生物多样性 ›› 2008, Vol. 16 ›› Issue (3): 245-254. DOI: 10.3724/SP.J.1003.2008.07396
马文章1,2, 刘文耀1,3,*(), 杨礼攀1,2, 杨国平1
收稿日期:
2008-01-07
接受日期:
2008-04-01
出版日期:
2008-05-20
发布日期:
2008-05-20
通讯作者:
刘文耀
作者简介:
*E-mail:Liuwy@xtbg.ac.cn基金资助:
Wenzhang Ma1,2, Wenyao Liu1,3,*(), Lipan Yang1,2, Guoping Yang1
Received:
2008-01-07
Accepted:
2008-04-01
Online:
2008-05-20
Published:
2008-05-20
Contact:
Wenyao Liu
摘要:
作为热带、亚热带地区森林生态系统中重要组成成分之一的附生植物, 对环境变化尤为敏感。为了解附生植物对森林生境破碎化的响应, 本研究选择哀牢山北段徐家坝山地湿性常绿阔叶林因人为干扰而形成的四处森林边缘为研究对象, 分别在距离林缘10、20、40和80 m的位置设立样带, 采用树干取样法调查各样带内距地面0-2 m范围附生植物的物种组成、生物量和附生苔藓植物的生活型特征。结果显示: 边缘效应对单位面积附生植物的生物量和附生苔藓植物盖度均具有极显著的影响(P<0.001), 其中附生苔藓比附生蕨类对边缘效应更为敏感。边缘效应对附生苔藓植物的作用强度在各生活型之间存在着差异, 其中对丛集型(P=0.014)和交织型(P=0.030)的附生苔藓植物的影响最显著。综合附生植物在各样带的分布特征, 我们认为单位面积上附生植物的生物量和附生苔藓植物的盖度可作为指示森林边缘生境的重要指标。黄牛毛藓(Ditrichum pallidum)、角状刺枝藓(Wijkia hornschuchii)和格氏剪叶苔(Herbertus giraldianus)的分布具有指示林缘生境的意义, 而尖叶羽藓(Thuidium philibertii)、大羽藓(T. cymbifolium)、多疣麻羽藓(Claopodium pellucinerve)、波叶金枝藓(Pseudotrismegistia undulata)和扭叶藓(Trachypus bicolor)的分布局限于林内, 具有一定的保护价值。
马文章, 刘文耀, 杨礼攀, 杨国平 (2008) 边缘效应对山地湿性常绿阔叶林附生植物的影响. 生物多样性, 16, 245-254. DOI: 10.3724/SP.J.1003.2008.07396.
Wenzhang Ma, Wenyao Liu, Lipan Yang, Guoping Yang (2008) Edge effects on epiphytes in montane moist evergreen broad-leaved forest. Biodiversity Science, 16, 245-254. DOI: 10.3724/SP.J.1003.2008.07396.
生活型 Life form | 特征描述 Description |
---|---|
垫状型 Cushion | 主茎或多或少直立, 分枝从基部向上呈辐射状 Dome-shaped colonies formed by regeneration from a central point of origin, with stems more or less vertical |
树型 Dendroid | 主茎或横走或直立, 分枝呈树状展开 Stems creep or erect, branching resemble miniature tree |
扇形 Fan | 生于垂直的生长基质上, 分枝在一个平面上 Shoots rising from vertical substratum, branch repeatedly in the horizontal plane to form flattened photosynthetic surfaces |
平铺型 Mat | 主茎横生, 其上着生假根与生长基质紧密相连 Shoots creep over the substratum with rhizoids closely attached to |
悬垂型 Pendant | 分枝从树枝上向下悬垂生长 Main shoots hang down form the point of attachment |
丛集型 Turf | 主茎直立, 平行聚集生长, 分支少, 常占据大面积区域 Many loosely or closely packed vertical stem with limited branching, occupying a large area |
交织型 Weft | 主茎与分枝疏松交织 Stems and branches loosely interwoven |
