子午岭主要乔木树种更新特性的研究
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摘要
黄土高原为世界最大的黄土堆积区,气候较干旱、植被稀疏、水土流失严重。除自然因素外,与人类活动,特别是滥垦滥伐,破坏天然植被等社会因素有密切关系,所以植被的重建与恢复对黄土高原的生态环境恢复至关重要。子午岭地处黄土高原腹地,森林覆盖率88.3%,是黄土高原生态脆弱地区保存最完整、最典型、最珍贵的天然次生森林生态系统。研究该地区主要乔木树种的更新特性对黄土高原植被恢复与重建有重要的现实意义,对西部的水土保持和环境的保护、改善有重要的指导和实践意义。
通过实地考察和野外植物群落调查并采用Levins生态位宽度指数对子午岭地区主要乔木树种,即建群种辽东栎(Quercus liaotungensis)、油松(Pinus Tabulaeformis)和主要乔木树种的幼苗、幼树和成树在不同坡向的辽东栎林、油松+辽东栎混交林和人工油松林三种群落中的生态位宽度和生态位重叠及幼苗与土壤养分和主要树种幼苗之间的关系进行了分析研究。结果显示:
(1)辽东栎和油松的更新生态位宽度在不同的群落中各不相同,成树的生态位宽度与群落类型相一致。辽东栎幼苗和幼树在阳坡和阴坡的人工油松林中的生态位宽度均较大,分别是0.91、0.95和0.98、0.94,与其他两个群落的生态位宽度差异显著(P＜0.05),表明辽东栎幼苗和幼树能较好的适应人工油松林的环境条件;在阳坡,油松幼苗的生态位宽度在辽东栎林最大是0.79,和其它两个群落的差异极显著(P＜0.01);油松幼树在阴坡人工油松林最大为0.63,阳坡人工油松林中油松幼树的生态位宽度最大是1.00,阴坡油松幼树的生态位宽度却在油松+辽东栎混交林最大是0.83,和其它两个群落的差异极其显著(P＜0.01)。表明油松幼苗既能适应阳坡的辽东栎林环境,也能适应阴坡的人工油松林环境;油松幼树不但能适应阳坡的人工油松林环境,而且也能适应阳坡的辽东栎林环境。人工种植油松林有利于该地区的植被恢复,人工油松林和油松+辽东栎混交林会在该地区存在较长时间,而阳坡的油松+辽东栎混交林存在时间会更长。
(2)辽东栎和油松幼苗、幼树和成树的生态位重叠大多数情况下与生态位宽度表现一致;但当生态位宽度大时其生态位重叠不一定大。辽东栎幼苗在阳坡和阴坡的人工油松林中的生态位重叠均较大,分别是0.68和0.70,与在阳坡其他两个群落差异显著(P＜0.05),在阴坡仅与辽东栎林差异显著(P＜0.05);辽东栎幼树在阴坡的生态位重叠并不显著(P＞0.05)。在阳坡,油松幼苗和幼树的生态位重叠在辽东栎林和人工油松林中较大,分别为0.56和0.65;油松幼苗和幼树在阴坡人工油松林和油松+辽东栎混交林中较大,为0.64和0.61。
(3)其他乔木的更新生态位宽度在三个群落类型阴阳坡的分析表明葛萝槭(Acergrosseri)、茶条槭(A.ginnala)、鹅耳枥(Carpinus turczaninowii)和漆树(Toxicodendronvernicifluum)的幼苗、幼树和成树的生态位宽度较大,由于其分布较广,是常见的伴生种。茶条槭幼苗的生态位宽度和辽东栎幼苗的生态位宽度在三个群落阴阳坡表现不同,即其阳坡的生态位宽度大于阴坡的,而辽东栎幼苗是阳坡的大于阴坡的;由于这两者幼苗的分布较广,所以两者幼苗生态位重叠较大。油松幼苗和茶条槭幼苗的生态位重叠较辽东栎幼苗与茶条槭幼苗生态位重叠小,但油松幼苗和辽东栎幼苗生态位重叠较大。
(4)乔木幼苗与土壤养分的分析表明,影响不同乔木幼苗更新的土壤因子不同。影响辽东栎幼苗的土壤因子是有机质,随着土壤中有机质含量的升高,其密度在缓慢减小,主要集中在有机质为5～8g/kg的地段,当有机质含量达到10g/kg时,辽东栎幼苗密度稳定在一个水平上。影响油松幼苗更新的主要土壤因子是速效钾,随着土壤中速效钾含量的升高,其密度在也在升高。天然油松林速效钾含量较人工油松林中高,建议在人工造林林种植油松时,应当注意调查种植地的速效钾含量。
