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植物多样性作为群落的重要特征,是生态系统结构、功能及其稳定性的基础[1-2]。随着生态学研究的不断深入,植物多样性格局及形成机制成为生态学科的热点问题之一[3-4],是理解区域生物多样性维持机制的重要内容,同时对生物多样性的保护和管理有着重要意义[5-8]。研究植物多样性,有助于认识群落的组成、变化和发展趋势,同时也可反映群落及其环境的保护状况[9-10]。胡桃楸Juglans mandshurica是北京唯一的国家保护高大乔木,在生物多样性相对匮乏的北京地区具有特殊的意义。松山国家级自然保护区是主要的胡桃楸林分布区,受到生态旅游及麻核桃Juglans hopeiensis开发等人为活动的影响,生境严重破碎化,林下群落结构遭到破坏,植物多样性受到严重威胁[11]。胡桃楸林不同演替阶段群落结构存在差异[9-10]。本研究对不同林龄阶段的胡桃楸林下植被进行调查,目的在于了解松山国家级自然保护区胡桃楸林下群落的植物多样性及变化规律,找到群落与环境因子和人为干扰等因素的互作机制,为松山国家级自然保护区多样性保护与恢复工作提供理论依据。
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调查记录胡桃楸林下植物43科78种,以菊科Asteraceae,蔷薇科Rosaceae,豆科Leguminosae,毛茛科Ranunculaceae和百合科Liliaceae等为主,其中木本植物18科26种,草本植物25科52种。同时还对胡桃楸林内灌木层和草本层植物的重要值进行了统计(表 1与表 2)。
表 1 灌木层主要组成物种及其重要值
Table 1. Main species composition and important values of the shrub layer
物种 林龄 幼龄林 中龄林 近熟林 成熟林 山楂叶悬钩子Rubus crataegifolius 0.247 0.104 0.065 0.095 荆条Vitex negundo var. heterophyila 0.221 0.217 山杏Armeniaca sibirica 0.184 0.147 暴马丁香Syringa reticulata var. mandshurica 0.256 0.267 红花锦鸡儿Caragana rosea 0.121 0.187 0.325 0.357 小叶鼠李Rhamnus parvif0lia 0.263 0.35 0.278 0.294 毛叶丁香Syringa pubescens 0.225 0.328 蚂蚱腿子Myripnois dioica 0.143 0.155 土庄绣线菊Spiraea pubescens 0.268 0.175 大果愉Ulmus macrocarpa 0.354 0.163 0.319 细叶小棄Berberis poiretii 0.262 三桠绣线菊Spiraea trilobata 0.134 小花搜疏Deutzia parviflora 0.229 0.367 雀儿舌头Lepopus chinensis 0.13 0.104 大花搜疏DeutEia grandiflora 0.183 小叶朴Celtis bungeana 0.287 0.147 卫矛Euonymus alatus 0.194 0.208 蘀叶蛇葡萄Ampelopsis humulifolia 0.058 0.092 0.109 0.064 表 2 草本层主要组成物种及其重要值
Table 2. Main species composition and important values of the herbage layer
物种 林龄 幼龄林 中龄林 近熟林 成熟林 披针叶薹草Carex lancifolia 0.102 0.158 0.094 大油芒Spodiopogon sibiricus 0.227 0.189 0.082 甲鸟妬草Commelina communis 0.065 0.055 0.074 0.067 东亚唐松草Thalictrum minus 0.087 0.091 0.107 紫菀Aster tataricus 0.074 0.053 0.082 车前Plantago asiatica 0.027 0.045 巴天酸模Rumex patientia 0.043 0.062 灰菜Chenopodium album 0.043 0.049 龙芽草Agrimonia pilosa 0.057 0.037 草乌Aconitum carmichaeli 0.097 0.086 萱草Hemerocallis fulva 0.075 0.055 益母草Leonurus artemisia 0.077 0.082 秋苦荬菜Paraxeris pinnatipartita 0.084 0.067 铁杆嵩Tripolium vulgare 0.063 0.089 石竹Dianthus chinensis 0.087 0.052 蝙蝠葛Menispermum dauricum 0.092 0.068 垂果南芥Arabis pendula 0.047 0.085 0.074 北马兜铃Aristolochia contorta 0.056 0.054 风毛菊Saussurea japonica 0.021 穿龙暑蓣Dioscorea nipponica 0.038 莓叶委陵菜Potentilla centigrana 0.044 黄精Polygonatum sibiricum 0.051 野青茅Deyeuxia arundinacea 0.095 宽叶薹草Carex siderosticta 0.095 北重楼Paris verticillata 0.087 鼠掌老鹤草Geranium sibiricum 0.028 0.082 毛哀Ranunculus japonicus 0.037 0.172 短尾铁线莲Clematis brevicaudata 0.054 0.052 蓝萼香茶菜Rabdosia japonica 0.107 0.118 大叶铁线莲Clematis heracleifolia 0.101 0.069 白屈菜Chelidonium majus 0.057 0.104 灌木层中,葎叶蛇葡萄Ampelopsis humulifolia,小叶鼠李Rhamnus parvifolia,红花锦鸡儿Caragana rosea和山楂叶悬钩子Rubus crataegifolius在不同林龄林下都有分布。