岷江上游植被群落特征研究
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摘要
岷江上游位于青藏高原和四川盆地的过渡地带,是我国一个重要的大尺度、复合型生态过渡带,也是一个生态系统脆弱区,其自然环境的复杂性、生态系统的脆弱性、经济发展的边缘性、社会文化的过渡性在我国都有典型的代表性。该区是世界生物多样性的热点地区和我国生物多样性保护的关键地区之一。近几十年来,由于恶劣的自然环境和过度的人类干扰,导致区域植被退化,引起了生物多样性丧失等一系列生态环境问题,对整个岷江流域乃至长江上游的生态安全和区域可持续发展构成严重的威胁。加之该区生态环境的脆弱性,一旦破坏后极易形成逆行演替,极难恢复,尽快开展植被群落特征的基础理论研究具有重要意义。
本文选取岷江上游干旱河谷和岷江源区为研究对象,通过对该地区植被的大量群落学调查,综合TWINSPAN、DCCA、典型相关分析、主成分分析等多元统计分析方法和层次分析法(AHP),围绕环境梯度的变化与植被数量特征的响应和相互关系,进行了植被分类、群落分布与环境因子的关系、种群生态位、群落多样性、群落复杂性和群落稳定性等植物群落特征的研究,得出以下主要结论:
(1)DCCA排序分析表明,岷江上游地区,植物群落的分布受土壤养分、水分及地形因子共同的影响,表现为DCCA各排序轴与土壤水分、养分和微地形呈现较高的相关性。地形因子中的海拔、坡形、坡度和坡向,土壤因子中的速效P、全P、土壤含水量、全N、pH值对植物群落分布有着较大的影响。
(2)将土壤养分、水分因子作为资源梯度轴来研究种群及其种组生态位,较以往研究中以群落为梯度研究生态位更能反映种群对环境资源的利用和适应能力。主要灌木和草本种群的生态位宽度反映了种群对环境的适应能力和分布均匀程度。小花滇紫草、三花莸、白刺花、多花胡枝子、马鞍羊蹄甲、绵枣儿、铁扫帚、翻白委陵菜、黄背草、密生苔草、高山绣线菊、尖叶栒子、峨眉蔷薇、金露梅、窄叶鲜卑花,披针苔草、珠光香青、毛莨、小柴胡、圆穗蓼有较高的生态位宽度,具有较强的生态适应性,形成了该地区的优势种群;而翠蓝绣线菊、檀子栎、对节刺、猪毛菜、紫花野茅、歧茎蒿、虱子草、沙柳、喜阴悬钩子、沙棘、灰苞蒿、珠芽蓼、细弱草莓等种群,生态位宽度较低。生态位宽度较小的物种并非不是群落的优势种,取决于种群的分布范围和分布的均匀程度。
(3)干旱河谷植物群落随海拔的增加,草本层和灌木层的α多样性和β多样性均表现为“高-低-高”的变化趋势。草本层的α多样性和β多样性高于灌木层,草本层和灌木层均在1400～1600 m和2000～2200 m两个海拔段有较高的α多样性和β多样性,反映了中海拔地区的生物多样性较低,低海拔和较高海拔地区多样性较高;不同群系类型的多样性表现为华帚菊-小黄素馨灌丛、金花小檗-忍冬灌丛、绣线菊灌丛有着较高的α多样性和β多样性,而西南野丁香灌丛、莸灌丛、小马鞍羊蹄甲-白刺花灌丛的群落α多样性和β多样性较低;不同坡向和坡形,α多样性和β多样性均表现为:阴坡>半阴半阳坡>阳坡、凹坡>平坡>凸坡;不同坡位,下坡位>上坡位>中坡位。灌木层和草本层α多样性在不同坡位上的多样性大小均为上坡位>下坡位>中坡位,而β多样性表现为下坡位>上坡位>中坡位。
岷江源区植物群落在2900-3100 m、3500-3700 m海拔段有较高的a多样性,2700-2900m、3100-3400m、3700-3900m海拔段的多样性较低。不同植被类型的a多样性表现为森林群系>灌丛群系>草甸群系。柳灌丛、窄叶鲜卑花灌丛、绣线菊灌丛、锦鸡儿灌丛的草本层多样性较高,沙棘灌丛和小果小檗灌丛较低。草甸群系中,白茅草甸和苔草草甸多样性较高,蒿草草甸和高山草甸的物种多样性较低。锦鸡儿灌丛灌木层多样性较高,沙棘灌丛的多样性最低。岷江源区植物群落物种组成受海拔梯度的影响强烈,不同海拔梯度的物种组成差异明显,群落β多样性受草本层变化的影响较大。在不同群落类型过渡的2900～3100m和3500～3700m这两个海拔段,物种平均替代速率均较高,群落异质性较高。而在2700～2900m、3200～3300m、3800～3900m海拔段,生境异质性较低,物种平均替代速率也较低。灌木层β多样性沿海拔梯度的变化相对平稳,最高值出现在2900m处,最低值出现在3300 m处。
β多样性与土壤水分和养分之间呈现显著的二次曲线关系,表明物种替代速率最大的植物群落,其样地的土壤养分和水分含量处于中间水平,并非养分、水分含量越高,物种周转速率就越高。
(4)岷江上游灌丛地上生物量与海拔、土壤含水量、速效K、有机质、全P、水解N、全N呈显著的正相关关系,与坡度和pH值呈显著的负相关关系,其中海拔、土壤水分和速效K对灌丛地上生物量的影响最为显著。灌丛地上生物量随物种多样性和物种丰富度的增加而增加,与物种多样性、物种丰富度呈极显著的线性正相关关系。