(5)辽东栎幼苗与山楂(Crataegus pinnatifid)幼苗、杜梨(Pyrus betulaefolia)幼苗和漆树幼苗成呈正相关(分别为:r=0.65,P＜0.01;r=0.43,P＜0.01;r=0.42,P＜0.01),辽东栎幼苗、山楂幼苗、和杜梨幼苗都与有机质呈负关联(分别为:r=-0.39,P＜0.01;r=-0.42,P＜0.05;r=-0.38,P＜0.05)。这表明,土壤养分对乔木的更新有着非常重要的影响,对群落的组成和结构起关键作用。油松幼苗与茶条槭幼苗呈负相关(r=-0.54,P＜0.01),同时油松幼苗和茶条槭幼苗都与速效钾含量之间存在着一定的相关(分别为:r=0.42,P＜0.01;r=-0.48,P＜0.01)。
(6)辽东栎幼苗和油松幼苗的数量随着年龄得增长逐渐降低,分别在9和8年时基本达到稳定状态,其死亡率和损失度分别在6年、8年和3年、7年达到最高。其死亡原因有待进一步深入研究。
The Loess Plateau is the largest accumulative loess area in the world,but the climate is arid,the vegetation is sparse,and the loss of water and soil is serious in this area.This situation is closely related to human activity,especially estrepement and deforestation,except natural factors.So it is imperative under the situation to accelerate the recovery and restoration.Ziwuling Mountain,which is located in the hinterland of the Loess Plateau,the forest coverage rate is 88.3%,and the vegetation is the best preserved,the most typical,the most valuable natural secondary forest ecosystems on the ecologically fragile areas of the Loess Plateau. Ziwuling Mountain forest region has crucial effect on adjusting regional climate,maintaining ecological balance and promoting ecologically sustainable economic development on the Loess Plateau.So the research of forest community regeneration has significant ecological meaning for recovery and restoration of degenerative ecosystem.
Based on field survey to Ziwuling Mountain,adopting Levins index and Pianka index,the generational(seedling,sapling and adult) niche breadth and overlap of Quercus liaotungensis,Pinus tabulaeformis and other arbor tree species were studied in different community types,which was Q. liaotungensis forest,P.tabulaeformis+Q.liaotungensis mixed forest and artificial P.tabulaeformis forest on sunny and shady slope.The relationship between arbor tree seedlings and soil nutrient,and the relationship among arbor tree seedlings were also studied.