红花锦鸡儿与小叶鼠李在4种林龄林下都有较大优势,而山楂叶悬钩子和葎叶蛇葡萄则存在差异。荆条Vitex negundo var. heterophylla和山杏Armeniaca sibirica分布在幼龄林及中龄林中,毛叶丁香Syringa pubescens,小花溲疏Deutzia parviflora,暴马丁香Syringa reticulata var. mandshurica和土庄绣线菊Spiraea pubescens等在近熟林与成熟林中分布,细叶小檗Berberis poiretii和大花溲疏Deutzia grandiflora仅在成熟林中可见。整体而言,幼龄林及中龄林的灌木种类较少,随着林龄的增加,灌木层植物种类逐渐增加,存在明显的变化趋势。保护区内胡桃楸林下灌木群落以小花溲疏、毛叶丁香、小叶鼠李、红花锦鸡儿等为主要优势种。
草本层中鸭跖草Commelina communis在不同林龄下均有分布,但重要值存在差异。不同林龄的胡桃楸林下草本群落组成差异显著,随着林龄增加,大油芒Spodiopogon sibiricus,披针叶薹草Carex lancifolia,蝙蝠葛Menispermum dauricum等逐渐被北重楼Paris verticillata,鼠掌老鹳草Geranium sibiricum,蓝萼香茶菜Rabdosia japonica等取代,形成较稳定的顶级群落。优势种上,幼龄林和中龄林主要是披针叶薹草,大油芒,草乌Aconitum carmichaeli和龙芽草Agrimonia pilosa等,近熟林和成熟林则以蓝萼香茶菜和白屈菜Chelidonium majus等为主。整体而言,随着林龄的增加,草本层植物种类减少的趋势明显。
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通过单因素方差分析,对不同林龄胡桃楸林下灌木层物种多样性进行差异显著性分析。结果显示:不同演替阶段的灌木层Simpson指数存在显著差异(P<0.05),随着林龄增加,灌木层物种变化明显,指数值不断增加(图 2和表 3);而Shannon-Wiener多样性指数和均匀度指数分别呈极显著(P<0.001) 和显著(P<0.05) 的差异,随着林龄的增加而逐渐增加。这表明,随着演替阶段的深入,灌木的物种多样性表现出不断递增的规律。
图 2 不同林龄胡桃楸林下灌木层物种多样性指数
Figure 2. Species diversity index of shrub layers under forest at different ages of manchurian walnut plantations
表 3 林龄对不同层物种多样性的影响
Table 3. One-way ANOVA for shrub and herbage diversity index aifect by forest age
指数 层次 自由度 平方和 均方 F P R 草本层 3 73.233 24.408 13.708 <0.001 灌木层 3 18.633 6.211 26.617 <0.001 D 草本层 3 0.270 0.090 4.084 0.047 灌木层 3 0.350 0.120 5.133 0.009 H 草本层 3 0.772 0.257 12.832 <0.001 灌木层 3 1.528 0.509 16.236 0.002 J 草本层 3 0.390 0.130 5.263 0.008 灌木层 3 0.270 0.110 5.370 0.004 M 草本层 3 1.280 0.427 15.472 <0.001 灌木层 3 1.070 0.296 9.823 0.007 -
单因素方差分析显示,不同林龄下草本层的Simpson多样性指数存在显著差异(P<0.05),并随林龄的增加,呈现逐渐降低的规律(图 3与表 3);Shannon-Wiener多样性指数差异更为显著(P<0.001),随着林龄增加,多样性指数下降的趋势更加明显;不同演替阶段的草本层均匀度指数也存在显著差异(P<0.001),但变化趋势是上升的。这表明,随着演替阶段的深入,草本植物有逐渐退出生态系统的趋势,最后形成一个稳定的顶级群落,这种变化规律与Bohanek等人提出的观点是一致的[17]。
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通过单因素方差分析,灌木层的物种丰富度差异极显著(P<0.001),与幼龄林相比,中龄林灌木层物种丰富度增加了20%以上,近熟林比中龄林又增加了近43%(图 4),最后在成熟林阶段达到了最高值,但与近熟林相比,成熟林的物种丰富度增加比例不到9%。这表明:从胡桃楸群落演替早期到达到稳定状态的期间,物种丰富度先快速增加,到中期增速达到最快,而到了后期阶段,随着群落逐渐稳定,植物物种的替换速率也随之下降。草本层的变化规律与灌木层呈现完全相反的趋势。随着胡桃楸林龄的增加,林下草本层植物的种类逐渐减少(图 4)。不同林龄胡桃楸林下草本层物种丰富度存在极显著差异(P<0.001),在演替初期,大量的草本植物出现,形成早期的先锋群落,但随着林龄增加,群落中草本植物不断退出群落,尽管有部分新的植物进入,但在整体上,最后的稳定状态的顶级群落与早期相比,草本层物种丰富度下降非常明显。
