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作者简介:

廖苗(1995-),硕士,研究方向为植物系统发育,(E-mail)liaomiao63@yeah.net。

通讯作者:

唐光大,博士,副教授,研究方向为植物系统分类、植物资源和植被动态等,(E-mail)gdtang@scau.edu.cn。

中图分类号:Q949.7

文献标识码:A

文章编号:1000-3142(2022)10-1717-13

DOI:10.11931/guihaia.gxzw202108036

参考文献
ALI SI, KHATOON S, 1982. Genus Vincetoxicum Von Wolf (Asclepiadaceae) in Pakistan [J]. Pak J Bot, 14(1): 61-68.
参考文献
BACKER CA, BAKHUIZEN VAN DEN BRINK RC, 1965. Flora of Java (spermatophytes only) Vol. II Angiospermae, families 111-160 [M]. Groningen: N. V. P. Noordhoff: 259-260.
参考文献
CHASE MW, CHRISTENHUSZ MJM, FAY MF, et al. , 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV [J]. Biol J Linn Soc, 181(1): 1-20.
参考文献
CHUN WY, TSIANG Y, 1941. Merrillanthus, a new genus of Asclepiadaceae from Hainan [J]. Sunyatsenia, 6(2): 105-108.
参考文献
ENDRESS ME, BRUYNS PV, 2000. A revised classification of the Apocynaceae s. l. [J]. Bot Rev, 66(1): 1-56.
参考文献
ENDRESS ME, LIEDE-SCHUMANN S, MEVE U, 2007. Advances in Apocynaceae: The enlightenment, an introduction [J]. Ann Mo Bot Gard, 94(2): 259-267.
参考文献
ENDRESS ME, LIEDE-SCHUMANN S, Meve U, 2014. An updated classification for Apocynaceae [J]. Phytotaxa, 159(3): 175-194.
参考文献
ENDRESS ME, MEVE U, MIDDLETON DJ, et al. , 2018. Apocynaceae [M]//KADEREIT JW, BITTRICH V. Flowering plants. Eudicots: Apiales, Gentianales (except Rubiaceae) 15. Cham: Springer International Publishing: 207-411.
参考文献
FISHBEIN M, LIVSHULTZ T, STRAUB SCK, et al. , 2018. Evolution on the backbone: Apocynaceae phylogenomics and new perspectives on growth forms, flowers, and fruits [J]. Am J Bot, 105(3): 495-513.
参考文献
FORSTER PI, 1991. A taxonomic revision of Cynanchum L. (Asclepiadaceae: Asclepiadoideae) in Australia [J]. Austrobaileya, 3(3): 443-466.
参考文献
FORSTER PI, 1994. A taxonomic revision of Tylophora (Asclepiadaceae: Marsdenieae) in Papuasia [J]. Aust Syst Bot, 7(5): 485-505.
参考文献
GILBERT MG, STEVENS WD, LI PT, 1995. Notes on the Asclepiadaceae of China [J]. Novon, 5(1): 1-16.
参考文献
GUINDON S, DUFAYARD J, LEFORT V, et al. , 2010. Newalgorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3. 0 [J]. Syst Biol, 59(3): 307-321.
参考文献
HSU T, HUANG W, CHUNG S, 2021. Vincetoxicum (Apocynaceae: Asclepiadoideae) in Taiwan: Two new combinations and a new record [J]. Phytotaxa, 478(2): 287-290.
参考文献
JIANG(TSIANG) Y, LI BT(PT), 1977. Flora Reipublicae Popularis Sinicae: Vol. 63 [M]. Beijing: Science Press: 249-575. [蒋英, 李秉滔, 1977. 中国植物志: 第63卷 [M]. 北京: 科学出版社: 249-575. ]
参考文献
JIN JJ, YU WB, YANG JB, et al. , 2020. GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes [J]. Genome Biol, 21(1): 241.
参考文献
KATOH K, STANDLEY DM, 2013. MAFFT multiple sequence alignment software Version 7: improvements in performance and usability [J]. Mol Biol Evol, 30(4): 772-780.
参考文献
KIDYOO A, KIDYOO M, 2018. A new species of Hoya (Marsdenieae), three new combinations and two new names in Vincetoxicum (Asclepiadeae) from Thailand [J]. Taiwania, 63(1): 25-31.
参考文献
LI PT, GILBERT MG, STEVENS WD, 1995. Asclepiadaceae [M]//WU ZY, RAVEN PH, HONG DY. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press: 189-270.
参考文献
LIEDE S, 1996. Cynanchum-Rhodostegiella-Vincetoxicum-Tylophora (Asclepiadaceae): New considerations on an old problem [J]. Taxon, 45(2): 193-211.
参考文献
LIEDE S, 2001. Subtribe Astephaninae (Apocynaceae—Asclepiadoideae) reconsidered: New evidence based on cpDNA spacers [J]. Ann Missouri Bot Gard, 88(4): 657-668.
参考文献
LIEDE-SCHUMANN S, KHANUM R, MUMTAZ AS, et al. , 2016. Going west — A subtropical lineage (Vincetoxicum, Apocynaceae: Asclepiadoideae) expanding into Europe [J]. Mol Phylogenet Evol, 94(A): 436-446.
参考文献
LIEDE-SCHUMANN S, KONG H, MEVE U, et al. , 2012. Vincetoxicum and Tylophora (Apocynaceae: Asclepiadoideae: Asclepiadeae)—two sides of the same medal: Independent shifts from tropical to temperate habitats [J]. Taxon, 61(4): 803-825.
参考文献
LIEDE-SCHUMANN S, MEVE U, 2018. Vincetoxicum (Apocynaceae—Asclepiadoideae) expanded to include Tylophora and allies [J]. Phytotaxa, 369(3): 129-184.
参考文献
MARKGRAF F, 1972. Asclepiadaceae [M]//TUTIN TG, HEYWOOD VH, BURGES NA, et al. Flora Europaea: Volume 3. Diapensiacea to Myoporaceae. Cambridge: Cambridge University Press: 71.
参考文献
MEVE U, OMLOR R, LIEDE-SCHUMANN S, 2002. A new combination in Tylophora (Apocynaceae, Asclepiadoideae) from the Philippines [J]. Syst Geogr Plant, 72(1/2): 27-32.
参考文献
MINH BQ, NGUYEN MAT, VON HAESELER A, 2013. Ultrafast approximation for phylogenetic bootstrap [J]. Mol Biol Evol, 30(5): 1188-1195.
参考文献
NGUYEN LT, SCHMIDT HA, VON HAESELER A, et al. , 2015. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies [J]. Mol Biol Evol, 32(1): 268-274.
参考文献
QIU SX, LI DZ, ZHANG ZX, et al. , 1989. Chemotaxonomy of Cynanchum and its allied genera with notes on the generic characteristics of Vincetoxicum [J]. Acta Bot Yunnan, 11(1): 41-50. [邱声祥, 李德铢, 张壮鑫, 等, 1989. 鹅绒藤属及其近缘属的化学分类——兼论Vincetoxicum的恢复问题 [J]. 云南植物研究, 11(1): 41-50. ]
参考文献
QU XJ, MOORE MJ, LI DZ, et al. , 2019. PGA: A software package for rapid, accurate, and flexible batch annotation of plastomes [J]. Plant Methods, 15(1): 1-12.
参考文献
RAO H, WANG XJ, MA JX, et al. , 2018. Characterization of the complete chloroplast genome of Biondia chinensis (Apocynaceae: Asclepiadoideae: Asclepiadeae), a rare and threatened liana endemic to China [J]. Mitochon DNA Part B Resour, 3(2): 763-764.
参考文献
RAPINI A, VAN DEN BERG C, LIEDE-SCHUMANN S, 2007. Diversification of Asclepiadoideae (Apocynaceae) in the New World [J]. Ann Mo Bot Gard, 94(2): 407-422.
参考文献
RAMBAUT A, 2012. FigTree version 1. 4. 0. http: //tree. bio. ed. ac. uk/software/figtree/.
参考文献
SCHLECHTER R, 1905. Asclepiadaceae [J]. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 36(Beibl. 82): 91.
参考文献
SHAH SA, SULTAN A, WEN J, et al. , 2021. Taxonomy of Vincetoxicum s. str. (Asclepiadoideae, Apocynaceae) from southern Asia including three new species and resurrected names [J]. PhytoKeys, 179: 35-73.
参考文献
WICK RR, SCHULTZ MB, ZOBEL J, et al. , 2015. Bandage: interactive visualization of de novo genome assemblies [J]. Bioinformatics, 31(20): 3350-3352.
参考文献
XIONG WJ, LEE SY, LIU PP, et al. , 2019. Complete chloroplast genome of Vincetoxicum hainanense (Apocyna-ceae: Asclepiadoideae), an endangered liana endemic to China [J]. Mitochon DNA Part B Resour, 4(2): 3608-3609.
目录contents