(1) The results showed that the generational niche breadth of Q.liaotungensis and P. tabulaeformis was different in these community types.The niche breadth of adult was the same as community.The niche breadths of Q.liaotungensis' seedling and sapling were the highest in artificial P.tabulaeformis forest on sunny slope and shady slope,which are 0.91,0.95,0.98 and 0.94. There was significant difference among other community.It indicated the growth of Q. liaotungensis' seedling and sapling was inclined to the environmental condition of artificial P. tabulaeformis forest.The niche breadth of Q.liaotungensis' seedling was the highest in Q. liaotungensis forest on sunny slope and artificial P.tabulaeformis forest on shady slope,which are 0.79 and 0.63.There was significant difference among other community.The niche breadth of Q. liaotungensis' sapling was the highest in artificial P.tabulaeformis forest on suuny slope and in P. tabulaeformis+Q.liaotungensis mixed forest on shady slope,which are 1.00 and 0.83.There was significant difference among other community.The growth of P.tabulaeformis' seedling was adaptable not only to the environment condition of Q.liaotungensis forest on sunny slope but to that of artificial P.tabulaeformis forest on shady slope.The growth of P.tabulaeformis' sapling was adaptable not only to the environment condition of artificial P.tabulaeformis forest but to that of Q. liaotungensis forest on sunny slope.As a whole,artificial P.tabulaeformis forest had an important effect on the restoration of natural vegetation in this region.Artificial P.tabulaeformis forest and P. tabulaeformis+Q.liaotungensis mixed forest would have existed over a long period of time,while P.tabulaeformis+Q.liaotungensis mixed forest on sunny slope would have even more.
(2) The niche overlap of Q.liaotungensis' seedling and sapling niche breadth is consistent in most cases,but some times is not consisten.The niche overlap of Q.liaotungensis' seedling were higher in artificial P.tabulaeformis forest on sunny slope and shady slope,which are 0.68 and 0.70. There was significant difference among other community on sunny slope,but there was significant difference between Q.liaotungensis' forest shady slope.On sunny slope,the niche overlap of P. tabulaeformis' seedling and sapling were higher in artificial P.tabulaeformis forest,which are 0.56 and 0.65.There was significant difference among other community,but there was significant difference between Q.liaotungensis' forest shady slope.On shady slope,the niche overlap of P. tabulaeformis' seedling and sapling were higher in P.tabulaeformis+Q.liaotungensis mixed forest, which are 0.64 and 0.61.
(3) The niche breadth of Acer grosseri,A.ginnala,Carpinus turczaninowii and Toxicodendron vernicifluum's seedling,sapling and adult were higher than other trees in different community types on sunny slope and shady slope.They are the common companion species and have a wide distribution.The niche breadth of A.ginnala's seedling,which the sunny slope niche breadth was higher than shady slope in different community types,was contrary to Q.liaotungensis' seedling in different community.Because both A.ginnala's and Q.liaotungensis' seedling have a wide distribution,so the niche overlap between two was more higher than the niche overlap between P. tabulaeformi and A.ginnala's.But the he niche overlap between P.tabulaeformi and Q. liaotungensis' seedling was higher.
(4) The analysis between tree seedlings and soil nutrient showed that the factor of influence on different trees seedling were difference.The factor of soil nutrient influence on Q.liaotungensis' seedling was organic matter.Along with the content of organic matter in soil increased its density decreases slowly,when the content of organic matter was 10g/kg,Q.liaotungensis' seedling density was to keep in a stable leve.And Q.liaotungensis' seedling density mainly concentrated in the organic matter for the 5～8 g/kg.The factor of soil nutrient influence on P.tabulaeformis seedling was available potassium.The content of available potassium in natural P.tabulaeformis forest was higher than that of artificial P.tabulaeformis forest,so when P.tabulaeformis as a restoration should be paid attention to investigating the available potassium content of cultivation.
(5) Q.liaotungensis' seedlings were significantly positive correlated with Crataegus pinnatifid seedling,Pyrus betulaefolia seedlings and Toxicodendron vernicifluum seedlings(r=0.65,P＜0.01; r=0.43,P＜0.01;r=0.42,P＜0.01).They were all negatively associated with the organic matter(r =-0.39,P＜0.01;r=-0.42,P＜0.05;r=-0.38,P＜0.05).This shows that the soil nutrients were very important to tree regeneration,but also play a important role in the community composition and structure.P.tabulaeformis seedlings were negative correlated with correlation with A.ginnala seedlings(r=-0.54,P＜0.01).At the same time,P.tabulaeformis seedlings and A.ginnala seedlings were statistically significantly correlated with available potassium content(r=0.42,P＜0.01;r=-0.48,P＜0.01).
