滇中地区云南松天然林群落结构及天然更新规律
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摘要
在云南省中部禄丰县雕林山省级自然保护区,在海拔2200-2450m的阳坡和半阳破,选择暗红壤、黄红壤和红壤上生长且长期封育恢复的云南松天然林以及择伐后更新不同年限的云南松天然次生林为研究对象,采用典型取样法设置样地,开展了云南松纯林和混交林群落的植物种类组成、植物多样性、群落结构、优势树种种群动态及林地土壤特征,云南松在不同亚类红壤和群落内的生长速率、针叶N、P含量与养分再吸收效率,以及云南松天然林及天然次生林的天然更新特点研究。主要研究结论如下:
(1)植物种类数量呈现暗红壤>红壤>黄红壤的变化趋势。暗红壤上植物以禾本科、蔷薇科、菊科、蝶形花科和杜鹃花科种类最多,亚热带代表科山茶科、壳斗科的种类在混交林中明显增多;红壤上纯林的属、种数减少较少,混交林中禾本科种类和蕨类植物明显增加,菊科种类相对减少;黄红壤上植物种类相比暗红壤降低40%以上,禾本科种类明显减少,混交林乔木层杜鹃花科和壳斗科种类明显增加。云南松是群落乔木层重要值最高的树种,乔、灌木优势树种均不同程度含有半湿润常绿阔叶林的部分种类,其中壳斗科树种在不同土壤上存在种类替代现象:暗红壤上为白穗石栎,黄红壤上为麻栎、黄毛青冈,红壤上为槲栎。有害入侵生物紫茎泽兰已侵占了群落草本层,成为群落草本层的优势种类,但对黄红壤上群落草本层的侵害稍轻。
(2)乔木层、灌木层和草本层物种丰富度均存在显著差异,乔木层最高的是暗红壤上云南松混交林及黄红壤云南松纯林,灌木层和草本层最高的是暗红壤上云南松纯林和云南松混交林。乔、灌、草3个层次的Shannon-Wiener指数呈现暗红壤>红壤>黄红壤的变化趋势,但纯林与混交林无明显变化规律。Simpson指数在乔木层表现为云南松混交林>云南松纯林,灌木层为暗红壤>红壤>黄红壤的变化趋势,Pielou指数仅在乔木层存在显著差异,最高的是云南松混交林。
(3)所有林木平均胸径均呈现暗红壤>红壤>黄红壤,云南松纯林>云南松混交林的变化趋势。云南松平均胸径在云南松纯林内呈现为暗红壤>黄红壤>红壤的变化趋势,而在云南松混交林内则为暗红壤>红壤>黄红壤。云南松纯林大树密度平均为10-11株/600m~2,云南松混交林为8-13株/600m~2,云南松大树(DBH≥30.0cm)密度与所有树种大树密度存在较好的协同性。云南松纯林枯立木密度为暗红壤>黄红壤>红壤,云南松混交林为黄红壤>暗红壤>黄红壤。云南松立木密度呈现出云南松纯林>云南松混交林,红壤>黄红壤>暗红壤的变化规律,总的胸高断面积趋势为暗红壤>黄红壤>红壤,云南松混交林>云南松纯林。林地土壤总N含量呈现云南松混交林>云南松林,暗红壤>红壤>黄红壤的明显梯度变化规律,土壤总P、速效N、速效P也存在显著变化,并对群落分布起到较大作用;土壤C/N比为黄红壤>红壤>暗红壤,黄红壤是暗红壤的2.14倍;N/P比为云南松混交林略大于云南松林,暗红壤显著大于黄红壤和红壤。
(4)暗红壤上群落林木层分为4层,为云南松单优群落,云南松混交林的群落高度大于云南松纯林;黄红壤和红壤上群落林木层只分为3层,除了黄红壤上的混交林为云南松、黄毛青冈共优群落外,均为云南松单优群落。云南松混交林胸径级分布呈倒“J”形,云南松纯林则呈倒“S”形。云南松在暗红壤和红壤上的云南松混交林内为稳定种群,在红壤上云南松纯林内为增长种群,其余为衰退种群。除红壤上云南松混交林的滇油杉、黄毛青冈等树种为间歇型种群外,其它群落内的优势树种或属于稳定种群,或属于增长种群,利于群落稳定与演替。
(5)云南松针叶N含量与土壤总N梯度变化一致。90%以上云南松个体生长发育受N元素限制。云南松针叶Nr大于Pr,与多数研究Pr>Nr的结果相反,揭示云南松具有较高的N养分保存能力;HGR和DGR呈现暗红壤>黄红壤>红壤,云南松林>云南松混交林的变化趋势,VGR为云南松混交林>云南松林。Nr与土壤全N、全P、速效N、速效K成正相关,Pr与速效K成显著正相关。HGR和DGR与速效N成显著正相关,VGR与总N、总P及速效N成显著正相关,总体上土壤总N、速效N、容重和自然含水量显著影响云南松高、胸径和材积的生长速率。
(6)长期封育云南松天然林更新密度呈现为云南松混交林>云南松林,暗红壤>红壤>黄红壤的变化趋势,壳斗科、山茶科、杜鹃花科树种更新密度差异显著,可能导致不同群落趋向不同的种类组成。云南松更新密度和更新频度以红壤云南松纯林和云南松混交林最高。云南松是稀疏云南松林和云南松择伐迹地更新频度最高的树种,具有不同大小级的幼苗幼树。择伐更新20年后的云南松天然次生林,Ⅳ级小树、Ⅴ级小树和Ⅰ级幼苗更新密度显著大于稀疏云南松林和择伐更新10年的云南松天然次生林。地被物基质影响云南松的天然更新,65%以上的云南松幼苗更新基质为裸露的矿质土壤,森林地被物和石块也占一定比例,枯落物和粗木质残余物的比例极低。
根据上述研究结果,可以通过增加土壤总N含量、降低土壤C/N比以促进群落内云南松的生长发育,并可通过引入林火干扰体系、创造林窗及矿质土壤裸露促进云南松的天然更新,以较低成本恢复云南松在群落内的优势地位,维持区域群落长期稳定性,维护滇中地区异质性的森林景观。
