蕨类植物的鳞片特征及演化I: 凤尾蕨科

doi:10.11983/CBB19118

植物学报 ›› 2020, Vol. 55 ›› Issue (2): 163-176.DOI: 10.11983/CBB19118

蕨类植物的鳞片特征及演化I: 凤尾蕨科

顾钰峰^1,^2,³,金冬梅¹,刘保东²,戴锡玲³,严岳鸿^1,^*()

1 中国科学院上海辰山植物园, 上海辰山植物科学研究中心, 上海 201602
2 哈尔滨师范大学, 黑龙江省普通高等学校植物生物学重点实验室, 哈尔滨 150025
3 上海师范大学生命科学学院, 上海 200234

收稿日期:2019-07-01 接受日期:2019-10-24 出版日期:2020-03-01 发布日期:2020-02-12
通讯作者: 严岳鸿
基金资助:
国家自然科学基金(31800450);生态环境部生物多样性调查与评估(No.2019HJ2096001006);上海市绿化和市容管理局科技攻关(No.F112422)

Morphology Characters and Evolution of Ferns Scale Ι: Pteridaceae

Yufeng Gu^1,^2,³,Dongmei Jin¹,Baodong Liu²,Xiling Dai³,Yuehong Yan^1,^*()

1 Shanghai Chenshan Plant Science Research Center, Shanghai Chenshan Botanical Garden, Chinese Academy of Sciences, Shanghai 201602, China
2 Key Laboratory of Plant Biology, College of Heilongjiang Province, Harbin Normal University, Harbin 150025, China
3 College of Life Sciences, Shanghai Normal University, Shanghai 200234, China

Received:2019-07-01 Accepted:2019-10-24 Online:2020-03-01 Published:2020-02-12
Contact: Yuehong Yan

1. 附表1 材料信息.pdf(71KB)

摘要/Abstract

摘要： 鳞片是蕨类植物体表常见附属物, 是蕨类植物非常显著的分类学特征。凤尾蕨科(Pteridaceae)在蕨类植物系统发育中的位置关系多次发生了变化, 不同学者对该科中包含的类群也有着不同的观点。通过对该科76种植物的鳞片进行取材, 利用解剖镜观察拍照, 对各属鳞片特征进行描述。结果显示, 鳞片特征在不同的属和亚科之间具有明显的形态差异。用GenBank数据库中的rbcL基因序列对所研究物种进行系统发育重建, 并对鳞片的边缘特征和筛孔类型进行祖先性状重建, 结果表明, 全缘型鳞片和均质型鳞片是凤尾蕨科鳞片的祖征性状, 非全缘类型和透明筛孔类型的鳞片是在后期演化过程中形成的特征。此外, 透明和不透明类型的筛孔其形成可能与蕨类植物生活环境中的光照强度有关。

关键词: 形态解剖学, 分类学, 鳞片, 系统发育, 凤尾蕨科

Abstract: Scales are epidermal appendages on the rhizomes and leaves of many ferns. Features of scales play an important role in classification of ferns. The phylogenetic position and delimitation of Pteridaceae were treated differently by different authors. Here we collected scales of 76 fern species in Pteridaceae, and observed them under a dissecting microscope. By comparing the morphological characters, we found that scales are different among genus and subfamilies. We reconstructed a phylogenetic tree with the plastid rbcL sequence of the species in this study downloaded from GenBank database and reconstruced the ancestral state for two selected characters (margin of the scale and mesh type). The results suggested that homogeneous scale and entire margin were plesiomorphic characters, while non-entire margin and transparent mesh were evolved late in evolutionary process. We also speculated that the formation of transparent or non-transparent mesh may relate to the light intensity in the habitats.

Key words: morphology and anatomy, classification, scales, phylogeny, Pteridaceae

顾钰峰,金冬梅,刘保东,戴锡玲,严岳鸿. 蕨类植物的鳞片特征及演化I: 凤尾蕨科. 植物学报, 2020, 55(2): 163-176.

Yufeng Gu,Dongmei Jin,Baodong Liu,Xiling Dai,Yuehong Yan. Morphology Characters and Evolution of Ferns Scale Ι: Pteridaceae. Chinese Bulletin of Botany, 2020, 55(2): 163-176.

