植物生态学报 ›› 2016, Vol. 40 ›› Issue (3): 187-199.DOI: 10.17521/cjpe.2015.0343
所属专题: 青藏高原植物生态学:群落生态学; 生物多样性
• 研究论文 • 下一篇
孔彬彬1, 卫欣华1, 杜家丽1, 李英年2, 朱志红1,*()
收稿日期:
2015-09-08
修回日期:
2016-02-06
出版日期:
2016-04-11
发布日期:
2016-03-25
通讯作者:
朱志红
基金资助:
Bin-Bin KONG1, Xin-Hua WEI1, Jia-Li DU1, Ying-Nian LI2, Zhi-Hong ZHU1,*()
Received:
2015-09-08
Revised:
2016-02-06
Online:
2016-04-11
Published:
2016-03-25
Contact:
Zhi-Hong ZHU
摘要:
强烈扰动和环境胁迫对植物群落的物种多样性(SD)和功能多样性(FD)有重要影响, 但SD和FD随时间的变化模式及其关系至今少有研究。该研究通过对高寒矮生嵩草(Kobresia humilis)草甸为期7年(2007-2013年)的刈割(3个水平: 不刈割、留茬3 cm和留茬1 cm)和施肥(2个水平: 施肥和不施肥)控制实验探讨了SD和FD随时间的变化模式及其关系。研究结果显示: (1)刈割显著增加了SD和FD, 施肥处理则降低了SD, 对FD增加仅有微弱影响; (2)各处理群落的SD随着时间而下降, FD则随时间增加; (3)随着刈割强度的增加, SD(x)-FD(y)关系由正线性相关变为无相关, 斜率大小为slopel ≥ slopem > slopeh (下标l、m和h分别表示轻度、中度和重度扰动强度), 施肥并不会改变此关系形式和斜率; (4)刈割与施肥对SD和FD的互作效应都不显著, 且不施肥群落SD(x)-FD(y)关系的斜率也为slopel ≥ slopem > slopeh。上述结果说明, 刈割并不一定导致植物功能性状的趋同构建, 也能引发趋异构建, 而施肥引起的强烈种间竞争也并未显著增强趋异构建过程, 这与植物群落构建理论的预测不完全一致。与施肥相比, 刈割扰动是决定群落中SD-FD时间关系形式的主要因素, 并决定着SD-FD关系斜率的变化。
孔彬彬, 卫欣华, 杜家丽, 李英年, 朱志红. 刈割和施肥对高寒草甸物种多样性和功能多样性时间动态及其关系的影响. 植物生态学报, 2016, 40(3): 187-199. DOI: 10.17521/cjpe.2015.0343
Bin-Bin KONG, Xin-Hua WEI, Jia-Li DU, Ying-Nian LI, Zhi-Hong ZHU. Effects of clipping and fertilization on the temporal dynamics of species diversity and functional diversity and their relationships in an alpine meadow. Chinese Journal of Plant Ecology, 2016, 40(3): 187-199. DOI: 10.17521/cjpe.2015.0343
图1 研究区样地设计图(左)及副区处理样方设置图(右)。I、II、III、IV、V代表不同区组。F, 施肥; NF, 不施肥。HC, 重度刈割; MC, 中度刈割; NC, 不刈割。
Fig. 1 The design of treatments (left) and layout of subplot (right). I, II, III, IV, and V represent different blocks. F, fertilizing; NF, no fertilizing. HC, heavy clipping; MC, moderate clipping; NC, no clipping.