(5)用物种重要值代替物种多度,应用通讯理论中的霍夫曼编码方法研究群落复杂性及其与环境因子之间的一般关系,并揭示了群落复杂性与多样性、均匀度之间的一般关系。干旱河谷植物群落,在高海拔和低海拔段有较高的复杂性,中海拔段复杂性较低;不同坡位、坡形及坡向,群落总复杂性和结构复杂性,均表现为上坡位>下坡位>中坡位,凹坡>平破>凸坡,阴坡>半阴半阳坡>阳坡。不同群系的复杂性表现为华帚菊-小黄素馨灌丛的总复杂性最高,西南野丁香灌丛、驼绒藜灌丛的总复杂性最低。源区植物群落,在2900-3100m和3500-3700m两个海拔段有较高的总复杂性。不同群系的总复杂性,表现为森林群系>灌丛群系>草甸群系。云杉林的总复杂性最高,蒿草草甸的总复杂性最低。灌丛群系中,锦鸡儿灌丛的总复杂性最高,沙棘灌丛的总复杂性最低。草甸群系中,苔草草甸和白茅草甸群系总复杂性较高,蒿草草甸和高山草甸总复杂性较低。与β多样性随土壤养分变化规律相似,群落复杂性与土壤水分和养分之间也呈现显著的二次曲线关系,但源区和干旱河谷中影响群落总复杂性的土壤因子不尽相同。群落总复杂性并不是随土壤养分和水分含量的增加而呈现简单的线性增加关系。
总复杂性与多样性及物种丰富度呈现出极显著的正相关关系,均匀度与结构复杂性有极显著的负相关关系。结构复杂性作为群落总复杂性与多样性的区分,在均匀度较高的群落中,群落总复杂性和多样性差别较小,结构复杂性较低。但也存在特殊情况,即当群落中物种数量较少时,即使较低的均匀度也不会使群落的结构复杂性达到最高,结构复杂性也会相对较低。结构复杂性对群落内物种数的变化较为敏感,均匀度和物种数共同影响着群落的复杂性。
(6)将源于系统决策的层次分析法引入到群落稳定性评价研究,将植被盖度、多样性、复杂性、演替度等反映植被稳定性的数量特征指标结合气候、地形、土壤等外部环境因子,评价了岷江上游典型植物群落的稳定性,弥补了以往群落稳定性研究中只考虑植被数量特征的不足。源区植被群系中,云杉林、柳灌丛、窄叶鲜卑花灌丛、冷杉林、绣线菊灌丛有较高的稳定性值,沙棘和小果小檗灌丛稳定性值较低。草甸群系中,苔草草甸和白茅草草甸具有较高的稳定性值,蒿草草甸和高山草甸稳定性值最低。干旱河谷植被群系中,绣线菊灌丛、瑞香灌丛、小花滇紫草灌丛、小马鞍羊蹄甲-白刺花灌丛、莸灌丛、橿子栎灌丛有较高的稳定性值。
长期的外界干扰下,岷江上游群落发生逆行演替。干旱河谷灌丛逆行演替模式为:檀子栎、绣线菊灌丛→华帚菊-小黄素馨灌丛、小花滇紫草灌丛→驼绒藜灌丛、莸灌丛、白刺花灌丛;源区逆行演替模式为云杉、冷杉针叶林→鲜卑花、绣线菊灌丛→锦鸡儿、小檗灌丛→白茅草甸、蒿草草甸。
The upper reaches of the Minjiang River, located between the Qinghai-Tibet plateau to the Sichuan Basin, is a fragile as well as an important large-scale and compound ecological transition zone , which is a typical case in terms of its complexity of natural environment, fragility of ecosystem, marginalizatioin of economical development and transitionality of social-culture in China, hence becoming one of the hot spots of world biodiversity conservation and key areas of domestic biodiversity conservation. A long history of anthropogenic influences combined with severe natural environments has resulted in its regional degraded vegetations, bringing out a series of ecological problems such as loss of biodiversity and posing a serious threat to the whole drainage basin of the Minjiang River even the ecological security and regional sustainable development in the upper reaches of Yangtze River. The readiness of regressive succession in ecological environment after being damaged and the slim possibility of restoration caused by fragility of ecological environment, makes an urgent call for a fundamental theoretical research on biodiversity conservation and restoration.