表1 附生苔藓植物生活型的划分依据(参考张晋昆1; Bates, 1998)
Table 1 Classification of life forms in epiphytic bryophytes
生活型 Life form | 特征描述 Description |
---|---|
垫状型 Cushion | 主茎或多或少直立, 分枝从基部向上呈辐射状 Dome-shaped colonies formed by regeneration from a central point of origin, with stems more or less vertical |
树型 Dendroid | 主茎或横走或直立, 分枝呈树状展开 Stems creep or erect, branching resemble miniature tree |
扇形 Fan | 生于垂直的生长基质上, 分枝在一个平面上 Shoots rising from vertical substratum, branch repeatedly in the horizontal plane to form flattened photosynthetic surfaces |
平铺型 Mat | 主茎横生, 其上着生假根与生长基质紧密相连 Shoots creep over the substratum with rhizoids closely attached to |
悬垂型 Pendant | 分枝从树枝上向下悬垂生长 Main shoots hang down form the point of attachment |
丛集型 Turf | 主茎直立, 平行聚集生长, 分支少, 常占据大面积区域 Many loosely or closely packed vertical stem with limited branching, occupying a large area |
交织型 Weft | 主茎与分枝疏松交织 Stems and branches loosely interwoven |
距林缘距离 Distance to forest edge | 优势种类 Dominant species | 频度(%) Frequency | 宿主数 No. of hosts | 盖度 Coverage (% ± SE) | 重要值 IV | 生活型 Life form |
---|---|---|---|---|---|---|
10 m | 多疣悬藓 Neodicladiella pendula | 21.9 | 6 | 26.0 ± 0.7 | 41.2 | 悬垂型 Pendant |
粗垂藓 Chrysocladium phaeum | 15.6 | 5 | 13.7 ± 0.6 | 23.1 | 悬垂型 Pendant | |
小叶鞭苔 Bazzania ovistipula | 25.0 | 7 | 9.4 ± 0.2 | 20.8 | 平铺型 Mat | |
树平藓 Homaliodendron flabellatum | 28.1 | 6 | 2.3 ± 0.1 | 12.6 | 扇型 Fan | |
疏叶丝带藓 Floribundaria walkeri | 25.0 | 7 | 1.5 ± 0.1 | 10.5 | 悬垂型 Pendant | |
粗齿稚尾藓 Cyathophorum adiantum | 9.4 | 3 | 3.4 ± 0.1 | 7.6 | 扇型 Fan | |
刺齿合叶苔 Scapania ciliata | 6.3 | 2 | 3.1 ± 0.4 | 6.2 | 丛集型 Turf | |
绢藓一种 Entodonsp. | 6.3 | 4 | 3.0 ± 0.2 | 6.0 | 交织型 Weft | |
短肋羽藓 Thuidium kanedae | 12.5 | 4 | 0.2 | 4.5 | 交织型 Weft | |
赤茎小锦藓 Brotherella erythrocaulis | 9.4 | 4 | 0.8 | 4.2 | 交织型 Weft | |
20 m | 刺齿合叶苔 Scapania ciliata | 19.5 | 8 | 19.0 ± 0.3 | 18.2 | 丛集型 Turf |
南亚白发藓Leucobryum neilgherrense | 46.3 | 14 | 6.7 ± 0.1 | 16.7 | 垫状型 Cushion | |
树形羽苔 Plagiochila arbuscula | 24.4 | 7 | 13.8 ± 0.2 | 15.9 | 丛集型 Turf | |
卵叶羽苔 P. ovilifolia | 7.3 | 4 | 19.1 ± 0.9 | 15.2 | 扇型 Fan | |
小叶鞭苔 Bazznia ovistipula | 24.4 | 20 | 10.4 ± 0.1 | 13.5 | 平铺型 Mat | |