Based on pure Yunnan Pine (Pinus yunnanensis) forest (PPY)and mixed Ynanan Pine forest(MPY) after long enclosure for restoration, which distributed on dark-red soil(D), yellow-red soil(Y) and red soil(R) respectively with the same site conditions and located the south to west aspect at the elevation of 2200-2450m in Diaolinshan Natural Reserve, Middle Yunnan Plateau, Yunnan, China, as well as natural second-growth forests at different regeneration ages after selective logging, the characteristics of species composition, plant diversity, community structure and natural regeneration, together with dominant tree species’population dynamics, soil properties in PPY and MPY, growth rates and leaf stoichiometry of P. yunnanensis in different soils and communities were studied by representatively sampling. The main results indicate as follows:
(1) The number of plant species in communities was in the order of D>R>Y. In dark-red soil, most of species were the family of Gramineae, Rosaceae, Compositae, Papilionaceae and Ericaceae, and the species of the family of Theaceae, Fagaceae and Aquifoliaceae increased in MPY apparently. Compared with dark-red soil, the number of species on yellow-red soil decreased by over 40%, but the species of Ericaceae and Fagaceae increased in MPY apparently. As for the red soil, the number of genus and species increased apparently in PPY, especially those of Gramineae and ferns increased apparently, but that of Compositae decreased relatively. P. yunnanensis was the dominant tree species in all communities with the highest important value. Other dominant tree and shrub species were partly attached to subtropical semi-humid evergreen broad-leaved forests to some extent, and the dominant species of Fagaceaea showed species replacement phenomena on different soils, namely Lithocarpus leucostachyus on dark-red soil, Quercus acutissima, Cyclobalanopsis delavayi on yellow-red soil, and Q. aliena on red soil. The harmful Eupatorium adenophorum invaded and occupied the herb layers in all communities and did little harm to forests on yellow-red soil.