Selecting the upper reaches region of Minjiang River as subject, revolving around the mutual responds and relationships between the environmental gradient and vegetation characteristics, this paper carries on the research on the vegetation classification, the connection between community distribution and environmental factors, population niche, community diversity, community complexity and community stability by investigating a large number of vegetation communities in this region and combining multivariate statistics such as TWINSPAN, DCCA, canonical correlation analysis, principal component analysis and AHP. The main conclusions this paper comes to are as follows:
(1) DCCA ordination analysis shows that in the upper reaches region of the Minjiang River, the distribution of plant communities under the mixed influence of soil nutrient and water content demonstrates a high correlation with soil water content, nutrient and micro-topographigraphy, as illustrated in each axis of DCCA. Altitude, slope shape, slope degree, direction as micro topographical factors and available P, total P, soil water content, total N and soil pH as soil factors can produce a significant influence on the distribution of plant community.
(2) Compared with previous studies in the gradient of community, this research on population niche in the resource gradient of soil nutrient and water content can generate a clearer demonstration of the ability of using and adaptability to environmental resources of populations. The niche breadth of main shrubs and herbs can reflect the adaptability to the environment and evenness of distribution of populations. Onosma farrerii, Caryopteris terniflora, Sorphora vrcifolia, Lespedeza floribunda, Bauhinia faberi, Scilla scilloides, Indigofera bungeana, Potentilla leuconota,Themeda triandra, Carex crebra, Spiraea alpina、 Cotoneaster acuminatus, Rosa omeiensis, Potentilla fruticosa, Sibiraea angustata, Carex lanceolata, Anaphalis margaritacea, Ranunculus japonicus, Bupleurum tenue, Polygonum phaerostachyum have higher niche width and stronger adaptability, hence becoming a dominant populations in this region; while Spiraea henryi, Quercus cocciferoides, Sageretia pycnophylla, Salsola collina, Calamagrostis purpurea, Artemisia igniaria, Tragus berteronianus，Salix cheilophilla, Rubus mesogaeus, Hippophae rhamnoides, Artemisia roxlourghiana, Polygonum viviparum,Fragaria gracilis have lower niche. Not all populations with lower niche are dominant populations. This peculiarity depends on the range and evenness of distribution of the certain population. Populations with higher niche breadth always have higher niche overlap, but not all populations with smaller niche breadth have lower niche overlap.
(3) Among the plant communities in arid valley, with an increase of altitude, both the a diversity and β diversity of herbs layer and shrubs layer display a "high-low-high" trend. The α diversity and β diversity of herbs layer are higher than those of shrubs layer. Both the herbs layer and shrubs layer have higher a diversity and β diversity at an altitude range of 1400～1600m and 2000～2200m, which is a potent proof that a low biodiversity occurs in regions of a medium altitude, while a high one exists in regions of high and low altitudes.The diversity varies in different formations: Form. Pertya sinensis, Jasminum humile .Form. Berberis wilsonae, Lonicera japonica, Form. Spiraea Spp.have higher a diversity and β diversity. Form. Leptodermis purdomi, Form. Caryopteris spp. and Form. Bauhinia faberi , Sorphora vrcifolia have lower α diversity and β diversity; both a diversity and β diversity in different slope directions and shapes: shaded >slope half-shaded> slope sunny slope, concave slope> straight> convex slope; in different positions: lower slope> upper slope> middle slope . The a diversity of shrubs layer and herbs layer alike in different slope positions: upper slope> lower slope> middle slope, while the β diversity, lower slope> upper slope> middle slope.
Among the plant communities in origin region of the Minjiang River, with an increase of altitude, diversity of both the herbs layer and shrubs layer display symmetrical changing trends that they are almost the same at altitude ranges of 2700-3300m and 3300-3900m. The altitude ranges of 2900-3100m and 3500-3700m see a higher diversity, while those of 2700-2900m, 3100-3400, 3700-3900m see a lower one. The maximum diversity of shrubs layer and herbs layer occurs at the altitudes of 3000m and 3600m respectively. The diversity in different vegetation types follows the trend: forests formations >shrubs formations >meadow formations. The diversity in herb layer of Form. Salix spp., Form. Sibiraea angustata, Form. Spiraea spp.and Form. Caragana spp.are at high level, while Form. Hippophae rhamnoides and Form. Berberis amurensis are at lower level.In meadow formations, the diversity of Form. Imperata cylindrica and Form. Carex spp.are higher than that of Form. Sub-alpine and Form. Kobresia spp. As for shrub layer , Form. Caragana spp. has the highest diversity, Form. Hippophae rhamnoides has the lowest diversity. Diversity in herb layer is higher than that of shrub layer.