树平藓 Homaliodendron flabellatum | 34.2 | 10 | 3.5 | 11.3 | 扇型 Fan | |
青毛藓 Dicranodontum denudatum | 9.8 | 6 | 10.8 ± 0.3 | 10.0 | 丛集型 Turf | |
黄牛毛藓 Ditrichum pallidum | 19.5 | 9 | 6.9 ± 0.1 | 9.8 | 丛集型 Turf | |
扁萼苔 Radula complanata | 9.8 | 4 | 10.5 ± 0.4 | 9.8 | 平铺型 Mat | |
绢藓一种 Entodon sp. | 7.3 | 5 | 7.1 ± 0.3 | 6.8 | 交织型 Weft | |
40 m | 南亚白发藓 Leucobryum neilgherrense | 39.4 | 17 | 14.8 ± 0.3 | 18.6 | 垫状型 Cushion |
异形凤尾藓 Fissidens anomalus | 30.3 | 11 | 16.3 ± 0.4 | 17.8 | 扇型 Fan | |
树平藓 Homaliodendron flabellatum | 57.6 | 19 | 7.5 ± 0.1 | 17.4 | 扇型 Fan | |
小叶鞭苔 Bazzania ovistipula | 27.3 | 13 | 12.6 ± 0.3 | 14.6 | 平铺型 Mat | |
细指叶苔 Lepidozia trichodes | 6.1 | 9 | 11.6 ± 0.9 | 9.3 | 交织型 Weft | |
侧枝匍灯藓 Plagiomnium maximoviczii | 21.2 | 8 | 6.7 ± 0.3 | 9.2 | 平铺型 Mat | |
树形羽苔 Plagiochila arbuscula | 21.2 | 13 | 5.5 ± 0.1 | 8.3 | 丛集型 Turf | |
赤茎小锦藓 Brotherella erythrocaulis | 27.3 | 11 | 1.8 | 7.0 | 平铺型 Mat | |
平藓 Neckera pennata | 24.2 | 9 | 2.4 | 6.8 | 树型 Dendroid | |
双齿鞭苔 Bazzania bidentula | 3.3 | 6 | 7.5 | 6.0 | 平铺型 Mat | |
80 m | 小叶鞭苔 B. ovistipula | 41.5 | 15 | 41.0 ± 0.4 | 30.0 | 平铺型 Mat |
南亚白发藓 Leucobryum neilgherrense | 43.9 | 16 | 24.9 ± 0.3 | 22.1 | 垫状型 Cushion | |
树平藓 Homaliodendron flabellatum | 68.3 | 22 | 10.2 ± 0.1 | 19.6 | 扇型 Fan | |
卵叶羽苔 Plagiochila ovilifolia | 14.6 | 10 | 27.2 ± 0.5 | 17.2 | 扇型 Fan | |
赤茎小锦藓 Brothrella erythrocaulis | 31.7 | 10 | 9.6 ± 0.2 | 11.6 | 平铺型 Mat | |
狭叶羽苔 Plagiochila trabeculata | 9.8 | 4 | 12.6 ± 0.2 | 8.6 | 扇型 Fan | |
树形羽苔 P. arbuscula | 22.0 | 12 | 7.1 ± 0.2 | 8.3 | 丛集型 Turf | |
细指叶苔 Leptozia trichodes | 14.6 | 10 | 9.7 ± 0.1 | 8.1 | 交织型 Weft | |
疏叶丝带藓 Floribundaria walkeri | 22.0 | 10 | 4.0 ± 0.1 | 6.7 | 悬垂型 Pendant | |
短肋羽藓 Thudium kanedae | 24.4 | 9 | 1.7 | 6.0 | 交织型 Weft |
表2 所有林缘样带内附生苔藓植物重要值居前10位的优势物种
距林缘距离 Distance to forest edge | 优势种类 Dominant species | 频度(%) Frequency | 宿主数 No. of hosts | 盖度 Coverage (% ± SE) | 重要值 IV | 生活型 Life form |
---|---|---|---|---|---|---|
10 m | 多疣悬藓 Neodicladiella pendula | 21.9 | 6 | 26.0 ± 0.7 | 41.2 | 悬垂型 Pendant |
粗垂藓 Chrysocladium phaeum | 15.6 | 5 | 13.7 ± 0.6 | 23.1 | 悬垂型 Pendant | |