    摘要

    秦岭藤属(Biondia Schltr.)是中国特有属,约13种,驼峰藤属(Merrillanthus Chun & Tsiang)是仅在我国与柬埔寨分布的单种属,目前两个属均被归并到白前属(Vincetoxicum Wolf),由于取样和系统发育分析尚存一些欠缺,因此其系统位置和归属尚需进一步确认。该文使用2个核糖体基因序列片段(ITS、ETS)、5个叶绿体基因序列片段(psbA-trnH、trnG、trnL、trnL-F、trnT-L),以及二者的合并数据,重建娃儿藤亚族(Tylophorinae)(包含属的模式秦岭藤 [B. chinensis Schltr.=V. shaanxiense (Schltr.) Meve & Liede]与驼峰藤 [M. hainanensis Chun & Tsiang=V. hainanense (Chun & Tsiang) Meve, H. H. Kong & Liede])的系统发育树。结果表明:秦岭藤属与驼峰藤属均包含在白前属中,秦岭藤与青龙藤 [B. henryi (Warb.) Tsiang & Li=V. henryi (Warb.) Meve & Liede]互为姐妹类群,并与V. kawaroense Meve & Liede聚为一支,而黑水藤 [B. insignis Tsiang=V. insigne (Tsiang) Meve, H. H. Kong & Liede]在另一分支中(亚热带分支Subtropical Clade);驼峰藤属与V. cissoides (Blume) Kuntze和V. philippicum Meve, Omlor & Liede聚为一支。核糖体和叶绿体基因数据均支持秦岭藤属与驼峰藤属归并入白前属,但秦岭藤属为多系,尚需收集更多种类和数据合并分析,深入探讨归并后白前属的种间系统关系和位置。

    Abstract

    Biondia Schltr., containing about 13 species, is endemic to China, and Merrillanthus Chun & Tsiang is a monotypic genus distributing only in China and Cambodia. These two genera have been included in Vincetoxicum Wolf, but their phylogenetic positions and taxonomic status need further study because of insufficient samples and phylogenetic analysis. We reconstructed the phylogenetic trees of Tylophorinae based on two ribosomal gene sequence data (ITS, ETS) and five chloroplast gene sequence data (psbA-trnH, trnG, trnL, trnL-F, trnT-L) individually and combined data, which contained two types of species samples [including B. chinensis Schltr.=V. shaanxiense (Schltr.) Meve & Liede and M. hainanensis Chun & Tsiang=V. hainanense (Chun & Tsiang) Meve, H. H. Kong & Liede]. The results showed that Biondia and Merrillanthus were nested inside the Vincetoxicum; the type species B. chinensis Schltr.=V. shaanxiense (Schltr.) Meve & Liede were sister to B. henryi (Warb.) Tsiang & Li=V. henryi (Warb.) Meve & Liede, and formed a clade with V. kawaroense Meve & Liede; while B. insignis Tsiang=V. insigne (Tsiang) Meve, H. H. Kong & Liede belonged to the Subtropical Clade; Merrillanthus formed a clade with V. cissoides (Blume) Kuntze and V. philippicum Meve, Omlor & Liede. Our results support that Biondia and Merrillanthus should be included in Vincetoxicum, but Biondia is polyphyletic, more species samples and data need to be collected and analysed to further explore their phylogenetic positions about Vincetoxicum.