变异来源1) Source of variance1) | 自由度 Degree of freedom (m, n)2) | SD | FD | |||||
---|---|---|---|---|---|---|---|---|
F-test | p | F-test | p | |||||
协变量 Covariable | D2007 | 1, 1 | 21.098 | <0.001** | 8.431 | 0.005* | ||
主区 Whole plot | C B | 2, 2 2, 2 | 19.802 0.220 | <0.001** 0.803 | 21.050 1.134 | <0.001** 0.326 | ||
副区 Subplot | F C × F C × B | 1, 1 2, 2 4, 4 | 6.856 0.313 3.341 | 0.010* 0.732 0.013* | 3.490 1.868 2.479 | 0.065† 0.160 0.049* |
表1 2008-2013年施肥、刈割对高寒草甸物种多样性(SD)和功能多样性(FD)影响的协方差分析
Table 1 Multivariate ANCOVA for the effects of clipping and fertilizing on the species diversity (SD) and functional diversity (FD) in the alpine meadow during 2008-2013
变异来源1) Source of variance1) | 自由度 Degree of freedom (m, n)2) | SD | FD | |||||
---|---|---|---|---|---|---|---|---|
F-test | p | F-test | p | |||||
协变量 Covariable | D2007 | 1, 1 | 21.098 | <0.001** | 8.431 | 0.005* | ||
主区 Whole plot | C B | 2, 2 2, 2 | 19.802 0.220 | <0.001** 0.803 | 21.050 1.134 | <0.001** 0.326 | ||
副区 Subplot | F C × F C × B | 1, 1 2, 2 4, 4 | 6.856 0.313 3.341 | 0.010* 0.732 0.013* | 3.490 1.868 2.479 | 0.065† 0.160 0.049* |
图2 刈割和施肥对物种多样性和功能多样性的影响(平均值±标准误差)。FD, 功能多样性; SD, 物种多样性。F、HC、MC、NC、NF分别表示施肥、重度刈割、中度刈割、不刈割和不施肥处理。标准误差上方相同的字母表示处理间差异不显著(p > 0.05), 不同字母表示处理间差异显著(p < 0.05)。
Fig. 2 Effects of clipping and fertilizing on the species diversity (SD) and functional diversity (FD) (mean ± SE) in the alpine meadow community. F, HC, MC, NC, and NF represent fertilizing, heavy clipping, moderate clipping, no clipping, and no fertilizing, respectively. The same letter above standard error bars indicates no difference among treatments (p > 0.05), while different letters indicate significant differences between treatments (p < 0.05).
群落1) Community1) | SD | FD | 自由度 Degree of freedom (m, n)2) | |||
---|---|---|---|---|---|---|
F | p | F | p | |||
NC-NF NC-F MC-NF MC-F HC-NF HC-F | 10.319 132.360 14.392 91.636 13.920 55.714 | 0.002* <0.001** 0.001** <0.001** <0.001** <0.001** | 11.652 1.310 28.214 10.884 40.676 24.987 | 0.001* 0.255 <0.001** 0.001** <0.001** <0.001** | (5,10) |
表2 物种多样性(SD)和功能多样性(FD)在2008-2013年间差异性的方差分析
Table 2 ANOVA for the differences of species diversity (SD) and functional diversity (FD) during 2008-2013
群落1) Community1) | SD | FD | 自由度 Degree of freedom (m, n)2) | |||
---|---|---|---|---|---|---|
F | p | F | p | |||
NC-NF NC-F MC-NF MC-F HC-NF HC-F | 10.319 132.360 14.392 91.636 13.920 55.714 | 0.002* <0.001** 0.001** <0.001** <0.001** <0.001** | 11.652 1.310 28.214 10.884 40.676 24.987 | 0.001* 0.255 <0.001** 0.001** <0.001** <0.001** | (5,10) |
图3 六种群落物种多样性(SD)和功能多样性(FD)在2008-2013年间的线性关系。F、HC、MC、NC、NF分别表示施肥、重度刈割、中度刈割、不刈割和不施肥处理。图中○为物种多样性, ●为功能多样性; *,不同年份之间差异显著(p < 0.05); **,不同年份之间差异极显著(p < 0.001)。
Fig. 3 The changes in species diversity (SD) and functional diversity (FD) of six communities with time during 2008-2013. F, HC, MC, NC, and NF represent fertilizing, heavy clipping, moderate clipping, no clipping, and no fertilizing, respectively. Circles and dots indicates SD and FD, respectively. *, significant differences (p < 0.05) between years; **, significant differences (p < 0.001) between years.
