摘要
本研究主要应用群落数量统计学方法对永安天宝岩国家级自然保护区天然柳杉群落学特征进行研究,内容包括自然保护区天然柳杉群落的物种组成及植物区系特征、群落主要优势种的数量特征、物种多样性特征、群落主要物种的空间分布格局特征、群落乔木层径级与高度级结构分布特征、乔木层主要种群生态位特征、种间关联度分析、种内竞争和种间竞争对比研究以及柳杉种群的生命过程等方面进行研究,主要研究结果如下:
(1)群落物种组成及区系特征方面,天然柳杉群落物种共112种(含变种),隶属53科88属,其中木本植物58种,草本植物54种。群落乔木层有常绿类15科27属33种,占乔木层总种数量的82.5%;落叶类6科6属7种占乔木层的17.5%,灌木层(下木层)常绿类26科34属44种,占下木层总种数量的77.8%,落叶类10科12属14种,占下木层总种数量的22.2%。群落主要物种科的分布类型共有11类,包含科数最多的是泛热带分布,其次为世界分布;属的地理分布包含17种分布类型,其中世界分布数量最多,其次为泛热带分布。
(2)群落乔木层主要物种数量特征研究表明,乔木层重要值较大的物种主要为柳杉( Cryptomeria fortunei )、马尾松( Pinus massoniana Lamb )、柃木(Eurya japonica Thunb)、长苞铁杉(Tsuga longibracteata)、栎树(Quercus palustris)等,其中柳杉的重要值最大,且与其他物种相差悬殊,在群落中是明显的优势树种。灌木层重要值较大的物种为柃木(Eurya japonica Thunb)、钝齿冬青(Ilex crenata Thunb)、鸭脚茶(Bredia sinensis (Diels))、杜鹃(Rhododendron moulmainense)、柳杉(Cryptomeria fortunei )等且许多为乔木层更新个体,优势种不明显;草本层最优势种群为铁芒萁( Dicranopteris linearis )、鳞毛蕨( Dryopteris podophylla )和芒草(Miscanthus sinensis),其他物种重要值分布较为均匀。
(3)采用6个多样性指数对不同海拔柳杉群落物种多样性进行研究并比较,结果表明,高海拔柳杉林群落物种多样性为灌木层>草本层>乔木层,中海拔和低海拔表现为草本层>乔木层>灌木层;不同海拔物种多样性对比发现,不同海拔上看,随着海拔的变化,乔木层:高海拔>中海拔>低海拔,灌木层:高海拔>低海拔>中海拔,草本层物种多样性情况受海拔影响差别不大,没有明显差异。高海拔具有良好的水热条件,而中低海拔地区的人为破坏使得该海拔柳杉群落物种的多样性显著降低。
(4)径级与高度级结构分布研究结果表明,从乔木层整体径级分布上看群落表现为典型正金字塔形状的潜力较大的增长型群落。中海拔和低海拔径级分布趋势表现较为一致,都为先增长而后持续下降的单峰分布。高、中海拔乔木层各径级分享度分布状况较为相似,都表现出先下降后上升,而后持续下降趋于零的趋势。高度级方面,分布状况基本表现为随着高度级级数的增大个体数量先增后减而后再增再减的双峰结构。中海拔和低海拔高度级分布趋势极为相似,且3个海拔群落乔木层高度级分享度曲线与其高度级数量分布曲线极为一致。
(5)群落乔木层主要物种空间分布格局分析结果表明:采取不同的空间分布指标种类和尺度进行分析,得出的结果有所差异,但影响较小。总体上看,采用多种方法分析,天然柳杉群落的柳杉、马尾松和柃木种群都为聚集分布。通过不同海拔柳杉种群空间分布格局对比发现,在高海拔和中海拔属于聚集分布,低海拔属于均匀分布。用Iwao方程进行回归,建立回归方程:M*=0.660+0.972 x (R=0.962)也表明柳杉种群符合聚集分布。
(6)应用Hegyi单木竞争指数模型对比分析了不同干扰强度下天然柳杉种群各种竞争指数的变化,并建立天然柳杉种群胸径与竞争指数之间的关系模型。结果表明:随着干扰强度的增大,天然柳杉林种群总竞争指数和种内竞争指数均呈明显递减趋势,即CI(1)>CI(2)>CI(3)、CIintra(1)>CIintra(2)>CIintra3),而种间竞争指数却有明显增长。竞争强度和对象木胸径的关系服从幂函数关系( CI = 161.1639 ?D1.1126),该预测模型能很好地预测柳杉种群所受的竞争强度。
(7)运用生态位宽度、生态位重叠值以及生态位相似比例等指标对天然柳杉群落中12个主要种群的生态位相关特征进行测定与分析。研究结果表明:天宝岩保护区天然柳杉群落的主要种群大多数为广生态位,狭生态位型物种相对较少些,且生态位宽度最大的4种物种为柳杉(Cryptromeria fortunei)>闽粤栲(Castanopsis fargesii Franch)>细叶青冈(Cyclobalanopsis gracilis)>甜槠(Castanopsis eyrei),群落物种间的生态位重叠度与相似度也相对较大,说明此群落中存在较大种间竞争。从柳杉种群来看,柳杉的生态位宽度最大,与木荷的生态位重叠和生态位相似度最大。
(8)在相邻格子法抽样调查的基础上,根据2×2联列表数据,采用方差分析、卡方检验、W检验、联结系数(AC)和Jaccard指数(JI)等指标,分析了天然柳杉群落乔木层15个优势种群的总体联结性、种对间联结显著性和关联系数。结果表明,整体上看,群落中15个优势种群间的总体联结性呈极显著正相关。成对物种种间联结性研究发现,除少数种对外,绝大多数种对间的联结性不显著相关。在联结性显著的种对中,11-12(细叶青冈-甜槠)和10-14(绒毛润楠-青冈)等6个种对种内存在显著正联结,2-4(马尾松-长苞铁杉)和7-9(黄山松-厚皮香)2个种对种内存在显著负联结。其他种对未达到显著水平,种对间的相互独立性较强。
(9)采用空间代替时间的方法,以种群生命表及生存分析理论为基础,编制种群静态生命表,并分析生存率、积累死亡率和死亡密度等函数曲线变化,进而分析种群数量的动态变化。结果表明,柳杉种群存活曲线经统计检验属于DeeveyⅡ型曲线,认为种群具有前期薄弱、中期稳定、后期衰退的特点。柳杉种群的存活率单调下降,累计死亡率单调上升,存活率与累计死亡率的变化率趋势相同,皆是前期高于后期。死亡率和亏损率变化基本一致,均出现2个高峰,都在第3龄级和第5龄级达到最高。生存分析中引入生命表中的几个函数能较好地说明柳杉种群的结构和动态变化。
Based on the methods of community Mathematics &Applied Statistics,the natural Cryptomeria fortunei community characteristics was researched in Yong’an Tian baoyan national nature reserve,including the species composition and the characters of flora,the quantity characteristics of dominant species,species diversity features,the spatial distribution patterns of the main species,the distribution of the high scales and diameter scales and various kinds of model fittings, main species’niche characteristics and inter-specific association in tree layer,intra- and inter- specific competition comparison,and static life table research.The main results were as following:
