中国薄叶卷柏复合群的物种划分

doi:10.17520/biods.2021273

生物多样性 ›› 2021, Vol. 29 ›› Issue (12): 1607-1619. DOI: 10.17520/biods.2021273 cstr: 32101.14.biods.2021273

• 研究报告: 植物多样性 • 上一篇下一篇

中国薄叶卷柏复合群的物种划分

张梦华¹^,², 张宪春¹^,^*()

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.中国科学院大学, 北京 100049

收稿日期:2021-07-09 接受日期:2021-11-03 出版日期:2021-12-20 发布日期:2021-12-16
通讯作者: 张宪春
作者简介:*E-mail: zhangxc@ibcas.ac.cn
基金资助:
国家植物标本资源库项目(E0117G1001)

Species delimitation of the Selaginella delicatula group in China

Menghua Zhang¹^,², Xianchun Zhang¹^,^*()

1 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093
2 University of Chinese Academy of Sciences, Beijing 100049

Received:2021-07-09 Accepted:2021-11-03 Online:2021-12-20 Published:2021-12-16
Contact: Xianchun Zhang

摘要/Abstract

摘要：

卷柏属(Selaginella)是石松类植物中最大的属, 也是分类难度较大的类群之一。该属的物种划分主要基于形态特征, 但许多近缘种在形态上很难区分。近年来, 已有大量分子证据被用于各植物类群的分类学研究, 但目前未发现一套适合卷柏属物种鉴定的分子标记。薄叶卷柏复合群(S. delicatula group)是卷柏属下鉴定较为困难的类群, 包括了薄叶卷柏(S. delicatula)、黑顶卷柏(S. picta)和瓦氏卷柏(S. wallichii) 3个物种, 主要分布于亚洲的热带和亚热带地区。为了探讨薄叶卷柏复合群内物种的亲缘关系和评估不同分子标记在卷柏属分类学研究中的应用价值, 本研究对该复合群物种进行广泛取样, 共收集到73个个体, 并选取3个叶绿体基因(rbcL, psbA和atpI)和2个核基因(26S nrDNA和pgiC)片段进行系统树的构建及叶绿体单倍型分析。研究发现, 基于叶绿体和核基因构建的系统发生关系存在冲突: 叶绿体基因树上薄叶卷柏个体分为两个分支(A和B), 薄叶卷柏B分支与薄叶卷柏A-S. picta分支呈姐妹关系, 并且rbcL单倍型分析结果也表明薄叶卷柏A和B两个分支存在明显分化; 而核基因结果则支持该复合群3个物种各自的单系性, 其中, S. delicatula分支与S. picta分支为姐妹群, S. wallichii与S. delicatula-S. picta分支为姐妹关系。在对复合群分布区大量标本的观察以及野外群体调查的基础上, 评估了植株茎和枝的分枝方式、孢子叶、营养叶(侧叶、中叶和腋叶)和孢子表面纹饰等形态性状的分类学价值。结果表明, 薄叶卷柏A和B分支的样本仅在植株分枝方式和大孢子表面纹饰上存在差异, 但无法依靠小孢子表面纹饰、孢子叶穗和营养叶形态等特征进行区分。基于现有证据, 薄叶卷柏复合群至少可划分为薄叶卷柏、黑顶卷柏和瓦氏卷柏3种, 但彻底澄清该复合群的物种划分还需要获取模式标本产地的材料和细胞学证据。最后, 建议在未来卷柏属的分类学研究集中于该属分类复杂的复合群, 结合使用形态学、细胞学、分子生物学(同时使用核基因和叶绿体分子标记)及地理分布等整合证据来进行物种划分。