(2) Species richness in all layers showed significant difference among different communities. MPY on dark-red soil and PPY on yellow-red soil presented the highest values in tree layer, whearas MPY and PPY on dark-red soil were the highest in shrub and herb layers. Shannon-Wiener index of 3 layers showed significant difference with the same tendency of D>R>Y, but no constant order between MPY and PPY. Simpson index indicated the order of MPY>PPY in tree layer and D>R>Y in shrub layer, and Pielou index showed significant difference in tree and shrub layers.
(3)Mean DBH of all trees was in the order of D>R>Y and PPY>MPY. Mean DBH of P. yunnanensis showed the tendency of D>Y>R in PPY, but indiceted the order of D>R>Y in MPY. The density of P. yunnanensis stumpage presented the tendency of PPY>MPY and R>Y>D, and total BA was in the order of MPY>PPY and D>Y>R. It was clear that soil total N changed along the gradients of MPY>PPY and D>R>Y. The ratio of C/N was changed as D>R>Y, and that of N/P as MPY>PY slightly, D>Y and R remarkably. Thus it was certain that PY and MPY changed with total N gradient on different soils. Soil total P, available N, available P changed remarkably and played an important role in the distribution of communities.
(4) On dark-red soil, the forests were featured as uni-dominant tree of P. yunnanensis and divided into 4 layers. Community height was in the order of MPY>PY. As for yellow-red soil and red soil, the forests were divided into 3 layers, but MPY on yellow-red soil featured as co-dominant trees of P. yunnanensis and Cyclobalanopsis delavayi. DBH distribution types of all tree layer individuals in MPY showed a typical reverse“J”shape and in PY a similar reverse“S”shape. P. yunnanensis population structure in MPY on dark-red and red soils was characterized as constant type, in PPY on red soil as incremental type and others as recessionary type. Except that Keteleeria evelyniana, Cy. delavayi, Q. aliena, Castanopsis delavayi and L. dealbatus in MPY on red soil were featured as interim population type, others characterized as constant or incremental population types.
(5) Leaf N of P. yunnanensis showed the same tendency along soil total N gradients. Over 90% individuals were N limited according to the threshold of limiting nutrients. The result of Nr>Pr tendency of leaf nutrient resorption efficiency was contrary to those in most study cases, thus it revealed that Yunnan Pine had higher ability of conserve N nutrient. Height and DBH growth rates showed the tendency of D>Y>R and PPY>MPY, but volume growth rate was in the order of MPY>PPY. There was positive correlation between leaf Nr and soil total N, total P, available N and available K. As for leaf Pr, there was positively remarkable correlation to available K, and positive correlation to soil total N, total P and available N. As far as the growth rates were concerned, HGR and DGR indicated remarkably positive correlation with available N, but VGR showed the same correlation with total N, total P and available N. As a whole, soil total N, available N, bulk density and natural water content affected HGR, DGR and VGR remarkably.
(6)Regeneration density in natural forests after long enclosure for restoration revealed the tendency of MPY>PY and D>R>Y. In the meanwhile regeneration density and percentage of Fagaceae, Theaceae and Ericaceae differed significantly among different soils and communities, which may result in different succession tendency and species composition. It was indicated that regeneration density and frequency of P. yunnanensis in PPY and MPY on red soil were higher significantly than those in other communities, but was very poor in other communities. P. yunnanensis ranked the top tree species about regeneration frequency in sparse forest and logging areas, and its seedlings and saplings at different size classes may result in forests with hetero-aged structure. The densities ofⅣsmall trees,Ⅴsmall trees andⅠseedlings in second-growth forests after 20 years logging were higher significantly than that of forests after 10 years logging and in sparse P. yunnanensis forest. Understory substrates affected natural regeneration of P. yunnanensis. Over 60% of P. yunnanensis seedlings were found growing on the mineral soil, others were found on forest floor, rock, litter and CWD.
