植物研究 ›› 2023, Vol. 43 ›› Issue (3): 447-460.doi: 10.7525/j.issn.1673-5102.2023.03.015
田琴1,2, 邵成艳1,3, 段涵宁1, 杨晨璇1,4, 李璐1()
收稿日期:
2022-07-26
出版日期:
2023-05-20
发布日期:
2023-06-06
通讯作者:
李璐
E-mail:lilu@swfu.edu.cn
作者简介:
田琴(1994—),女,硕士研究生,研究方向为植物分类学。
基金资助:
Qin TIAN1,2, Chengyan SHAO1,3, Hanning DUAN1, Chenxuan YANG1,4, Lu LI1()
Received:
2022-07-26
Online:
2023-05-20
Published:
2023-06-06
Contact:
Lu LI
E-mail:lilu@swfu.edu.cn
About author:
E-mail:lilu@swfu.edu.cn.Supported by:
摘要:
堇菜属(Viola)植物种类多、分布广、形态变异大,物种鉴定和分类系统争议较大。因此,为解决该属 物种的分类学问题,以中国云南产的8种堇菜属植物为研究对象,利用显微镜和石蜡切片技术观察叶形态解剖特征,结果如下:(1)叶型可分为4种:肾形、戟形、卵形和三深裂;(2)叶片边缘和叶脉处稀疏分布着单细胞单列毛状体,可分为3类:短柱状、中柱状和长柱状;(3)叶表皮细胞垂周壁式样有平直-弓形、浅波状和深波状,气孔器类型有平列型、横列型和无规则型;气孔多为椭圆形,少数为近圆形;(4)叶中脉处上下表皮突起有均等型和不等型;(5)叶柄横切面轮廓和主维管束形态相关,前者为椭圆形、半圆形和近圆形,后者为浅U型、深U型和圆形。分析结果表明,堇菜属的叶型、表皮毛被、叶表皮细胞形态、气孔类型、叶中脉处的上下表皮形态、叶柄横切面及主维管束轮廓等,在种间差异明显,可以用作属内物种划分和近似种间的区别,支持长萼堇菜(V. inconspicua)、早开堇菜(V. prionantha)和紫花地丁(V. philippica)是一组有近缘关系的独立种。
中图分类号:
田琴, 邵成艳, 段涵宁, 杨晨璇, 李璐. 中国云南8种堇菜属植物的叶形态解剖特征及分类学意义[J]. 植物研究, 2023, 43(3): 447-460.
Qin TIAN, Chengyan SHAO, Hanning DUAN, Chenxuan YANG, Lu LI. Morph-anatomy of Leaf and Taxonomic Insights of Eight Viola species from Yunnan,China[J]. Bulletin of Botanical Research, 2023, 43(3): 447-460.
表1
8种堇菜属植物的基本信息[2]
编号 Number | 物种 Species | 亚属 Subgenus | 组 Section | 叶形 Leaf shape |
---|---|---|---|---|
1 | 双花堇菜 V. biflora | 二裂花柱亚属 Subgen. Dischidium | 短距堇菜组 Sect. Brevicalcaratae | 肾形、近圆形,或宽卵形 Reniform,Suborbicular, Broad ovate |
2 | 灰叶堇菜 V. delavayi | 宽卵形或三角状卵形 Broad ovate,or triangular | ||
3 | 长萼堇菜 V. inconspicua | 堇菜亚属 Subgen. Viola | 合生托叶组 Sect. Adnatae | 三角状卵形或戟形 Triangular or Hastate |
4 | 紫花地丁 V. philippica | 三角状卵形 Triangular-ovate | ||
5 | 早开堇菜 V. prionantha | 长圆状卵形 Oblong-ovate | ||
6 | 七星莲 V.diffusa | 蔓茎堇菜组Sect. Diffusae | 长圆状卵形 Oblong-ovate | |
7 | 南山堇菜 V. chaerophylloides | 裂叶堇菜组Sect. Pinnatae | 3-全裂叶 | |
8 | 香堇菜 V. odorata | 堇菜组 Sect. Viola | 圆形或肾形至宽卵形 Orbicular,Reniform,Ovate broadly or Ovate-cordate |
图版Ⅱ
8种堇菜属植物的3类叶表皮毛被类型1~3.短柱状毛(1,2.南山堇菜(中脉,叶缘);3.香堇菜);4~6.中柱状毛(4.紫花地丁;5.双花堇菜;6长萼堇菜);7~9.长柱状毛(7.早开堇菜;8.七星莲;9.灰叶堇菜)Plate Ⅱ Three types of leaf trichome in eight Viola species1-3.Short-column trichome(1-2.V. chaerophylloides(midrib,ridge);3.V. odorata);4-6.Medium-column trichome(4.V. philippica;5.V. biflora;6.V. inconspicua);7-9.Long-column trichome(7.V. prionantha;8.V. diffusa;9.V. delavayi)
图版Ⅲ
8种堇菜属植物的叶表皮特征1~2.双花堇菜(上表皮,下表皮);3~4.早开堇菜(上表皮,下表皮);5~6.香堇菜(上表皮,下表皮);7~8.灰叶堇菜(上表皮,下表皮);9~10.长萼堇菜(上表皮,下表皮);11~12.紫花地丁(上表皮,下表皮);13~14.七星莲(上表皮,下表皮);15.南山堇菜(上表皮)Plate Ⅲ Morphological characteristics of leaf epidermis in eight Viola species1, 2. Viola biflora(Adaxial epidermis,Abaxial epidermis); 3, 4. V. prionantha(Adaxial epidermis,Abaxial epidermis); 5, 6. V. odorata(Adaxial epidermis,Abaxial epidermis); 7, 8. V.delavayi(Adaxial epidermis,Abaxial epidermis); 9, 10. V. inconspicua(Adaxial epidermis,Abaxial epidermis); 11, 12. V. philippica(Adaxial epidermis,Abaxial epidermis); 13, 14. V.diffusa(Adaxial epidermis,Abaxial epidermis); 15. V. chaerophylloides(Adaxial epidermis)
