12-20 Published:2023-06-08
Contact: **(Liu YB, liuyb@craes.org.cn; Xiang QP, qpxiang@ibcas.ac.cn)
About author:^*Contributed equally to this work
Supported by:
Supported by the Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment of China(2019HJ2096001006)

摘要/Abstract

摘要：

物种是生物学中最基本的分类单元, 对于珍稀濒危物种而言, 物种的正确界定对于制定保护措施至关重要。分布于中国亚热带地区的百山祖冷杉(Abies beshanzuensis)、资源冷杉(A. ziyuanensis)和大院冷杉(A. dayuanensis)种群极小, 且分类上长期存在争议。现行分类依据形态和地理分布将大院冷杉归并到资源冷杉, 又将资源冷杉处理为百山祖冷杉的变种, 但分子证据不足。该研究对百山祖冷杉、资源冷杉和大院冷杉分布区内8个种群的23株个体进行靶向捕获测序, 获得了60个单拷贝核基因中的805个单核苷酸多态性位点。种群遗传结构和历史动态分析表明, 这些个体可分为两个谱系, 分别对应于百山祖冷杉和资源冷杉, 其中资源冷杉在2.35 Ma前与百山祖冷杉和大院冷杉的共同祖先分化, 而大院冷杉与百山祖冷杉亲缘关系更近, 形成一个谱系。百山祖冷杉、资源冷杉和大院冷杉的遗传多样性水平整体较低, 种群间存在较为明显的遗传分化(遗传分化指数0.083-0.208)。由于未检测到分化后的谱系间遗传交流, 推测分布区的碎片化是谱系间遗传交流受阻进而产生分化的主要原因。生态位比较分析结果显示百山祖冷杉、资源冷杉和大院冷杉这群濒危冷杉分布区的年平均气温和最冷季平均气温显著高于东亚分布的非濒危冷杉物种, 因此, 该研究认为全球气候变暖是导致其濒危的关键因素。综合以上研究结果, 该研究对百山祖冷杉、资源冷杉和大院冷杉这群植物进行新的分类处理, 将大院冷杉处理为百山祖冷杉的异名, 确认资源冷杉种的地位。在对其濒危机制深入理解的基础上, 建议在西南的横断山和秦巴山地区尝试迁地栽培实验, 同时就地人工辅助育种。

关键词: 大院冷杉, 溯祖分析, 生态位, 种群遗传学, 物种保护, 物种划分, 靶向捕获测序

Abstract:

Aims “Species” is the basic taxonomical unit in biology. The accuracy of endangered species delimitation largely determines the efficiency and conservation strategies. Abies beshanzuensis, A. ziyuanensis and A. dayuanensis, a group of endangered firs endemic to the subtropical region of China, have extremely small populations and controversial taxonomy. According to morphological differences and geographical distribution, A. dayuanensis has been considered as the synonym of A. ziyuanensis, whereas A. ziyuanensis is currently treated as a variety of A. beshanzuensis. However, no genetic evidence has been provided to test this classification hypothesis.
Methods We performed target enrichment sequencing on 23 individuals from eight extant populations and obtained 805 single nucleotide polymorphic sites from 60 single-copy nuclear genes for population genetic analyses.
Important findings Results of population genetic structure and demographic history analyses showed that this endangered fir group has diverged into two lineages corresponding to A. beshanzuensis and A. ziyuanensis. Abies ziyuanensis first diverged from the common ancestor of A. beshanzuensis and A. dayuanensis, the latter two split apart later and more closely related, forming the other lineage. The genetic diversity is low within these populations. Strong genetic differentiation was found among populations (fixation index, F_ST= 0.083-0.208). Gene flow is not detected after lineages split, probably because of the fragmented habitats which blocked effective genetic exchanges among populations. Ecological niche comparison showed that the average annual temperature and the temperature of the coldest quarter were significantly higher in the habitats of these endangered firs compared to those of non-threatened firs in East Asia. Therefore, we inferred that global warming is likely to push this group of endangered firs to the edge of extinction. According to the significant population differentiation, we provide genetic evidence that A. dayuanensis is a synonym of A. beshanzuensis, and resurrect A. ziyuanensis as a separate species. Based on the niche comparison results, we highlight the necessity to carry out ex suit conservation in the Hengduan Mountains and the Qingling-Bashan Mountains and conduct in suit regeneration.

