天宝岩长苞铁杉林倒木的基础特征及其环境效应研究
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摘要
天宝岩国家级自然保护区内的长苞铁杉是我国特有珍稀濒危植物，在促进森林演替、维持生态系统平衡稳定以及涵养水源方面，发挥着十分重要的作用，然而，其立地条件贫瘠及更新困难的问题已成为困扰长苞铁杉保护和重建的关键问题。倒木是森林生态系统中重要的结构性和功能性的组成要素，不仅能够为森林生物多样性提供生境，而且还是重要的营养库，在森林营养物质循环、能量流动、更新演替及土壤形成等方面扮演着重要角色，这些特殊的生态功能，为长苞铁杉林的系统稳定及天然更新提供了一条富有挑战的新思路。
本研究在倒木数量特征调查的基础上，对倒木的养分贮量、能量贮量及环境因子（包括地形、干扰、林分结构、气象因子及倒木接触处土壤微环境）进行跟踪监测研究，探讨长苞铁杉林倒木在森林生态系统物质循环及能量流动中的生态功能，为后期苗床实验的选址和倒木的选择，提供理论和实践依据，也为森林资源的管理和保护，提供了借鉴依据。研究结果显示：
（1）天宝岩长苞铁杉林内倒木主要由低腐烂等级、0~10cm径级及1~3m长度级倒木构成，数量密度随径级及长度级的升高而降低，体积随径级升高而升高，倒木的密度、总体体积贮量及覆盖面积呈现出随海拔下降而升高的趋势；就腐烂等级而言，倒木密度随腐烂等级升高而下降，体积贮量和覆盖面积在腐烂等级间呈近似正态分布，腐烂等级及林型对倒木数量特征分布的影响较为显著；研究区倒木的体积贮量为18.14m~3/hm~2，后期苗床实验中，若仅考虑最适幼苗着生的第Ⅳ腐烂等级，则可用于苗床实验的倒木体积仅有3.37m~3/m~2。
（2）不同腐烂等级间倒木全C、全K含量差异不显著（P>0.05），全N、全P及C/N差异极显著（P<0.01）；全N、全P含量随腐烂等级升高大致呈递增趋势，C/N随腐烂等级升高而递减，高腐烂等级倒木养分贮量大于低腐烂等级；C总贮量2.316t/hm~2，N总贮量9.832kg/hm~2，P总贮量1.154kg/hm~2，K总贮量1.713kg/hm~2，长苞铁杉纯林内养分贮量最小，长苞铁杉+毛竹混交林内最大，其中，C贮量的分布服从长苞铁杉纯林<长苞铁杉+阔叶树<长苞铁杉+猴头杜鹃<长苞铁杉+毛竹，氮磷钾贮量的分布服从长苞铁杉纯林<长苞铁杉+猴头杜鹃<长苞铁杉+阔叶树<长苞铁杉+毛竹，不同养分元素的贮量随倒木分解大致呈下降趋势，但释放量较小，说明倒木分解缓慢，其养分的归还也较慢，在参与森林生态系统的生物地球化学循环中，对维持森林稳定与可持续发展中扮演着十分重要的角色，特别对于珍稀濒危植物长苞铁杉贫瘠的立地条件而言，显得更为重要。
（3）综合燃烧指数及熵权系数法的评价结果显示，天宝岩长苞铁杉林内倒木的燃烧性随腐烂等级升高而降低，长苞铁杉纯林内倒木的燃烧性及潜在能量最小，而长苞铁杉+毛竹混交林内倒木的燃烧性及潜在能量最大，研究区倒木属于低燃烧性可燃物，其产生火灾的可能性也较小，在森林经营的过程中可保留现有倒木，而暂不做清除或部分移除措施，这有利于最大限度地发挥倒木的生态功能。
（4）采用方差分析、相关分析、通径分析及因子分析对倒木接触处土壤特性展开研究，结果显示，经倒木覆盖后土壤容重、pH值降低、总孔隙度提高、碳氮钾养分含量提高，土壤蛋白酶、脲酶、纤维素酶、蔗糖酶及酸性磷酸酶5种酶的活性显著升高，总体上提高了森林土壤的生产力。随腐烂等级升高，倒木接触处土壤有机碳含量升高，腐烂等级间有机碳含量差异显著，其他土壤特性在腐烂等级间差异不显著，季节及林型对倒木接触处土壤特性变化存在显著影响，土壤特性指标之间显著相关，其中，土壤有机碳与所选理化性质指标极显著相关，全氮与所选土壤酶指标极显著。有机碳、全N及C/N比3者成为影响该区5种酶活性的主要指标。综合看来，倒木的覆盖有利于提高土壤肥力质量，促进土壤环境的物质循环和能量流动。
（5）采用冗余分析（redundancy analysis，RDA）排序分析方法探讨倒木与环境的生态关系，结果表明，海拔是该研究区倒木各项特征分布的主要影响因素；所选环境因子中，海拔对倒木数量特征、养分贮量及能量贮量分布的影响效应最大，降雨量对碳贮量、氮贮量、磷贮量及潜在能量释放的影响最小。除人为干扰外，长苞铁杉林内倒木主要来源于雪灾或风害的干扰影响，较少产生于林木自然稀疏与衰老。越往南坡，高腐烂等级相对增加，土壤酶活性升高，倒木养分和能量的释放增加，说明温度的升高，会加速该区倒木的分解，缩短了倒木作为苗床使用的时间，后期苗床实验中倒木的放置不应过于靠南坡。坡度对倒木数量特征及接触处土壤酶活性的影响最小，对倒木养分及能量的释放影响效应中等；有幼苗着床的倒木主要出现在中坡位，第Ⅳ腐烂等级倒木上的生物多样性更丰富，也更有利于幼苗的着生。
因此，综合上述研究可以得出，天宝岩长苞铁杉林内倒木的存在，既有利于系统养分的固定，也提高了土壤的生产力及异质性，参与了生态系统的物质循环与能量流动，促进森林生态系统的稳定性发展；后期苗床实验中，若选择第Ⅳ腐烂等级倒木作为更新苗床，可置于中坡位，其坡度要求不大，但应注意倒木的放置不应过于靠南坡。
Tsuga longibracteata W. C. Cheng, a rare and endangered coniferous species endemic to China,plays an important role in promoting forest succession,maintaining the balance and stability offorest ecosystems, and water conservation. However, barren site condition and poor naturalregeneration were critical problem for the protection and restoration of Tsuga longibracteata forest.Fallen logs play important roles in maintaining the structural and functional integrity of forestecosystems;they provide habitat for forest species, act as a long-term nutrient store, and contributeto nutrient cycling,energy