(6) The number of Q.liaotungensis' seedlings and P.tabulaeformis seedlings were gradually decreased as the age increasing,which respectively at the age of nine and eight to achieve a stability state.The mortality rate and killing valuable of Q.liaotungensisa seedling was higher at the age of six and eight,while the mortality rate and killing valuable of P.tabulaeformis seedling was higher at the age of three and seven.But the cause of their death needs further research.
通过实地考察和野外植物群落调查并采用Levins生态位宽度指数对子午岭地区主要乔木树种,即建群种辽东栎(Quercus liaotungensis)、油松(Pinus Tabulaeformis)和主要乔木树种的幼苗、幼树和成树在不同坡向的辽东栎林、油松+辽东栎混交林和人工油松林三种群落中的生态位宽度和生态位重叠及幼苗与土壤养分和主要树种幼苗之间的关系进行了分析研究。结果显示:
(1)辽东栎和油松的更新生态位宽度在不同的群落中各不相同,成树的生态位宽度与群落类型相一致。辽东栎幼苗和幼树在阳坡和阴坡的人工油松林中的生态位宽度均较大,分别是0.91、0.95和0.98、0.94,与其他两个群落的生态位宽度差异显著(P＜0.05),表明辽东栎幼苗和幼树能较好的适应人工油松林的环境条件;在阳坡,油松幼苗的生态位宽度在辽东栎林最大是0.79,和其它两个群落的差异极显著(P＜0.01);油松幼树在阴坡人工油松林最大为0.63,阳坡人工油松林中油松幼树的生态位宽度最大是1.00,阴坡油松幼树的生态位宽度却在油松+辽东栎混交林最大是0.83,和其它两个群落的差异极其显著(P＜0.01)。表明油松幼苗既能适应阳坡的辽东栎林环境,也能适应阴坡的人工油松林环境;油松幼树不但能适应阳坡的人工油松林环境,而且也能适应阳坡的辽东栎林环境。人工种植油松林有利于该地区的植被恢复,人工油松林和油松+辽东栎混交林会在该地区存在较长时间,而阳坡的油松+辽东栎混交林存在时间会更长。
(2)辽东栎和油松幼苗、幼树和成树的生态位重叠大多数情况下与生态位宽度表现一致;但当生态位宽度大时其生态位重叠不一定大。辽东栎幼苗在阳坡和阴坡的人工油松林中的生态位重叠均较大,分别是0.68和0.70,与在阳坡其他两个群落差异显著(P＜0.05),在阴坡仅与辽东栎林差异显著(P＜0.05);辽东栎幼树在阴坡的生态位重叠并不显著(P＞0.05)。在阳坡,油松幼苗和幼树的生态位重叠在辽东栎林和人工油松林中较大,分别为0.56和0.65;油松幼苗和幼树在阴坡人工油松林和油松+辽东栎混交林中较大,为0.64和0.61。
(3)其他乔木的更新生态位宽度在三个群落类型阴阳坡的分析表明葛萝槭(Acergrosseri)、茶条槭(A.ginnala)、鹅耳枥(Carpinus turczaninowii)和漆树(Toxicodendronvernicifluum)的幼苗、幼树和成树的生态位宽度较大,由于其分布较广,是常见的伴生种。茶条槭幼苗的生态位宽度和辽东栎幼苗的生态位宽度在三个群落阴阳坡表现不同,即其阳坡的生态位宽度大于阴坡的,而辽东栎幼苗是阳坡的大于阴坡的;由于这两者幼苗的分布较广,所以两者幼苗生态位重叠较大。油松幼苗和茶条槭幼苗的生态位重叠较辽东栎幼苗与茶条槭幼苗生态位重叠小,但油松幼苗和辽东栎幼苗生态位重叠较大。
(4)乔木幼苗与土壤养分的分析表明,影响不同乔木幼苗更新的土壤因子不同。影响辽东栎幼苗的土壤因子是有机质,随着土壤中有机质含量的升高,其密度在缓慢减小,主要集中在有机质为5～8g/kg的地段,当有机质含量达到10g/kg时,辽东栎幼苗密度稳定在一个水平上。影响油松幼苗更新的主要土壤因子是速效钾,随着土壤中速效钾含量的升高,其密度在也在升高。天然油松林速效钾含量较人工油松林中高,建议在人工造林林种植油松时,应当注意调查种植地的速效钾含量。
(5)辽东栎幼苗与山楂(Crataegus pinnatifid)幼苗、杜梨(Pyrus betulaefolia)幼苗和漆树幼苗成呈正相关(分别为:r=0.65,P＜0.01;r=0.43,P＜0.01;r=0.42,P＜0.01),辽东栎幼苗、山楂幼苗、和杜梨幼苗都与有机质呈负关联(分别为:r=-0.39,P＜0.01;r=-0.42,P＜0.05;r=-0.38,P＜0.05)。这表明,土壤养分对乔木的更新有着非常重要的影响,对群落的组成和结构起关键作用。油松幼苗与茶条槭幼苗呈负相关(r=-0.54,P＜0.01),同时油松幼苗和茶条槭幼苗都与速效钾含量之间存在着一定的相关(分别为:r=0.42,P＜0.01;r=-0.48,P＜0.01)。
(6)辽东栎幼苗和油松幼苗的数量随着年龄得增长逐渐降低,分别在9和8年时基本达到稳定状态,其死亡率和损失度分别在6年、8年和3年、7年达到最高。其死亡原因有待进一步深入研究。
The Loess Plateau is the largest accumulative loess area in the world,but the climate is arid,the vegetation is sparse,and the loss of water and soil is serious in this area.This situation is closely related to human activity,especially estrepement and deforestation,except natural factors.So it is imperative under the situation to accelerate the recovery and restoration.Ziwuling Mountain,which is located in the hinterland of the Loess Plateau,the forest coverage rate is 88.3%,and the vegetation is the