Under a great influence of altitude gradient, species compositions in plant communities in the origin area of Minjiang River show significant differences and the β diversity is more easily affected by the change of herbs layer. Along the altitude gradient, there are higher species turnover rates at the altitude ranges of 2900-3100m and 3500-3700m, and the community has a higher heterogeneity. While at the altitude ranges of 2700-2900m, 3200-3300m, 3800-3900m,, the community has a lower heterogeneity and species turnover rate. The change of β diversity in shrubs layer is comparatively stable along the altitude gradient, with its climax occurring at an altitude of 2900m and the minimum at 3300m.
A significant quadratic curve relationship between the β diversity and soil moisture and nutrient shows that the soil nutrient and water content of the plant community with the highest species turnover rate are at a medium level, instead of a high level of nutrient and water content. (4) The very significant positive relationship exist between the aboveground biomass of shrub communities and the altitude, available K, soil moisture, organic matter, total P, hydrolyazble N, total N, available N in the upper reaches of the Minjiang River, while the very significant negative relationship exist between slope gradient and pH value, among which altitude, available K and soil moisture play a leading role in affecting the aboveground shrub biomass. The aboveground biomass of shrub communities increases with the raise of species diversity and species richness and has a very positive linear relationship with species diversity and species richness.
(5) Huffman coding method of communication theory is used to study the plant complexity and the relationship with environmental factors and reveal that between plant community complexity and diversity , evenness. Both total and structural community complexity vary with degree, shape and aspect of slope, with an order of upper slope>middle slope>lower slope, shaded slope> half-shaded slope>sunny slope, and concave slope>straight slope>convex slope. Form. Pertyasinensis,Jasminumhumile have the highest total complexity, and Form Leptodermispurdomi and Form Ceratoidesarborescens communities have the lowest total complexity, while Form Quercnscocciferoides and Form Onosma farrerii communities have the highest structural complexity.
The plant formations in origin area have a high total complexity at the altitude ranges of 2900-3100m and 3500-3700m. The total complexity is as the following: Forest plant formations>shrub formations>meadow forations. Form. Picea asperata is ranked as the highest in total complexity, Form. Kobresia spp have a lowest one . In shrub formations, Form. Caragana spp.have the highest total complexity, Form. Hippophae rhamnoides have the lowest one. In meadow formations, the total complexity of Form. Carex spp. and Form. Imperata cylindrica are at high level, Form. Kobresia spp.and Form. Sub-alpine are at lower level. Similar to the correlation of β diversity with soil nutrient, The total community complexity had significant quadratic correlations with soil water content and nutrient. However, the soil factors that influence total community complexity in origin area and arid valley are different, therefore, it does not grow in a simple linear way with the increase of soil nutrient and water content.
Total community complexity is positively correlated with community diversity, evenness and species richness, while structural complexity is negatively correlated with community evenness. As two components of total community complexity, structural complexity is more sensitive than diversity to the change of species in the community, which is not only related to community evenness, but also to community richness. In the communities with a high evenness, the difference between total complexity and diversity is not quite obvious, and structural complexity is lower. However, the exceptional case exists in which a low evenness cannot push the low structural complexity of the communities with a low species richness to its climax. The relative contribution of structural complexity and diversity to total complexity would be different for different study area or ecosystems.
(6) The stability of typical plant formations in the upper reaches of Minjiang River is analysed by using AHP methods, combining quantitative character indices reflecting vegetation stability, such as vegetation coverage, diversity, complexity and degree of succession with external environmental factors such as climate, topography and soil character, hence supplementing the insufficiency that previous researches on vegetation stability assessment take the only vegetation character into account. The result indicates: the plant formations of origin region, Form. Picea asperata, Form. Salix spp., Form. Sibiraea angustata, Form. Abies faxoniana, Form. Spiraea spp. Have higher stability, while Hippophae rhamnoides and Form. Berberis amurensis have lower one. Form. Imperata cylindrical and Form. Imperata cylindrica have a higher stability, Form. Kobresia spp.and Form. Sub-alpine have lower one. In the plant formations of the arid valley region, Form. Spiraea spp,Form. Daphne spp., Form. Onosma farrerii, Form. Bauhinia faberi, Sorphora vrcifolia, Form. Caryopteris spp., Form. Quercus cocciferoides have a higher stability.