  • 夹竹桃科(Apocynaceae)在Angiosperm Phylogeny Group IV系统(APG IV)中与钩吻科(Gelsemiaceae)、龙胆科(Gentianaceae)、马钱科(Loganiaceae)和茜草科(Rubiaceae)共同属于龙胆目(Gentianales)(Chase et al.,2016),包含传统界定的狭义夹竹桃科(Apocynaceae s.s.)和萝藦科(Asclepiadaceae)两大类群,约5 350种,主要分布在热带和亚热带地区(Endress et al.,2018)。夹竹桃科目前已建立了相对稳定的分子演化系统(Fishbein et al.,2018),一些修订的属作了归并或位置调整,目前总属数由422个(Endress &;Bruyns,2000)缩减到378个(Endress et al.,2018)。

  • 白前属(Vincetoxicum Wolf)隶属于夹竹桃科马利筋族(Asclepiadeae)娃儿藤亚族(Tylophorinae),范围变化较大(Endress et al.,2018)。传统的白前属通常茎直立,少数上部缠绕,须根簇生,花冠辐状,有5个肉质的副花冠裂片,乳汁透明(邱声祥等,1989;Liede,1996)。白前属与鹅绒藤属(Cynanchum L.)具有相似的副花冠、合蕊柱、花粉器等特征,部分学者认为白前属应归并在鹅绒藤属中(蒋英和李秉滔,1977;Forster,1991;Li et al.,1995),但有学者认为白前属应作为独立的属(Markgraf,1972;Ali &;Khatoon,1982)。邱声祥等(1989)结合化学成分、形态、染色体数目和分布区等证据,提出白前属应从鹅绒藤属中独立出来,Liede(1996)基于形态与化学证据也支持该观点,并认为其与娃儿藤属(Tylophora R. Br.)的关系最近。

  • 然而,分子系统学研究显示白前属与鹅绒藤属亲缘关系较远(Liede,2001;Rapini et al.,2007),支持白前属与娃儿藤属关系更近,但它们都是非单系的,而秦岭藤属(Biondia Schltr.)、Blyttia Arn.、Diplostigma K. Schum.、Goydera Liede、Pleurostelma Baill.和Rhyncharrhena F. Muell.6个属都嵌套在Vincetoxicum-Tylophora分支中(Liede-Schumann et al.,2012,2016)。娃儿藤属和上述6个属,以及形态上相近的一些属均被归并在白前属中,增加了大量的新组合或新名称(Kidyoo &;Kidyoo,2018;Liede-Schumann &;Meve,2018;Hsu et al.,2021;Shah et al.,2021)。原主要分布在中国的3个属,即驼峰藤属(Merrillanthus Chun &;Tsiang)、白水藤属(Pentastelma Tsiang &;Li)和箭药藤属(Belostemma Wall. ex Wight)也归并在白前属中。新定义的白前属有150多种,通常有透明乳汁;花小,花冠辐状,少数钟状、坛状、长圆锥状;副花冠由肉质分离的雄蕊上的裂片组成,或由合生的雄蕊和片状的雄蕊间的部分组成环状;花粉块柄直立、水平或斜上,分布于热带、亚热带非洲、亚洲、欧亚大陆等区域(Endress et al.,2018)。

  • 秦岭藤属约13种,是中国特有属,分布于我国东部和西南部;驼峰藤属仅有驼峰藤(Merrillanthus hainanensis Chun et Tsiang)1种,分布于我国的广东、海南以及柬埔寨(蒋英和李秉滔,1977;Li et al.,1995)。在Liede-Schumann等(2012,2016)的系统发育分析中,秦岭藤属包含2个种,即黑水藤(Biondia insignis Tsiang)和青龙藤 [B. henryi(Warb. ex Schltr. et Diels)Tsiang et P. T. Li]的样品,缺少属的模式秦岭藤(B. chinensis Schltr.)的样品,同时缺乏驼峰藤属的样品。部分学者报道了秦岭藤与驼峰藤的叶绿体基因组数据,并进行了简易的系统发育分析,其中秦岭藤显示与Vincetoxicum rossicum(Kleopow)Barbar.是姐妹类群,但系统树中仅包含白前属1个种的数据(Rao et al.,2018);驼峰藤则显示与黑水藤是姐妹类群,但系统发育树中的取样太少(Xiong et al.,2019),这2个种是否属于白前属,或属于白前属哪个分支等问题依旧不清楚。

  • 本研究收集了秦岭藤与驼峰藤的数据,其中新增驼峰藤的分子测序,秦岭藤的数据参考使用Rao等(2018)的测序结果,并结合白前属其他物种的分子数据(Liede-Schumann et al.,2016),开展了系统发育分析,拟进一步明确这2个属的系统位置和归属。