小叶鞭苔 Bazzania ovistipula | 25.0 | 7 | 9.4 ± 0.2 | 20.8 | 平铺型 Mat | |
树平藓 Homaliodendron flabellatum | 28.1 | 6 | 2.3 ± 0.1 | 12.6 | 扇型 Fan | |
疏叶丝带藓 Floribundaria walkeri | 25.0 | 7 | 1.5 ± 0.1 | 10.5 | 悬垂型 Pendant | |
粗齿稚尾藓 Cyathophorum adiantum | 9.4 | 3 | 3.4 ± 0.1 | 7.6 | 扇型 Fan | |
刺齿合叶苔 Scapania ciliata | 6.3 | 2 | 3.1 ± 0.4 | 6.2 | 丛集型 Turf | |
绢藓一种 Entodonsp. | 6.3 | 4 | 3.0 ± 0.2 | 6.0 | 交织型 Weft | |
短肋羽藓 Thuidium kanedae | 12.5 | 4 | 0.2 | 4.5 | 交织型 Weft | |
赤茎小锦藓 Brotherella erythrocaulis | 9.4 | 4 | 0.8 | 4.2 | 交织型 Weft | |
20 m | 刺齿合叶苔 Scapania ciliata | 19.5 | 8 | 19.0 ± 0.3 | 18.2 | 丛集型 Turf |
南亚白发藓Leucobryum neilgherrense | 46.3 | 14 | 6.7 ± 0.1 | 16.7 | 垫状型 Cushion | |
树形羽苔 Plagiochila arbuscula | 24.4 | 7 | 13.8 ± 0.2 | 15.9 | 丛集型 Turf | |
卵叶羽苔 P. ovilifolia | 7.3 | 4 | 19.1 ± 0.9 | 15.2 | 扇型 Fan | |
小叶鞭苔 Bazznia ovistipula | 24.4 | 20 | 10.4 ± 0.1 | 13.5 | 平铺型 Mat | |
树平藓 Homaliodendron flabellatum | 34.2 | 10 | 3.5 | 11.3 | 扇型 Fan | |
青毛藓 Dicranodontum denudatum | 9.8 | 6 | 10.8 ± 0.3 | 10.0 | 丛集型 Turf | |
黄牛毛藓 Ditrichum pallidum | 19.5 | 9 | 6.9 ± 0.1 | 9.8 | 丛集型 Turf | |
扁萼苔 Radula complanata | 9.8 | 4 | 10.5 ± 0.4 | 9.8 | 平铺型 Mat | |
绢藓一种 Entodon sp. | 7.3 | 5 | 7.1 ± 0.3 | 6.8 | 交织型 Weft | |
40 m | 南亚白发藓 Leucobryum neilgherrense | 39.4 | 17 | 14.8 ± 0.3 | 18.6 | 垫状型 Cushion |
异形凤尾藓 Fissidens anomalus | 30.3 | 11 | 16.3 ± 0.4 | 17.8 | 扇型 Fan | |
树平藓 Homaliodendron flabellatum | 57.6 | 19 | 7.5 ± 0.1 | 17.4 | 扇型 Fan | |
小叶鞭苔 Bazzania ovistipula | 27.3 | 13 | 12.6 ± 0.3 | 14.6 | 平铺型 Mat | |
细指叶苔 Lepidozia trichodes | 6.1 | 9 | 11.6 ± 0.9 | 9.3 | 交织型 Weft | |
侧枝匍灯藓 Plagiomnium maximoviczii | 21.2 | 8 | 6.7 ± 0.3 | 9.2 | 平铺型 Mat | |
树形羽苔 Plagiochila arbuscula | 21.2 | 13 | 5.5 ± 0.1 | 8.3 | 丛集型 Turf | |
赤茎小锦藓 Brotherella erythrocaulis | 27.3 | 11 | 1.8 | 7.0 | 平铺型 Mat | |
平藓 Neckera pennata | 24.2 | 9 | 2.4 | 6.8 | 树型 Dendroid | |
双齿鞭苔 Bazzania bidentula | 3.3 | 6 | 7.5 | 6.0 | 平铺型 Mat | |
80 m | 小叶鞭苔 B. ovistipula | 41.5 | 15 | 41.0 ± 0.4 | 30.0 | 平铺型 Mat |
南亚白发藓 Leucobryum neilgherrense | 43.9 | 16 | 24.9 ± 0.3 | 22.1 | 垫状型 Cushion | |
树平藓 Homaliodendron flabellatum | 68.3 | 22 | 10.2 ± 0.1 | 19.6 | 扇型 Fan | |
