朱秀玉 王 東

(華中師范大學生命科學學院, 武漢 430079)

Typha × glauca Godr., 香蒲屬(香蒲科)中國新記錄雜種及其形態(tài)特征

朱秀玉 王 東

(華中師范大學生命科學學院, 武漢 430079)

香蒲屬(Typha L.)為多年生水生或沼生草本植物, 種間存在十分普遍的雜交現(xiàn)象, 其中一些雜種在濕地生態(tài)系統(tǒng)中有重要的作用。在查閱大量臘葉標本基礎上, 結合野外居群生物學工作, 作者發(fā)現(xiàn)中國一新記錄雜種, 即T. × glauca Godr. (T. angustifolia L. × T. latifolia L.), 并新擬“粉綠香蒲”作該雜種的中文名。對粉綠香蒲的形態(tài)特征進行了研究, 討論了其與親本水燭和寬葉香蒲的區(qū)別, 并給出檢索表。

香蒲屬; 水燭; 寬葉香蒲; 粉綠香蒲; 新記錄

香蒲屬(Typha L.)隸屬香蒲科(Typhaceae), 世界約24種[1—4], 分布于南北兩半球的溫帶和熱帶地區(qū)。我國約有12種, 南北廣泛分布[5,6]。該屬種間存在十分普遍的雜交現(xiàn)象, 歐洲、美洲等地已經(jīng)報道有7個雜種[7—13], 其中T. angustifolia × T. latifolia (= T. × glauca Godr.)、T. angustifolia × T. domingensis和T. domingensis × T. latifolia (= T. × provincialis Camus)[8]等3個雜種經(jīng)過細胞遺傳學實驗確認。在歐洲和北美等地, 香蒲屬雜種是濕地尤其是在嚴重人為干擾或易受較大水位波動影響生境中的優(yōu)勢植物, 它們在濕地生態(tài)系統(tǒng)中的作用一直受到植物學家和生態(tài)學家的關注[9—11,14—19]。

我們在研究香蒲屬標本過程中, 發(fā)現(xiàn)采自新疆西部的10份標本, 其形態(tài)特征屬于雜種T. × glauca Godr.的范疇。該雜種以前在國內(nèi)未見報道, 屬于中國的一分布新記錄, 本文予以報道。

1 材料與方法

1.1 臘葉標本研究

研究存放中國科學院植物研究所植物標本館(PE)、中國科學院華南植物園植物標本館(IBSC)、中國科學院成都生物研究所植物標本館(CDBI)、中國科學院武漢植物園植物標本館(HIB)、武漢大學植物標本館(WH)、四川大學植物標本館(SZ)等所有館藏香蒲標本以及我們野外采集的香蒲標本[存放華中師范大學植物標本館(CCNU)], 尤其是館藏標本中被鑒定為水燭(T. angustifolia L.)和寬葉香蒲(T. latifolia L.)的標本。形態(tài)特征對比研究包括外部形態(tài)、花、果實、種子及花粉等。

1.2 形態(tài)觀察和測量

用直接觀察法研究植物的營養(yǎng)器官和繁殖器官的形態(tài)特征。在NIKON SMZ-1500體視顯微鏡下觀察花、果實和種子, 在OLYMPUS BX51型光學顯微鏡下觀察花粉粒, 并分別測量大小和拍照。取臘葉標本的花藥, 并經(jīng)冰醋酸浸泡、醋酸酐處理、離心沉淀淘洗后制作裝片, 用于花粉粒的觀察和大小測量。選取10個以上樣本用于各項數(shù)據(jù)在顯微鏡下性狀的測量, 求平均值。

2 分類與形態(tài)學描述

粉綠香蒲(雜種) 新擬 圖版Ⅰ: 6—11, 20

Typha × glauca Godr. (T. angustifolia L. ×T.latifolia L.). Fl. Lorr. 3: 20, 1844.

Synonymy: T. latifolia var. elongata Dudley in Bul. Cornell Univ. (Sci.) 2: 102, 1886; T. angustifolia var. elongata (Dudley) Wiegand in Rhodora 26: 1, 1924.