  • 1 材料与方法

  • 1.1 类群取样和分子序列数据来源

  • 秦岭藤(凭证标本:ZJB-2017-152-1,保存于陕西师范大学)数据来自Rao等(2018)中的浅层测序基因数据。对驼峰藤(凭证标本:LHB-AP17,保存于华南农业大学)进行叶片取样,经硅胶干燥后,将样品放入干冰中冷藏,快递至北京诺禾致源生物有限公司武汉分公司进行基因组总DNA的提取、小片段文库的建库与测序。具体操作如下:首先,DNA经检测合格后,先随机打断为350 bp左右的文库,再进行PE150双端测序,得到原始数据(raw data),经质控后得到最终的有效数据(clean data)10 Gb。然后,将秦岭藤的已有浅层测序数据与本次测序的驼峰藤的数据用GetOrganelle1.7(Jin et al.,2020)进行组装,设置参数为默认;使用Bandage 0.8(Wick et al.,2015)和Geneious Prime2019(https://www.geneious.com/)对组装得到的fastg文件进行可视化和序列提取,得到最终的秦岭藤的核糖体基因组序列,以及驼峰藤的叶绿体基因组与核糖体基因组序列。最后,分别以无油樟[Amborella trichopoda Baill.(AJ506156)]和罗布麻[Apocynum venetum L.(MT313688)]的叶绿体基因组作参考,先利用Plastid Genome Annotator(PGA)(Qu et al.,2019)对驼峰藤的叶绿体基因组进行注释,再根据log文档在Geneious Prime2019中作进一步的手动校正,得到叶绿体基因组的注释信息;以Asclepias coulteri A.Gray(JN665084)的核糖体基因组nrDNA连续片段(18S+ITS1+5.8S+ITS2+26S)和Vincetoxicum biglandulosum(Endl.)Kuntze(LN880610)的核糖体基因序列片段(ETS)为参考,在Geneious Prime2019中对秦岭藤与驼峰藤的核糖体基因组进行注释,并利用Excract选项提取出所需的核糖体基因序列片段与叶绿体基因序列片段,并将数据上传至GenBank(https://www.ncbi.nlm.nih.gov/)。

  • 基于已发表的娃儿藤亚族分子系统学研究数据(Liede-Schumann et al.,2016),补充驼峰藤和秦岭藤的DNA数据,选取鹅绒藤亚族(Cynanchinae)中的3个种为外类群,分别利用5个叶绿体基因序列片段(psbA-trnH、trnG、trnL、trnL-F、trnT-L)(137个种)、2个核糖体基因序列片段(ETS、ITS)(136个种),以及二者的合并数据构建系统发育树(139个种)。序列详细信息见表1,样品凭证标本详细信息见文献Liede-Schumann等(2016)。部分样品序列数据不全,相关序列矩阵中以缺失数据形式补齐。

  • 1.2 序列比对和拼接

  • 采用MAFFT软件(Katoh &;Standley,2013)单独对每个片段进行序列比对,多片段数据在Geneious Prime2019中进行序列拼接,得到联合矩阵数据。

  • 1.3 系统发育分析

  • 首先,利用IQ-TREE(Nguyen et al.,2015)基于最大似然法(maximum likelihood method,ML)对序列进行系统发育分析,IQ-TREE中ModelFinder按照BIC准则(Bayesian Information Criterion,BIC)自动测试,并选择出最佳替代模型,其中核糖体基因序列片段合并数据使用模型为TVM+F+R3、叶绿体基因序列片段合并数据使用模型为K3Pu+F+R3、核糖体与叶绿体基因序列片段合并数据使用模型为TVM+F+R3;然后,进行1 000次的SH-aLRT检验(Guindon et al.,2010)和超快自展值检验(ultrafast bootstrap approach,UFboot)(Minh et al.,2013),每个分支含有SH-aLRT和UFboot 2个支持率,如果SH-aLRT≥80%且UFboot≥95%,则视为得到较好支持,其分支结果可信;最后,将所得系统树在Figtree1.4.2(Rambaut,2012)中查看。

  • 表1 取样物种和序列GenBank编号

  • Table1 List of taxa sampled and GenBank accession numbers of sequences

  • 续表1

  • 续表1

  • 续表1

  • 续表1

  • 注: — 表示信息的缺失;加粗字体表示新增物种,它们的叶绿体基因组序列片段对应的序列号提供的是完整叶绿体基因组序列号,所用片段从中进行了截取。

  • Note: — indicates the absence of information; taxa in bold are newly added for the present study, their GenBank accession numbers corresponding to the plastid DNA data are provided for the complete chloroplast genome, and the plastid DNA data used were extracted from the complete chloroplast genome.

  • 2 结果与分析

  • 基于核糖体基因序列片段(ETS、ITS)构建的系统发育树显示,娃儿藤亚族中白前属是非单系的,Pentatropis R. Br. ex Wight &;Arn.嵌套在白前属中,并与Vincetoxicum apiculatum + V. sylvaticum + V. tenuipedunculatum成姐妹分支,支持率分别为SH-aLRT=82.8%、UFboot=82%(82.8/82)(图1);基于叶绿体基因序列片段以及合并数据构建的系统树都显示娃儿藤亚族分为2个大分支,Pentatropis R. Br. ex Wight &;Arn.与白前属呈姐妹关系,支持率分别为99.8/100、99.9/100(图2,图3)。

  • 3组分子矩阵数据所建系统树都高度支持驼峰藤属与V. cissoidesTylophora cissoides)和V. philippicumT. parviflora)聚为一支,图1、图2、图3中支持率分别为100/100、97.9/100、100/100;秦岭藤属中的3个种均嵌套在白前属分支中,但在系统树中非单系,其中在基于核糖体基因序列片段(ETS、ITS)所建的系统树与合并数据所建的系统树中,秦岭藤与青龙藤V. henryiBiondia henryi)互为姐妹类群,支持率分别为97.7/100、97.8/100(图1,图3),并与V. kawaroenseT. japonica)聚为一支,支持率分别为97.1/100、97.8/100,在基于叶绿体基因序列片段构建的系统树中,秦岭藤与青龙藤聚为一支,但支持率较低(31.3/67)(图2),而黑水藤在3组系统树中均显示在亚热带分支(Subtropical Clade)中(图1、图2、图3中支持率分别为100/100、93.8/100、100/100),并与V. villosumT. villosa)聚为一支(图1、图2、图3中支持率分别为74.3/100、0/96、72.6/100)。

