生物多样性 ›› 2008, Vol. 16 ›› Issue (2): 110-117. DOI: 10.3724/SP.J.1003.2008.07349
收稿日期:
2007-11-14
接受日期:
2008-01-21
出版日期:
2008-03-20
发布日期:
2008-02-20
通讯作者:
包维楷
作者简介:
*E-mail:baowk@cib.ac.cn基金资助:
Received:
2007-11-14
Accepted:
2008-01-21
Online:
2008-03-20
Published:
2008-02-20
Contact:
Weikai Bao
摘要:
为评估皆伐迹地地表苔藓组成与特点, 同时为苔藓生物多样性的保护与恢复提供理论依据, 作者于2006年7-10月调查了四川省阿坝藏族羌族自治州地区壤塘二林场云杉(Picea likiangensisvar. rubescens)林皆伐迹地自然恢复过程中苔藓植物和维管植物的组成和结构; 采用方差分析法对苔藓层结构参数进行差异性检验, 对苔藓层结构参数和环境因子作非参数Spearman相关性分析, 并对显著性因子进行回归分析。结果表明: (1)在9-27年植被自然恢复过程中, 不同阶段苔藓植物群落优势物种组成发生了更替。一些个体较小、易产生孢子、喜阳、抗干扰的顶蒴藓物种侵入并占据优势; 而一些出现于早期采伐迹地、喜湿、对环境变化较敏感的侧蒴藓物种消失。(2) 随着迹地植被自然恢复, 苔藓植物群落物种丰富度、多样性指数和均匀度指数总体上呈下降趋势; 迹地上样方间苔藓层盖度和生物量波动较大, 但随着自然恢复进程而迅速减少, 群落结构趋向简单化; (3)维管植物发育程度与苔藓层发育关系密切; 草本层生物量对苔藓盖度和种丰富度起主导作用, 草本层高度是对苔藓生物量有显著影响的主导因子。综合分析表明, 早期采伐迹地的自然恢复进程中由维管植物群落结构变化导致的生境差异对苔藓植物发育影响较大。
闫晓丽, 包维楷 (2008) 高海拔区云杉林皆伐迹地早期自然恢复进程中苔藓组成与发育程度评估. 生物多样性, 16, 110-117. DOI: 10.3724/SP.J.1003.2008.07349.
Xiaoli Yan, Weikai Bao (2008) Evaluation of species composition and development of bryophyte community during early natural recovery progress of high-altitude spruce cutovers. Biodiversity Science, 16, 110-117. DOI: 10.3724/SP.J.1003.2008.07349.
样地 Site | 森林采伐时间 Felling time | 海拔 Elevation (m) | 坡向 Aspect | 坡度 Slope degree | 土壤类型 Soil type |
---|---|---|---|---|---|
十五支沟 Shiwuzhigou (CT98) | 1998 | 3,554 | N-E15° | 30° | 山地棕壤 Mountain brown soil |
十二支沟 Shierzhigou (CT96) | 1996 | 3,654 | N-W15° | 50° | |
十支沟 Shizhigou (CT85) | 1985 | 3,587 | N-E30° | 45° | |
九支沟 Jiuzhigou (CT80) | 1980 | 3,593 | N-E30° | 50° |
表1 4个采伐迹地样地的基本信息
Table 1 Basic information about the four clear-cutting cutovers studied in Rangtang County, Aaba region, Sichuan Province
样地 Site | 森林采伐时间 Felling time | 海拔 Elevation (m) | 坡向 Aspect | 坡度 Slope degree | 土壤类型 Soil type |
---|---|---|---|---|---|
十五支沟 Shiwuzhigou (CT98) | 1998 | 3,554 | N-E15° | 30° | 山地棕壤 Mountain brown soil |
十二支沟 Shierzhigou (CT96) | 1996 | 3,654 | N-W15° | 50° | |
十支沟 Shizhigou (CT85) | 1985 | 3,587 | N-E30° | 45° | |
九支沟 Jiuzhigou (CT80) | 1980 | 3,593 | N-E30° | 50° |
采伐时间 Felling time | ||||
---|---|---|---|---|
调查项目 | 1998 | 1996 | 1985 | 1980 |
恢复时间 Restoration age (year) | 9 | 11 | 22 | 27 |
灌木盖度 Shrub coverage (%) | 4.17±1.57a | 5.9±1.48a | 4.1±1.78a | 12.5±4.79b |
草本盖度 Herb coverage (%) | 50.63±4.84a | 47.28±4.68a | 61.77±4.96b | 69±3.68b |