best preserved,the most typical,the most valuable natural secondary forest ecosystems on the ecologically fragile areas of the Loess Plateau. Ziwuling Mountain forest region has crucial effect on adjusting regional climate,maintaining ecological balance and promoting ecologically sustainable economic development on the Loess Plateau.So the research of forest community regeneration has significant ecological meaning for recovery and restoration of degenerative ecosystem.
Based on field survey to Ziwuling Mountain,adopting Levins index and Pianka index,the generational(seedling,sapling and adult) niche breadth and overlap of Quercus liaotungensis,Pinus tabulaeformis and other arbor tree species were studied in different community types,which was Q. liaotungensis forest,P.tabulaeformis+Q.liaotungensis mixed forest and artificial P.tabulaeformis forest on sunny and shady slope.The relationship between arbor tree seedlings and soil nutrient,and the relationship among arbor tree seedlings were also studied.
(1) The results showed that the generational niche breadth of Q.liaotungensis and P. tabulaeformis was different in these community types.The niche breadth of adult was the same as community.The niche breadths of Q.liaotungensis' seedling and sapling were the highest in artificial P.tabulaeformis forest on sunny slope and shady slope,which are 0.91,0.95,0.98 and 0.94. There was significant difference among other community.It indicated the growth of Q. liaotungensis' seedling and sapling was inclined to the environmental condition of artificial P. tabulaeformis forest.The niche breadth of Q.liaotungensis' seedling was the highest in Q. liaotungensis forest on sunny slope and artificial P.tabulaeformis forest on shady slope,which are 0.79 and 0.63.There was significant difference among other community.The niche breadth of Q. liaotungensis' sapling was the highest in artificial P.tabulaeformis forest on suuny slope and in P. tabulaeformis+Q.liaotungensis mixed forest on shady slope,which are 1.00 and 0.83.There was significant difference among other community.The growth of P.tabulaeformis' seedling was adaptable not only to the environment condition of Q.liaotungensis forest on sunny slope but to that of artificial P.tabulaeformis forest on shady slope.The growth of P.tabulaeformis' sapling was adaptable not only to the environment condition of artificial P.tabulaeformis forest but to that of Q. liaotungensis forest on sunny slope.As a whole,artificial P.tabulaeformis forest had an important effect on the restoration of natural vegetation in this region.Artificial P.tabulaeformis forest and P. tabulaeformis+Q.liaotungensis mixed forest would have existed over a long period of time,while P.tabulaeformis+Q.liaotungensis mixed forest on sunny slope would have even more.