关键词: 26S nrDNA, pgiC, 叶绿体分子标记, 石松类植物, 薄叶卷柏复合群

Abstract

Aims: Species delimitation of Selaginella remains largely based on morphological characters, however, many closely related species are morphologically indistinguishable from each other. This study aims to (1) explore phylogenetic relationships within the S. delicatula group; and (2) evaluate the utility of nuclear and chloroplast DNA markers for the classification of Selaginella.
Methods: A total of 73 individuals were sampled, covering the geographical distribution and morphological variation of this group. We sequenced three chloroplast markers (rbcL, psbA, and atpI) and two nuclear markers (26S nrDNA and pgiC) to infer the phylogenetic relationships and haplotype network.
Results: We detected incongruence within S. delicatula-S. picta clade between chloroplast and nuclear phylogenies. The chloroplast dataset indicated S. delicatula is not monophyletic, with S. delicatula B sister to S. delicatula A-S. picta. In contrast, nuclear dataset discovered the monophyly of all three species, with S. delicatula sister to S. picta and S. wallichii sister to the S. delicatula-S. picta clade. Morphological characters, including main stem and branches, sporophylls, sterile leaves (ventral, dorsal, and axillary leaves) and spores were carefully observed and assessed. Morphological comparisons indicated that the samples of S. delicatula A and B clades are only distinct in branch patterns and the micromorphology of megaspores surface, not differential in morphology of leaves, strobili and microspores.
Conclusion: As a result of available evidence, three taxa were recognized in the S. delicatula group. It is necessary to perform further studies with cytological evidence and more samples of S. delicatula from type locality in order to obtain a better understanding of the species delimitation. We suggested that the taxonomic work of Selaginella should focus on the closely related species and multiple lines of evidence should be considered, including morphological characters, cytological evidence, molecular data (nuclear and chloroplast DNA markers), and geographical distribution.

Key words: 26S nrDNA, pgiC, chloroplast markers, lycophytes, Selaginella delicatula group

张梦华, 张宪春 (2021) 中国薄叶卷柏复合群的物种划分. 生物多样性, 29, 1607-1619. DOI: 10.17520/biods.2021273.

Menghua Zhang, Xianchun Zhang (2021) Species delimitation of the Selaginella delicatula group in China. Biodiversity Science, 29, 1607-1619. DOI: 10.17520/biods.2021273.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2021273

https://www.biodiversity-science.net/CN/Y2021/V29/I12/1607

图/表 8

表1 用于扩增和测序的引物

Table 1 Primers used in amplification and sequencing

分子标记 Marker	引物 Primer	正反向 Direction	序列 Sequence (5′-3′)	参考文献 Reference
26S	26S 60F	正向 Forward	TTTAAGCATATCACTAAGCGGAGG	Korall & Kenrick (2004)
26S	26S 380F	正向 Forward	CCGCGAGGGAAAGATGAAAAGGAC	Korall & Kenrick (2004)
26S	26S 1160R	反向 Reverse	CCAGTTCTGCTTACCAAAAATGGCCC	Korall & Kenrick (2004)
pgiC	SWPGIC-1666F	正向 Forward	VTTYGCTTTYTGGGAYTGGG	Weststrand & Korall (2016a)
pgiC	SWPGIC-2523R	反向 Reverse	GTCGTGGTTRCTSACAATCTC	Weststrand & Korall (2016a)
rbcL	rbcL-192F	正向 Forward	CACGTGGACTACCGTTTGGA	Shalimov et al (2019)
rbcL	rbcL-1324R	反向 Reverse	TACCCTCAAGAGCGGGATCA	Shalimov et al (2019)
psbA	psbA-169F	正向 Forward	CCRGTAGATATTGACGGTATTC	Shalimov et al (2019)
psbA	psbA-1026R	反向 Reverse	ATCTRGWGGGAAGTTGTGAGC	Shalimov et al (2019)
atpI	atpI-119F	正向 Forward	CYCAGGTTCATGGACAAGTAC	Shalimov et al (2019)
atpI	atpI-540R	反向 Reverse	GRGTATYGGGGTTGGTTG	Shalimov et al (2019)

图1 基于核基因26S nrDNA + pgiC串联数据矩阵构建的薄叶卷柏复合群最大似然树。最大似然值和后验概率值分别标注在分支上(ML/PP), (-)表示无支持率。

Fig. 1 Maximum likelihood tree of the Selaginella delicatula group based on the nuclear dataset (26S nrDNA + pgiC). Maximum parsimony bootstrap values and Bayesian posterior probabilities are shown on the branches (ML/PP). A dash (-) indicates no support value.

图2 基于叶绿体基因atpI + psbA + rbcL串联数据矩阵构建的薄叶卷柏复合群最大似然树。最大似然值和后验概率值分别标注在分支上(ML/PP), (-)表示无支持率。

Fig. 2 Maximum likelihood tree of the Selaginella delicatula group based on the chloroplast dataset (atpI + psbA + rbcL). Maximum parsimony bootstrap values and Bayesian posterior probabilities are shown on the branches (ML/PP). A dash (-) indicates no support value.

