摘"要: "半耳箬竹[Indocalamus semifalcatus (H. R. Zhao et Y. L. Yang) T. P. Yi]在原始文獻(xiàn)中僅有部分營養(yǎng)器官的描述。該文通過野外居群調(diào)查、室內(nèi)體視解剖和掃描電子顯微鏡(SEM)觀察,新增了半耳箬竹的花器官描述和葉下表皮微形態(tài)特征,完善了其營養(yǎng)器官的性狀描述,更新了其地理分布。結(jié)果表明:(1)繁殖器官性狀:花序為圓錐狀,小穗及小穗軸密被白色短柔毛,穎片、外稃及內(nèi)稃光滑無毛,雄蕊3枚,花藥紫紅色,柱頭2,白色,羽毛狀。(2)營養(yǎng)體性狀:稈高達(dá)4.5 m,徑達(dá)2 cm,籜耳半鐮形或微弱,籜片直立緊貼稈,葉耳微弱或無,葉舌上具較發(fā)達(dá)的纖毛;葉片兩面同色且無毛。(3)葉下表皮微形態(tài)特征:氣孔器凹陷不可見,8~10個長乳突平鋪覆蓋氣孔,硅質(zhì)體馬鞍形,未見有大毛和刺毛。(4)新分布區(qū)域1個,即貴州省貴陽市觀音山。該種與箬葉竹(I. longiauritus Hand.-Mazz.)最為相似,主要區(qū)別在于該種的籜鞘和葉鞘上具有半鐮形的籜耳或籜耳缺失,稈高達(dá)4.5 m,徑達(dá)2 cm。
關(guān)鍵詞: 竹類, 箬竹屬, 描述, SEM, 地理分布
中圖分類號: "Q949
文獻(xiàn)標(biāo)識碼: "A
文章編號: "1000-3142(2024)04-0650-07
Morphological supplementary description of Indocalamus
semifalcatus (H. R. Zhao et Y. L. Yang)
T. P. Yi (Bambusoideae)
GAO Liqin1,2, LI Yonglong1, CUI Ling1, YANG Guangyao1, ZHANG Wengen1*
( 1. Jiangxi Provincial Key Laboratory for Bamboo Germplasm Resources and Utilization, Jiangxi Agricultural University,
Nanchang 330045, China; 2. Jiangxi Academy of Forestry, Nanchang 330032, China )
Abstract: "In the protologue of Indocalamus semifalcatus (H. R. Zhao et Y. L. Yang) T. P. Yi, there were only parts of vegetative organs reported. Based on field population investigation, indoor stereoscopic anatomy and scanning electron microscope (SEM), its reproductive organs, complete vegetative organs and micromorphological features of the abaxial leaf epidermis, were described and illustrated herein, and its geographical distribution was also updated. The results were as follows: (1) Reproductive organ features: panicle, spikelet and spike densely white pubescent, glume, pelea and lemma both smooth and glabrous, stamens 3, anthers purple, stigmas 2, white, feathery. (2) Vegetative features: culms up to 4.5 m tall, up to 2 cm in diameter, culm sheath auricle semifalcate or weak, culm sheath erect close to culm, foliage auricle weak or absent, ligule margin with densely ciliate; both sides of leaf blade homochromatic and glabrous. (3) Micromorphologic characteristics of the abaxial leaf: stomatal apparatus invisible, eight to ten elongated papillae covered the stomatal apparatus, silica bodies saddle-shaped, micro-hairs and macro-hairs both absent. (4) The new distribution area was Guanyin Mountain of Guiyang in Guizhou. The species is the most similar to I. longiauritus Hand.-Mazz., differed by culm sheath and leaf sheath with auricles semifalcate or rare, culm up to 4.5 m tall and 2 cm in diameter.