  • 图1 基于2个核糖体基因序列片段(ETS、ITS)构建的娃儿藤亚族的系统发育树

  • Fig.1 Phylogeny tree of Tylophorinae constructed by two ribosomal gene sequence fragments (ETS, ITS)

  • 3 讨论与结论

  • 3.1 秦岭藤属的系统发育与属级分类地位

  • 在秦岭藤属建立时,Schlechter(1905)指出其花粉块下垂,与牛奶菜属(Marsdenia R. Br.)的直立花粉块存在明显差异;花冠坛状或近钟状,与娃儿藤属的花冠5深裂、辐状或广辐状有别,并认为秦岭藤属应放在马利筋亚族(Asclepiadinae)中。Gilbert等(1995)认为秦岭藤属的特点是叶狭窄,叶正面具有微小的苍白点,花小,花冠缩小,通常呈环状,花冠管多数较发达,并观察到娃儿藤属的一些种也有相似的苍白点,说明秦岭藤属与娃儿藤属形态性状存在交叉现象。娃儿藤属已被归并至白前属中(Liede-Schumann et al.,2012;Liede-Schumann &;Meve,2018;Endress et al.,2018),本研究的分子系统结果支持秦岭藤属归并入白前属的分类处理。

  • 图2 基于5个叶绿体基因序列片段(psbA-trnH、trnG、trnL、trnL-F、trnT-L)构建的娃儿藤亚族的系统发育树

  • Fig.2 Phylogeny tree of Tylophorinae constructed by five plastid gene sequence fragments (psbA-trnH, trnG, trnL, trnL-F, trnT-L)

  • 图3 基于5个叶绿体基因序列片段(psbA-trnH、trnG、trnL、trnL-F、trnT-L)和 2个核糖体基因序列片段(ETS、ITS)构建的娃儿藤亚族的系统发育树

  • Fig.3 Phylogeny tree of Tylophorinae constructed by five plastid gene sequence fragments (psbA-trnH, trnG, trnL, trnL-F, trnT-L) and two ribosomal gene sequence fragments (ETS, ITS)

  • 本研究分子系统分析结果显示,秦岭藤属的3个种都在白前属内,其中秦岭藤与青龙藤聚为一支,而黑水藤与V. villosumT. villosa)聚为一支,并在亚热带分支中,但它们在形态上差异较大,黑水藤的枝、叶柄均被单列短柔毛,叶线状披针形,除叶面中脉被微毛外,无毛,花冠近钟状,无毛,而V. villosum的枝、叶柄均密被柔毛,叶卵形或卵状长圆形,两面具毛,花冠辐状,具长柔毛;地理分布方面,前者分布于我国贵州、湖南、四川、西藏和云南(Li et al.,1995),后者分布于爪哇岛(Backer &;Bakhuizen,1965)。有关黑水藤与近缘种在形态和地理分布上差异较大,但亲缘关系较近的情况尚需进一步采集更多样品开展深入探究。

  • 3.2 驼峰藤属的系统位置与属级分类地位

  • 驼峰藤属在发表时被认为与徐长卿属(Pycnostelma Bunge ex Decne.)最相近(Chun &;Tsiang,1941),后者现已被归并至白前属中(Liede-Schumann &;Meve,2018),两者的副花冠都贴生于雄蕊,它们在生活型、副花冠裂片的形状、花冠裂片生长样式和果实形状大小等方面存在明显区别,其中驼峰藤属为木质藤本,副花冠裂片背部隆起,基部增厚,花冠裂片连生,未开花时裂片顶端向内连生,蓇葖果单生,纺锤形,大型;而徐长卿属为直立草本,副花冠向先端加厚,花冠深裂,果实披针形,较小(Chun &;Tsiang,1941);驼峰藤属的花冠裂片与醉魂藤属(Heterostemma Wight &;Arn.)和球兰属(Hoya R. Br.)类似,其膜质心形叶片与夜来香属(Telosma Coville)相似,驼峰藤属的花粉块下垂,另3个属的花粉块直立,可以明显区分(Chun &;Tsiang,1941)。蒋英和李秉滔(1977)发现驼峰藤属与印度产的原Iphisia Wight &;Arn=Vincetoxicum Wolf较相近,不同之处在于驼峰藤属的花冠裂片端部未开放时互相粘合,花粉块柄斜行上升,且花粉块完全倒垂,而Iphisia的花粉块柄先横平,次上升,后斜下。驼峰藤属的花部结构与宜昌娃儿藤 [T. augustiniana(Hemsl.)Craib=V. augustinianum(Hemsl.)Meve &;Liede]十分相似,但驼峰藤属的花更大。此外,由于驼峰藤属的蓇葖果大(长9~12 cm,直径3.5~4 cm),有较厚的纤维状中果皮,而娃儿藤属通常果皮薄,具有纸质的蓇葖果壁,因此驼峰藤属被保留为独立的属(Gilbert et al.,1995)。本研究未收集到宜昌娃儿藤的数据,但研究结果显示驼峰藤属与其他2个娃儿藤属V. cissoidesT. cissoides)和V. philippicumT. parviflora)聚为一支,这2个种果实均较大(前者长10~12 cm,直径3~5 cm;后者长13~15 cm,直径3~4 cm),叶大小与叶形与驼峰藤属很相似;地理分布上,V. cissoides广泛分布于印度尼西亚东部和新几内亚(Forster,1994),V. philippicum仅分布于菲律宾(Meve et al.,2002),而驼峰藤属分布于我国的广东、海南以及柬埔寨(Li et al.,1995)。因此,仅凭果实大小、花粉块柄的生长方式、叶子的形状与大小作为娃儿藤亚族内属的鉴别特征可能并不够充分,但这些特征可作为白前属内种间的鉴别特征。依据花粉块下垂的特征,驼峰藤属在最初建立时被置于马利筋族马利筋亚族中(Chun &;Tsiang,1941),自Endress和Bruyns(2000)合并夹竹桃科与萝藦科后,该属也被置于马利筋族中,但先后被置于鹅绒藤亚族(Endress et al.,2007)和娃儿藤亚族中(Endress et al.,2014;Endress et al.,2018)。本研究支持当前驼峰藤属归并至白前属,并放置在娃儿藤亚族中。