草本高度 Herb height (cm) | 18.58±1.36a | 23.17±1.7b | 35.03±2.36c | 39.9±2.11d |
草本生物量 Herb biomass (g/m2) | 94.85±9.86a | 100.78±11.69a | 167.53±16.99b | 216.59±16c |
草本丰富度 Herb richness (sp/plot) | 5.1±0.53a | 4.83±0.27a | 7.47±0.37b | 9.34±0.37c |
凋落物生物量 Litter biomass(g/m2) | 87.23±27.78ab | 68.18±16.38a | 95.24±15.97ab | 98.5±13.2b |
表2 迹地植被自然恢复进程中维管植物群落结构特征(平均值±标准误, n = 30)
Table 2 Characteristics of vascular plant community structure during natural recovery succession (mean ± SE, n = 30)
采伐时间 Felling time | ||||
---|---|---|---|---|
调查项目 | 1998 | 1996 | 1985 | 1980 |
恢复时间 Restoration age (year) | 9 | 11 | 22 | 27 |
灌木盖度 Shrub coverage (%) | 4.17±1.57a | 5.9±1.48a | 4.1±1.78a | 12.5±4.79b |
草本盖度 Herb coverage (%) | 50.63±4.84a | 47.28±4.68a | 61.77±4.96b | 69±3.68b |
草本高度 Herb height (cm) | 18.58±1.36a | 23.17±1.7b | 35.03±2.36c | 39.9±2.11d |
草本生物量 Herb biomass (g/m2) | 94.85±9.86a | 100.78±11.69a | 167.53±16.99b | 216.59±16c |
草本丰富度 Herb richness (sp/plot) | 5.1±0.53a | 4.83±0.27a | 7.47±0.37b | 9.34±0.37c |
凋落物生物量 Litter biomass(g/m2) | 87.23±27.78ab | 68.18±16.38a | 95.24±15.97ab | 98.5±13.2b |
物种组成 Species composition | 采伐时间 Felling time | |||
---|---|---|---|---|
1998 | 1996 | 1985 | 1980 | |
苔类 Liverworts | ||||
蛇苔 Conocephalum conicum (L.) Dum. | - | 0 | 0.06 | 0.05 |
剪叶苔 Herbertus aduncus (Dicks.) Gray | 0 | - | - | - |
叶苔一种 Jungermannia sp. | - | 0 | - | - |
羽苔一种 Plagiochila sp. | 0.05 | 0.02 | 0.02 | 0.02 |
藓类 Moss | ||||
顶蒴藓类 Acrocarpous moss | ||||
银藓一种 Anomobryum sp. | - | 0 | 0.02 | 0.01 |
溪边扭口藓 Barbula rivicola Broth. | 0.15 | - | - | - |
狭叶扭口藓 B. subcontorta Broth. | - | 0.03 | 0.01 | - |
珠藓 Bartramia halleriana Hedw. | 0.01 | - | - | - |
短月藓 Brachymeniumsp. | 0.01 | - | - | - |
高山红叶藓 Bryoerythrophyllum alpigenum (Vent.) Chen | - | 0 | - | - |
红叶藓 B. recurvirostrum (Hedw.) Chen | 0 | - | - | - |
比拉真藓 Bryum billardieri Schwaegr. | 0 | - | - | - |
银叶真藓 B. argenteum Hedw. | 0.03 | 0.05 | 0.01 | - |
真藓一种 Bryum sp. 1 | 0.02 | 0.03 | - | - |
真藓一种 Bryum sp. 2 | - | - | - | 0.01 |
真藓一种 Bryum sp. 3 | - | - | - | 0.01 |
青毛藓 Dicranodontium denudatum(Brid.) Britt. | - | 0 | - | - |
马氏曲尾藓 Dicranum mayrii Broth. | 0.01 | 0 | - | - |
日本葫芦藓 Funaria japonica Broth. | - | 0.01 | - | - |
湿地藓 Hyophila javanica (Nees et Blum.) Brid. | - | - | 0.01 | - |