(2) The niche overlap of Q.liaotungensis' seedling and sapling niche breadth is consistent in most cases,but some times is not consisten.The niche overlap of Q.liaotungensis' seedling were higher in artificial P.tabulaeformis forest on sunny slope and shady slope,which are 0.68 and 0.70. There was significant difference among other community on sunny slope,but there was significant difference between Q.liaotungensis' forest shady slope.On sunny slope,the niche overlap of P. tabulaeformis' seedling and sapling were higher in artificial P.tabulaeformis forest,which are 0.56 and 0.65.There was significant difference among other community,but there was significant difference between Q.liaotungensis' forest shady slope.On shady slope,the niche overlap of P. tabulaeformis' seedling and sapling were higher in P.tabulaeformis+Q.liaotungensis mixed forest, which are 0.64 and 0.61.
(3) The niche breadth of Acer grosseri,A.ginnala,Carpinus turczaninowii and Toxicodendron vernicifluum's seedling,sapling and adult were higher than other trees in different community types on sunny slope and shady slope.They are the common companion species and have a wide distribution.The niche breadth of A.ginnala's seedling,which the sunny slope niche breadth was higher than shady slope in different community types,was contrary to Q.liaotungensis' seedling in different community.Because both A.ginnala's and Q.liaotungensis' seedling have a wide distribution,so the niche overlap between two was more higher than the niche overlap between P. tabulaeformi and A.ginnala's.But the he niche overlap between P.tabulaeformi and Q. liaotungensis' seedling was higher.
(4) The analysis between tree seedlings and soil nutrient showed that the factor of influence on different trees seedling were difference.The factor of soil nutrient influence on Q.liaotungensis' seedling was organic matter.Along with the content of organic matter in soil increased its density decreases slowly,when the content of organic matter was 10g/kg,Q.liaotungensis' seedling density was to keep in a stable leve.And Q.liaotungensis' seedling density mainly concentrated in the organic matter for the 5～8 g/kg.The factor of soil nutrient influence on P.tabulaeformis seedling was available potassium.The content of available potassium in natural P.tabulaeformis forest was higher than that of artificial P.tabulaeformis forest,so when P.tabulaeformis as a restoration should be paid attention to investigating the available potassium content of cultivation.
(5) Q.liaotungensis' seedlings were significantly positive correlated with Crataegus pinnatifid seedling,Pyrus betulaefolia seedlings and Toxicodendron vernicifluum seedlings(r=0.65,P＜0.01; r=0.43,P＜0.01;r=0.42,P＜0.01).They were all negatively associated with the organic matter(r =-0.39,P＜0.01;r=-0.42,P＜0.05;r=-0.38,P＜0.05).This shows that the soil nutrients were very important to tree regeneration,but also play a important role in the community composition and structure.P.tabulaeformis seedlings were negative correlated with correlation with A.ginnala seedlings(r=-0.54,P＜0.01).At the same time,P.tabulaeformis seedlings and A.ginnala seedlings were statistically significantly correlated with available potassium content(r=0.42,P＜0.01;r=-0.48,P＜0.01).
(6) The number of Q.liaotungensis' seedlings and P.tabulaeformis seedlings were gradually decreased as the age increasing,which respectively at the age of nine and eight to achieve a stability state.The mortality rate and killing valuable of Q.liaotungensisa seedling was higher at the age of six and eight,while the mortality rate and killing valuable of P.tabulaeformis seedling was higher at the age of three and seven.But the cause of their death needs further research.
引文
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