Key words: Bamboo, Indocalamus, description, SEM, geographical distribution
箬竹屬(Indocalamus Nakai)是一類灌木狀木本竹子,地下莖復(fù)軸型,葉片寬大常用于食品包裝(耿伯介和王正平,1996;Wang amp; Stapleton,2006)或竹葉黃酮提?。ㄙ嚍缘?,2013;李夏冰,2017;張亞蘭等,2022)。本屬很多竹種對鉻、鉛、鎘、銅等重金屬污染有較強的耐受性和抗性(楊勝香等,2012;馬迎莉等,2019;張穎等,2022),抗寒和抗旱性強(李娟和高健,2016;劉思奇,2019),適用于礦區(qū)等重金屬污染嚴(yán)重土壤的修復(fù)。
箬竹屬是由日本學(xué)者中井猛之進(jìn)于1925年建立的,屬建立的核心依據(jù)是源自與赤竹屬(Sasa Makino amp; Shibata)花器官的比較,即箬竹屬雄蕊3、柱頭2而明顯有別于赤竹屬花器官雄蕊6、柱頭3 (Nakai, 1925)。屬建立至今,發(fā)表在屬下的有效雙名共計78個,經(jīng)耿以禮、趙奇僧、趙惠如、楊雅玲、Vorontsova等國內(nèi)外竹類分類學(xué)家不斷地研究,目前最新資料記錄該屬包含33種。除了Indocalamus petelotii (A. Camus) Ohrnb.分布于越南,該屬其余種均分布于中國(耿以禮, 1959;趙奇僧等,1980;趙惠如和楊雅玲,1985;楊雅玲,1987; 楊雅玲和趙惠如,1990;Vorontsova et al., 2017),但有花部性狀描述的種類僅占該屬的1/4 (Wang amp; Stapleton,2006)??梢?,完善箬竹屬種類花部性狀的工作任重而道遠(yuǎn)。
半耳箬竹 [Indocalamus semifalcatus (H. R. Zhao et Y. L. Yang) T. P. Yi] (2000: 26),基名為Indocalamus longiauritus var. semifalcatus H. R. Zhao et Y. L. Yang (1985: 464),均無生殖器官性狀特征的描述且營養(yǎng)器官描述較為簡單,其基名的原文僅有特征集要“A typo culmorum vaginarum auriculis et foliorum auriculis omnibus semifalcatis, laminis subtus secus costam utrinque glabris differt”(本變種與原變種的區(qū)別在于籜耳和葉耳均為半鐮形,葉片下面的中脈兩側(cè)無毛) (趙惠如和楊雅玲,1985)。其模式標(biāo)本“趙惠如無號”(H. R. Zhao s. n.; N)館藏于南京大學(xué),采自四川省灌縣(1988年5月改為都江堰市)二王廟附近。2000年,該分類群被易同培先生提升為種(易同培,2000),已收錄在《中國竹類圖志》(易同培等,2008)和《中國竹亞科屬種檢索表》(易同培等,2009)。
為了完善該分類群的描述和特征集要,2019—2020年,筆者在半耳箬竹模式產(chǎn)地四川省灌縣進(jìn)行了比較詳細(xì)的野外居群調(diào)查,在崇州市三郎鎮(zhèn)意外發(fā)現(xiàn)正在開花的半耳箬竹居群,在貴州省貴陽市觀音山見到成片的野生群落。基于室內(nèi)解剖和葉下表皮電鏡掃描觀察,本文在此對半耳箬竹的形態(tài)描述和地理分布予以補充完善,并增加葉下表皮微形態(tài)特征。
半耳箬竹"圖版I、圖版Ⅱ和圖版Ⅲ
Indocalamus semifalcatus[KG2mm] (H. R. Zhao et Y. L.
Yang) T. P. Yi in J. Bamboo. Res. 19(1): 26. 2000. ≡ Indocalamus longiauritus var. semifalcatus H. R. Zhao et Y. L. Yang in Acta. Phytotax. Sin. 23(6): 464. 1985. Type: — China. Sichuan, Dujiangyan, Guanxian, Erwangmiao, 12 Oct. 1979, H. R. Zhao s. n.
(holotype: N).
