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Typha × Glauca Godr., 香蒲属(香蒲科) 中国新记录杂种及其形态特征

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Typha × Glauca Godr., 香蒲属(香蒲科) 中国新记录杂种及其形态特征 第37卷 第1期 水生生物学报 Vol. 37, No.1 2013 年 1 月 ACTA HYDROBIOLOGICA SINICA Jan., 2 0 1 3 doi: 10.7541/2013.29 Typha × glauca Godr., 香蒲属(香蒲科) 中国新记录杂种及其形态特征 朱秀玉 王 东 (华中师范大学生命科学学院, 武汉 430079) 摘要: 香蒲属(Typha L.)为多年生水生或沼生草本植物, 种间存在十分普遍的杂交现象, 其中一些杂种在湿 地生态系统中有重要的作用。在查阅大量腊叶标本基础上, 结合野外居群生物学工作, 作者发现中国一新记 录杂种, 即 T. × glauca Godr. (T. angustifolia L. × T. latifolia L.), 并新拟“粉绿香蒲”作该杂种的中文名。对粉 绿香蒲的形态特征进行了研究, 讨论了其与亲本水烛和宽叶香蒲的区别, 并给出检索表。 关键词: 香蒲属; 水烛; 宽叶香蒲; 粉绿香蒲; 新记录 中图分类号: Q949.7 文献标识码: A 文章编号: 1000-3207(2013)01-0029-05 香蒲属(Typha L.)隶属香蒲科(Typhaceae), 世界 中国科学院成都生物研究所植物标本馆(CDBI)、中 约 24 种[1—4], 分布于南北两半球的温带和热带地 国科学院武汉植物园植物标本馆(HIB)、武汉大学植 区。我国约有 12 种, 南北广泛分布[5, 6]。该属种间 物标本馆(WH)、四川大学植物标本馆(SZ)等所有馆 存在十分普遍的杂交现象, 欧洲、美洲等地已经报 藏香蒲标本以及我们野外采集的香蒲标本[存放华 道有 7 个杂种[7—13], 其中 T. angustifolia × T. latifolia 中师范大学植物标本馆(CCNU)], 尤其是馆藏标本 (= T. × glauca Godr.)、T. angustifolia × T. domingensis 中被鉴定为水烛(T. angustifolia L.)和宽叶香蒲(T. 和 T. domingensis × T. latifolia (= T. × provincialis latifolia L.)的标本。形态特征对比研究包括外部形 Camus)[8]等 3 个杂种经过细胞遗传学实验确认。在 态、花、果实、种子及花粉等。 欧洲和北美等地, 香蒲属杂种是湿地尤其是在严重 1.2 形态观察和测量 人为干扰或易受较大水位波动影响生境中的优势植 用直接观察法研究植物的营养器官和繁殖器官 物, 它们在湿地生态系统中的作用一直受到植物学 的形态特征。在 NIKON SMZ-1500 体视显微镜下观 家和生态学家的关注[9—11, 14—19]。 察花、果实和种子, 在 OLYMPUS BX51 型光学显微 我们在研究香蒲属标本过程中, 发现采自新疆 镜下观察花粉粒, 并分别测量大小和拍照。取腊叶 西部的 10 份标本, 其形态特征属于杂种 T. × glauca 标本的花药, 并经冰醋酸浸泡、醋酸酐处理、离心 Godr.的范畴。该杂种以前在国内未见报道, 属于中 沉淀淘洗后制作装片, 用于花粉粒的观察和大小测 国的一分布新记录, 本文予以报道。 量。选取 10 个以上样本用于各项数据在显微镜下性 状的测量, 求平均值。 1 材料与方法 2 分类与形态学描述 1.1 腊叶标本研究 研究存放中国科学院植物研究所植物标本馆 粉绿香蒲(杂种) 新拟 图版Ⅰ: 6—11, 20 (PE)、中国科学院华南植物园植物标本馆(IBSC)、 Typha × glauca Godr. (T. angustifolia L. ×T. 收稿日期: 2011-12-06; 修订日期: 2012-04-13 基金项目: 国家自然科学基金项目(30870151); 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开放课题(LSEB2011- 06); 国家自然科学基金委重大国际合作项目(31110103911); 中国科学院战略性先导科技专项—应对气候变化的碳收 支认证及相关问题(XDA05050206)项目资助 作者简介: 朱秀玉(1984—), 女, 山东临沂人; 硕士研究生; 主要从事植物分类学和生态学研究。E-mail: [email protected] 通信作者: 王东, E-mail: [email protected] 30 水生生物学报 37 卷 latifolia L.). Fl. Lorr. 3: 20, 1844. 学者认为该分类群应作为种来处理外[7, 20], 多数欧 Synonymy: T. latifolia var. elongata Dudley in 美学者根据来自形态学、杂交、血清学、电泳、酶 Bul. Cornell Univ. (Sci.) 2: 102, 1886; T. angustifolia 和分子等方面的研究证据 都支持它是杂种的观 var. elongata (Dudley) Wiegand in Rhodora 26: 1, 1924. , [8, 12, 13, 21—24] 多年生水生或沼生草本, 雌雄同株。地上茎直 点 。