卵叶羽苔 Plagiochila ovilifolia | 14.6 | 10 | 27.2 ± 0.5 | 17.2 | 扇型 Fan | |
赤茎小锦藓 Brothrella erythrocaulis | 31.7 | 10 | 9.6 ± 0.2 | 11.6 | 平铺型 Mat | |
狭叶羽苔 Plagiochila trabeculata | 9.8 | 4 | 12.6 ± 0.2 | 8.6 | 扇型 Fan | |
树形羽苔 P. arbuscula | 22.0 | 12 | 7.1 ± 0.2 | 8.3 | 丛集型 Turf | |
细指叶苔 Leptozia trichodes | 14.6 | 10 | 9.7 ± 0.1 | 8.1 | 交织型 Weft | |
疏叶丝带藓 Floribundaria walkeri | 22.0 | 10 | 4.0 ± 0.1 | 6.7 | 悬垂型 Pendant | |
短肋羽藓 Thudium kanedae | 24.4 | 9 | 1.7 | 6.0 | 交织型 Weft |
距林缘距离 Distance to forest edge | 种类 Species | 总株数 Total no. | 平均簇数 (簇/宿主) Average no. of individual | 出现频率 Frequency (%) | 重要值 IV | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|
10 m | 小果露蕨 Mecodium microsorum | 111 | 27.8 | 6.3 | 8.0 | ||||||
棕鳞瓦韦 Lepisorus scolopendrium | 4 | 2.0 | 6.3 | 6.6 | |||||||
20 m | 小果露蕨 Mecodium microsorum | 279 | 69.8 | 14.6 | 18.0 | ||||||
棕鳞瓦韦 Lepisorus scolopendrium | 6 | 3.0 | 7.3 | 7.7 | |||||||
高山条蕨 Oleandra wallichii | 3 | 3.0 | 3.0 | 3.1 | |||||||
黑鳞假瘤蕨 Phymatopsis ebenipes | 1 | 1.0 | 2.4 | 2.5 | |||||||
书带蕨 Vittaria flexuosa | 2 | 0.7 | 2.4 | 2.5 | |||||||
40 m | 小果露蕨 Mecodium microsorum | 159 | 39.8 | 15.2 | 17.0 | ||||||
友水龙骨 Polypodiodes amoena | 3 | 0.8 | 9.1 | 9.3 | |||||||
雨蕨 Gymnogrammitis dardiformis | 1 | 1.0 | 9.1 | 9.2 | |||||||
书带蕨 Vittaria flexuosa | 5 | 2.5 | 6.1 | 6.4 | |||||||
胎生铁角蕨 Asplenium viviparum | 4 | 1.3 | 6.1 | 6.2 | |||||||
褐柄剑蕨 Loxogramme duclouxii | 1 | 1.0 | 6.1 | 6.2 | |||||||
棕鳞瓦韦 Lepisorus scolopendrium | 3 | 1.0 | 3.0 | 3.2 | |||||||
书带蕨 Vittaria flexuosa | 2 | 1.0 | 3.0 | 3.1 | |||||||
80 m | 小果露蕨 Mecodium microsorum | 54 | 13.5 | 12.2 | 13.0 | ||||||
友水龙骨 Polypodiodes amoena | 4 | 1.3 | 9.8 | 10.0 | |||||||
雨蕨 Gymnogrammitis dardiformis | 2 | 2.0 | 9.8 | 10.0 | |||||||
褐柄剑蕨 Loxogramme duclouxii | 1 | 0.5 | 2.4 | 2.5 | |||||||
栗柄水龙骨 Polypodiodes microrhizoma | 1 | 1.0 | 2.4 | 2.5 | |||||||
书带蕨 Vittaria flexuosa | 2 | 2.0 | 2.4 | 2.5 | |||||||
胎生铁角蕨 Asplenium viviparum | 1 | 1.0 | 2.4 | 2.5 |
Table 2 所有林缘工作样带内的附生蕨类植物
Table 3 The epiphytic ferns across all edge transects