Description. Rhizomes leptomorph. Culms 1.2-4.5 m tall; internodes terete, 15-76 cm long, 0.8-2.0 cm in diameter, hollow, sparsely strigose, and with a ring of brown velvet at the infranodal region; wall 1.0-3.0 mm thick; supranodal ridge slightly raised; intranodes 7-11 mm long; culm branches from the base at nodes 3 to 5, usually solitary, nearly as thick as culms. Culm sheaths purple-green, persistent, usually shorter than internodes, leathery; dark brown strigose and white tomentose, base with raised corky ring; margin densely reddish-brown
ciliate; auricles short semifalcate, purple-green, becoming brown when dry, occasionally inconspicuous or absent; oral setae radiate, brown, ca. 0.5-1 cm; ligules 0.5-1.0 mm tall, truncate, margin sparsely or not "ciliate; blades purple-green, narrowly triangular to ovate-lanceolate, base abruptly rounded, apex acuminate. Foliage leaves 3-7 per ultimate branch; leaf sheath rigid, glabrous or abaxially initially finely strigose, outer margin smooth; auricles developed or inconspicuous; oral setae radiate, short; ligules truncate, 2-3 mm tall, margin densely ciliate; blades oblong-lanceolate, 30.0-50.0 × 5.0-9.0 cm, both surfaces glabrous, longitudinal veins 10-15 pairs, base cuneate, apex long-acuminate, margin entire. Panicles 8-25 cm, loose and spread, axis densely white tomentose. Spikelets greenish or straw-colored at maturity, 2-6 cm; florets 3-5. Rachilla internodes compressed-clavate, 4-5.5 mm, angular, densely white tomentose. Glumes 2, lanceolate; first glume 3-5 mm, second glume 8-12 mm; lemma oblong-lanceolate, apex awnlike, smooth hairless, the first lemma subequal to the second lemma, 1-1.2 cm long; lodicules 3, oblong-lanceolate, ciliate, distally sparsely pilose. Anthers purple, 8-9 mm; filaments white, 8-12 mm; stigmas 2, white, feathery, ca. 5-7 mm. Caryopsis unknown.
Phenology. New shoots Apr. -May, fl. Jul. -Aug.
Distribution and habitat. Indocalamus semifal-catus is endemic to China, distributed in Guangxi, Sichuan, Fujian and Guizhou, cultivated in Zhejiang. It usually grows in mountain slopes, hillsides or roadsides at elevations of 600-1 100 m.
Leaf micromorphology. One to two rows of stomatal apparatuses are usually distributed between veins (Plate IV: A). Stomatal apparatuses are sub-orbicular; eight to ten elongated papillae overarch the stomatal apparatus, and short papillae are rare (Plate IV: C, D); micro-hairs distributed in the intercostal region (Plate IV: B); saddle-shaped silica bodies (Plate IV: B) are distributed in the costal and intercostal regions.
Additional specimens examined. China. Sichuan: Guanxian, Erwangmiao, T. P. Yi 76008 (SIFS?。?L. Q. Gao et L. Cui SCGX1904 (JXAU?。?L. Q. Gao "et al. "SCGX2022 (JXAU?。? Chongzhou, Sanlangzhen, L. Q. Gao et L. Cui SCCZ1905 (FL., JXAU?。?L. Q. Gao "et al. "SCCZ2023 (FL., JXAU?。? Guizhou: Guiyang, Guanyinshan, L. Q. Gao "et al. "GZGY2024 (JXAU?。? Guangxi: Shanglin, Damingshan, L. Q. Gao et al. GXSL2025 (JXAU?。?