据文献记载, 粉绿香蒲广泛分布于 [7, 9, 11—13, 15, 16] 立, 粗壮, 高约 2—3 m。叶条形, 粉绿色, 叶片长 欧洲、北美等地 , 如它在北美东北部大 73—140 cm, 宽 10—16 mm; 叶鞘耳状抱茎。雌雄 西洋沿岸到中部广阔的草原地带不但分布较广, 而 [9, 11] 花序分离或紧密连接, 相距(0—)1—12 mm; 雄花序 且数量多 。种加词“glauca”意指叶片粉绿色, 因 长 25—40 mm, 雄花由 1 枚或 2 枚雄蕊组成, 花药长 此本文新拟“粉绿香蒲”作该杂种的中文名。 约 2—3 mm, 长矩圆形, 花粉粒单体、二合体、三合 产我国新疆的粉绿香蒲与国外分布的植物在形 体或四合体; 雌花序长 26—43 cm, 宽 19—22 mm, 态上有一定的差异, 表现在叶片(宽度)、雌花序(大 深棕色; 雌花具小苞片, 小苞片长约 7.6 mm, 浅褐 小)、雌花序与雄花序的间距、柱头(形态、大小)等 色至先端白色, 比柱头窄; 孕性雌花柱头长 1.2— 性状方面(表 1)。在叶(叶片宽度、叶鞘形态)、雌花 1.6 mm, 宽 0.1—0.2 mm, 条状披针形, 子房狭披针 序(颜色、大小)、雌花(大小、孕性雌花的柱头、丝 形, 长约 1.2 mm; 不孕雌花子房长 0.9—1.5 mm, 近 状毛、小苞片以及不孕雌花的子房形态)等许多重要 倒圆锥形, 先端圆形。花果期 6—9 月。 的分种特征方面, 粉绿香蒲表现出介于宽叶香蒲和 China. Xinjiang(新疆): Huocheng(霍城), 生于 水烛两种之间的中间类型的性状。一方面, 它的雌 积水塘、人工沟渠中。2010–8–13, D. Wang(王东) 花序深棕色, 雌雄花序有时相互靠近或紧密连接, 6248, 6249(CCNU); Tacheng(塔城), 生于水塘中。 叶片具渐狭的鞘, 孕性雌花柱头条状披针形, 不孕 2010–8–9, D. Wang(王东)6223–1(CCNU)。中国分布 雌花子房倒圆锥形, 与宽叶香蒲的特征接近(图版Ⅰ: 新记录。欧洲(英国、法国)、北美(美国、加拿大) 2, 5, 7, 11); 另一方面, 它的叶片有时具耳状叶鞘, 等地也有分布。 孕性雌花具小苞片, 雌雄花序有时远离等特征, 又 与水烛的许多特征相符(图版Ⅰ: 9, 14)。同时它又有 3 讨论 明显的可鉴别特征, 如花粉粒除单体、四合体外, 还 粉绿香蒲最早是由法国学者 D. A. Godron 于 有二体、三体出现, 花粉粒常败育; 柱头条状披针形 1844 年在法国东北部 Lorraine 地区植物志 Flore de (比宽叶香蒲的卵状披针形柱头窄, 但比水烛的线形 Lorraine 中命名的一种植物。作者在原描述中指出 柱头宽); 雌花小苞片浅褐色至先端白色(宽叶香蒲 该植物在外部形态上与宽叶香蒲相近, 与水烛有明 雌花无小苞片, 水烛小苞片浅褐色至深褐色), 比柱 显区别。1886 年美国学者 W. R. Dudley 在美国纽约 头窄等(图版Ⅰ: 6—11, 20)。上述区别特征国外很多 州 Cayuga 湖泊发现一种与宽叶香蒲形态相似的植 学者都有记述[7— 9, 12,13]。另外, 粉绿香蒲与 T. × 物, 但因其雌花序长达 30 cm, 他把这种植物定名为 provincialis Camus 不同, 后者通常无耳状叶鞘, 在 T. latifolia var. elongata, 1924 年美国学者 K. M. 叶片基部和叶鞘的近轴面有黏液状腺体等; 与 T. Wiegand 在复查 Cayuga 湖泊水生植物时, 根据 T. angustifolia × T. domingensis 也不同, 后者花粉单 latifolia var. elongata 在外部形态上更接近于水烛而 体、小苞片急尖至渐尖等[8]。目前四合体花粉粒在 将其命名为 T. angustifolia var. elongata, 这两个变 香蒲属中见于宽叶香蒲、小香蒲(T. minima Funck)、 种名称后来被证明都属于粉绿香蒲[7]。除少数北美 T. alelseevii Mavrodiev、T. tzvelevii Mavrodiev、粉 表 1 粉绿香蒲的形态变异 Tab. 1 Comparison of morphological characters of Typha × glauca between our study and other studies 形态特征 本研究 Hotchkiss & Dozier Smith Grace & Harrison Kuehn & White Morphological characters This study (1949) (1967) (1986) (1999) 叶宽 Leaf width (mm) 10—16 6—12 5—8.5 5—19 6—21 雌花序长 Pistillate spike length (mm) 260—430 100—250 150—300 —— 100—281 雌花序宽 Pistillate spike width (mm) 19—22 18—25 19—20 14—27 6—25 雌雄花序间距 Spike interval (mm) 0—12 0—40 4—33 0—33 0—42 柱头宽 Stigma width (mm) 0.1—0.2 —— —— —— 0.5—0.8 1 期 朱秀玉等: Typha × glauca Godr., 香蒲属(香蒲科)中国新记录杂种及其形态特征 31 绿香蒲和 T. × provincialis, 而二体和三体花粉粒则 水烛和宽叶香蒲在我国西北、东北、华北、华中和 仅见于粉绿香蒲[1, 2, 4, 7, 13]。 