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从表 4物种多样性半变异函数模型及方差分析结果可知,松山胡桃楸林4种多样性指数中,均匀度指数和Simpson多样性指数为球面模型,Shannon-Wiener多样性指数和丰富度指数则更符合指数模型,半变异函数模型能够很好地反映物种多样性指数的空间结构特性。在分析模型中块金值表征由于取样误差和小于取样尺度上随机引起的空间异质性,基台值越大,总的空间异质性越高;块金与基台的比值表示由系统变量引起的空间相关性,比值小于0.25表示空间相关性强烈,0.25~0.75表示中等的空间相关性,大于0.75表示空间相关性较弱[18];4种多样性指数比值都为0.25~0.75,因此研究区的物种多样性为中等的空间相关性。
表 4 物种多样性半变异函数模型及其相关参数
Table 4. Parameters and test values of theoretical variogram models for species diversity indexes
指数 模型(C0) 块金 基台 块金/基台[C0/(C0+C1)] 变程 决定系数 D 球面模型0.023 0.072 0.319 13.2 0.67 H 指数模型0.017 0.047 0.361 7.6 0.64 J 球面模型0.035 0.084 0.416 11.2 0.76 M 指数模型0.027 0.067 0.403 8.4 0.65 同时也说明:在大于研究尺度,小于变程的范围内结构性因子所占比例为0.584~0.681,随机性因子所占比例为0.319~0.416,说明尽管结构性因子仍然占主导地位,但诸如人为砍伐、旅游开发等人为活动引起的空间异质性起着非常重要的作用,人为活动的干扰对胡桃楸植物多样性有着显著影响。
Undergrowth diversity at different ages of Juglans mandshurica forests in the Songshan National Nature Reserve, Beijing
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摘要: 以北京松山国家级自然保护区胡桃楸Juglans mandshurica林为研究对象,采用典型抽样法研究林下植物物种组成与多样性。结果表明:胡桃楸林灌木层植物种类随林龄增加而逐渐增加,草本层植物种类则逐渐减少,存在明显的时空动态;幼龄林和中龄林草本层以披针叶薹草Carex lancifolia,大油芒Spodiopogon sibiricus,草乌Aconitum carmichaeli和龙芽草Agrimonia pilosa等为优势种,近熟林和成熟林则以蓝萼香茶菜Rabdosia japonica和白屈菜Chelidonium majus等为优势种;幼龄林和中龄林灌木层以小叶鼠李Rhamnus parvifolia和荆条Vitex negundo var. heterophylla等为优势种,近熟林和成熟林则以红花锦鸡儿Caragana rosea,小花溲疏Deutzia parviflora,毛叶丁香Syringa pubescens和暴马丁香Syringa reticulata var. mandshurica等为优势种。不同林龄胡桃楸林下灌木层与草本层4个物种多样性指数存在显著差异(P < 0.05):灌木层物种多样性指数中,随着林龄的增加,Simpson指数从0.598 0增加到0.787 9,Shannon-Wiener指数从1.183 9增加到1.835 6,Pielou均匀度指数从0.682 0增加到0.839 0,Margalef丰富度指数从0.979 0增加到1.614 9,均呈现连续增加的趋势;草本层均匀度指数随着林龄的增加从0.884 6增加到0.987 2,表现为连续递增趋势,其余指数中,Simpson指数从0.843 6减少到0.759 4,Shannon-Wiener指数从2.120 9减少到1.673 9,丰富度指数从1.951 7减少到1.347 1,均随着林龄的增加呈持续递减的趋势。不同林龄胡桃楸林下灌木层和草本层物种丰富度均存在极显著差异(P < 0.001),由幼龄林到成熟林,灌木层物种丰富度从3.385 7逐渐增加到6.340 0,在成熟林阶段达到最高值;草本层的变化规律与灌木层呈现完全相反的趋势,随着林龄的增加,林下草本层植物的种类逐渐减少。用半变异函数模型模拟,均匀度指数和Simpson指数为球面模型,Shannon-Wiener指数和丰富度指数则更符合指数模型,研究区的物种多样性为中等的空间相关性,结构性因子所占比例为0.584~0.681,随机性因子所占比例为0.319~0.416。建议该区科学合理地规划景区建设及道路开发,并对当地居民的种子采集等活动进行管理。Abstract: To effectively protect the Juglans mandshurica forest and prevent it's further habitat deterioration in Beijing, the study attempted to clarify the distribution characteristics of the undergrowth plant diversity. Shrub and herb plant species composition and diversity were researched at different forest ages of Juglans mandshurica forest in the Songshan National Nature Reserve by the typical sampling method that twenty-four 20 m×20 m plots were set up in different typical stands, and one 5 m×5 m shrub sample and one 1 m×1 m herb sample were made at each quadrangle and center of each arbor