图3 基于叶绿体基因rbcL构建的薄叶卷柏复合群(薄叶卷柏Selaginella delicatula、瓦氏卷柏S. wallichii和黑顶卷柏S. picta)单倍型网络关系(A)和地理分布(B)。

Fig. 3 Chloroplast rbcL gene haplotype network of the Selaginella delicatula group (A) and their distributions (B). CQ, Chongqing; FJ, Fujian; GD, Guangdong; GX, Guangxi; GZ, Guizhou; HB, Hubei; HK, Hong Kong; HN, Hainan; HuN, Hunan; SC, Sichuan; TW, Taiwan; XZ, Xizang; YN, Yunnan; MY, Myanmar; VT, Vietnam; ML, Malaysia.

图4 薄叶卷柏复合群孢子叶穗和营养叶片形态。(A-D)黑顶卷柏, 西藏, B. S. Li & S. Z. Cheng 02630 (PE); (E-H)瓦氏卷柏, 云南, X. C. Zhang & S. Y. Dong 1421 (PE); (I-L)薄叶卷柏B, 海南, S. Y. Dong 826 (PE); (M-P)薄叶卷柏A, 湖南, Y. H. Yan 7765 (PE); (A, E, I, M)孢子叶穗, 标尺 = 1 mm; (B, F, J, N)侧叶; (C, G, K, O)中叶; (D, H, L, P)腋叶。标尺 = 0.2 mm。

Fig. 4 Strobili and sterile leaves of Selaginella delicatula group. (A-D) S. picata, Xizang, B. S. Li & S. Z. Cheng 02630 (PE); (E-H) S. wallichii, Yunnan, X. C. Zhang & S. Y. Dong 1421 (PE); (I-L) S. delicatula B, Hainan, S. Y. Dong 826 (PE); (M-P) S. delicatula A, Hunan, Y. H. Yan 7765 (PE). (A, E, I, M) Strobili, Scale bars = 1 mm; (B, F, J, N) Ventral leaves; (C, G, K, O) Dorsal leaves; (D, H, L, P) Axillary leaves. Scale bars = 0.2 mm.

图5 薄叶卷柏复合群腊叶标本。(A)薄叶卷柏A, 中国贵州, X. C. Zhang et al. 6849 (PE); (B)薄叶卷柏B, 靠近新几内亚的Pisang岛, Gaudichaud 13 (Lectotype, G); (C)瓦氏卷柏, 马来西亚槟榔屿和新加坡, Dr. Wallich 128 (Isotype, E); (D)黑顶卷柏, 印度, Griffith sn (Syntype, K)。

Fig. 5 Specimens of Selaginella delicatula group. (A) S. delicatula A, Guizhou, X. C. Zhang et al. 6849 (PE); (B) S. delicatula B, Pisang Island near New Guinea, Gaudichaud 13 (Lectotype, G); (C) S. wallichii, Penang and Singapore, Dr. Wallich 128 (Isotype, E); (D) S. picta, India, Griffith sn (Syntype, K).

图6 薄叶卷柏复合群孢子形态。(A-E)薄叶卷柏A (A-C大孢子, D-E小孢子), 贵州, X. C. Zhang et al 6849 (PE); (F-K)薄叶卷柏B (F-I大孢子, J-K小孢子), 海南, X. C. Zhang 2867 (PE); (L-O)黑顶卷柏(L-M大孢子, N-O小孢子), 云南, Z. Y. Li & R. Wei WR295 (PE); (P)瓦氏卷柏(大孢子), 广西, Shiwandashan Exped. 050 (PE); (Q-R)瓦氏卷柏(小孢子), 广西, X. C. Zhang & Z. Y. Guo 7882 (PE)。

Fig. 6 Spores morphology of Selaginella delicatula group. (A-E) S. delicatula A (A-C Megaspores, D-E Microspores), Guizhou, X. C. Zhang et al. 6849 (PE); (F-K) S. delicatula B (F-I Megaspores, J-K Microspores), Hainan, X. C. Zhang 2867 (PE); (L-O) S. picta (L-M Megaspores, N-O Microspores), Yunnan, Z. Y. Li & R. Wei WR295 (PE); (P) S. wallichii (Megaspores), Guangxi, Shiwandashan Exped. 050 (PE); (Q-R) S. wallichii (Microspores), Guangxi, X. C. Zhang & Z. Y. Guo 7882 (PE).

图7 基于薄叶卷柏复合群(薄叶卷柏 Selaginella delicatula,瓦氏卷柏 S. wallichii和黑顶卷柏 S. picta) 6个形态特征的主成分分析

Fig. 7 Principal components plots based on six morphological characters of the Selaginella delicatula group

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中国薄叶卷柏复合群的物种划分

Species delimitation of the Selaginella delicatula group in China

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