华南等地均有广泛的分布, 两物种也有较大的分布 根据文献记载, 粉绿香蒲在北美是由于 20 世纪 重叠区, 理论上推测粉绿香蒲在我国也应该有较广 水烛从大西洋沿岸向其他地区迅速扩散并与乡土种 的分布, 但我们检查了我国主要植物标本馆馆藏中 宽叶香蒲杂交产生的, 在其亲本水烛和宽叶香蒲分 所有鉴定为水烛和宽叶香蒲的标本, 却仅在我们采 布的重叠区普遍存在[9, 12]。它依赖营养繁殖, 具有较强 自新疆的标本中发现有粉绿香蒲, 该杂种在我国的 的竞争优势与入侵扩张能力和较高的净初级生产力, 地理分布范围尚有待于进一步研究。下面给出粉绿 这不仅增加了底泥中有机物含量, 也影响了湿地植物 香蒲与亲本近缘种的分种检索表, 以更好的了解其 [8—10, 14—16, 18, 19] 群落的结构和功能以及沼泽演替方向 。 鉴别特征。 分种检索表 1. 雌花无小苞片; 柱头披针形或卵状披针形; 花粉四合体; 叶宽 5—25 mm; 叶通常与花序等长或稍超出 .......................................................................................................................................... 宽叶香蒲 T. latifolia 1. 雌花具小苞片; 柱头条形、线形或条状披针形; 花粉粒全部或部分单体; 叶明显超出花序。 2. 雌花序浅棕色; 柱头条形或线形; 小苞片先端浅棕色至深棕色, 比柱头宽; 花粉粒单体; 叶宽 3—8 mm ........................................................................................................................................水烛 T. angustifolia 2. 雌花序深棕色; 柱头条状披针形; 小苞片先端灰白色或近于无色, 比柱头窄; 花粉粒常单体、二体、 三体和四合体混合; 叶宽 10—16 mm........................................................................粉绿香蒲 T. × glauca 致谢: 类分析在东北水生植物香蒲属分类研究中的应用. 水生 感谢中国科学院植物研究所植物标本馆(PE)马 生物学报, 1998, 22(1): 86—89] [7] Hotchkiss N, Dozier H L. Taxonomy and distribution of 欣堂先生和田希娅女士、中国科学院华南植物园植 North American cattails [J]. American Midland Naturalist, 物标本馆(IBSC)杨亲二研究员、中国科学院武汉植 1949, 4l(1): 237—254 物园植物标本馆(HIB)李建强研究员、中国科学院成 [8] Smith S G. Experimental and natural hybrids in North America Typha (Typhaceae) [J]. American Midland Natu- 都生物研究所植物标本馆(CDBI)高信芬研究员、武 ralist, 1967, 78(2): 257—287 汉大学植物标本馆(WH)汪小凡教授和四川大学植 [9] Grace J B, Harrison J S. The Biology of Canadian Weeds. 73. 物标本馆(SZ)何兴金教授在检查标本中给予的热情 Typha latifolia L., Typha angustifolia L. and Typha × glauca Godr. [J]. Canadian Journal of Plant Science, 1986, 66(2): 帮助。 361—379 [10] Smith S G. Typha: its taxonomy and the ecological signi- 参考文献: ficance of hybrids [J]. Archiv für Hydrobiologie, 1987, 27(1): 129—138 [1] Graebner P. Typhaceae. In: Engler A. Das Pflanzenreich [M]. [11] Thieret J W, Luken J O. The Typhaceae in the southeastern Leipzig. 1900, 8(2): 8—16 United States [J]. Harvard Papers in Botany, 1996, 8, 27—56 [2] Mavrodiev E V. 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(TYPHACEAE) NEWLY RECORDED IN CHINA AND ITS MORPHOLOGICAL CHARACTERISTICS ZHU Xiu-Yu and WANG Dong (School of Life Sciences, Central China Normal University, Wuhan 430079, China) Abstract: Typha L.
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