  • 3.3 白前属的系统发育

  • 虽然基于核糖体基因序列和叶绿体基因序列合并数据的系统发育分析显示白前属为单系,但其含有的信息位点依旧较少,白前属属内的种间系统发育关系支持率较低。基于高通量测序的系统发育分析将更清晰地重建白前属的系统发育分支关系。由于白前属的分类变动较大,因此非常有必要基于基因组学研究结果与形态特征分析进行全面的分类学修订。

  • 致谢 承蒙广东中山树木园廖浩斌老师在样品采集上提供帮助,陕西师范大学邹嘉宾老师提供分子数据,以及武汉植物园张彩飞老师协助数据获取,特此致谢。

  • 参考文献

    • ALI SI, KHATOON S, 1982. Genus Vincetoxicum Von Wolf (Asclepiadaceae) in Pakistan [J]. Pak J Bot, 14(1): 61-68.

    • BACKER CA, BAKHUIZEN VAN DEN BRINK RC, 1965. Flora of Java (spermatophytes only) Vol. II Angiospermae, families 111-160 [M]. Groningen: N. V. P. Noordhoff: 259-260.

    • CHASE MW, CHRISTENHUSZ MJM, FAY MF, et al. , 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV [J]. Biol J Linn Soc, 181(1): 1-20.

    • CHUN WY, TSIANG Y, 1941. Merrillanthus, a new genus of Asclepiadaceae from Hainan [J]. Sunyatsenia, 6(2): 105-108.

    • ENDRESS ME, BRUYNS PV, 2000. A revised classification of the Apocynaceae s. l. [J]. Bot Rev, 66(1): 1-56.

    • ENDRESS ME, LIEDE-SCHUMANN S, MEVE U, 2007. Advances in Apocynaceae: The enlightenment, an introduction [J]. Ann Mo Bot Gard, 94(2): 259-267.

    • ENDRESS ME, LIEDE-SCHUMANN S, Meve U, 2014. An updated classification for Apocynaceae [J]. Phytotaxa, 159(3): 175-194.

    • ENDRESS ME, MEVE U, MIDDLETON DJ, et al. , 2018. Apocynaceae [M]//KADEREIT JW, BITTRICH V. Flowering plants. Eudicots: Apiales, Gentianales (except Rubiaceae) 15. Cham: Springer International Publishing: 207-411.

    • FISHBEIN M, LIVSHULTZ T, STRAUB SCK, et al. , 2018. Evolution on the backbone: Apocynaceae phylogenomics and new perspectives on growth forms, flowers, and fruits [J]. Am J Bot, 105(3): 495-513.

    • FORSTER PI, 1991. A taxonomic revision of Cynanchum L. (Asclepiadaceae: Asclepiadoideae) in Australia [J]. Austrobaileya, 3(3): 443-466.

    • FORSTER PI, 1994. A taxonomic revision of Tylophora (Asclepiadaceae: Marsdenieae) in Papuasia [J]. Aust Syst Bot, 7(5): 485-505.

    • GILBERT MG, STEVENS WD, LI PT, 1995. Notes on the Asclepiadaceae of China [J]. Novon, 5(1): 1-16.

    • GUINDON S, DUFAYARD J, LEFORT V, et al. , 2010. Newalgorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3. 0 [J]. Syst Biol, 59(3): 307-321.

    • HSU T, HUANG W, CHUNG S, 2021. Vincetoxicum (Apocynaceae: Asclepiadoideae) in Taiwan: Two new combinations and a new record [J]. Phytotaxa, 478(2): 287-290.

    • JIANG(TSIANG) Y, LI BT(PT), 1977. Flora Reipublicae Popularis Sinicae: Vol. 63 [M]. Beijing: Science Press: 249-575. [蒋英, 李秉滔, 1977. 中国植物志: 第63卷 [M]. 北京: 科学出版社: 249-575. ]

    • JIN JJ, YU WB, YANG JB, et al. , 2020. GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes [J]. Genome Biol, 21(1): 241.

    • KATOH K, STANDLEY DM, 2013. MAFFT multiple sequence alignment software Version 7: improvements in performance and usability [J]. Mol Biol Evol, 30(4): 772-780.

    • KIDYOO A, KIDYOO M, 2018. A new species of Hoya (Marsdenieae), three new combinations and two new names in Vincetoxicum (Asclepiadeae) from Thailand [J]. Taiwania, 63(1): 25-31.

    • LI PT, GILBERT MG, STEVENS WD, 1995. Asclepiadaceae [M]//WU ZY, RAVEN PH, HONG DY. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press: 189-270.

    • LIEDE S, 1996. Cynanchum-Rhodostegiella-Vincetoxicum-Tylophora (Asclepiadaceae): New considerations on an old problem [J]. Taxon, 45(2): 193-211.

    • LIEDE S, 2001. Subtribe Astephaninae (Apocynaceae—Asclepiadoideae) reconsidered: New evidence based on cpDNA spacers [J]. Ann Missouri Bot Gard, 88(4): 657-668.

    • LIEDE-SCHUMANN S, KHANUM R, MUMTAZ AS, et al. , 2016. Going west — A subtropical lineage (Vincetoxicum, Apocynaceae: Asclepiadoideae) expanding into Europe [J]. Mol Phylogenet Evol, 94(A): 436-446.

    • LIEDE-SCHUMANN S, KONG H, MEVE U, et al. , 2012. Vincetoxicum and Tylophora (Apocynaceae: Asclepiadoideae: Asclepiadeae)—two sides of the same medal: Independent shifts from tropical to temperate habitats [J]. Taxon, 61(4): 803-825.