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图/表 8

图1 凤尾蕨科植物的生境 (A) 凤了蕨; (B) 粗梗水蕨; (C) 红秆凤尾蕨; (D) 蜈蚣草; (E) 银粉背蕨; (F) 旱蕨; (G) 书带蕨; (H) 铁线蕨。Bars=20 cm

Figure1 The habitats of Pteridaceae (A) Coniogramme japonica; (B) Ceratopteris pteridoides; (C) Pteris amoena; (D) P. vittata; (E) Aleuritopteris argentea; (F) Pellaea nitidula; (G) Haplopteris flexuosa; (H) Adiantum capillus-veneris. Bars=20 cm

图2 珠蕨亚科的鳞片形态 (A) 尾尖凤了蕨; (B) 黑轴凤了蕨; (C) 峨眉凤了蕨; (D) 凤了蕨; (E) 井冈山凤了蕨; (F) 普通凤了蕨; (G) 疏网凤了蕨; (H) 直角凤了蕨; (I) 高山珠蕨。Bars=0.5 mm

Figure 2 Scale morphology of Cryptogrammoideae (A) Coniogramme caudiformis; (B) C. robusta; (C) C. emeiensis; (D) C. japonica; (E) C. jinggangshanensis; (F) C. intermedia; (G) C. wilsonii; (H) C. procera; (I) Cryptogramma brunoniana. Bars=0.5 mm

图3 水蕨亚科的鳞片形态 (A) 卤蕨; (B) 水蕨。Bars=0.5 mm

Figure 3 Scale morphology of Ceratopteridoideae (A) Cystopteris fragilis; (B) Ceratopteris thalictroides. Bars=0.5 mm

图4 凤尾蕨亚科的鳞片形态 (A) 蜈蚣草; (B) 粉叶蕨; (C) 翠蕨; (D) 竹叶蕨; (E) 金粉蕨; (F) 栗柄金粉蕨; (G) 野雉尾金粉蕨; (H) 西南凤尾蕨; (I) 凤尾蕨; (J) 岩凤尾蕨; (K) 狭叶凤尾蕨; (L) 鸡爪凤尾蕨; (M) 指叶凤尾蕨; (N) 刺齿半边旗; (O) 红秆凤尾蕨; (P) 多羽凤尾蕨; (Q) 条纹凤尾蕨; (R) 全缘凤尾蕨; (S) 溪边凤尾蕨; (T) 三色凤尾蕨; (U) 有刺凤尾蕨; (V) 紫轴凤尾蕨; (W) 栗柄凤尾蕨; (X) 狭羽凤尾蕨。Bars=0.5 mm

Figure 4 Scale morphology of Pteridoideae (A) Pteris vittata; (B) Pityrogramme calomelanos; (C) Anogramma microphylla; (D) Taenitis blechnoides; (E) Onychium siliculosum; (F) O. japonicum var. lucidum; (G) O. japonicum; (H) P. wallichiana; (I) P. cretica var. nervosa; (J) P. deltodon; (K) P. henryi; (L) P. gallinopes; (M) P. dactylina; (N) P. dispar; (O) P. amoena; (P) P. decrescens; (Q) P. cadieri; (R) P. insignis; (S) P. excelsa; (T) P. aspericaulis var. tricolor; (U) P. setuloso-costulata; (V) P. aspericaulis; (W) P. plumbea; (X) P. stenophylla. Bars=0.5 mm

图5 碎米蕨亚科的鳞片形态 (A) 欧洲金毛裸蕨; (B) 川西金毛裸蕨; (C) 金毛裸蕨; (D) 耳羽金毛裸蕨; (E) 黑心蕨; (F) 泽泻蕨; (G) 隐囊蕨; (H) 中华隐囊蕨; (I)薄叶碎米蕨; (J) 毛轴碎米蕨; (K) 平羽碎米蕨; (L) 银粉背蕨; (M) 金粉背蕨; (N) 粉背蕨; (O) 西藏粉背蕨; (P) 旱蕨; (Q) 滇西旱蕨。Bars=0.5 mm