多年生水生或沼生草本, 雌雄同株。地上莖直立, 粗壯, 高約2—3 m。葉條形, 粉綠色, 葉片長73—140 cm, 寬10—16 mm; 葉鞘耳狀抱莖。雌雄花序分離或緊密連接, 相距(0—)1—12 mm; 雄花序長25—40 mm, 雄花由1枚或2枚雄蕊組成, 花藥長約2—3 mm, 長矩圓形, 花粉粒單體、二合體、三合體或四合體; 雌花序長26—43 cm, 寬19—22 mm,深棕色; 雌花具小苞片, 小苞片長約7.6 mm, 淺褐色至先端白色, 比柱頭窄; 孕性雌花柱頭長1.2—1.6 mm, 寬0.1—0.2 mm, 條狀披針形, 子房狹披針形, 長約1.2 mm; 不孕雌花子房長0.9—1.5 mm, 近倒圓錐形, 先端圓形?；ü?—9月。

China. Xinjiang(新疆): Huocheng(霍城), 生于積水塘、人工溝渠中。2010–8–13, D. Wang(王東) 6248, 6249(CCNU); Tacheng(塔城), 生于水塘中。2010–8–9, D. Wang(王東)6223–1(CCNU)。中國分布新記錄。歐洲(英國、法國)、北美(美國、加拿大)等地也有分布。

3 討論

粉綠香蒲最早是由法國學者D. A. Godron于1844年在法國東北部Lorraine地區(qū)植物志Flore de Lorraine中命名的一種植物。作者在原描述中指出該植物在外部形態(tài)上與寬葉香蒲相近, 與水燭有明顯區(qū)別。1886年美國學者W. R. Dudley在美國紐約州Cayuga湖泊發(fā)現(xiàn)一種與寬葉香蒲形態(tài)相似的植物, 但因其雌花序長達30 cm, 他把這種植物定名為T. latifolia var. elongata, 1924年美國學者K. M. Wiegand在復查Cayuga湖泊水生植物時, 根據(jù)T. latifolia var. elongata在外部形態(tài)上更接近于水燭而將其命名為T. angustifolia var. elongata, 這兩個變種名稱后來被證明都屬于粉綠香蒲[7]。除少數(shù)北美學者認為該分類群應作為種來處理外[7,20], 多數(shù)歐美學者根據(jù)來自形態(tài)學、雜交、血清學、電泳、酶和分子等方面的研究證據(jù), 都支持它是雜種的觀點[8,12,13,21—24]。據(jù)文獻記載, 粉綠香蒲廣泛分布于歐洲、北美等地[7,9,11—13,15,16], 如它在北美東北部大西洋沿岸到中部廣闊的草原地帶不但分布較廣, 而且數(shù)量多[9,11]。種加詞“glauca”意指葉片粉綠色, 因此本文新擬“粉綠香蒲”作該雜種的中文名。

產(chǎn)我國新疆的粉綠香蒲與國外分布的植物在形態(tài)上有一定的差異, 表現(xiàn)在葉片(寬度)、雌花序(大小)、雌花序與雄花序的間距、柱頭(形態(tài)、大小)等性狀方面(表 1)。在葉(葉片寬度、葉鞘形態(tài))、雌花序(顏色、大小)、雌花(大小、孕性雌花的柱頭、絲狀毛、小苞片以及不孕雌花的子房形態(tài))等許多重要的分種特征方面, 粉綠香蒲表現(xiàn)出介于寬葉香蒲和水燭兩種之間的中間類型的性狀。一方面, 它的雌花序深棕色, 雌雄花序有時相互靠近或緊密連接,葉片具漸狹的鞘, 孕性雌花柱頭條狀披針形, 不孕雌花子房倒圓錐形, 與寬葉香蒲的特征接近(圖版Ⅰ: 2, 5, 7, 11); 另一方面, 它的葉片有時具耳狀葉鞘,孕性雌花具小苞片, 雌雄花序有時遠離等特征, 又與水燭的許多特征相符(圖版Ⅰ: 9, 14)。同時它又有明顯的可鑒別特征, 如花粉粒除單體、四合體外, 還有二體、三體出現(xiàn), 花粉粒常敗育; 柱頭條狀披針形(比寬葉香蒲的卵狀披針形柱頭窄, 但比水燭的線形柱頭寬); 雌花小苞片淺褐色至先端白色(寬葉香蒲雌花無小苞片, 水燭小苞片淺褐色至深褐色), 比柱頭窄等(圖版Ⅰ: 6—11, 20)。上述區(qū)別特征國外很多學者都有記述[7—9,12,13]。另外, 粉綠香蒲與T. × provincialis Camus不同, 后者通常無耳狀葉鞘, 在葉片基部和葉鞘的近軸面有黏液狀腺體等; 與T. angustifolia × T. domingensis也不同, 后者花粉單體、小苞片急尖至漸尖等[8]。目前四合體花粉粒在香蒲屬中見于寬葉香蒲、小香蒲(T. minima Funck)、T. alelseevii Mavrodiev、T. tzvelevii Mavrodiev、粉綠香蒲和T. × provincialis, 而二體和三體花粉粒則僅見于粉綠香蒲[1,2,4,7,13]。