flow, regeneration succession and soil formation. These functionsprovide evidence for the importance of logs for forest ecosystem. A better knowledge of logs couldhelp in solving the problem of stability and regeneration in Tsuga longibracteata forest. Base onthe investigation on fallen logs quantity and quality,nutrient stock,energy stock and environmentalfactors（including topographic factors, human disturbance, stand structure, meteorological factors andsoil microenvironment under fallen logs）were track monitoring, the present study try to demonstratethe logs ecological function in nutrient cycling and energy flow,and found the right site for seedbedstudy.It also contribute to a reference basis for conservation and sustainable management of Tsugalongibracteata forest. The results showed that:
（1） The fallen logs in Tsuga longibracteata forest in Tianbaoyan National Nature Reserve,mainly belonged to slightly decayed and small-classed logs,that is,they mainly distributed at therange of decay class I~II, diameter class0~10cm, length class1~3m.Logs density were decreasedwhile volume increased with the increased of diameter class（length class）, Logs density, volumeand covered area were increased with the decreased of elevation.Logs density were decreased withthe increased of decay class, volume and covered area distribution were close to normal distribution.fallen logs quantity and quality were differed significantly among decay class and forest types （P<0.05）. fallen logs volume in this area was18.14m~3/hm~2, and the selective volume was only3.37m~3/hm~2when we chose class IV for seedbed study.
（2） The content of C, K were not significantly in different decay classes（P>0.05）, whileN, P and C/N were extremely significantly（P<0.01）; With the increased of the decayclass, thecontent of N, P displayed the increased trend, however C/N decreased; logs nutrient stocks inhighly decay class were larger than slightly decayed; and the total C stock was2.316t/hm~2,while N stock was9.832kg/hm~2,P stock was1.154kg/hm~2,K stock was1.713kg/hm~2. Nutrient stocks waslowest in Tsuga longibracteata pure forest, and largest in Tsuga longibracteata+Phyllostachysedulis forest, C stock in different forest types showed the following trend: Tsuga longibracteatapure forest （3）Burnable logs index and burnable entropy results showed that fallen logs combustibility weredecreased with the increased of decay class, logs in Tsuga longibracteata pure forest were thelowest combustibility and potential energy, whereas highest in Tsuga longibracteata+Phyllostachysedulis forest; in this area,logs were low combustibility and had lower ignition potential,so in theforest management,retain but not remove the current logs could optimize meet their function.