sample. Diversity analysis included richness index (M), Shannon-Wiener diversity index (H), and Simpson diversity index (D). Results showed that as forestsincreased in age, shrub species and quantities increased, and herb species and quantities decreased. The most important herb plant species in the young and middle-aged forests were Carex lancifolia, Spodiopogon sibiricus, Aconitum carmichaelii, and Agrimonia pilosa; whereas, in the nearly mature forests and mature forests Rabdosia japonica and Chelidonium majus were predominant. Shrubs were dominated in the young forests and middle-age forests by Rhamnus parvifolia and Vitex negundo var. heterophylla but in the nearly mature forests and mature forests by Caragana rosea, Deutzia parviflora, Syringa pubescens, and Syringa reticulata var. mandshurica. There were significant differences in species diversity indices between shrub layer and herb layer in Juglans mandshurica forest at different ages(P < 0.05). In the shrub layer, with the increase of age, Simpson index increased from 0.598 0 to 0.787 9, the Shannon-Wiener index is 1.183 9 to 1.835 6, the Pielou evenness index is 0.682 0 to 0.839 0, and the Margalef richness index is 0.979 0 to 1.614 9, showing a continuous increase trend; In the herb layer, the Pielou evenness index increased from 0.884 6 to 0.987 2, and the Simpson index decreased from 0.843 6 to 0.759 4, and the Shannon-Wiener index decreased from is 2.120 9 to 1.673 9, while the Margalef richness index is 1.951 7 to 1.347 1, and decreased with the increase of stand age. Shrub layer and herb layer of Juglans mandshurica forest under different forest ages was significantly different(P < 0.001). The species richness of shrub layer increased from 3.385 7 to 6.340 0 from young forest to mature forest, and reached the highest value in mature forest stage. The change of herb layer was opposite to that of shrub layer. With the increase of age, the species of the herb layer in the undergrowth layer decreased gradually. Using the semi-variogram model, Pielou evenness index and Simpson index were spherical model, Shannon-Wiener index and Margalef richness index were more consistent with it. The species diversity of the study area was moderate spatial correlation, the proportion of structural factors is 0.584-0.681, and the proportion of random factors is 0.319-0.416. For spatial variation of species diversity, human factors were dominant. Therefore, reasonable planning and development activities at the local level should be undertaken with local residents conducting seed collection and other management activities.