图4 不同处理群落物种多样性和功能多样性在时间序列(2008-2013年)上的关系。F、HC、MC、NC、NF分别表示施肥、重度刈割、中度刈割、不刈割和不施肥处理。
Fig. 4 Relationships between species diversity and functional diversity in different treatment communities (2008-2013). F, HC, MC, NC, and NF represent fertilizing, heavy clipping, moderate clipping, no clipping, and no fertilizing, respectively.
物种名称 Species name | 性状及其分类状态 Trait name and their classification states | ||||
---|---|---|---|---|---|
生长型 Growth form | 植物类群 Plant groups | 生活周期 Life cycle | 植物倾斜度 Plant inclination | 主要繁殖方式 Main reproduction modes | |
1 =散生 Scattered 2 =丛生 Bunched 3 =密丛生 Closely bunched | 1 =禾草科 Gramineae 2 =莎草科 Cyperaceae 3 =豆科 Leguminosae 4 =杂类草 Forbs | 0 =非多年生 Not perennial 1 =多年生 Perennial | 1 =伏卧状 Prostrate 2 =莲座状 Rosette 3 =直立 Erect | 0 =有性繁殖 Sexual reproduction 1 =营养繁殖 Vegetative propagation | |
披碱草 Elymus nutans | 2 | 1 | 1 | 3 | 0 |
异针茅 Stipa aliena | 3 | 1 | 1 | 3 | 1 |
矮生嵩草 Kobresia humilis | 3 | 2 | 1 | 2 | 1 |
糙喙薹草 Carex scabrirostris | 2 | 2 | 1 | 2 | 1 |
羊茅 Festuca ovina | 3 | 1 | 1 | 3 | 1 |
早熟禾 Poa annua | 2 | 1 | 0 | 3 | 0 |
| 3 | 1 | 1 | 3 | 0 |
圆萼刺参 Morina chinensis | 1 | 4 | 1 | 2 | 0 |
麻花艽 Gentiana straminea | 1 | 4 | 1 | 1 | 0 |
美丽风毛菊 Saussurea pulchra | 1 | 4 | 1 | 1 | 0 |
萎软紫菀 Aster flaccidus | 1 | 4 | 1 | 2 | 0 |
肉果草 Lancea tibetica | 1 | 4 | 1 | 1 | 0 |
钝苞雪莲 Saussurea nigrescens | 1 | 4 | 1 | 2 | 0 |
高山唐松草 Thalictrum alpinum | 1 | 4 | 1 | 2 | 0 |
雪白委陵菜 Potentilla nivea | 1 | 4 | 1 | 2 | 1 |
芸香叶唐松草 Thalictrum rutifolium | 1 | 4 | 1 | 3 | 0 |
高山豆 Tibetia himalaica | 1 | 3 | 1 | 2 | 1 |
委陵菜 Potentilla chinensis | 1 | 4 | 1 | 2 | 1 |
短腺小米草 Euphrasia regelii | 1 | 4 | 1 | 3 | 0 |
青海苜蓿 Medicago archiducrisnicolai | 1 | 3 | 1 | 2 | 1 |
湿地繁缕 Stellaria uda | 1 | 4 | 0 | 2 | 1 |
钝裂银莲花 Anemone obtusiloba | 1 | 4 | 1 | 2 | 1 |
皱边喉毛花 Comastoma polycladum | 1 | 4 | 1 | 3 | 1 |
海乳草 Glaux maritima | 1 | 4 | 1 | 3 | 1 |
二裂委陵菜 Potentilla bifurca | 1 | 4 | 1 | 2 | 1 |
蒌蒿 Artemisia selengensis | 1 | 4 | 1 | 3 | 0 |
蒲公英 Taraxacum mongolicum | 1 | 4 | 1 | 1 | 1 |
灰果蒲公英 Taraxacum maurocarpum | 1 | 4 | 1 | 1 | 1 |