(1)Analyzed from the quantity characteristics of the main species in tree layer,the results showed that the main species with a relatively larger importance value in the tree layer were Cryptomeria fortunei , Pinus massoniana Lamb , Eurya japonica Thunb ,Tsuga longibracteata W.C.Cheng,Quercus palustris,etc.The species with the largest importance value was Cryptomeria fortunei which reached 97.087 percent and much more than other plants.The main species with a larger importance value in the shrub layer were Eurya japonica Thunb , Ilex crenata Thunb , Bredia sinensis ( Diels ) ,Rhododendron moulmainense Hook,Cryptomeria fortunei,etc and the dominant species were not obviously;the most dominant species in herb layer were Dicranopteris linearis, Dryopteris podophylla and Miscanthus sinensis while the rest of all distributed relatively evenly.
(2)Analyzed the different species compositions of the studied community,the results showed that the natural Cryptomeria fortunei community had 112 kinds of species in all,which belonging to 53 families and 88 genus,including 58 kinds of woody plants and 54 kinds of herbaceous plants.There were 33 kinds of species(82.5%)which belong to 15 families and 27 genus in the tree layer were evergreen trees;7 species(17.5%) were deciduous trees belonging to 6 families and 6 genus;44 species(77.8%)belonging to 26 families and 34 genus in the shrub layer were evergreen plants while 14 kinds(22.2%) were deciduous plants belonging to 10 families and 12 genus.The analysis of the flora features showed that there were 11 kinds geographical distribution patterns of the families in natural Cryptomeria fortunei community,The distribution types which took up the most families were pan-tropical distribution,and world distribution was in the second place.According to the geographical distribution types of genus,the community had 17 tpyes, the most dominant type of which was world distribution and pan-tropical distribution closely followed.
(3)6 species diversity indexes were used to estimate Cryptomeria fortunei community species diversity in different elevation,and the results showed that the order of diversity value in high elevation was shrub layer>herb layer>tree layer,and herb layer>tree layer>shrub layer in mid- and low-elevation.With the change of the elevation,the diversity order in tree layer was:high-elevation>mid elevation>low-elevation,in shrub layer was:high-elevation > low-elevation > mid elevation,and there was no significant difference in herb layer.It concluded that at high-elevation there were better thermal and humidity condition , while the conspicuous decline of species richness at mid-elevation and low-elevation due to serious human disturbance.