Under a constant external disturbance, regressive succession of the vegetation occurs in the upper reaches of the Minjiang River. Regressive succession model of the vegetation in the arid valley is: Form. Quercus cocciferoides, Form. Spiraea Spp.→Form. Pertya sinensis, Jasminum humile, Form. Onosma farrerii→Form. Ceratoides arborescens, Form. Caryopteris spp. Form. Bauhinia faberi , Sorphora vrcifolia; the regressive succession model of the origin region is Form. Abies faxoniana、 Form. Picea asperata→Form. Sibiraea angustata→Form. Caragana spp., Form. Berberis amurensis→Form. Imperata cylindrical, Form. Kobresia spp.
本文选取岷江上游干旱河谷和岷江源区为研究对象,通过对该地区植被的大量群落学调查,综合TWINSPAN、DCCA、典型相关分析、主成分分析等多元统计分析方法和层次分析法(AHP),围绕环境梯度的变化与植被数量特征的响应和相互关系,进行了植被分类、群落分布与环境因子的关系、种群生态位、群落多样性、群落复杂性和群落稳定性等植物群落特征的研究,得出以下主要结论:
(1)DCCA排序分析表明,岷江上游地区,植物群落的分布受土壤养分、水分及地形因子共同的影响,表现为DCCA各排序轴与土壤水分、养分和微地形呈现较高的相关性。地形因子中的海拔、坡形、坡度和坡向,土壤因子中的速效P、全P、土壤含水量、全N、pH值对植物群落分布有着较大的影响。
(2)将土壤养分、水分因子作为资源梯度轴来研究种群及其种组生态位,较以往研究中以群落为梯度研究生态位更能反映种群对环境资源的利用和适应能力。主要灌木和草本种群的生态位宽度反映了种群对环境的适应能力和分布均匀程度。小花滇紫草、三花莸、白刺花、多花胡枝子、马鞍羊蹄甲、绵枣儿、铁扫帚、翻白委陵菜、黄背草、密生苔草、高山绣线菊、尖叶栒子、峨眉蔷薇、金露梅、窄叶鲜卑花,披针苔草、珠光香青、毛莨、小柴胡、圆穗蓼有较高的生态位宽度,具有较强的生态适应性,形成了该地区的优势种群;而翠蓝绣线菊、檀子栎、对节刺、猪毛菜、紫花野茅、歧茎蒿、虱子草、沙柳、喜阴悬钩子、沙棘、灰苞蒿、珠芽蓼、细弱草莓等种群,生态位宽度较低。生态位宽度较小的物种并非不是群落的优势种,取决于种群的分布范围和分布的均匀程度。
(3)干旱河谷植物群落随海拔的增加,草本层和灌木层的α多样性和β多样性均表现为“高-低-高”的变化趋势。草本层的α多样性和β多样性高于灌木层,草本层和灌木层均在1400～1600 m和2000～2200 m两个海拔段有较高的α多样性和β多样性,反映了中海拔地区的生物多样性较低,低海拔和较高海拔地区多样性较高;不同群系类型的多样性表现为华帚菊-小黄素馨灌丛、金花小檗-忍冬灌丛、绣线菊灌丛有着较高的α多样性和β多样性,而西南野丁香灌丛、莸灌丛、小马鞍羊蹄甲-白刺花灌丛的群落α多样性和β多样性较低;不同坡向和坡形,α多样性和β多样性均表现为:阴坡>半阴半阳坡>阳坡、凹坡>平坡>凸坡;不同坡位,下坡位>上坡位>中坡位。灌木层和草本层α多样性在不同坡位上的多样性大小均为上坡位>下坡位>中坡位,而β多样性表现为下坡位>上坡位>中坡位。
岷江源区植物群落在2900-3100 m、3500-3700 m海拔段有较高的a多样性,2700-2900m、3100-3400m、3700-3900m海拔段的多样性较低。不同植被类型的a多样性表现为森林群系>灌丛群系>草甸群系。柳灌丛、窄叶鲜卑花灌丛、绣线菊灌丛、锦鸡儿灌丛的草本层多样性较高,沙棘灌丛和小果小檗灌丛较低。草甸群系中,白茅草甸和苔草草甸多样性较高,蒿草草甸和高山草甸的物种多样性较低。锦鸡儿灌丛灌木层多样性较高,沙棘灌丛的多样性最低。岷江源区植物群落物种组成受海拔梯度的影响强烈,不同海拔梯度的物种组成差异明显,群落β多样性受草本层变化的影响较大。在不同群落类型过渡的2900～3100m和3500～3700m这两个海拔段,物种平均替代速率均较高,群落异质性较高。而在2700～2900m、3200～3300m、3800～3900m海拔段,生境异质性较低,物种平均替代速率也较低。灌木层β多样性沿海拔梯度的变化相对平稳,最高值出现在2900m处,最低值出现在3300 m处。
β多样性与土壤水分和养分之间呈现显著的二次曲线关系,表明物种替代速率最大的植物群落,其样地的土壤养分和水分含量处于中间水平,并非养分、水分含量越高,物种周转速率就越高。
(4)岷江上游灌丛地上生物量与海拔、土壤含水量、速效K、有机质、全P、水解N、全N呈显著的正相关关系,与坡度和pH值呈显著的负相关关系,其中海拔、土壤水分和速效K对灌丛地上生物量的影响最为显著。灌丛地上生物量随物种多样性和物种丰富度的增加而增加,与物种多样性、物种丰富度呈极显著的线性正相关关系。