Key words: Abies dayuanensis, coalescent analysis, ecological niche, population genetics, species conservation, species delimitation, targeted sequencing

杨玲, 梁思琪, 潘佳明, 韦金鑫, 丁涛, 蒋日红, 邵毅贞, 张宪春, 刘勇波, 向巧萍. 濒危植物百山祖冷杉和资源冷杉的物种划分及其遗传资源的保护. 植物生态学报, 2023, 47(12): 1629-1645. DOI: 10.17521/cjpe.2022.0295

YANG Ling, LIANG Si-Qi, PAN Jia-Ming, WEI Jin-Xin, DING Tao, JIANG Ri-Hong, SHAO Yi-Zhen, ZHANG Xian-Chun, LIU Yong-Bo, XIANG Qiao-Ping. Species delimitation and genetic conservation of the endangered firs Abies beshanzuensis and A. ziyuanensis. Chinese Journal of Plant Ecology, 2023, 47(12): 1629-1645. DOI: 10.17521/cjpe.2022.0295

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0295

https://www.plant-ecology.com/CN/Y2023/V47/I12/1629

图/表 12

表1 百山祖冷杉-资源冷杉谱系各种群的地理信息及采样数

Table 1 Geographic information and samples size of Abies beshanzuensis-A. ziyuanensis lineage

组群 Group	种群 Population	编号 Code	数量 Number	经度 Longitude (° E)	纬度 Latitude (° N)	海拔 Altitude (m)
百山祖 Baishanzu (BSZ)	浙江庆元县百山祖 Baishanzu, Qingyuan, Zhejiang	BSZ	3	119.20	27.75	1 750
大院 Dayuan (DY)	湖南炎陵县大院农场 Dayuan, Yanling, Hunan	DY	3	114.07	26.45	1 478
大院 Dayuan (DY)	江西井冈山市平水山 Pingshuishan, Jinggangshan, Jiangxi	JGS	1	114.11	26.50	1 676
	广西全州县大云山 Dayunshan, Quanzhou, Guangxi	QZ	3	110.96	26.38	1 801
	广西资源县银竹老山 Yinzhulaoshan, Ziyuan, Guangxi	YZ	5	110.56	26.25	1 753
资源 Ziyuan (ZY)	湖南城步县金童山 Jintongshan, Chengbu, Hunan	CB	4	110.58	26.28	1 646
	湖南江永县千家峒 Qianjiadong, Jiangyong, Hunan	QJD	3	111.30	25.57	1 800
	湖南新宁县舜皇山 Shunhuangshan, Xinning, Hunan	SHS	1	110.56	26.22	1 778

图1 基于805个核基因单核苷酸多态性位点构建的百山祖冷杉-资源冷杉谱系的邻接树(A)和主成分(PC)分析图(B)。树枝上方数字为大于50%的自展值(1 000次重复)。种群编号见表1。

Fig. 1 Neighbor-joining phylogenetic tree (A) and PCA result (B) of Abies beshanzuensis-A. ziyuanensis lineage constructed using 805 nuclear single nucleotide polymorphisms. Numbers on branches are bootstrap value > 50% (1 000 replications). Population codes are shown in Table 1.

图2 基于805个核基因单核苷酸多态性位点使用SplitsTree构建的百山祖冷杉-资源冷杉谱系的系统发生网络关系。种群编号见表1。

Fig. 2 Phylogenetic network of Abies beshanzuensis-A. ziyuanensis lineage based on 805 nuclear single nucleotide polymorphisms using SplitsTree. Population codes are shown in Table 1.

图3 基于805个核基因单核苷酸多态性位点使用ADMIXTURE分析的百山祖冷杉-资源冷杉谱系的祖先成分划分结果(K = 2)。种群编号见表1。

Fig. 3 Ancestral assignment of Abies beshanzuensis-A. ziyuanensis lineage based on 805 nuclear single nucleotide polymorphisms using ADMIXTURE (K = 2). Population codes are shown in Table 1.

表2 百山祖冷杉-资源冷杉谱系各种群的种群遗传学参数

Table 2 Population genetic parameter of Abies beshanzuensis-A. ziyuanensis lineage

种群 Population	多态位点比例 Proportion of polymorphic markers	核苷酸多样性 Nucleotide diversity	期望杂合度 Expected heterozygosity	观测杂合度 Observed heterozygosity	Tajima的D值 Tajima’s D
BSZ	0.372 7	0.098 6	0.072 6	0.128 2	0.391 3
DY	0.444 7	0.115 3	0.063 0	0.104 3	0.307 4
QJD	0.438 5	0.108 0	0.066 6	0.099 1	0.185 1
CB	0.447 2	0.108 7	0.076 3	0.130 8	0.391 7
ZY	0.555 3	0.117 2	0.101 1	0.157 9	0.154 3
QZ	0.473 3	0.125 2	0.109 1	0.182 9	0.377 7