根狀莖細(xì)長型。稈高1.2~4.5 m,稈徑8~20 mm,節(jié)間長15~76 cm,幼稈節(jié)下被稀疏疣基刺毛和一圈棕色的絨氈毛。稈壁厚1~3 mm;稈環(huán)略隆起,較籜環(huán)略高;節(jié)內(nèi)長度7~11 mm;稈自基部第3~5節(jié)處開始分枝,稈每節(jié)1分枝,與主稈近等大或略小。新鮮稈籜紫紅色帶綠色,革質(zhì),背面被棕黃色伏貼的疣基刺毛,尤以基部為甚,靠近上部光滑無毛,宿存或脫落,基部常具一圈或半圈木栓質(zhì)圈;籜耳半鐮形,紫色略帶綠色,干后呈棕色,偶微弱或缺失;鞘口疏生0.5~1 cm的短繸毛。每小枝具葉片3~7枚;葉耳微弱或無,鞘口繸毛稀疏且短;葉舌高2~3 mm,上面具較發(fā)達(dá)的纖毛;葉片長30~50 cm,寬5~9 cm,兩面無毛,葉綠色,葉兩面同色,次脈10~15對(圖版I)。圓錐花序疏松開展,長8~25 cm,花序軸密被白色絨毛;小穗長2~6 cm,綠色或成熟后呈稻草色,含3~5朵小花;小穗軸節(jié)間長4~5.5 mm,呈扁棒狀,有縱棱,密被白色短絨毛;穎片2,披針形,第一穎長3~5 mm,第二穎長8~12 mm;外稃卵狀披針形,長10~12 mm,光滑無毛,具8(10)脈;內(nèi)稃與外稃近等長,上部被微毛,具2 脊;鱗被3,卵狀披針形,長2~2.5 mm,邊緣具纖毛;雄蕊3枚,花藥紫紅色,長8~9 mm,花絲白色,長8~12 mm;柱頭2,白色,羽毛狀,長5~7 mm (圖版Ⅱ,圖版Ⅲ)。穎果未見。筍期4—5 月,花期6—8月。
葉下表皮的氣孔器呈近圓形,周圍被8~10個長乳突且有規(guī)律地環(huán)繞;長乳突先端相接且平鋪覆蓋氣孔,基部偶具1~2個短乳突分枝;短乳突分布稀疏;未見大毛、刺毛分布;微毛伏貼,在脈間廣泛分布;硅質(zhì)體馬鞍形(圖版IV)。
A. 居群和生境; B. 地下莖; C. 筍; D. 節(jié)下; E. 籜片及籜鞘; F, G. 籜耳、籜舌及繸毛; H. 葉鞘、葉舌及葉基等; I. 節(jié); J. 分枝、髓心等。比例尺: 1 m (A); 1 cm (B-J)。
A. Population and habitat; B. Rhizome; C. Shoot; D. Infranodal region on culm; E. Culm blade and sheath; F, G. Sheath auricle, ligule and setae; H. Leaf sheath, ligule and leaf basal, etc.; I. Node; J. Mature culm and its longitudinal section, etc. Scale bars: 1 m (A); 1 cm (B-J).
A. 花序; B. 小穗和小花; C. 小花; D. 第一穎(左)和第二穎(右); E. 內(nèi)稃(左)和外稃(右); F. 雄蕊; G. 漿片(左)和 雌蕊(右)。比例尺: 1 cm (A, B); 5 mm (C-G)。
A. Inflorescence; B. Spikelets and florets; C. Floret; D. The first glume (left) and the second glume (right); E. Palea (left) and lemma (right); F. Anthers; G. Lodicules (left) and stigmas (right). Scale bars: 1 cm (A, B); 5 mm (C-G).
圖版 Ⅱ"半耳箬竹花形態(tài)特征
Plate Ⅱ"Floral morphological characteristics of Indocalamus semifalcatus
A. 花序; B. 小花; C. 內(nèi)稃(左)和外稃(右); D. 漿片; E. 雌蕊。繪畫:郭蓉 (引自標(biāo)本L. Q. Gao et al. SCGX2022, JXAU)。
A. Inflorescence; B. Floret; C. Palea (left) and lemma (right); D. Lodicules; E. Stigmas. Illustrated by GUO Rong (from specimen L. Q. Gao "et al. "SCGX2022, JXAU).