(5)用物种重要值代替物种多度,应用通讯理论中的霍夫曼编码方法研究群落复杂性及其与环境因子之间的一般关系,并揭示了群落复杂性与多样性、均匀度之间的一般关系。干旱河谷植物群落,在高海拔和低海拔段有较高的复杂性,中海拔段复杂性较低;不同坡位、坡形及坡向,群落总复杂性和结构复杂性,均表现为上坡位>下坡位>中坡位,凹坡>平破>凸坡,阴坡>半阴半阳坡>阳坡。不同群系的复杂性表现为华帚菊-小黄素馨灌丛的总复杂性最高,西南野丁香灌丛、驼绒藜灌丛的总复杂性最低。源区植物群落,在2900-3100m和3500-3700m两个海拔段有较高的总复杂性。不同群系的总复杂性,表现为森林群系>灌丛群系>草甸群系。云杉林的总复杂性最高,蒿草草甸的总复杂性最低。灌丛群系中,锦鸡儿灌丛的总复杂性最高,沙棘灌丛的总复杂性最低。草甸群系中,苔草草甸和白茅草甸群系总复杂性较高,蒿草草甸和高山草甸总复杂性较低。与β多样性随土壤养分变化规律相似,群落复杂性与土壤水分和养分之间也呈现显著的二次曲线关系,但源区和干旱河谷中影响群落总复杂性的土壤因子不尽相同。群落总复杂性并不是随土壤养分和水分含量的增加而呈现简单的线性增加关系。
总复杂性与多样性及物种丰富度呈现出极显著的正相关关系,均匀度与结构复杂性有极显著的负相关关系。结构复杂性作为群落总复杂性与多样性的区分,在均匀度较高的群落中,群落总复杂性和多样性差别较小,结构复杂性较低。但也存在特殊情况,即当群落中物种数量较少时,即使较低的均匀度也不会使群落的结构复杂性达到最高,结构复杂性也会相对较低。结构复杂性对群落内物种数的变化较为敏感,均匀度和物种数共同影响着群落的复杂性。
(6)将源于系统决策的层次分析法引入到群落稳定性评价研究,将植被盖度、多样性、复杂性、演替度等反映植被稳定性的数量特征指标结合气候、地形、土壤等外部环境因子,评价了岷江上游典型植物群落的稳定性,弥补了以往群落稳定性研究中只考虑植被数量特征的不足。源区植被群系中,云杉林、柳灌丛、窄叶鲜卑花灌丛、冷杉林、绣线菊灌丛有较高的稳定性值,沙棘和小果小檗灌丛稳定性值较低。草甸群系中,苔草草甸和白茅草草甸具有较高的稳定性值,蒿草草甸和高山草甸稳定性值最低。干旱河谷植被群系中,绣线菊灌丛、瑞香灌丛、小花滇紫草灌丛、小马鞍羊蹄甲-白刺花灌丛、莸灌丛、橿子栎灌丛有较高的稳定性值。
长期的外界干扰下,岷江上游群落发生逆行演替。干旱河谷灌丛逆行演替模式为:檀子栎、绣线菊灌丛→华帚菊-小黄素馨灌丛、小花滇紫草灌丛→驼绒藜灌丛、莸灌丛、白刺花灌丛;源区逆行演替模式为云杉、冷杉针叶林→鲜卑花、绣线菊灌丛→锦鸡儿、小檗灌丛→白茅草甸、蒿草草甸。
The upper reaches of the Minjiang River, located between the Qinghai-Tibet plateau to the Sichuan Basin, is a fragile as well as an important large-scale and compound ecological transition zone , which is a typical case in terms of its complexity of natural environment, fragility of ecosystem, marginalizatioin of economical development and transitionality of social-culture in China, hence becoming one of the hot spots of world biodiversity conservation and key areas of domestic biodiversity conservation. A long history of anthropogenic influences combined with severe natural environments has resulted in its regional degraded vegetations, bringing out a series of ecological problems such as loss of biodiversity and posing a serious threat to the whole drainage basin of the Minjiang River even the ecological security and regional sustainable development in the upper reaches of Yangtze River. The readiness of regressive succession in ecological environment after being damaged and the slim possibility of restoration caused by fragility of ecological environment, makes an urgent call for a fundamental theoretical research on biodiversity conservation and restoration.