    • LIEDE-SCHUMANN S, MEVE U, 2018. Vincetoxicum (Apocynaceae—Asclepiadoideae) expanded to include Tylophora and allies [J]. Phytotaxa, 369(3): 129-184.

    • MARKGRAF F, 1972. Asclepiadaceae [M]//TUTIN TG, HEYWOOD VH, BURGES NA, et al. Flora Europaea: Volume 3. Diapensiacea to Myoporaceae. Cambridge: Cambridge University Press: 71.

    • MEVE U, OMLOR R, LIEDE-SCHUMANN S, 2002. A new combination in Tylophora (Apocynaceae, Asclepiadoideae) from the Philippines [J]. Syst Geogr Plant, 72(1/2): 27-32.

    • MINH BQ, NGUYEN MAT, VON HAESELER A, 2013. Ultrafast approximation for phylogenetic bootstrap [J]. Mol Biol Evol, 30(5): 1188-1195.

    • NGUYEN LT, SCHMIDT HA, VON HAESELER A, et al. , 2015. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies [J]. Mol Biol Evol, 32(1): 268-274.

    • QIU SX, LI DZ, ZHANG ZX, et al. , 1989. Chemotaxonomy of Cynanchum and its allied genera with notes on the generic characteristics of Vincetoxicum [J]. Acta Bot Yunnan, 11(1): 41-50. [邱声祥, 李德铢, 张壮鑫, 等, 1989. 鹅绒藤属及其近缘属的化学分类——兼论Vincetoxicum的恢复问题 [J]. 云南植物研究, 11(1): 41-50. ]

    • QU XJ, MOORE MJ, LI DZ, et al. , 2019. PGA: A software package for rapid, accurate, and flexible batch annotation of plastomes [J]. Plant Methods, 15(1): 1-12.

    • RAO H, WANG XJ, MA JX, et al. , 2018. Characterization of the complete chloroplast genome of Biondia chinensis (Apocynaceae: Asclepiadoideae: Asclepiadeae), a rare and threatened liana endemic to China [J]. Mitochon DNA Part B Resour, 3(2): 763-764.

    • RAPINI A, VAN DEN BERG C, LIEDE-SCHUMANN S, 2007. Diversification of Asclepiadoideae (Apocynaceae) in the New World [J]. Ann Mo Bot Gard, 94(2): 407-422.

    • RAMBAUT A, 2012. FigTree version 1. 4. 0. http: //tree. bio. ed. ac. uk/software/figtree/.

    • SCHLECHTER R, 1905. Asclepiadaceae [J]. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 36(Beibl. 82): 91.

    • SHAH SA, SULTAN A, WEN J, et al. , 2021. Taxonomy of Vincetoxicum s. str. (Asclepiadoideae, Apocynaceae) from southern Asia including three new species and resurrected names [J]. PhytoKeys, 179: 35-73.

    • WICK RR, SCHULTZ MB, ZOBEL J, et al. , 2015. Bandage: interactive visualization of de novo genome assemblies [J]. Bioinformatics, 31(20): 3350-3352.

    • XIONG WJ, LEE SY, LIU PP, et al. , 2019. Complete chloroplast genome of Vincetoxicum hainanense (Apocyna-ceae: Asclepiadoideae), an endangered liana endemic to China [J]. Mitochon DNA Part B Resour, 4(2): 3608-3609.

  • 参考文献

    • ALI SI, KHATOON S, 1982. Genus Vincetoxicum Von Wolf (Asclepiadaceae) in Pakistan [J]. Pak J Bot, 14(1): 61-68.

    • BACKER CA, BAKHUIZEN VAN DEN BRINK RC, 1965. Flora of Java (spermatophytes only) Vol. II Angiospermae, families 111-160 [M]. Groningen: N. V. P. Noordhoff: 259-260.

    • CHASE MW, CHRISTENHUSZ MJM, FAY MF, et al. , 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV [J]. Biol J Linn Soc, 181(1): 1-20.

    • CHUN WY, TSIANG Y, 1941. Merrillanthus, a new genus of Asclepiadaceae from Hainan [J]. Sunyatsenia, 6(2): 105-108.

    • ENDRESS ME, BRUYNS PV, 2000. A revised classification of the Apocynaceae s. l. [J]. Bot Rev, 66(1): 1-56.

    • ENDRESS ME, LIEDE-SCHUMANN S, MEVE U, 2007. Advances in Apocynaceae: The enlightenment, an introduction [J]. Ann Mo Bot Gard, 94(2): 259-267.

    • ENDRESS ME, LIEDE-SCHUMANN S, Meve U, 2014. An updated classification for Apocynaceae [J]. Phytotaxa, 159(3): 175-194.

    • ENDRESS ME, MEVE U, MIDDLETON DJ, et al. , 2018. Apocynaceae [M]//KADEREIT JW, BITTRICH V. Flowering plants. Eudicots: Apiales, Gentianales (except Rubiaceae) 15. Cham: Springer International Publishing: 207-411.

    • FISHBEIN M, LIVSHULTZ T, STRAUB SCK, et al. , 2018. Evolution on the backbone: Apocynaceae phylogenomics and new perspectives on growth forms, flowers, and fruits [J]. Am J Bot, 105(3): 495-513.

    • FORSTER PI, 1991. A taxonomic revision of Cynanchum L. (Asclepiadaceae: Asclepiadoideae) in Australia [J]. Austrobaileya, 3(3): 443-466.

    • FORSTER PI, 1994. A taxonomic revision of Tylophora (Asclepiadaceae: Marsdenieae) in Papuasia [J]. Aust Syst Bot, 7(5): 485-505.

    • GILBERT MG, STEVENS WD, LI PT, 1995. Notes on the Asclepiadaceae of China [J]. Novon, 5(1): 1-16.