距林缘距离 Distance to forest edge | 种类 Species | 总株数 Total no. | 平均簇数 (簇/宿主) Average no. of individual | 出现频率 Frequency (%) | 重要值 IV | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|
10 m | 小果露蕨 Mecodium microsorum | 111 | 27.8 | 6.3 | 8.0 | ||||||
棕鳞瓦韦 Lepisorus scolopendrium | 4 | 2.0 | 6.3 | 6.6 | |||||||
20 m | 小果露蕨 Mecodium microsorum | 279 | 69.8 | 14.6 | 18.0 | ||||||
棕鳞瓦韦 Lepisorus scolopendrium | 6 | 3.0 | 7.3 | 7.7 | |||||||
高山条蕨 Oleandra wallichii | 3 | 3.0 | 3.0 | 3.1 | |||||||
黑鳞假瘤蕨 Phymatopsis ebenipes | 1 | 1.0 | 2.4 | 2.5 | |||||||
书带蕨 Vittaria flexuosa | 2 | 0.7 | 2.4 | 2.5 | |||||||
40 m | 小果露蕨 Mecodium microsorum | 159 | 39.8 | 15.2 | 17.0 | ||||||
友水龙骨 Polypodiodes amoena | 3 | 0.8 | 9.1 | 9.3 | |||||||
雨蕨 Gymnogrammitis dardiformis | 1 | 1.0 | 9.1 | 9.2 | |||||||
书带蕨 Vittaria flexuosa | 5 | 2.5 | 6.1 | 6.4 | |||||||
胎生铁角蕨 Asplenium viviparum | 4 | 1.3 | 6.1 | 6.2 | |||||||
褐柄剑蕨 Loxogramme duclouxii | 1 | 1.0 | 6.1 | 6.2 | |||||||
棕鳞瓦韦 Lepisorus scolopendrium | 3 | 1.0 | 3.0 | 3.2 | |||||||
书带蕨 Vittaria flexuosa | 2 | 1.0 | 3.0 | 3.1 | |||||||
80 m | 小果露蕨 Mecodium microsorum | 54 | 13.5 | 12.2 | 13.0 | ||||||
友水龙骨 Polypodiodes amoena | 4 | 1.3 | 9.8 | 10.0 | |||||||
雨蕨 Gymnogrammitis dardiformis | 2 | 2.0 | 9.8 | 10.0 | |||||||
褐柄剑蕨 Loxogramme duclouxii | 1 | 0.5 | 2.4 | 2.5 | |||||||
栗柄水龙骨 Polypodiodes microrhizoma | 1 | 1.0 | 2.4 | 2.5 | |||||||
书带蕨 Vittaria flexuosa | 2 | 2.0 | 2.4 | 2.5 | |||||||
胎生铁角蕨 Asplenium viviparum | 1 | 1.0 | 2.4 | 2.5 |
两两样带之间 Between two transects | 10-20 m | 10-40 m | 10-80 m | 20-40 m | 20-80 m | 40-80 m |
---|---|---|---|---|---|---|
S?rensen 相似性系数 S?rensen’s similarity coefficient | 0.641 | 0.701 | 0.506 | 0.741 | 0.627 | 0.683 |
共有种数目 Number of common species | 25 | 27 | 20 | 30 | 26 | 28 |
表4 林缘各样带间附生植物共有种与相似性系数
Table 4 The number of common species and S?rensen’s similarity coefficient between transects
两两样带之间 Between two transects | 10-20 m | 10-40 m | 10-80 m | 20-40 m | 20-80 m | 40-80 m |
---|---|---|---|---|---|---|
S?rensen 相似性系数 S?rensen’s similarity coefficient | 0.641 | 0.701 | 0.506 | 0.741 | 0.627 | 0.683 |
共有种数目 Number of common species | 25 | 27 | 20 | 30 | 26 | 28 |
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