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Key words:
- forest ecology /
- Juglans mandshurica /
- shrub /
- herb /
- species diversity /
- Songshan in Beijing
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表 1 灌木层主要组成物种及其重要值
Table 1. Main species composition and important values of the shrub layer
物种 林龄 幼龄林 中龄林 近熟林 成熟林 山楂叶悬钩子Rubus crataegifolius 0.247 0.104 0.065 0.095 荆条Vitex negundo var. heterophyila 0.221 0.217 山杏Armeniaca sibirica 0.184 0.147 暴马丁香Syringa reticulata var. mandshurica 0.256 0.267 红花锦鸡儿Caragana rosea 0.121 0.187 0.325 0.357 小叶鼠李Rhamnus parvif0lia 0.263 0.35 0.278 0.294 毛叶丁香Syringa pubescens 0.225 0.328 蚂蚱腿子Myripnois dioica 0.143 0.155 土庄绣线菊Spiraea pubescens 0.268 0.175 大果愉Ulmus macrocarpa 0.354 0.163 0.319 细叶小棄Berberis poiretii 0.262 三桠绣线菊Spiraea trilobata 0.134 小花搜疏Deutzia parviflora 0.229 0.367 雀儿舌头Lepopus chinensis 0.13 0.104 大花搜疏DeutEia grandiflora 0.183 小叶朴Celtis bungeana 0.287 0.147 卫矛Euonymus alatus 0.194 0.208 蘀叶蛇葡萄Ampelopsis humulifolia 0.058 0.092 0.109 0.064 表 2 草本层主要组成物种及其重要值
Table 2. Main species composition and important values of the herbage layer
物种 林龄 幼龄林 中龄林 近熟林 成熟林 披针叶薹草Carex lancifolia 0.102 0.158 0.094 大油芒Spodiopogon sibiricus 0.227 0.189 0.082 甲鸟妬草Commelina communis 0.065 0.055 0.074 0.067 东亚唐松草Thalictrum minus 0.087 0.091 0.107 紫菀Aster tataricus 0.074 0.053 0.082 车前Plantago asiatica 0.027 0.045 巴天酸模Rumex patientia 0.043 0.062 灰菜Chenopodium album 0.043 0.049 龙芽草Agrimonia pilosa 0.057 0.037 草乌Aconitum carmichaeli 0.097 0.086 萱草Hemerocallis fulva 0.075 0.055 益母草Leonurus artemisia 0.077 0.082 秋苦荬菜Paraxeris pinnatipartita 0.084 0.067 铁杆嵩Tripolium vulgare 0.063 0.089 石竹Dianthus chinensis 0.087 0.052 蝙蝠葛Menispermum dauricum 0.092 0.068 垂果南芥Arabis pendula 0.047 0.085 0.074 北马兜铃Aristolochia contorta 0.056 0.054 风毛菊Saussurea japonica 0.021 穿龙暑蓣Dioscorea nipponica 0.038 莓叶委陵菜Potentilla centigrana 0.044 黄精Polygonatum sibiricum 0.051 野青茅Deyeuxia arundinacea 0.095 宽叶薹草Carex siderosticta 0.095 北重楼Paris verticillata 0.087 鼠掌老鹤草Geranium sibiricum 0.028 0.082 毛哀Ranunculus japonicus 0.037 0.172 短尾铁线莲Clematis brevicaudata 0.054 0.052 蓝萼香茶菜Rabdosia japonica 0.107 0.118 大叶铁线莲Clematis heracleifolia 0.101 0.069 白屈菜Chelidonium majus 0.057 0.104 表 3 林龄对不同层物种多样性的影响
Table 3. One-way ANOVA for shrub and herbage diversity index aifect by forest age
指数 层次 自由度 平方和 均方 F P R 草本层 3 73.233 24.408 13.708 <0.001 灌木层 3 18.633 6.211 26.617 <0.001 D 草本层 3 0.270 0.090 4.084 0.047 灌木层 3 0.350 0.120 5.133 0.009 H 草本层 3 0.772 0.257 12.832 <0.001 灌木层 3 1.528 0.509 16.236 0.002 J 草本层 3 0.390 0.130 5.263 0.008 灌木层 3 0.270 0.110 5.370 0.004 M 草本层 3 1.280 0.427 15.472 <0.001 灌木层 3 1.070 0.296 9.823 0.007 表 4 物种多样性半变异函数模型及其相关参数
Table 4. Parameters and test values of theoretical variogram models for species diversity indexes
指数 模型(C0) 块金 基台 块金/基台[C0/(C0+C1)] 变程 决定系数 D 球面模型0.023 0.072 0.319 13.2 0.67 H 指数模型0.017 0.047 0.361 7.6 0.64 J 球面模型0.035 0.084 0.416 11.2 0.76 M 指数模型0.027 0.067 0.403 8.4 0.65 -
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