From the results above, there are some implications for restoration. Soil total N increment or soil C/N ratio reduction may be proposed to enhance the growth of P. yunnanensis. Fire disturbance regimes may be introduced to create gaps and naked mineral soil in order to facilitate natural regeneration of P. yunnanensis. Thus P. yunnanensis will keep its dominance in communnities at lower costs so that community stability and landscape heterogeneity may be maintained regionally.
(1)植物种类数量呈现暗红壤>红壤>黄红壤的变化趋势。暗红壤上植物以禾本科、蔷薇科、菊科、蝶形花科和杜鹃花科种类最多,亚热带代表科山茶科、壳斗科的种类在混交林中明显增多;红壤上纯林的属、种数减少较少,混交林中禾本科种类和蕨类植物明显增加,菊科种类相对减少;黄红壤上植物种类相比暗红壤降低40%以上,禾本科种类明显减少,混交林乔木层杜鹃花科和壳斗科种类明显增加。云南松是群落乔木层重要值最高的树种,乔、灌木优势树种均不同程度含有半湿润常绿阔叶林的部分种类,其中壳斗科树种在不同土壤上存在种类替代现象:暗红壤上为白穗石栎,黄红壤上为麻栎、黄毛青冈,红壤上为槲栎。有害入侵生物紫茎泽兰已侵占了群落草本层,成为群落草本层的优势种类,但对黄红壤上群落草本层的侵害稍轻。
(2)乔木层、灌木层和草本层物种丰富度均存在显著差异,乔木层最高的是暗红壤上云南松混交林及黄红壤云南松纯林,灌木层和草本层最高的是暗红壤上云南松纯林和云南松混交林。乔、灌、草3个层次的Shannon-Wiener指数呈现暗红壤>红壤>黄红壤的变化趋势,但纯林与混交林无明显变化规律。Simpson指数在乔木层表现为云南松混交林>云南松纯林,灌木层为暗红壤>红壤>黄红壤的变化趋势,Pielou指数仅在乔木层存在显著差异,最高的是云南松混交林。
(3)所有林木平均胸径均呈现暗红壤>红壤>黄红壤,云南松纯林>云南松混交林的变化趋势。云南松平均胸径在云南松纯林内呈现为暗红壤>黄红壤>红壤的变化趋势,而在云南松混交林内则为暗红壤>红壤>黄红壤。云南松纯林大树密度平均为10-11株/600m~2,云南松混交林为8-13株/600m~2,云南松大树(DBH≥30.0cm)密度与所有树种大树密度存在较好的协同性。云南松纯林枯立木密度为暗红壤>黄红壤>红壤,云南松混交林为黄红壤>暗红壤>黄红壤。云南松立木密度呈现出云南松纯林>云南松混交林,红壤>黄红壤>暗红壤的变化规律,总的胸高断面积趋势为暗红壤>黄红壤>红壤,云南松混交林>云南松纯林。林地土壤总N含量呈现云南松混交林>云南松林,暗红壤>红壤>黄红壤的明显梯度变化规律,土壤总P、速效N、速效P也存在显著变化,并对群落分布起到较大作用;土壤C/N比为黄红壤>红壤>暗红壤,黄红壤是暗红壤的2.14倍;N/P比为云南松混交林略大于云南松林,暗红壤显著大于黄红壤和红壤。
(4)暗红壤上群落林木层分为4层,为云南松单优群落,云南松混交林的群落高度大于云南松纯林;黄红壤和红壤上群落林木层只分为3层,除了黄红壤上的混交林为云南松、黄毛青冈共优群落外,均为云南松单优群落。云南松混交林胸径级分布呈倒“J”形,云南松纯林则呈倒“S”形。云南松在暗红壤和红壤上的云南松混交林内为稳定种群,在红壤上云南松纯林内为增长种群,其余为衰退种群。除红壤上云南松混交林的滇油杉、黄毛青冈等树种为间歇型种群外,其它群落内的优势树种或属于稳定种群,或属于增长种群,利于群落稳定与演替。
(5)云南松针叶N含量与土壤总N梯度变化一致。90%以上云南松个体生长发育受N元素限制。云南松针叶Nr大于Pr,与多数研究Pr>Nr的结果相反,揭示云南松具有较高的N养分保存能力;HGR和DGR呈现暗红壤>黄红壤>红壤,云南松林>云南松混交林的变化趋势,VGR为云南松混交林>云南松林。Nr与土壤全N、全P、速效N、速效K成正相关,Pr与速效K成显著正相关。HGR和DGR与速效N成显著正相关,VGR与总N、总P及速效N成显著正相关,总体上土壤总N、速效N、容重和自然含水量显著影响云南松高、胸径和材积的生长速率。
(6)长期封育云南松天然林更新密度呈现为云南松混交林>云南松林,暗红壤>红壤>黄红壤的变化趋势,壳斗科、山茶科、杜鹃花科树种更新密度差异显著,可能导致不同群落趋向不同的种类组成。云南松更新密度和更新频度以红壤云南松纯林和云南松混交林最高。云南松是稀疏云南松林和云南松择伐迹地更新频度最高的树种,具有不同大小级的幼苗幼树。择伐更新20年后的云南松天然次生林,Ⅳ级小树、Ⅴ级小树和Ⅰ级幼苗更新密度显著大于稀疏云南松林和择伐更新10年的云南松天然次生林。地被物基质影响云南松的天然更新,65%以上的云南松幼苗更新基质为裸露的矿质土壤,森林地被物和石块也占一定比例,枯落物和粗木质残余物的比例极低。
根据上述研究结果,可以通过增加土壤总N含量、降低土壤C/N比以促进群落内云南松的生长发育,并可通过引入林火干扰体系、创造林窗及矿质土壤裸露促进云南松的天然更新,以较低成本恢复云南松在群落内的优势地位,维持区域群落长期稳定性,维护滇中地区异质性的森林景观。