黄芪 Astragalus sp. | 1 | 3 | 1 | 2 | 1 |
甘肃棘豆 Oxytropis kansuensis | 1 | 3 | 1 | 2 | 1 |
黄花棘豆 Oxytropis ochrocephala | 1 | 3 | 1 | 2 | 1 |
宽苞棘豆 Oxytropis latibracteata | 1 | 3 | 1 | 2 | 1 |
矮火绒草 Leontopodium nanum | 2 | 4 | 1 | 1 | 1 |
美丽毛茛 Ranunculus pulchellus | 1 | 4 | 1 | 3 | 0 |
棉毛茛 Ranunculus membranaceus | 1 | 4 | 1 | 3 | 0 |
西伯利亚蓼 Polygonum sibiricum | 1 | 4 | 1 | 3 | 0 |
三脉梅花草 Parnassia trinervis | 1 | 4 | 1 | 3 | 0 |
细叶亚菊 Ajania tenuifolia | 1 | 4 | 1 | 3 | 0 |
刺芒龙胆 Gentiana aristata | 1 | 4 | 0 | 2 | 1 |
线叶龙胆 Gentiana lawrencei | 1 | 4 | 1 | 2 | 1 |
偏翅龙胆 Gentiana pudica | 1 | 4 | 0 | 2 | 1 |
双柱头针蔺 Trichophorum distigmaticum | 1 | 2 | 1 | 3 | 1 |
甘肃马先蒿 Pedicularis kansuensis | 1 | 4 | 0 | 3 | 1 |
琴盔马先蒿 Pedicularis lyrata | 1 | 4 | 0 | 3 | 1 |
乳白香青 Anaphalis lactea | 2 | 4 | 0 | 2 | 1 |
铃铃香青 Anaphalis hancockii | 2 | 4 | 0 | 2 | 1 |
湿生扁蕾 Gentianopsis paludosa | 1 | 4 | 0 | 3 | 1 |
珠芽蓼 Polygonum viviparum | 1 | 4 | 1 | 3 | 0 |
鸢尾 Iris tectorum | 1 | 4 | 1 | 3 | 0 |
西藏沙棘 Hippophae tibetana | 1 | 4 | 1 | 3 | 0 |
细蝇子草 Sliene gracilicaulis | 1 | 4 | 1 | 2 | 1 |
藏异燕麦 Helictotrichon tibeticum | 2 | 1 | 0 | 3 | 1 |
微孔草 Microula sikkimensis | 2 | 4 | 0 | 3 | 0 |
刺果猪殃殃 Galium echinocarpum | 1 | 4 | 0 | 2 | 1 |
双叉细柄茅 Ptilagrostis dichotoma | 3 | 1 | 0 | 3 | 1 |
箭叶橐吾 Ligularia sagitta | 1 | 4 | 1 | 3 | 0 |
黄帚橐吾 Ligularia virgaurea | 1 | 4 | 1 | 3 | 0 |
披针叶野决明 Thermopsis lanceolata | 1 | 4 | 1 | 3 | 0 |
草地老鹳草 Geranium pratense | 2 | 4 | 1 | 3 | 0 |
黑柴胡 Bupleurum smithii | 2 | 4 | 1 | 3 | 0 |
宽叶羌活 Notopterygium franchetii | 1 | 4 | 1 | 3 | 0 |
香薷 Elsholtzia ciliata | 1 | 4 | 0 | 3 | 0 |
蓝翠雀花 Delphinium caeruleum | 1 | 4 | 1 | 3 | 0 |
矮生忍冬 Lonicera rupicola | 2 | 4 | 1 | 3 | 0 |
长果婆婆纳 Veronica ciliata | 1 | 4 | 1 | 3 | 0 |
鳞茎堇菜 Viola bulbosa | 1 | 4 | 1 | 1 | 0 |
发草 Deschampsia cespitosa | 3 | 1 | 1 | 3 | 0 |
露蕊乌头 Aconitum gymnandrum | 1 | 4 | 0 | 3 | 0 |
小叶金露梅 Potentilla parvifolia | 1 | 4 | 0 | 3 | 0 |
鸟足毛茛 Ranunculus brotherusii | 1 | 4 | 1 | 3 | 0 |
四数獐牙菜 Swertia tetraptera | 1 | 4 | 1 | 3 | 1 |