薄齿藓 Leptodontium viticulosoides (P. Beauv.) Wijk et Marg. | - | 0 | - | - |
具缘提灯藓 Mnium marginatum (With.) P. Beauv. | 0 | 0.02 | 0.18 | - |
刺叶提灯藓 M. spinosum (Voit.) Schwaegr. | 0 | 0.02 | - | 0.02 |
曲背藓 Oncophorus wahlenbergii Brid. | 0.01 | - | 0.03 | 0.01 |
小酸土藓 Oxystegus cuspidatus (Doz.et Molk.) Chen | 0.03 | 0.02 | - | - |
酸土藓 O. cylindricus (Brid.) Hilp. | 0.01 | 0 | 0.02 | 0.06 |
拟白发藓 Paraleucobryum enerve (Ther.) Loesk. | 0.09 | 0.18 | 0.1 | 0.31 |
皱叶匍灯藓 Plagiomnium confertidens (Lindb.& Arn.) Kop. | 0.08 | 0.09 | 0.04 | 0.31 |
多蒴匍灯藓 P. medium (B.S.G.) Kop. | - | 0.01 | - | - |
侧立藓 Pleuroweisia schliephackei Limpr. | - | 0 | - | - |
长蒴丝瓜藓 Pohlia elongata Hedw. | 0.01 | 0.05 | 0.46 | 0.24 |
无齿藓 Pseudochorisatontium gymnostomum (Mitt.) C. Gao et al. | 0.01 | 0.01 | - | - |
韩氏无齿藓 P. hokinense (Besch.) C. Gao et al. | 0.08 | 0.01 | - | - |
四川无齿藓 P. setschwanicum (Broth.) C. | 0.03 | 0.01 | - | - |
黄砂藓 Racomitrium anomodontoides Card. | 0.01 | - | - | - |
阔边大叶藓 Rhodobryum laxe-limbatum (Ochi) Iwats.et T.Kop. | - | 0.01 | - | - |
长齿连轴藓 Schistidium trichodon (Brid.) Poelt | - | 0.01 | - | 0.01 |
泛生墙藓 Tortula muralis Hedw. | - | - | - | 0.03 |
树形疣灯藓 Trachycysis ussuriensis (Maack et Regel) T.Kop. | 0.10 | 0.04 | - | - |
皱叶毛口藓 Trichostomum crispulumBruch | - | 0.01 | - | - |
侧蒴藓类 Pleurocarpous moss | ||||
锦丝藓 Actinothuidium hookeri (Mitt.) Broth. | 0.23 | 0.22 | 0.09 | 0.05 |
勃氏青藓 Brachythecium brotheriPar. | 0.07 | 0.27 | 0.45 | 0.68 |
物种组成 Species composition | 采伐时间 Felling time | |||
1998 | 1996 | 1985 | 1980 | |
匙叶毛尖藓 Cirriphyllum cirrosum (Schwaegr.) Grout | - | 0.03 | - | 0.15 |
厚角绢藓 Entodon concinnus (De Not.) Par. | 0.42 | 0.38 | 0.58 | 0.14 |
穗枝赤齿藓 Erythrodontium julaceum (Schwaegr.) Par. | - | 0 | - | - |
密枝粗枝藓 Gollania turgens (C. Muell.) Ando | 0.06 | 0.09 | - | - |
狭叶小羽藓 Haplocladium angustifolium (Hampe et C. Muell.) Broth. | - | - | - | 0.01 |
塔藓 Hylocomium splendens (Hedw.) B.S.G. | 0.28 | 0.13 | - | - |
弯叶灰藓 Hypnum hamulosum B. S. G. | 0.04 | 0.02 | 0.05 | 0.02 |
细叉羽藓 Leptopterigynandrum tenellum Broth. | - | - | - | 0.02 |
波叶棉藓 Plagiothecium undulatum (Hedw.) B.S.G. | 0 | - | - | - |
高山拟金发藓 Polytrichastrum alpinum (Hedw.) G.Smith | 0.12 | 0.02 | 0.02 | - |
台湾拟金发藓 P. formosum (Hedw.) G.Smith | 0.01 | 0.1 | - | - |
霉疣拟金发藓 P. papillatumG.Sm. | 0.01 | - | - | - |