Selecting the upper reaches region of Minjiang River as subject, revolving around the mutual responds and relationships between the environmental gradient and vegetation characteristics, this paper carries on the research on the vegetation classification, the connection between community distribution and environmental factors, population niche, community diversity, community complexity and community stability by investigating a large number of vegetation communities in this region and combining multivariate statistics such as TWINSPAN, DCCA, canonical correlation analysis, principal component analysis and AHP. The main conclusions this paper comes to are as follows:
(1) DCCA ordination analysis shows that in the upper reaches region of the Minjiang River, the distribution of plant communities under the mixed influence of soil nutrient and water content demonstrates a high correlation with soil water content, nutrient and micro-topographigraphy, as illustrated in each axis of DCCA. Altitude, slope shape, slope degree, direction as micro topographical factors and available P, total P, soil water content, total N and soil pH as soil factors can produce a significant influence on the distribution of plant community.
(2) Compared with previous studies in the gradient of community, this research on population niche in the resource gradient of soil nutrient and water content can generate a clearer demonstration of the ability of using and adaptability to environmental resources of populations. The niche breadth of main shrubs and herbs can reflect the adaptability to the environment and evenness of distribution of populations. Onosma farrerii, Caryopteris terniflora, Sorphora vrcifolia, Lespedeza floribunda, Bauhinia faberi, Scilla scilloides, Indigofera bungeana, Potentilla leuconota,Themeda triandra, Carex crebra, Spiraea alpina、 Cotoneaster acuminatus, Rosa omeiensis, Potentilla fruticosa, Sibiraea angustata, Carex lanceolata, Anaphalis margaritacea, Ranunculus japonicus, Bupleurum tenue, Polygonum phaerostachyum have higher niche width and stronger adaptability, hence becoming a dominant populations in this region; while Spiraea henryi, Quercus cocciferoides, Sageretia pycnophylla, Salsola collina, Calamagrostis purpurea, Artemisia igniaria, Tragus berteronianus，Salix cheilophilla, Rubus mesogaeus, Hippophae rhamnoides, Artemisia roxlourghiana, Polygonum viviparum,Fragaria gracilis have lower niche. Not all populations with lower niche are dominant populations. This peculiarity depends on the range and evenness of distribution of the certain population. Populations with higher niche breadth always have higher niche overlap, but not all populations with smaller niche breadth have lower niche overlap.
(3) Among the plant communities in arid valley, with an increase of altitude, both the a diversity and β diversity of herbs layer and shrubs layer display a "high-low-high" trend. The α diversity and β diversity of herbs layer are higher than those of shrubs layer. Both the herbs layer and shrubs layer have higher a diversity and β diversity at an altitude range of 1400～1600m and 2000～2200m, which is a potent proof that a low biodiversity occurs in regions of a medium altitude, while a high one exists in regions of high and low altitudes.The diversity varies in different formations: Form. Pertya sinensis, Jasminum humile .Form. Berberis wilsonae, Lonicera japonica, Form. Spiraea Spp.have higher a diversity and β diversity. Form. Leptodermis purdomi, Form. Caryopteris spp. and Form. Bauhinia faberi , Sorphora vrcifolia have lower α diversity and β diversity; both a diversity and β diversity in different slope directions and shapes: shaded >slope half-shaded> slope sunny slope, concave slope> straight> convex slope; in different positions: lower slope> upper slope> middle slope . The a diversity of shrubs layer and herbs layer alike in different slope positions: upper slope> lower slope> middle slope, while the β diversity, lower slope> upper slope> middle slope.
Among the plant communities in origin region of the Minjiang River, with an increase of altitude, diversity of both the herbs layer and shrubs layer display symmetrical changing trends that they are almost the same at altitude ranges of 2700-3300m and 3300-3900m. The altitude ranges of 2900-3100m and 3500-3700m see a higher diversity, while those of 2700-2900m, 3100-3400, 3700-3900m see a lower one. The maximum diversity of shrubs layer and herbs layer occurs at the altitudes of 3000m and 3600m respectively. The diversity in different vegetation types follows the trend: forests formations >shrubs formations >meadow formations. The diversity in herb layer of Form. Salix spp., Form. Sibiraea angustata, Form. Spiraea spp.and Form. Caragana spp.are at high level, while Form. Hippophae rhamnoides and Form. Berberis amurensis are at lower level.In meadow formations, the diversity of Form. Imperata cylindrica and Form. Carex spp.are higher than that of Form. Sub-alpine and Form. Kobresia spp. As for shrub layer , Form. Caragana spp. has the highest diversity, Form. Hippophae rhamnoides has the lowest diversity. Diversity in herb layer is higher than that of shrub layer.