表3 百山祖冷杉-资源冷杉谱系种群间分子方差分析(AMOVA) (p < 0.01)

Table 3 Results of analysis of molecular variance (AMOVA) from populations of Abies beshanzuensis-A. ziyuanensis lineage (p < 0.01)

变异来源 Source of variation	自由度 df	方差和 Sum of squares	变异组分 Variance components	占总变异比例 Percentage of variation (%)
种群间 Among populations	4	723.79	5.57	6.17
种群内 Within populations	39	3 305.75	84.76	93.83
总计 Total	45	4 029.54	93.18

表4 百山祖冷杉-资源冷杉谱系种群间的遗传分化指数(FST)

Table 4 Fixation index (FST) between populations of Abies beshanzuensis-A. ziyuanensis lineage

种群 Population	BSZ	DY	QJD	CB	YZ	QZ
BSZ	-
DY	0.145 4	-
QJD	0.140 2	0.079 9	-
CB	0.184 1	0.114 0	0.075 8	-
YZ	0.168 6	0.113 7	0.055 5	0.088 8	-
QZ	0.207 5	0.169 1	0.110 1	0.126 3	0.083 1	-

图4 百山祖冷杉-资源冷杉谱系(A、B)和资源冷杉(C、D)种群间遗传距离与地理距离和环境距离的Mantel检验。

Fig. 4 Mantel test between genetic distance and geographic distance or environmental distance among populations of Abies beshanzuensis-A. ziyuanensis lineage (A, B) and A. ziyuanensis (C, D).

图5 FASTSIMCOAL2分析中百山祖冷杉-资源冷杉谱系的最优分化历史示意图。箱型宽度表示相对有效种群大小;T1、 T2分别代表谱系第一、二次分化时间; NANC代表共同祖先群体的有效群体大小; BSZ代表百山祖组群; DY代表大院组群; ZY代表资源组群。

Fig. 5 The best demographic model of Abies beshanzuensis-A. ziyuanensis lineage. The width of the boxes represents the relative effective population size; T1 and T2 represent the first and second time of lineage differentiation, respectively; NANC indicates ancestral population size; BSZ represents the Baishanzu group; DY represents the Dayuan group; ZY represents the Ziyuan group.

图6 东亚冷杉属植物的地理分布图(A)和基于19个生态因子的主成分(PC)分析(B)。EN, 中国亚热带地区濒危冷杉分布点; HH, 横断-喜马拉雅山脉冷杉分布点; JK, 朝鲜半岛和日本山区冷杉分布点; TW, 中国台湾山脉冷杉分布点。

Fig. 6 Geographical distribution map (A) and principal component (PC) analysis of 19 climatic factors (B) of firs in East Asia. EN, endangered firs distributed in subtropical mountains of China; HH, firs in Hengduan-Himalayan mountains; JK, firs in Japan-Korean peninsula; TW, firs in Taiwan mountains of China.

图7 4组东亚冷杉属植物的生态因子差异比较。EN, 中国亚热带地区濒危冷杉分布点; HH, 横断-喜马拉雅山脉冷杉分布点; JK, 朝鲜半岛和日本山区冷杉分布点; TW, 中国台湾山脉冷杉分布点。不同小写字母代表差异显著(p < 0.01)。

Fig. 7 Results of ANOVA and multiple comparative analysis for Abies in East Asia. EN, endangered firs distributed in subtropical mountains of China; HH, firs in Hengduan-Himalayan mountains; JK, firs in Japan-Korean peninsula; TW, firs in Taiwan mountains of China. Different lowercase letters indicate significant difference (p < 0.01).

表5 百山祖冷杉和资源冷杉的形态特征和地理分布表

Table 5 Morphological characteristics and geographical distribution of Abies beshanzuensis and A. ziyuanensis