弯叶金灰藓 Pylaisiella falcata (B.S.G.) Ando | 0 | - | - | - |
疣拟垂枝藓 Rhytidiadelphus triquetrus (Hedw.) Warnst. | 0.01 | 0.05 | 0.01 | 0.02 |
垂枝藓 Rhytidium rugosum (Hedw.) Kindb. | 0.29 | 0.01 | - | - |
三洋藓 Sanionia uncinata (Hedw.) Loeske | 0.09 | 0.03 | - | - |
大羽藓 Thuidium cymbifolium (Dozy et Molk.) Dozy et Molk. | 0.60 | 1.02 | 0.85 | 0.83 |
表3 迹地植被自然恢复进程中地表苔藓物种组成及其重要值
Table 3 Species composition and their important values of ground bryophyte during natural recovery succession
物种组成 Species composition | 采伐时间 Felling time | |||
---|---|---|---|---|
1998 | 1996 | 1985 | 1980 | |
苔类 Liverworts | ||||
蛇苔 Conocephalum conicum (L.) Dum. | - | 0 | 0.06 | 0.05 |
剪叶苔 Herbertus aduncus (Dicks.) Gray | 0 | - | - | - |
叶苔一种 Jungermannia sp. | - | 0 | - | - |
羽苔一种 Plagiochila sp. | 0.05 | 0.02 | 0.02 | 0.02 |
藓类 Moss | ||||
顶蒴藓类 Acrocarpous moss | ||||
银藓一种 Anomobryum sp. | - | 0 | 0.02 | 0.01 |
溪边扭口藓 Barbula rivicola Broth. | 0.15 | - | - | - |
狭叶扭口藓 B. subcontorta Broth. | - | 0.03 | 0.01 | - |
珠藓 Bartramia halleriana Hedw. | 0.01 | - | - | - |
短月藓 Brachymeniumsp. | 0.01 | - | - | - |
高山红叶藓 Bryoerythrophyllum alpigenum (Vent.) Chen | - | 0 | - | - |
红叶藓 B. recurvirostrum (Hedw.) Chen | 0 | - | - | - |
比拉真藓 Bryum billardieri Schwaegr. | 0 | - | - | - |
银叶真藓 B. argenteum Hedw. | 0.03 | 0.05 | 0.01 | - |
真藓一种 Bryum sp. 1 | 0.02 | 0.03 | - | - |
真藓一种 Bryum sp. 2 | - | - | - | 0.01 |
真藓一种 Bryum sp. 3 | - | - | - | 0.01 |
青毛藓 Dicranodontium denudatum(Brid.) Britt. | - | 0 | - | - |
马氏曲尾藓 Dicranum mayrii Broth. | 0.01 | 0 | - | - |
日本葫芦藓 Funaria japonica Broth. | - | 0.01 | - | - |
湿地藓 Hyophila javanica (Nees et Blum.) Brid. | - | - | 0.01 | - |
薄齿藓 Leptodontium viticulosoides (P. Beauv.) Wijk et Marg. | - | 0 | - | - |
具缘提灯藓 Mnium marginatum (With.) P. Beauv. | 0 | 0.02 | 0.18 | - |
刺叶提灯藓 M. spinosum (Voit.) Schwaegr. | 0 | 0.02 | - | 0.02 |
曲背藓 Oncophorus wahlenbergii Brid. | 0.01 | - | 0.03 | 0.01 |
小酸土藓 Oxystegus cuspidatus (Doz.et Molk.) Chen | 0.03 | 0.02 | - | - |
酸土藓 O. cylindricus (Brid.) Hilp. | 0.01 | 0 | 0.02 | 0.06 |
拟白发藓 Paraleucobryum enerve (Ther.) Loesk. | 0.09 | 0.18 | 0.1 | 0.31 |
皱叶匍灯藓 Plagiomnium confertidens (Lindb.& Arn.) Kop. | 0.08 | 0.09 | 0.04 | 0.31 |
多蒴匍灯藓 P. medium (B.S.G.) Kop. | - | 0.01 | - | - |
侧立藓 Pleuroweisia schliephackei Limpr. | - | 0 | - | - |
长蒴丝瓜藓 Pohlia elongata Hedw. | 0.01 | 0.05 | 0.46 | 0.24 |