（4） Soil properties under fallen logs were analyzed by variance analysis,correlationanalysis,path analysis and factor analysis,the results showed that soil bulk density and pH werelower, while porosity, SOC, TN, TK, protease, urease, invertase, cellulose and acid phosphataseactivity were higher in soil beneath fallen logs, which indicated that fallen logs were contributed toimprove the soil productivity; SOC were increased with the increased of decay class, SOC wereextremely significant difference in soils under fallen logs of all decay classes, while the other soilproperties were no significant difference; season were significant effects on most soil properties（P<0.05）; Soil properties showed a significant correlation between each other, Highly significantcorrelation was found between SOC and the selected physical and chemical properties,while thesame between TN and the five soil enzyme activities. SOC,TN and C/N were the most importantindex on the five soil enzyme activities.Comprehensively, fallen logs was contributed to improvesoil quality, and promote nutrient cycling and energy flow in soil environment.
（5） Redundancy analysis（RDA） were performed to determine therelationships between fallenlogs and environmental factors, and the results showed that fallen logs quantity and quality was strongly effected by elevation, precipitation have the lowest effect on the release of C,N and Pstock. Except for human disturbance, fallen logs were not mainly from self-thinning and senescence,but mostly origin from snow or wind disturbance.The highly decayed logs, soil enzymeactivities,nutrient and energy release stock were increased from northern to southern slope, whichindicated that elevated temperature will accelerate the decomposition of fallen logs in the studyarea, and thus shorten the duration of its function as a seedbed, so sampling logs that use forseeding experiment should not be overly concentrated on southern slopes in further study. Slopegradient had the least effect on logs quantity and quality and soil enzyme activity under fallenlogs,but middle effect on nutrient and energy release,seedbed logs were found on middleslope,biodiversity was richer on class Ⅳ, which were advantage for seedlings colonization.
Thus,all the results suggested that fallen logs on Tsuga longibracteata forest floor acted aslong-term storage for nutrients,contributed to soil fertility and heterogeneity, the accumulation oflogs would lead to accelerated nutrient cycling and energy flow,which would promoted the stabilityof forest ecosystem.In addition, if the class Ⅳwas chooseen for seedbed experiment, they shouldput in middle slope,and without considering slope gradient, but not be overly concentrated onsouthern slope.
本研究在倒木数量特征调查的基础上，对倒木的养分贮量、能量贮量及环境因子（包括地形、干扰、林分结构、气象因子及倒木接触处土壤微环境）进行跟踪监测研究，探讨长苞铁杉林倒木在森林生态系统物质循环及能量流动中的生态功能，为后期苗床实验的选址和倒木的选择，提供理论和实践依据，也为森林资源的管理和保护，提供了借鉴依据。研究结果显示：
（1）天宝岩长苞铁杉林内倒木主要由低腐烂等级、0~10cm径级及1~3m长度级倒木构成，数量密度随径级及长度级的升高而降低，体积随径级升高而升高，倒木的密度、总体体积贮量及覆盖面积呈现出随海拔下降而升高的趋势；就腐烂等级而言，倒木密度随腐烂等级升高而下降，体积贮量和覆盖面积在腐烂等级间呈近似正态分布，腐烂等级及林型对倒木数量特征分布的影响较为显著；研究区倒木的体积贮量为18.14m~3/hm~2，后期苗床实验中，若仅考虑最适幼苗着生的第Ⅳ腐烂等级，则可用于苗床实验的倒木体积仅有3.37m~3/m~2。
（2）不同腐烂等级间倒木全C、全K含量差异不显著（P>0.05），全N、全P及C/N差异极显著（P<0.01）；全N、全P含量随腐烂等级升高大致呈递增趋势，C/N随腐烂等级升高而递减，高腐烂等级倒木养分贮量大于低腐烂等级；C总贮量2.316t/hm~2，N总贮量9.832kg/hm~2，P总贮量1.154kg/hm~2，K总贮量1.713kg/hm~2，长苞铁杉纯林内养分贮量最小，长苞铁杉+毛竹混交林内最大，其中，C贮量的分布服从长苞铁杉纯林<长苞铁杉+阔叶树<长苞铁杉+猴头杜鹃<长苞铁杉+毛竹，氮磷钾贮量的分布服从长苞铁杉纯林<长苞铁杉+猴头杜鹃<长苞铁杉+阔叶树<长苞铁杉+毛竹，不同养分元素的贮量随倒木分解大致呈下降趋势，但释放量较小，说明倒木分解缓慢，其养分的归还也较慢，在参与森林生态系统的生物地球化学循环中，对维持森林稳定与可持续发展中扮演着十分重要的角色，特别对于珍稀濒危植物长苞铁杉贫瘠的立地条件而言，显得更为重要。