表1 粉綠香蒲的形態(tài)變異Tab. 1 Comparison of morphological characters of Typha × glauca between our study and other studies

根據(jù)文獻記載, 粉綠香蒲在北美是由于20世紀水燭從大西洋沿岸向其他地區(qū)迅速擴散并與鄉(xiāng)土種寬葉香蒲雜交產(chǎn)生的, 在其親本水燭和寬葉香蒲分布的重疊區(qū)普遍存在[9,12]。它依賴營養(yǎng)繁殖, 具有較強的競爭優(yōu)勢與入侵擴張能力和較高的凈初級生產(chǎn)力,這不僅增加了底泥中有機物含量, 也影響了濕地植物群落的結構和功能以及沼澤演替方向[8—10,14—16,18,19]。水燭和寬葉香蒲在我國西北、東北、華北、華中和華南等地均有廣泛的分布, 兩物種也有較大的分布重疊區(qū), 理論上推測粉綠香蒲在我國也應該有較廣的分布, 但我們檢查了我國主要植物標本館館藏中所有鑒定為水燭和寬葉香蒲的標本, 卻僅在我們采自新疆的標本中發(fā)現(xiàn)有粉綠香蒲, 該雜種在我國的地理分布范圍尚有待于進一步研究。下面給出粉綠香蒲與親本近緣種的分種檢索表, 以更好的了解其鑒別特征。

分種檢索表

1. 雌花無小苞片; 柱頭披針形或卵狀披針形; 花粉四合體; 葉寬5—25 mm; 葉通常與花序等長或稍超出..........................................................................................................................................寬葉香蒲 T. latifolia

1. 雌花具小苞片; 柱頭條形、線形或條狀披針形; 花粉粒全部或部分單體; 葉明顯超出花序。

2. 雌花序淺棕色; 柱頭條形或線形; 小苞片先端淺棕色至深棕色, 比柱頭寬; 花粉粒單體; 葉寬3—8 mm ........................................................................................................................................水燭 T. angustifolia

2. 雌花序深棕色; 柱頭條狀披針形; 小苞片先端灰白色或近于無色, 比柱頭窄; 花粉粒常單體、二體、三體和四合體混合; 葉寬10—16 mm........................................................................粉綠香蒲 T. × glauca

致謝:

感謝中國科學院植物研究所植物標本館(PE)馬欣堂先生和田希婭女士、中國科學院華南植物園植物標本館(IBSC)楊親二研究員、中國科學院武漢植物園植物標本館(HIB)李建強研究員、中國科學院成都生物研究所植物標本館(CDBI)高信芬研究員、武漢大學植物標本館(WH)汪小凡教授和四川大學植物標本館(SZ)何興金教授在檢查標本中給予的熱情幫助。

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TYPHA × GLAUCA GODR., A HYBRID OF TYPHA L. (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)