无齿藓 Pseudochorisatontium gymnostomum (Mitt.) C. Gao et al. | 0.01 | 0.01 | - | - |
韩氏无齿藓 P. hokinense (Besch.) C. Gao et al. | 0.08 | 0.01 | - | - |
四川无齿藓 P. setschwanicum (Broth.) C. | 0.03 | 0.01 | - | - |
黄砂藓 Racomitrium anomodontoides Card. | 0.01 | - | - | - |
阔边大叶藓 Rhodobryum laxe-limbatum (Ochi) Iwats.et T.Kop. | - | 0.01 | - | - |
长齿连轴藓 Schistidium trichodon (Brid.) Poelt | - | 0.01 | - | 0.01 |
泛生墙藓 Tortula muralis Hedw. | - | - | - | 0.03 |
树形疣灯藓 Trachycysis ussuriensis (Maack et Regel) T.Kop. | 0.10 | 0.04 | - | - |
皱叶毛口藓 Trichostomum crispulumBruch | - | 0.01 | - | - |
侧蒴藓类 Pleurocarpous moss | ||||
锦丝藓 Actinothuidium hookeri (Mitt.) Broth. | 0.23 | 0.22 | 0.09 | 0.05 |
勃氏青藓 Brachythecium brotheriPar. | 0.07 | 0.27 | 0.45 | 0.68 |
物种组成 Species composition | 采伐时间 Felling time | |||
1998 | 1996 | 1985 | 1980 | |
匙叶毛尖藓 Cirriphyllum cirrosum (Schwaegr.) Grout | - | 0.03 | - | 0.15 |
厚角绢藓 Entodon concinnus (De Not.) Par. | 0.42 | 0.38 | 0.58 | 0.14 |
穗枝赤齿藓 Erythrodontium julaceum (Schwaegr.) Par. | - | 0 | - | - |
密枝粗枝藓 Gollania turgens (C. Muell.) Ando | 0.06 | 0.09 | - | - |
狭叶小羽藓 Haplocladium angustifolium (Hampe et C. Muell.) Broth. | - | - | - | 0.01 |
塔藓 Hylocomium splendens (Hedw.) B.S.G. | 0.28 | 0.13 | - | - |
弯叶灰藓 Hypnum hamulosum B. S. G. | 0.04 | 0.02 | 0.05 | 0.02 |
细叉羽藓 Leptopterigynandrum tenellum Broth. | - | - | - | 0.02 |
波叶棉藓 Plagiothecium undulatum (Hedw.) B.S.G. | 0 | - | - | - |
高山拟金发藓 Polytrichastrum alpinum (Hedw.) G.Smith | 0.12 | 0.02 | 0.02 | - |
台湾拟金发藓 P. formosum (Hedw.) G.Smith | 0.01 | 0.1 | - | - |
霉疣拟金发藓 P. papillatumG.Sm. | 0.01 | - | - | - |
弯叶金灰藓 Pylaisiella falcata (B.S.G.) Ando | 0 | - | - | - |
疣拟垂枝藓 Rhytidiadelphus triquetrus (Hedw.) Warnst. | 0.01 | 0.05 | 0.01 | 0.02 |
垂枝藓 Rhytidium rugosum (Hedw.) Kindb. | 0.29 | 0.01 | - | - |
三洋藓 Sanionia uncinata (Hedw.) Loeske | 0.09 | 0.03 | - | - |
大羽藓 Thuidium cymbifolium (Dozy et Molk.) Dozy et Molk. | 0.60 | 1.02 | 0.85 | 0.83 |
采伐时间 Felling time | ||||
---|---|---|---|---|
1998 | 1996 | 1985 | 1980 | |
丰富度 Richness | 38 | 42 | 19 | 22 |
Shannon-Wiener指数 Shannon-Wiener index | 2.37 | 1.76 | 1.79 | 1.71 |
均匀度指数 Evenness index | 0.65 | 0.47 | 0.61 | 0.55 |
表4 迹地植被自然恢复进程中苔藓植物群落多样性指数
Table 4 Diversity indices of bryophyte community during natural recovery succession
采伐时间 Felling time | ||||