Figure 5 Scale morphology of Cheilanthoideae (A) Paragymnopteris marantae; (B) Pa. bipinnata; (C) Pa. vestita; (D) Pa. bipinnata var. auriculata; (E) Doryopteris concolor; (F) Hemionitis arifolia; (G) Notholaena hirsuta; (H) N. chinensis; (I) Cheilosoria tenuifolia; (J) Ch. chusana; (K) Ch. patula; (L) Aleuritopteris argentea; (M) Al. chrysophylla; (N) Al. pseudofarinosa; (O) Al. subvillosa var. tibetica; (P) Pellaea nitidula; (Q) P. mairei. Bars=0.5 mm

图6 书带蕨亚科的鳞片形态 (A) 连孢一条线蕨; (B) 美叶车前蕨; (C) 长柄车前蕨; (D) 姬书带蕨; (E) 剑叶书带蕨; (F) 书带蕨; (G) 曲鳞书带蕨; (H) 带状书带蕨; (I) 平肋书带蕨; (J) 线叶书带蕨; (K) 灰背铁线蕨; (L) 扇叶铁线蕨; (M) 条裂铁线蕨; (N) 铁线蕨; (O) 肾盖铁线蕨; (P) 冯氏铁线蕨; (Q) 长盖铁线蕨; (R) 半月形铁线蕨; (S) 鞭叶铁线蕨; (T) 假鞭叶铁线蕨; (U) 梅山铁线蕨; (V) 仙霞铁线蕨; (W) 白垩铁线蕨; (X) 小铁线蕨。Bars=0.5 mm

Figure 6 Scale morphology of Vittarioideae (A) Monogramma paradoxa; (B) Antrophyum callifolium; (C) A. obovatum; (D) Haplopteris anguste-elongata; (E) H. amboinensis; (F) H. flexuosa; (G) H. plurisulcata; (H) H. doniana; (I) H. fudzinoi; (J) H. linearifolia; (K) Adiantum myriosorum; (L) Ad. flabellulatum; (M) Ad. capillus-veneris var. dissectum; (N) Ad. capillus-veneris; (O) Ad. erythrochlamys; (P) Ad. fengianum; (Q) Ad. fimbriatum; (R) Ad. philippense; (S) Ad. caudatum; (T) Ad. malesianum; (U) Ad. meishanianum; (V) Ad. juxtapositum; (W) Ad. gravesii; (X) Ad. mariesii. Bars=0.5 mm

图7 基于叶绿体基因rbcL序列用最大似然法构建的凤尾蕨科系统发生树用MrBayes 3.2.7软件基于贝叶斯信息准则(BIC)在jModelTest最优模型进行BI分析。每条链跑样1 000 000代, 每1 000代进行抽样。各分支上数字代表支持率。A: 珠蕨亚科; B: 水蕨亚科; C: 凤尾蕨亚科; D: 碎米蕨亚科; E: 书带蕨亚科

Figure 7 Phylogenetic tree of Pteridaceae obtained from the maximum likelihood analysis of the plastid rbcL sequence The BI analysis was performed in MrBayes 3.2.7 based on the best model Bayesian information criterion (BIC) implemented in jModelTest. Each chains was run for 1 000 000 generations, and was sampled every 1 000 generations. Numbers on branches indicate support values. A: Cryptogrammoideae; B: Ceratopteridoideae; C: Pteridoideae; D: Cheilanthoideae; E: Vittarioideae

图8 鳞片边缘和筛孔类型的祖先性状在Bayes中利用叶绿体基因rbcL序列在最大似然法(ML)下重建系统树系统树中只采用凤尾蕨科的物种。饼状图显示每个节点缺失的百分比和每个节点的状态在所有树中似然的平均值。(L) 筛孔类型; (R) 鳞片边缘。A: 珠蕨亚科; B: 水蕨亚科; C: 凤尾蕨亚科; D: 碎米蕨亚科; E: 书带蕨亚科

Figure 8 Ancestral state reconstruction for margin of scales and mesh type characters optimized onto the best tree obtained in the plateau phase of the Bayesian with the maximum likelihood (ML) analysis of plastid gene rbcL sequence Only species of Pteridaceae were used for the phylogeny. Pie charts show the percentage of node absence and the average likelihood received by each state across all the trees possessing that node. (L) Mesh type; (R) Margin of scales. A: Cryptogrammoideae; B: Ceratopteridoideae; C: Pteridoideae; D: Cheilanthoideae; E: Vittarioideae

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蕨类植物的鳞片特征及演化I: 凤尾蕨科

Morphology Characters and Evolution of Ferns Scale Ι: Pteridaceae

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