(4)The research of DBH and high scale distribution showed that the DBH distribution was a classic pyramidal shapes which had a great potential in communities development.The curves of the mid- and low- elevation increased first and then decreased immediately.The share rate of every DBH scale in high- and mid- elevation was very similar which decreased first and then increased and last decreased closely to zero.The high scale distribution shaps in mid- and low elevation were very similar,and the share rate curves of high scales in tree layer were very similar with the quantity distribution of high scales in the three elevation.
(5)The research of main species spatial distribution patterns in tree layer of the community showed that It would have a little affection to the distribution patterns if different indexes and scales were chosed to be used.The comparison of the spatial distribution patterns of the main species between the natural Cryptomeria fortunei community and planted Cryptomeria fortunei community showed that,in general, Cryptomeria fortunei, Pinus massoniana,and Eurya japonica Thunb all followed aggregated distribution pattern under different analysis methods,which was very consistent with the research formerly.From the comparison of the spatial distribution patterns of Cryptomeria fortunei population in different elevation,the results showed that it was aggregated distribution pattern in high and mid- elevation,while even distribution pattern in low elevation.The function mode M*=0.660+0.972 x(R=0.962)was found using the Iwao equation which also indicated that the population followed the aggregated distribution pattern.
(6)The Hegyi’s signal trees competition index model was used to research the changes of the competition index of Cryptromeria fortunei populations under disturbance of different intensities,and the relationship between competition index and DBH has been studied,and the relationship model was established.The results showed that with the increase of the disturbance intensities,the general and the intra-specific competition intensities of the investigated three sample plots decreased obviously.The order of the general and intra-pecific competition intensity was CI(1)> CI(2)> CI(3) and CIintra(1)> CIintra(2)> CIintra(3),although the inter-pecific competition intensity didn’t increase obviously.The relationship between competition intensity and DBH closely follow the power function model: CI = 161.1639 ?D1.1126,and the model can be well used to predict the competition intensity of Cryptromeria fortunei populations in this area.
(7)Niche characteristics of 12 major populations of natural Cryptromeria fortunei forest in Tianbaoyan National Nature Reserve were analyzed by quantitative method. The niche characteristics include niche breadth , niche similarity proportion and niche overlap. The results showed that most of the populations investigated in the natural Cryptromeria fortunei forest had wide niche, and the populations with narrow niche relatively less.The first four populations with the widest niche were Cryptromeria fortunei,>Castanopsis fargesii Franch>Cyclobalanopsis gracilis>Castanopsis eyrei . The niche overlap and similarity proportion of the 12 dominant species were comparably big,which indicated that the intra species competition was very serious in the community investigated.From the niche characteristics of Cryptromeria fortunei population,we can see that Cryptromeria fortunei had the widest niche breadth,and also had the biggest niche overlap and niche similarity proportion with Schima superba Gardn et Champ.
(8)A series of methods including the analysis of variance,chi-square test,W test,association coefficient(AC)and Jaccard coefficient and so on were calculated based upon a 2×2 contingency table to determine the overall association,the statistical significance and the association coefficient of the each species pair of 15 main species in arborous layer of Cryptromeria fortunei communities in Yong’an Tianbaoyan national nature reserve,Fujian province.The results showed that the overall association was significantly positive correlation among fifteen main populations and the inter-specific association was not obviously correlated among most of species pairs except only a few of them.6 Species pairs like 11-12( Cyclobalanopsis gracilis-Castanopsis eyrei ) and 10-14 ( Machilus velutina- Cyclobalanopsis glauca)were significantly positive correlation.2 Species pairs like 2-4( Pinus massoniana- Tsuga longibracteata ) and 7-9 ( Pinus taiwanensis Hayata- Ternstroemia gymnanthera )were significantly negative correlation.But most of them didn't reach a significant level,and furthermore the mutual independence of them was very strong between each species pair.
(9)An ecological investigation was made on the natural Cryptromeria fortunei population in Tianbaoyan national nature reserve of Fujian Province by using checkerboard frame method and based on the theory of population life table and survival analysis,the life table of the population was established by the method of spatial series substituting for time series.The population dynamic variation was analyzed on survival rate,accumulated mortality rate and mortality density curves.The results showed that in general,the survival curve was DeeveyⅡpattern,and the population grew weakly in earlier period and stably in middle age period but declined in old age period.The survival rate of the population decreased monotonically,whereas the accumulative mortality rate was in adverse.The mortality rate curve and the survival rate curve had the similar variation rate both of which was bigger in the earlier period than that in the later period.The variation trend between mortality rate and loss rate was basically similar both of which existed 2 peaks which was NO.3 and NO.5.Several functions of life table were used well in survival analysis to explain clearly the structure and dynamic variation of Cryptromeria fortunei population.
引文
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