Typha L. (Typhaceae), one of the most common aquatic plants found in marshes and shallow water, is almost cosmopolitan. However, the distribution of approximately 24 species has two main centers: Eurasia and North America. To date 12 species have been reported in China. Typha was described by Linnaeus in 1753, and its taxonomy remains confusing because of the variability in its reproductive and vegetative characteristics, and its frequent hybridization. Seven hybrids of Typha species were reported from North America and Europe but none of them has been recorded in China. We have discovered a hybrid cattail, Typha × glauca Godr. (T. angustifolia L. × T. latifolia L.), through extensive field collections and herbarium studies. That was a newly recorded taxon for China. We studied the hybrid T. × glauca in the field and examined Typha collections from the following herbaria including PE, IBSC, CDBI, HIB, WU, SZ, and CCNU. Both vegetative and reproductive characteristics were measured for available specimens of the hybrid and parental species from China. Pollen grains of Typha species were collected from our field collections, and were examined under a light microscope at × 1000 magnification. Morphological features of the hybrid and parental species was illustrated and discussed, and an identification key for them was provided. The hybrid was morphologically variable and generally intermediate to the parental species. The most readily diagnostic features of the hybrid were: 1. leaves were glaucous; 2. pollen was typically a mixture of monads, dyads, triads, and tetrads (vs. tetrads in T. latifolia and monads in T. angustifolia), often abortive; 3. pistillate bracteoles were narrower than the stigma (vs. broader in T. angustifolia and absent in T. latifolia), and 4. the stigmas were liner-lanceolate (vs. linear in T. angustifolia, and lanceolate to ovate-lanceolate in T. latifolia). Importantly, the presence /or absence of dyads and triads of pollen grains could be used to distinguish the hybrid and parental species, implying that pollen morphology is of importance in taxonomic treatment of genus Typha. In North America, Typha × glauca is a well-known cattail hybrid and can occur wherever T. angustifolia and T. latifolia grow together and often out-competes the parental species in habitats that are heavily disturbed or subject to high-magnitude water level fluctuations. The hybrid plants could interfere with wetland communities by forming large monospecific stands, out-competing native species, and altering substrate characteristics, suggesting that the appearance of hybrid plants may be used as indicators of disturbance and, perhaps, of wetland health. After checking the available specimens assigned to either T. angustifolia or T. latifolia that were deposited in the Herbaria of China, we found that T. × glauca was confined to the Xinjiang Province of China. Given the paucity of field collections on Typha plants over the past several decades and thus resultant relatively limited information on the hybrid in China, the distributional range and its ecological role of the hybrid cattail need to be studied further. In summary, we concluded that the field identification characters of the leaves and inflorescences may be used with some degree of reliability to tentatively separate T. × glauca from the parental species, but microscopic floral characters should be used to confirm the identification. To positively identify T. × glauca and parental species microscopic floral characters must be used.

Typha L.; T. angustifolia L.; T. latifolia L.; T. × glauca; New record

Q949.7

A

1000-3207(2013)01-0029-05

10.7541/2013.29

圖版I PlateⅠ

1—5: 寬葉香蒲(1. 孕性雌花; 2. 柱頭; 3. 孕性子房; 4. 絲狀毛; 5. 不孕子房); 6—11: 粉綠香蒲(6. 孕性雌花; 7. 柱頭; 8. 孕性子房; 9.小苞片; 10. 絲狀毛; 11. 不孕子房); 12—17: 水燭(12. 孕性雌花; 13. 柱頭; 14. 孕性子房; 15. 小苞片; 16. 絲狀毛; 17. 不孕子房); 18—20: 花粉粒形態(tài)(18. 寬葉香蒲, 示四合體; 19. 水燭, 示單體; 20. 粉綠香蒲, 示單體、二體、三體、四體)
1—5: Typha latifolia (1. fertile flower; 2. stigma; 3. fertile ovary; 4. hairs; 5. sterile ovary); 6—11; T. × glauca (6. fertile flower; 7. stigma; 8. fertile ovary; 9. pistillate bracteoles; 10. hairs; 11. sterile ovary); 12—17: T. angustifolia (12. fertile flower; 13. stigma; 14. fertile ovary; 15. pistillate bracteoles; 16. hairs; 17. sterile ovary); 18—20: pollen grains (18. tetrad in T. latifolia; 19. monads in T. angustifolia; 20. the mixture of monad, dyad, triad, and tetrad in T. × glauca

2011-12-06;

2012-04-13

國家自然科學基金項目(30870151); 中國科學院植物研究所系統(tǒng)與進化植物學國家重點實驗室開放課題(LSEB2011-06); 國家自然科學基金委重大國際合作項目(31110103911); 中國科學院戰(zhàn)略性先導科技專項—應對氣候變化的碳收支認證及相關問題(XDA05050206)項目資助

朱秀玉(1984—), 女, 山東臨沂人; 碩士研究生; 主要從事植物分類學和生態(tài)學研究。E-mail: wetlandplant@163.com

王東, E-mail: dongwang.cn@gmail.com