（3）综合燃烧指数及熵权系数法的评价结果显示，天宝岩长苞铁杉林内倒木的燃烧性随腐烂等级升高而降低，长苞铁杉纯林内倒木的燃烧性及潜在能量最小，而长苞铁杉+毛竹混交林内倒木的燃烧性及潜在能量最大，研究区倒木属于低燃烧性可燃物，其产生火灾的可能性也较小，在森林经营的过程中可保留现有倒木，而暂不做清除或部分移除措施，这有利于最大限度地发挥倒木的生态功能。
（4）采用方差分析、相关分析、通径分析及因子分析对倒木接触处土壤特性展开研究，结果显示，经倒木覆盖后土壤容重、pH值降低、总孔隙度提高、碳氮钾养分含量提高，土壤蛋白酶、脲酶、纤维素酶、蔗糖酶及酸性磷酸酶5种酶的活性显著升高，总体上提高了森林土壤的生产力。随腐烂等级升高，倒木接触处土壤有机碳含量升高，腐烂等级间有机碳含量差异显著，其他土壤特性在腐烂等级间差异不显著，季节及林型对倒木接触处土壤特性变化存在显著影响，土壤特性指标之间显著相关，其中，土壤有机碳与所选理化性质指标极显著相关，全氮与所选土壤酶指标极显著。有机碳、全N及C/N比3者成为影响该区5种酶活性的主要指标。综合看来，倒木的覆盖有利于提高土壤肥力质量，促进土壤环境的物质循环和能量流动。
（5）采用冗余分析（redundancy analysis，RDA）排序分析方法探讨倒木与环境的生态关系，结果表明，海拔是该研究区倒木各项特征分布的主要影响因素；所选环境因子中，海拔对倒木数量特征、养分贮量及能量贮量分布的影响效应最大，降雨量对碳贮量、氮贮量、磷贮量及潜在能量释放的影响最小。除人为干扰外，长苞铁杉林内倒木主要来源于雪灾或风害的干扰影响，较少产生于林木自然稀疏与衰老。越往南坡，高腐烂等级相对增加，土壤酶活性升高，倒木养分和能量的释放增加，说明温度的升高，会加速该区倒木的分解，缩短了倒木作为苗床使用的时间，后期苗床实验中倒木的放置不应过于靠南坡。坡度对倒木数量特征及接触处土壤酶活性的影响最小，对倒木养分及能量的释放影响效应中等；有幼苗着床的倒木主要出现在中坡位，第Ⅳ腐烂等级倒木上的生物多样性更丰富，也更有利于幼苗的着生。
因此，综合上述研究可以得出，天宝岩长苞铁杉林内倒木的存在，既有利于系统养分的固定，也提高了土壤的生产力及异质性，参与了生态系统的物质循环与能量流动，促进森林生态系统的稳定性发展；后期苗床实验中，若选择第Ⅳ腐烂等级倒木作为更新苗床，可置于中坡位，其坡度要求不大，但应注意倒木的放置不应过于靠南坡。
Tsuga longibracteata W. C. Cheng, a rare and endangered coniferous species endemic to China,plays an important role in promoting forest succession,maintaining the balance and stability offorest ecosystems, and water conservation. However, barren site condition and poor naturalregeneration were critical problem for the protection and restoration of Tsuga longibracteata forest.Fallen logs play important roles in maintaining the structural and functional integrity of forestecosystems;they provide habitat for forest species, act as a long-term nutrient store, and contributeto nutrient cycling,energy flow, regeneration succession and soil formation. These functionsprovide evidence for the importance of logs for forest ecosystem. A better knowledge of logs couldhelp in solving the problem of stability and regeneration in Tsuga longibracteata forest. Base onthe investigation on fallen logs quantity and quality,nutrient stock,energy stock and environmentalfactors（including topographic factors, human disturbance, stand structure, meteorological factors andsoil microenvironment under fallen logs）were track monitoring, the present study try to demonstratethe logs ecological function in nutrient cycling and energy flow,and found the right site for seedbedstudy.It also contribute to a reference basis for conservation and sustainable management of Tsugalongibracteata forest. The results showed that:
（1） The fallen logs in Tsuga longibracteata forest in Tianbaoyan National Nature Reserve,mainly belonged to slightly decayed and small-classed logs,that is,they mainly distributed at therange of decay class I~II, diameter class0~10cm, length class1~3m.Logs density were decreasedwhile volume increased with the increased of diameter class（length class）, Logs density, volumeand covered area were increased with the decreased of elevation.Logs density were decreased withthe increased of decay class, volume and covered area distribution were close to normal distribution.fallen logs quantity and quality were differed significantly among decay class and forest types （P<0.05）. fallen logs volume in this area was18.14m~3/hm~2, and the selective volume was only3.37m~3/hm~2when we chose class IV for seedbed study.