美花草 Callianthemum pimpinelloides | 1 | 4 | 1 | 3 | 0 |
车前 Plantago asiatica | 1 | 4 | 1 | 1 | 0 |
附录 计算功能多样性指数的定性性状及其赋值分类状态
Supplement Qualitative functional traits and their categories for calculating functional diversity
物种名称 Species name | 性状及其分类状态 Trait name and their classification states | ||||
---|---|---|---|---|---|
生长型 Growth form | 植物类群 Plant groups | 生活周期 Life cycle | 植物倾斜度 Plant inclination | 主要繁殖方式 Main reproduction modes | |
1 =散生 Scattered 2 =丛生 Bunched 3 =密丛生 Closely bunched | 1 =禾草科 Gramineae 2 =莎草科 Cyperaceae 3 =豆科 Leguminosae 4 =杂类草 Forbs | 0 =非多年生 Not perennial 1 =多年生 Perennial | 1 =伏卧状 Prostrate 2 =莲座状 Rosette 3 =直立 Erect | 0 =有性繁殖 Sexual reproduction 1 =营养繁殖 Vegetative propagation | |
披碱草 Elymus nutans | 2 | 1 | 1 | 3 | 0 |
异针茅 Stipa aliena | 3 | 1 | 1 | 3 | 1 |
矮生嵩草 Kobresia humilis | 3 | 2 | 1 | 2 | 1 |
糙喙薹草 Carex scabrirostris | 2 | 2 | 1 | 2 | 1 |
羊茅 Festuca ovina | 3 | 1 | 1 | 3 | 1 |
早熟禾 Poa annua | 2 | 1 | 0 | 3 | 0 |
| 3 | 1 | 1 | 3 | 0 |
圆萼刺参 Morina chinensis | 1 | 4 | 1 | 2 | 0 |
麻花艽 Gentiana straminea | 1 | 4 | 1 | 1 | 0 |
美丽风毛菊 Saussurea pulchra | 1 | 4 | 1 | 1 | 0 |
萎软紫菀 Aster flaccidus | 1 | 4 | 1 | 2 | 0 |
肉果草 Lancea tibetica | 1 | 4 | 1 | 1 | 0 |
钝苞雪莲 Saussurea nigrescens | 1 | 4 | 1 | 2 | 0 |
高山唐松草 Thalictrum alpinum | 1 | 4 | 1 | 2 | 0 |
雪白委陵菜 Potentilla nivea | 1 | 4 | 1 | 2 | 1 |
芸香叶唐松草 Thalictrum rutifolium | 1 | 4 | 1 | 3 | 0 |
高山豆 Tibetia himalaica | 1 | 3 | 1 | 2 | 1 |
委陵菜 Potentilla chinensis | 1 | 4 | 1 | 2 | 1 |
短腺小米草 Euphrasia regelii | 1 | 4 | 1 | 3 | 0 |
青海苜蓿 Medicago archiducrisnicolai | 1 | 3 | 1 | 2 | 1 |
湿地繁缕 Stellaria uda | 1 | 4 | 0 | 2 | 1 |
钝裂银莲花 Anemone obtusiloba | 1 | 4 | 1 | 2 | 1 |
皱边喉毛花 Comastoma polycladum | 1 | 4 | 1 | 3 | 1 |
海乳草 Glaux maritima | 1 | 4 | 1 | 3 | 1 |
二裂委陵菜 Potentilla bifurca | 1 | 4 | 1 | 2 | 1 |
蒌蒿 Artemisia selengensis | 1 | 4 | 1 | 3 | 0 |
蒲公英 Taraxacum mongolicum | 1 | 4 | 1 | 1 | 1 |
灰果蒲公英 Taraxacum maurocarpum | 1 | 4 | 1 | 1 | 1 |
黄芪 Astragalus sp. | 1 | 3 | 1 | 2 | 1 |
甘肃棘豆 Oxytropis kansuensis | 1 | 3 | 1 | 2 | 1 |
黄花棘豆 Oxytropis ochrocephala | 1 | 3 | 1 | 2 | 1 |