Under a great influence of altitude gradient, species compositions in plant communities in the origin area of Minjiang River show significant differences and the β diversity is more easily affected by the change of herbs layer. Along the altitude gradient, there are higher species turnover rates at the altitude ranges of 2900-3100m and 3500-3700m, and the community has a higher heterogeneity. While at the altitude ranges of 2700-2900m, 3200-3300m, 3800-3900m,, the community has a lower heterogeneity and species turnover rate. The change of β diversity in shrubs layer is comparatively stable along the altitude gradient, with its climax occurring at an altitude of 2900m and the minimum at 3300m.
A significant quadratic curve relationship between the β diversity and soil moisture and nutrient shows that the soil nutrient and water content of the plant community with the highest species turnover rate are at a medium level, instead of a high level of nutrient and water content. (4) The very significant positive relationship exist between the aboveground biomass of shrub communities and the altitude, available K, soil moisture, organic matter, total P, hydrolyazble N, total N, available N in the upper reaches of the Minjiang River, while the very significant negative relationship exist between slope gradient and pH value, among which altitude, available K and soil moisture play a leading role in affecting the aboveground shrub biomass. The aboveground biomass of shrub communities increases with the raise of species diversity and species richness and has a very positive linear relationship with species diversity and species richness.
(5) Huffman coding method of communication theory is used to study the plant complexity and the relationship with environmental factors and reveal that between plant community complexity and diversity , evenness. Both total and structural community complexity vary with degree, shape and aspect of slope, with an order of upper slope>middle slope>lower slope, shaded slope> half-shaded slope>sunny slope, and concave slope>straight slope>convex slope. Form. Pertyasinensis,Jasminumhumile have the highest total complexity, and Form Leptodermispurdomi and Form Ceratoidesarborescens communities have the lowest total complexity, while Form Quercnscocciferoides and Form Onosma farrerii communities have the highest structural complexity.
The plant formations in origin area have a high total complexity at the altitude ranges of 2900-3100m and 3500-3700m. The total complexity is as the following: Forest plant formations>shrub formations>meadow forations. Form. Picea asperata is ranked as the highest in total complexity, Form. Kobresia spp have a lowest one . In shrub formations, Form. Caragana spp.have the highest total complexity, Form. Hippophae rhamnoides have the lowest one. In meadow formations, the total complexity of Form. Carex spp. and Form. Imperata cylindrica are at high level, Form. Kobresia spp.and Form. Sub-alpine are at lower level. Similar to the correlation of β diversity with soil nutrient, The total community complexity had significant quadratic correlations with soil water content and nutrient. However, the soil factors that influence total community complexity in origin area and arid valley are different, therefore, it does not grow in a simple linear way with the increase of soil nutrient and water content.
Total community complexity is positively correlated with community diversity, evenness and species richness, while structural complexity is negatively correlated with community evenness. As two components of total community complexity, structural complexity is more sensitive than diversity to the change of species in the community, which is not only related to community evenness, but also to community richness. In the communities with a high evenness, the difference between total complexity and diversity is not quite obvious, and structural complexity is lower. However, the exceptional case exists in which a low evenness cannot push the low structural complexity of the communities with a low species richness to its climax. The relative contribution of structural complexity and diversity to total complexity would be different for different study area or ecosystems.
(6) The stability of typical plant formations in the upper reaches of Minjiang River is analysed by using AHP methods, combining quantitative character indices reflecting vegetation stability, such as vegetation coverage, diversity, complexity and degree of succession with external environmental factors such as climate, topography and soil character, hence supplementing the insufficiency that previous researches on vegetation stability assessment take the only vegetation character into account. The result indicates: the plant formations of origin region, Form. Picea asperata, Form. Salix spp., Form. Sibiraea angustata, Form. Abies faxoniana, Form. Spiraea spp. Have higher stability, while Hippophae rhamnoides and Form. Berberis amurensis have lower one. Form. Imperata cylindrical and Form. Imperata cylindrica have a higher stability, Form. Kobresia spp.and Form. Sub-alpine have lower one. In the plant formations of the arid valley region, Form. Spiraea spp,Form. Daphne spp., Form. Onosma farrerii, Form. Bauhinia faberi, Sorphora vrcifolia, Form. Caryopteris spp., Form. Quercus cocciferoides have a higher stability.
Under a constant external disturbance, regressive succession of the vegetation occurs in the upper reaches of the Minjiang River. Regressive succession model of the vegetation in the arid valley is: Form. Quercus cocciferoides, Form. Spiraea Spp.→Form. Pertya sinensis, Jasminum humile, Form. Onosma farrerii→Form. Ceratoides arborescens, Form. Caryopteris spp. Form. Bauhinia faberi , Sorphora vrcifolia; the regressive succession model of the origin region is Form. Abies faxoniana、 Form. Picea asperata→Form. Sibiraea angustata→Form. Caragana spp., Form. Berberis amurensis→Form. Imperata cylindrical, Form. Kobresia spp.
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