（2） The content of C, K were not significantly in different decay classes（P>0.05）, whileN, P and C/N were extremely significantly（P<0.01）; With the increased of the decayclass, thecontent of N, P displayed the increased trend, however C/N decreased; logs nutrient stocks inhighly decay class were larger than slightly decayed; and the total C stock was2.316t/hm~2,while N stock was9.832kg/hm~2,P stock was1.154kg/hm~2,K stock was1.713kg/hm~2. Nutrient stocks waslowest in Tsuga longibracteata pure forest, and largest in Tsuga longibracteata+Phyllostachysedulis forest, C stock in different forest types showed the following trend: Tsuga longibracteatapure forest
（4） Soil properties under fallen logs were analyzed by variance analysis,correlationanalysis,path analysis and factor analysis,the results showed that soil bulk density and pH werelower, while porosity, SOC, TN, TK, protease, urease, invertase, cellulose and acid phosphataseactivity were higher in soil beneath fallen logs, which indicated that fallen logs were contributed toimprove the soil productivity; SOC were increased with the increased of decay class, SOC wereextremely significant difference in soils under fallen logs of all decay classes, while the other soilproperties were no significant difference; season were significant effects on most soil properties（P<0.05）; Soil properties showed a significant correlation between each other, Highly significantcorrelation was found between SOC and the selected physical and chemical properties,while thesame between TN and the five soil enzyme activities. SOC,TN and C/N were the most importantindex on the five soil enzyme activities.Comprehensively, fallen logs was contributed to improvesoil quality, and promote nutrient cycling and energy flow in soil environment.
（5） Redundancy analysis（RDA） were performed to determine therelationships between fallenlogs and environmental factors, and the results showed that fallen logs quantity and quality was strongly effected by elevation, precipitation have the lowest effect on the release of C,N and Pstock. Except for human disturbance, fallen logs were not mainly from self-thinning and senescence,but mostly origin from snow or wind disturbance.The highly decayed logs, soil enzymeactivities,nutrient and energy release stock were increased from northern to southern slope, whichindicated that elevated temperature will accelerate the decomposition of fallen logs in the studyarea, and thus shorten the duration of its function as a seedbed, so sampling logs that use forseeding experiment should not be overly concentrated on southern slopes in further study. Slopegradient had the least effect on logs quantity and quality and soil enzyme activity under fallenlogs,but middle effect on nutrient and energy release,seedbed logs were found on middleslope,biodiversity was richer on class Ⅳ, which were advantage for seedlings colonization.
Thus,all the results suggested that fallen logs on Tsuga longibracteata forest floor acted aslong-term storage for nutrients,contributed to soil fertility and heterogeneity, the accumulation oflogs would lead to accelerated nutrient cycling and energy flow,which would promoted the stabilityof forest ecosystem.In addition, if the class Ⅳwas chooseen for seedbed experiment, they shouldput in middle slope,and without considering slope gradient, but not be overly concentrated onsouthern slope.
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