    • GUINDON S, DUFAYARD J, LEFORT V, et al. , 2010. Newalgorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3. 0 [J]. Syst Biol, 59(3): 307-321.

    • HSU T, HUANG W, CHUNG S, 2021. Vincetoxicum (Apocynaceae: Asclepiadoideae) in Taiwan: Two new combinations and a new record [J]. Phytotaxa, 478(2): 287-290.

    • JIANG(TSIANG) Y, LI BT(PT), 1977. Flora Reipublicae Popularis Sinicae: Vol. 63 [M]. Beijing: Science Press: 249-575. [蒋英, 李秉滔, 1977. 中国植物志: 第63卷 [M]. 北京: 科学出版社: 249-575. ]

    • JIN JJ, YU WB, YANG JB, et al. , 2020. GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes [J]. Genome Biol, 21(1): 241.

    • KATOH K, STANDLEY DM, 2013. MAFFT multiple sequence alignment software Version 7: improvements in performance and usability [J]. Mol Biol Evol, 30(4): 772-780.

    • KIDYOO A, KIDYOO M, 2018. A new species of Hoya (Marsdenieae), three new combinations and two new names in Vincetoxicum (Asclepiadeae) from Thailand [J]. Taiwania, 63(1): 25-31.

    • LI PT, GILBERT MG, STEVENS WD, 1995. Asclepiadaceae [M]//WU ZY, RAVEN PH, HONG DY. Flora of China. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press: 189-270.

    • LIEDE S, 1996. Cynanchum-Rhodostegiella-Vincetoxicum-Tylophora (Asclepiadaceae): New considerations on an old problem [J]. Taxon, 45(2): 193-211.

    • LIEDE S, 2001. Subtribe Astephaninae (Apocynaceae—Asclepiadoideae) reconsidered: New evidence based on cpDNA spacers [J]. Ann Missouri Bot Gard, 88(4): 657-668.

    • LIEDE-SCHUMANN S, KHANUM R, MUMTAZ AS, et al. , 2016. Going west — A subtropical lineage (Vincetoxicum, Apocynaceae: Asclepiadoideae) expanding into Europe [J]. Mol Phylogenet Evol, 94(A): 436-446.

    • LIEDE-SCHUMANN S, KONG H, MEVE U, et al. , 2012. Vincetoxicum and Tylophora (Apocynaceae: Asclepiadoideae: Asclepiadeae)—two sides of the same medal: Independent shifts from tropical to temperate habitats [J]. Taxon, 61(4): 803-825.

    • LIEDE-SCHUMANN S, MEVE U, 2018. Vincetoxicum (Apocynaceae—Asclepiadoideae) expanded to include Tylophora and allies [J]. Phytotaxa, 369(3): 129-184.

    • MARKGRAF F, 1972. Asclepiadaceae [M]//TUTIN TG, HEYWOOD VH, BURGES NA, et al. Flora Europaea: Volume 3. Diapensiacea to Myoporaceae. Cambridge: Cambridge University Press: 71.

    • MEVE U, OMLOR R, LIEDE-SCHUMANN S, 2002. A new combination in Tylophora (Apocynaceae, Asclepiadoideae) from the Philippines [J]. Syst Geogr Plant, 72(1/2): 27-32.

    • MINH BQ, NGUYEN MAT, VON HAESELER A, 2013. Ultrafast approximation for phylogenetic bootstrap [J]. Mol Biol Evol, 30(5): 1188-1195.

    • NGUYEN LT, SCHMIDT HA, VON HAESELER A, et al. , 2015. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies [J]. Mol Biol Evol, 32(1): 268-274.

    • QIU SX, LI DZ, ZHANG ZX, et al. , 1989. Chemotaxonomy of Cynanchum and its allied genera with notes on the generic characteristics of Vincetoxicum [J]. Acta Bot Yunnan, 11(1): 41-50. [邱声祥, 李德铢, 张壮鑫, 等, 1989. 鹅绒藤属及其近缘属的化学分类——兼论Vincetoxicum的恢复问题 [J]. 云南植物研究, 11(1): 41-50. ]

    • QU XJ, MOORE MJ, LI DZ, et al. , 2019. PGA: A software package for rapid, accurate, and flexible batch annotation of plastomes [J]. Plant Methods, 15(1): 1-12.

    • RAO H, WANG XJ, MA JX, et al. , 2018. Characterization of the complete chloroplast genome of Biondia chinensis (Apocynaceae: Asclepiadoideae: Asclepiadeae), a rare and threatened liana endemic to China [J]. Mitochon DNA Part B Resour, 3(2): 763-764.

    • RAPINI A, VAN DEN BERG C, LIEDE-SCHUMANN S, 2007. Diversification of Asclepiadoideae (Apocynaceae) in the New World [J]. Ann Mo Bot Gard, 94(2): 407-422.

    • RAMBAUT A, 2012. FigTree version 1. 4. 0. http: //tree. bio. ed. ac. uk/software/figtree/.

    • SCHLECHTER R, 1905. Asclepiadaceae [J]. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 36(Beibl. 82): 91.

    • SHAH SA, SULTAN A, WEN J, et al. , 2021. Taxonomy of Vincetoxicum s. str. (Asclepiadoideae, Apocynaceae) from southern Asia including three new species and resurrected names [J]. PhytoKeys, 179: 35-73.

    • WICK RR, SCHULTZ MB, ZOBEL J, et al. , 2015. Bandage: interactive visualization of de novo genome assemblies [J]. Bioinformatics, 31(20): 3350-3352.

    • XIONG WJ, LEE SY, LIU PP, et al. , 2019. Complete chloroplast genome of Vincetoxicum hainanense (Apocyna-ceae: Asclepiadoideae), an endangered liana endemic to China [J]. Mitochon DNA Part B Resour, 4(2): 3608-3609.