---|---|---|---|---|
1998 | 1996 | 1985 | 1980 | |
丰富度 Richness | 38 | 42 | 19 | 22 |
Shannon-Wiener指数 Shannon-Wiener index | 2.37 | 1.76 | 1.79 | 1.71 |
均匀度指数 Evenness index | 0.65 | 0.47 | 0.61 | 0.55 |
采伐时间 Felling time | ||||
---|---|---|---|---|
1998 | 1996 | 1985 | 1980 | |
盖度 Coverage (%) | 48.53±5.79c | 34.16±4.13b | 10.9±2.13a | 10.7±2.1a |
丰富度 Richness | 6.53±0.41b | 5.24±0.32a | 5.53±0.4a | 5.03±0.32a |
有性生殖数 Sexual reproduction | 0.67±0.22ab | 0.41±0.11a | 0.87±0.10bc | 0.8±0.15b |
生物量 Biomass (g/m2) | 138.22±23.51b | 117.52±21.78b | 18.25±7.05a | 13.95±4.37a |
表5 迹地自然恢复进程中苔藓植物群落结构特征(平均值±标准误, n=30)
Table 5 Characteristics of bryophyte community structure during natural recovery succession (mean±SE, n= 30)
采伐时间 Felling time | ||||
---|---|---|---|---|
1998 | 1996 | 1985 | 1980 | |
盖度 Coverage (%) | 48.53±5.79c | 34.16±4.13b | 10.9±2.13a | 10.7±2.1a |
丰富度 Richness | 6.53±0.41b | 5.24±0.32a | 5.53±0.4a | 5.03±0.32a |
有性生殖数 Sexual reproduction | 0.67±0.22ab | 0.41±0.11a | 0.87±0.10bc | 0.8±0.15b |
生物量 Biomass (g/m2) | 138.22±23.51b | 117.52±21.78b | 18.25±7.05a | 13.95±4.37a |
苔藓层 Bryophyte layer | |||
---|---|---|---|
盖度 Coverage | 丰富度 Richness | 生物量 Biomass | |
灌木层盖度 Shrub coverage (%) | 0.226* | 0.054 | 0.197* |
草本层盖度 Herb coverage (%) | -0.341** | -0.230* | -0.329** |
草本层高度 Herb height (cm) | -0.516** | -0.145 | -0.559** |
草本层丰富度 Herb richness (sp/plot) | -0.289** | -0.079 | -0.379** |
草本层地上生物量 Herb aboveground biomass (g/m2) | -0.503** | -0.205* | -0.538** |
凋落物盖度 Litter coverage (%) | 0.174 | 0.201 | 0.053 |
凋落物生物量 Litter biomass (g/m2) | -0.245* | 0.093 | -0.201* |
表6 皆伐迹地地表苔藓与维管植物发育程度的非参数Spearman相关系数( n=116)
Table 6 Nonparametric Spearman correlation coefficient between ground bryophyte and vascular plant (n = 116)
苔藓层 Bryophyte layer | |||
---|---|---|---|
盖度 Coverage | 丰富度 Richness | 生物量 Biomass | |
灌木层盖度 Shrub coverage (%) | 0.226* | 0.054 | 0.197* |
草本层盖度 Herb coverage (%) | -0.341** | -0.230* | -0.329** |
草本层高度 Herb height (cm) | -0.516** | -0.145 | -0.559** |
草本层丰富度 Herb richness (sp/plot) | -0.289** | -0.079 | -0.379** |
草本层地上生物量 Herb aboveground biomass (g/m2) | -0.503** | -0.205* | -0.538** |
凋落物盖度 Litter coverage (%) | 0.174 | 0.201 | 0.053 |
凋落物生物量 Litter biomass (g/m2) | -0.245* | 0.093 | -0.201* |
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