Based on pure Yunnan Pine (Pinus yunnanensis) forest (PPY)and mixed Ynanan Pine forest(MPY) after long enclosure for restoration, which distributed on dark-red soil(D), yellow-red soil(Y) and red soil(R) respectively with the same site conditions and located the south to west aspect at the elevation of 2200-2450m in Diaolinshan Natural Reserve, Middle Yunnan Plateau, Yunnan, China, as well as natural second-growth forests at different regeneration ages after selective logging, the characteristics of species composition, plant diversity, community structure and natural regeneration, together with dominant tree species’population dynamics, soil properties in PPY and MPY, growth rates and leaf stoichiometry of P. yunnanensis in different soils and communities were studied by representatively sampling. The main results indicate as follows:
(1) The number of plant species in communities was in the order of D>R>Y. In dark-red soil, most of species were the family of Gramineae, Rosaceae, Compositae, Papilionaceae and Ericaceae, and the species of the family of Theaceae, Fagaceae and Aquifoliaceae increased in MPY apparently. Compared with dark-red soil, the number of species on yellow-red soil decreased by over 40%, but the species of Ericaceae and Fagaceae increased in MPY apparently. As for the red soil, the number of genus and species increased apparently in PPY, especially those of Gramineae and ferns increased apparently, but that of Compositae decreased relatively. P. yunnanensis was the dominant tree species in all communities with the highest important value. Other dominant tree and shrub species were partly attached to subtropical semi-humid evergreen broad-leaved forests to some extent, and the dominant species of Fagaceaea showed species replacement phenomena on different soils, namely Lithocarpus leucostachyus on dark-red soil, Quercus acutissima, Cyclobalanopsis delavayi on yellow-red soil, and Q. aliena on red soil. The harmful Eupatorium adenophorum invaded and occupied the herb layers in all communities and did little harm to forests on yellow-red soil.
(2) Species richness in all layers showed significant difference among different communities. MPY on dark-red soil and PPY on yellow-red soil presented the highest values in tree layer, whearas MPY and PPY on dark-red soil were the highest in shrub and herb layers. Shannon-Wiener index of 3 layers showed significant difference with the same tendency of D>R>Y, but no constant order between MPY and PPY. Simpson index indicated the order of MPY>PPY in tree layer and D>R>Y in shrub layer, and Pielou index showed significant difference in tree and shrub layers.