物种 Species	冬芽 Winter bud	球果 Seed cones				种翅 Seed wings	地理分布 Distribution	海拔 Altitude (m)
物种 Species	冬芽 Winter bud	形状 Shape	成熟时颜色 Color when ripe	苞鳞 Bract scales	中部种鳞 Middle seed scales	种翅 Seed wings	地理分布 Distribution	海拔 Altitude (m)
百山祖冷杉 A. beshanzuensis	卵圆形至圆锥形 Ovoid to conical	圆柱形 Cylindric	淡褐色或淡褐黄色 Light brown or brownish yellow	苞鳞上部宽7-8 mm, 中下部渐窄, 宽3-4 mm Bract 7-8 mm wide at upper part, middle and lower part gradually attenuate, 3-4 mm wide	扇状四边形或肾状四边形, 长1.8-2.4 cm, 宽2.5- 3 cm Flabellate- trapeziform or reniform- trapeziform, 1.8-2.4 cm × 2.5- 3 cm	种子(含种翅)长1.0-2.3 cm, 种翅与种子等长 Length of seed wing as long as the seed, including seed 1.0-2.3 cm	浙江(庆元县百山祖)、湖南(炎陵县大院农场)、江西(井冈山市平水山、遂川县南风面) Zhejiang: Qingyuan (Baishanzu), Hunan: Yanling (Dayuan); Jiangxi: Jinggangshan (Pingshuishan), Suichuan (Nanfengmian)	1 350-1 950
资源冷杉 A. ziyuanensis	圆锥形或圆锥状卵圆形 Conical or conical-ovoid	长椭圆状圆柱形 Long ellipsoid-cylindric	暗绿褐色或暗褐色 Dark greenish brown or dark brown	苞鳞上部宽9-10 mm, 中部及中下部明显变窄, 宽约3 mm Bract 9-10 mm wide at upper part, middle and lower part conspicuous attenuate, 3 mm wide	扇状四边形, 长2.3-2.5 cm, 宽3- 3.3 cm Flabellate- trapeziform, 2.3-2.5 cm × 3-3.3 cm	种子(含种翅)长2.4-3.1 cm, 种翅较种子长一倍以上 Length of seed wing two times that of the seed, including seed 2.4-3.1 cm	广西(资源县银竹老山、全州县)、湖南(新宁县舜皇山、城步县金童山、江永县千家峒) Guangxi: Ziyuan (Yinzhulaoshan), Quanzhou; Hunan: Xinning (Shunhuangshan), Chengbu (Jintongshan), Jiangyong (Qianjiadong)	1 600-1 800

表5 百山祖冷杉和资源冷杉的形态特征和地理分布表

Table 5 Morphological characteristics and geographical distribution of Abies beshanzuensis and A. ziyuanensis

物种 Species	冬芽 Winter bud	球果 Seed cones				种翅 Seed wings	地理分布 Distribution	海拔 Altitude (m)
物种 Species	冬芽 Winter bud	形状 Shape	成熟时颜色 Color when ripe	苞鳞 Bract scales	中部种鳞 Middle seed scales	种翅 Seed wings	地理分布 Distribution	海拔 Altitude (m)
百山祖冷杉 A. beshanzuensis	卵圆形至圆锥形 Ovoid to conical	圆柱形 Cylindric	淡褐色或淡褐黄色 Light brown or brownish yellow	苞鳞上部宽7-8 mm, 中下部渐窄, 宽3-4 mm Bract 7-8 mm wide at upper part, middle and lower part gradually attenuate, 3-4 mm wide	扇状四边形或肾状四边形, 长1.8-2.4 cm, 宽2.5- 3 cm Flabellate- trapeziform or reniform- trapeziform, 1.8-2.4 cm × 2.5- 3 cm	种子(含种翅)长1.0-2.3 cm, 种翅与种子等长 Length of seed wing as long as the seed, including seed 1.0-2.3 cm	浙江(庆元县百山祖)、湖南(炎陵县大院农场)、江西(井冈山市平水山、遂川县南风面) Zhejiang: Qingyuan (Baishanzu), Hunan: Yanling (Dayuan); Jiangxi: Jinggangshan (Pingshuishan), Suichuan (Nanfengmian)	1 350-1 950
资源冷杉 A. ziyuanensis	圆锥形或圆锥状卵圆形 Conical or conical-ovoid	长椭圆状圆柱形 Long ellipsoid-cylindric	暗绿褐色或暗褐色 Dark greenish brown or dark brown	苞鳞上部宽9-10 mm, 中部及中下部明显变窄, 宽约3 mm Bract 9-10 mm wide at upper part, middle and lower part conspicuous attenuate, 3 mm wide	扇状四边形, 长2.3-2.5 cm, 宽3- 3.3 cm Flabellate- trapeziform, 2.3-2.5 cm × 3-3.3 cm	种子(含种翅)长2.4-3.1 cm, 种翅较种子长一倍以上 Length of seed wing two times that of the seed, including seed 2.4-3.1 cm	广西(资源县银竹老山、全州县)、湖南(新宁县舜皇山、城步县金童山、江永县千家峒) Guangxi: Ziyuan (Yinzhulaoshan), Quanzhou; Hunan: Xinning (Shunhuangshan), Chengbu (Jintongshan), Jiangyong (Qianjiadong)	1 600-1 800

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濒危植物百山祖冷杉和资源冷杉的物种划分及其遗传资源的保护

Species delimitation and genetic conservation of the endangered firs Abies beshanzuensis and A. ziyuanensis

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