宽苞棘豆 Oxytropis latibracteata | 1 | 3 | 1 | 2 | 1 |
矮火绒草 Leontopodium nanum | 2 | 4 | 1 | 1 | 1 |
美丽毛茛 Ranunculus pulchellus | 1 | 4 | 1 | 3 | 0 |
棉毛茛 Ranunculus membranaceus | 1 | 4 | 1 | 3 | 0 |
西伯利亚蓼 Polygonum sibiricum | 1 | 4 | 1 | 3 | 0 |
三脉梅花草 Parnassia trinervis | 1 | 4 | 1 | 3 | 0 |
细叶亚菊 Ajania tenuifolia | 1 | 4 | 1 | 3 | 0 |
刺芒龙胆 Gentiana aristata | 1 | 4 | 0 | 2 | 1 |
线叶龙胆 Gentiana lawrencei | 1 | 4 | 1 | 2 | 1 |
偏翅龙胆 Gentiana pudica | 1 | 4 | 0 | 2 | 1 |
双柱头针蔺 Trichophorum distigmaticum | 1 | 2 | 1 | 3 | 1 |
甘肃马先蒿 Pedicularis kansuensis | 1 | 4 | 0 | 3 | 1 |
琴盔马先蒿 Pedicularis lyrata | 1 | 4 | 0 | 3 | 1 |
乳白香青 Anaphalis lactea | 2 | 4 | 0 | 2 | 1 |
铃铃香青 Anaphalis hancockii | 2 | 4 | 0 | 2 | 1 |
湿生扁蕾 Gentianopsis paludosa | 1 | 4 | 0 | 3 | 1 |
珠芽蓼 Polygonum viviparum | 1 | 4 | 1 | 3 | 0 |
鸢尾 Iris tectorum | 1 | 4 | 1 | 3 | 0 |
西藏沙棘 Hippophae tibetana | 1 | 4 | 1 | 3 | 0 |
细蝇子草 Sliene gracilicaulis | 1 | 4 | 1 | 2 | 1 |
藏异燕麦 Helictotrichon tibeticum | 2 | 1 | 0 | 3 | 1 |
微孔草 Microula sikkimensis | 2 | 4 | 0 | 3 | 0 |
刺果猪殃殃 Galium echinocarpum | 1 | 4 | 0 | 2 | 1 |
双叉细柄茅 Ptilagrostis dichotoma | 3 | 1 | 0 | 3 | 1 |
箭叶橐吾 Ligularia sagitta | 1 | 4 | 1 | 3 | 0 |
黄帚橐吾 Ligularia virgaurea | 1 | 4 | 1 | 3 | 0 |
披针叶野决明 Thermopsis lanceolata | 1 | 4 | 1 | 3 | 0 |
草地老鹳草 Geranium pratense | 2 | 4 | 1 | 3 | 0 |
黑柴胡 Bupleurum smithii | 2 | 4 | 1 | 3 | 0 |
宽叶羌活 Notopterygium franchetii | 1 | 4 | 1 | 3 | 0 |
香薷 Elsholtzia ciliata | 1 | 4 | 0 | 3 | 0 |
蓝翠雀花 Delphinium caeruleum | 1 | 4 | 1 | 3 | 0 |
矮生忍冬 Lonicera rupicola | 2 | 4 | 1 | 3 | 0 |
长果婆婆纳 Veronica ciliata | 1 | 4 | 1 | 3 | 0 |
鳞茎堇菜 Viola bulbosa | 1 | 4 | 1 | 1 | 0 |
发草 Deschampsia cespitosa | 3 | 1 | 1 | 3 | 0 |
露蕊乌头 Aconitum gymnandrum | 1 | 4 | 0 | 3 | 0 |
小叶金露梅 Potentilla parvifolia | 1 | 4 | 0 | 3 | 0 |
鸟足毛茛 Ranunculus brotherusii | 1 | 4 | 1 | 3 | 0 |
四数獐牙菜 Swertia tetraptera | 1 | 4 | 1 | 3 | 1 |
美花草 Callianthemum pimpinelloides | 1 | 4 | 1 | 3 | 0 |
车前 Plantago asiatica | 1 | 4 | 1 | 1 | 0 |
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