(3)Mean DBH of all trees was in the order of D>R>Y and PPY>MPY. Mean DBH of P. yunnanensis showed the tendency of D>Y>R in PPY, but indiceted the order of D>R>Y in MPY. The density of P. yunnanensis stumpage presented the tendency of PPY>MPY and R>Y>D, and total BA was in the order of MPY>PPY and D>Y>R. It was clear that soil total N changed along the gradients of MPY>PPY and D>R>Y. The ratio of C/N was changed as D>R>Y, and that of N/P as MPY>PY slightly, D>Y and R remarkably. Thus it was certain that PY and MPY changed with total N gradient on different soils. Soil total P, available N, available P changed remarkably and played an important role in the distribution of communities.
(4) On dark-red soil, the forests were featured as uni-dominant tree of P. yunnanensis and divided into 4 layers. Community height was in the order of MPY>PY. As for yellow-red soil and red soil, the forests were divided into 3 layers, but MPY on yellow-red soil featured as co-dominant trees of P. yunnanensis and Cyclobalanopsis delavayi. DBH distribution types of all tree layer individuals in MPY showed a typical reverse“J”shape and in PY a similar reverse“S”shape. P. yunnanensis population structure in MPY on dark-red and red soils was characterized as constant type, in PPY on red soil as incremental type and others as recessionary type. Except that Keteleeria evelyniana, Cy. delavayi, Q. aliena, Castanopsis delavayi and L. dealbatus in MPY on red soil were featured as interim population type, others characterized as constant or incremental population types.
(5) Leaf N of P. yunnanensis showed the same tendency along soil total N gradients. Over 90% individuals were N limited according to the threshold of limiting nutrients. The result of Nr>Pr tendency of leaf nutrient resorption efficiency was contrary to those in most study cases, thus it revealed that Yunnan Pine had higher ability of conserve N nutrient. Height and DBH growth rates showed the tendency of D>Y>R and PPY>MPY, but volume growth rate was in the order of MPY>PPY. There was positive correlation between leaf Nr and soil total N, total P, available N and available K. As for leaf Pr, there was positively remarkable correlation to available K, and positive correlation to soil total N, total P and available N. As far as the growth rates were concerned, HGR and DGR indicated remarkably positive correlation with available N, but VGR showed the same correlation with total N, total P and available N. As a whole, soil total N, available N, bulk density and natural water content affected HGR, DGR and VGR remarkably.
(6)Regeneration density in natural forests after long enclosure for restoration revealed the tendency of MPY>PY and D>R>Y. In the meanwhile regeneration density and percentage of Fagaceae, Theaceae and Ericaceae differed significantly among different soils and communities, which may result in different succession tendency and species composition. It was indicated that regeneration density and frequency of P. yunnanensis in PPY and MPY on red soil were higher significantly than those in other communities, but was very poor in other communities. P. yunnanensis ranked the top tree species about regeneration frequency in sparse forest and logging areas, and its seedlings and saplings at different size classes may result in forests with hetero-aged structure. The densities ofⅣsmall trees,Ⅴsmall trees andⅠseedlings in second-growth forests after 20 years logging were higher significantly than that of forests after 10 years logging and in sparse P. yunnanensis forest. Understory substrates affected natural regeneration of P. yunnanensis. Over 60% of P. yunnanensis seedlings were found growing on the mineral soil, others were found on forest floor, rock, litter and CWD.
From the results above, there are some implications for restoration. Soil total N increment or soil C/N ratio reduction may be proposed to enhance the growth of P. yunnanensis. Fire disturbance regimes may be introduced to create gaps and naked mineral soil in order to facilitate natural regeneration of P. yunnanensis. Thus P. yunnanensis will keep its dominance in communnities at lower costs so that community stability and landscape heterogeneity may be maintained regionally.
引文
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