圖版 Ⅲ"半耳箬竹花形態(tài)墨線圖
Plate Ⅲ"Flower morphology ink line
diagram of Indocalamus semifalcatus
本種與箬葉竹最為相似,但主要區(qū)別在于本種的籜鞘和葉鞘上具有半鐮形的籜耳或籜耳缺失,稈高可達(dá)4~5 m,稈徑可達(dá)2 cm。
地理分布: 四川的都江堰二王廟、崇州三郎鎮(zhèn);貴州貴陽觀音山;廣西上林縣大明山。
箬竹屬竹類營養(yǎng)體的典型特征是灌木狀竹類,稈高通常2 m左右,稈徑常不超過1.0 cm,如箬竹、箬葉竹等種(耿伯介和王正平,1996;Wang amp; Stapleton,2006),但在半耳箬竹的居群中發(fā)現(xiàn)該種稈高可達(dá)4.5 m,稈徑1.8~2 cm,在箬竹屬中實屬少見,更新了該屬中竹種稈高、稈徑的傳統(tǒng)認(rèn)知。該種營養(yǎng)體形態(tài)特征與箬葉竹最主要區(qū)別在于該種的籜鞘和葉鞘上具有半鐮形的籜耳或籜耳缺失,稈高達(dá)4.5 m,稈徑達(dá)2 cm。
半耳箬竹花序呈圓錐狀,疏松開展,生于主枝或側(cè)枝頂上,花序軸和小穗軸密被白色短絨毛,小穗含小花3~5朵、穎片2、鱗被3、雄蕊3枚、柱頭2裂且呈羽毛狀,這與箬葉竹無顯著差異。此外,其葉下表皮氣孔器呈近圓形、周圍被8~10個長乳突平鋪覆蓋氣孔的特征與箬葉竹十分相似,這與Zhang等(2014)和冀雪楠(2019)的研究結(jié)果一致。
形態(tài)學(xué)研究是經(jīng)典植物分類學(xué)的重要基礎(chǔ)。本文基于半耳箬竹的原始描述, 通過測量形態(tài)性
A. 葉下表皮; B. 微毛和硅質(zhì)體; C. 長乳突和短乳突; D. 氣孔器。 ep. 長乳突; sp. 短乳突; mi. 微毛; sb. 硅質(zhì)體; st. 氣孔器。
A. Abaxial epidermis; B. Micro-hairs and silica bodies; C. Elongated papillae and short papillae; D. Stomatal apparatus. ep. Elongated papillae; sp. Short papillae; mi. Micro-hairs; sb. Silica bodies; st. Stomatal apparatus.
圖版 IV"半耳箬竹葉下表皮微形態(tài)特征
Plate IV"Leaf epidermis micromorphological characteristics of Indocalamus semifalcatus under SEM
狀、掃描葉下表皮微形態(tài)、考察并明確了一部分地理分布點,完善并補充了營養(yǎng)體、花部性狀數(shù)據(jù)和圖譜,新增了葉下表皮微形態(tài)性狀,更新了其地理分布,使半耳箬竹的研究資料更加準(zhǔn)確,為物種鑒定、箬竹屬屬內(nèi)分類修訂及系統(tǒng)演化提供重要參考依據(jù)。
致謝"承蒙四川農(nóng)業(yè)大學(xué)楊林高級實驗師指導(dǎo),江西農(nóng)業(yè)大學(xué)林學(xué)院研究生郭蓉繪畫,特此表示感謝。
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(責(zé)任編輯"鄧斯麗)
DOI: 10.11931/guihaia.gxzw202305074
高麗琴, "李永龍, 崔齡, 等, 2024.
半耳箬竹(竹亞科)的形態(tài)補充描述 [J]. 廣西植物, 44(4): 650-656.GAO LQ, LI YL, CUI L, et al., 2024.
Morphological supplementary description of Indocalamus semifalcatus (H. R. Zhao et Y. L. Yang) T. P. Yi (Bambusoideae) [J]. Guihaia, 44(4): 650-656.
高麗琴等: 半耳箬竹(竹亞科)的形態(tài)補充描述
收稿日期: "2023-07-15"接受日期: 2023-08-30
基金項目: "國家自然科學(xué)基金(31960335); 江西省重點研發(fā)計劃(20192BBF60015)。
第一作者: 高麗琴(1988—),博士,助理研究員,主要從事植物分類、資源開發(fā)與利用研究,(E-mail)gaoliqin2017@126.com。
通信作者: "張文根,博士,副教授,從事植物分類、資源與進(jìn)化發(fā)育等方面研究,(E-mail)wgzhang@jxau.edu.cn。