表2
8种堇菜属植物的叶表皮形态特征
编号 No. | 种名 Species | 毛被(上/下表皮) Trichome(Ad/Ab) | 表皮细胞(上/下表皮) Epidermal | 气孔(上/下表皮) Stomata(Ad/Ab) | ||||
---|---|---|---|---|---|---|---|---|
长度 Length /μm | 类型 Type | 轮廓 Outline | 垂周壁 Anticlinal wall | 长宽比(形状) Ratio(Shape) | 气孔指数 SI | 气孔类型 Type | ||
1 | 双花堇菜 V. biflora | *249.45/— | 中柱状毛 Medium-colum | 多边形 Polygonal | 平直-弓形 Straight-arched | 1.31/1.29 椭圆形Elliptical | 0.15/0.30 | 不等 Anisocytic |
2 | 灰叶堇菜 V. delavayi | 903.69/676.39 | 长柱状毛 Long-column | *多边形/不规则型 Polygonal/Anomocytic | *平直-弓形/深波状 Straight-arched/Sinuolate | 1.27/1.28 椭圆形Elliptical | 0.04/0.27 | *不等;横列/不等;平列 Anisocytic;Diacytic/ Anisocytic;Paracytic |
3 | 长萼堇菜 V. inconspicua | *177.61/— | 中柱状毛 Medium-colum | 不规则型 Anomocytic | 浅波状 Sinuate | 1.26/1.09 近圆形Orbicular | 0.04/0.26 | *—/不等 —/Anisocytic |
4 | 紫花地丁 V. philippica | 219.61/162.88 | 中柱状毛 Medium-colum | 不规则型 Anomocytic | 深波状 Sinuolate | 1.43/1.36 椭圆形Elliptical | 0.10/0.13 | 不等 Anisocytic |
5 | 早开堇菜 V. prionantha | 311.89/201.49 | *长柱状毛/中柱状毛 Long-column/ Medium-colum | 不规则型 Anomocytic | *浅波状/深波状 Sinuate/Sinuolate | 1.24/1.38 椭圆形Elliptical | 0.07/0.29 | 不等 Anisocytic |
6 | 七星莲 V. diffusa | 630.46/310.82 | 长柱状毛 Long-column | 不规则型 Anomocytic | 深波状 Sinuolate | 1.31/1.24 椭圆形Elliptical | 0.09/0.24 | *不等;平列/不等 Anisocytic;Paracytic/ Anisocytic |
7 | 南山堇菜 V. chaerophylloides | 62.86/136.94 | 短状状毛 Short-colum | 不规则型 Anomocytic | *浅波状/深波状 Sinuate/Sinuolate | —/1.30 椭圆形Elliptical | —/0.19 | *—/不等 —/Anisocytic |
8 | 香堇菜 V. odorata | *67.14/— | 短状状毛 Short-colum | *多边形/不规则型 Polygonal/Anomocytic | *平直-弓形/深波状 Straight-arched/Sinuolate | 1.17/1.04 近圆形Orbicular | 0.14/0.23 | 不等 Anisocytic |
图版Ⅳ
8种堇菜属植物叶片主脉的横切面解剖特征1~4.均等型(1.香堇菜;2.紫花地丁;3.长萼堇菜;4.七星莲);5~8.不等型(5.灰叶堇菜;6.双花堇菜;7.早开堇菜;8.南山堇菜);C.晶体;E.表皮;Ph.韧皮部;Pp.栅栏组织;Sp:海绵组织;T.毛被;Xy.木质部Plate Ⅳ Cross sections of leaf midrib in eight Viola species1-4.Equal(1.Viola odorata;2.V. philippica;3.V. inconspicua;4.V. diffusa);5-8.Unequal(5.V. delavayi;6.V. biflora;7.V. prionantha;8.V. chaerophylloides);C.Crystal;E.Epidermis;Ph.Phloem;Pp.Palisade parenchyma;Sp.Spongy parenchyma;T.Trichome;Xy.Xylem
表3
8种堇菜属植物叶中脉和叶柄的横切面解剖特征
编号 Number | 种名 Species | 叶中脉横切面 Cross section of leaf midrib | 叶柄横切面 Cross section of petiole | |
---|---|---|---|---|
上/下表皮突起形态 Ad/Ab epidermis | 形态 Shape | 主维管束类型 Vascular bundle | ||
1 | 双花堇菜V. biflora | 不等型Unequal | 圆形Round outlines | 圆形Round outlines |
2 | 灰叶堇菜V.delavayi | 不等型Unequal | 近圆形Sub-orbicular outlines | 深U型Deep U-type |
3 | 长萼堇菜V. inconspicua | 均等型Equal | 椭圆形Elliptical1 outlines | 浅U型Shallow U-type |
4 | 紫花地丁V. philippica | 均等型Equal | 椭圆形Elliptical1 outlines | 浅U型Shallow U-type |
5 | 早开堇菜V. prionantha | 不等型Unequal | 椭圆形Elliptical1 outlines | 浅U型Shallow U-type |
6 | 七星莲V.diffusa | 均等型Equal | 椭圆形Elliptical1 outlines | 浅U型Shallow U-type |
7 | 南山堇菜V.chaerophylloides | 不等型Unequal | 近圆形Sub-orbicular outlines | 深U型Deep U-type |
8 | 香堇菜V. odorata | 均等型Equal | 近圆形Sub-orbicular outlines | 深U型Deep U-type |
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