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Volume 43 Issue 5
Oct.  2022
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DU Y, BAO W K. Research progress on Pinus densata forest[J]. Journal of Sichuan Forestry Science and Technology, 2022, 43(5): 1−10 doi: 10.12172/202208220004
Citation: DU Y, BAO W K. Research progress on Pinus densata forest[J]. Journal of Sichuan Forestry Science and Technology, 2022, 43(5): 1−10 doi: 10.12172/202208220004

Research Progress on Pinus densata Forest


doi: 10.12172/202208220004
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  • Corresponding author: baowk@cib.ac.cn
  • Received Date: 2022-08-22
    Available Online: 2022-09-16
  • Publish Date: 2022-10-26
  • Pinus densata forest is a unique forest type in southwest China, and it is an important carbon pool with high ecological conservation value. However, previous studies on community classification and structural characteristics, and ecosystem function of Pinus densata forest are scattered, and cannot provide systematic theoretical support for the protection, management and resource utilization of Pinus densata forest. We synthesized the community classification and structural characteristics, biomass and productivity, and water conservation capacity of Pinus densata forest based on relevant literature. Results showed that: (1) The forest could be classified into six formations, nine association groups and five associations; (2) The community biomass of mature forest was 81.24~318.79 t·hm−2, of which arbor layer was 79.39~311.53 t·hm−2, the carbon density of arbor layer was 49.543~103.24 t·hm−2, and the annual productivity was 5.48~18.07 t·hm−2·a−1, of which arbor layer was 4.29~14.23 t·hm−2; (3) The canopy interception rate of the forest was 24.32%~28.87%, and the maximum water-holding capacity of moss layer, litter layer and soil (0~30 cm) layer were 8.69 t·hm−2, 117.27 t·hm−2 and 380.98 t·hm−2. And clarified the research contents that need further attention: (1) The current research cannot form a complete Pinus densata forest classification system, supplementary field investigations is needed to complete the relevant understanding of community characteristics; (2) Recent researches on biomass, carbon storage and productivity were concentrated in the arbor layer, and the research areas were concentrated in Linzhi City and Shangri-La County, data of other regions, understory vegetation and underground parts should be supplemented, and the variation laws and driving factors of biomass and productivity along various geographic gradients should be explored; (3) Recent researches on water conservation in mature forests were lacking, and the water-holding capacity of soil layer and moss layer should be focused.
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Research Progress on Pinus densata Forest

doi: 10.12172/202208220004
  • 1. Chinese Academy of Sciences Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China
  • 2. University of Chinese Academy of Sciences, Beijing 100049, China
  • Corresponding author: baowk@cib.ac.cn

Abstract: Pinus densata forest is a unique forest type in southwest China, and it is an important carbon pool with high ecological conservation value. However, previous studies on community classification and structural characteristics, and ecosystem function of Pinus densata forest are scattered, and cannot provide systematic theoretical support for the protection, management and resource utilization of Pinus densata forest. We synthesized the community classification and structural characteristics, biomass and productivity, and water conservation capacity of Pinus densata forest based on relevant literature. Results showed that: (1) The forest could be classified into six formations, nine association groups and five associations; (2) The community biomass of mature forest was 81.24~318.79 t·hm−2, of which arbor layer was 79.39~311.53 t·hm−2, the carbon density of arbor layer was 49.543~103.24 t·hm−2, and the annual productivity was 5.48~18.07 t·hm−2·a−1, of which arbor layer was 4.29~14.23 t·hm−2; (3) The canopy interception rate of the forest was 24.32%~28.87%, and the maximum water-holding capacity of moss layer, litter layer and soil (0~30 cm) layer were 8.69 t·hm−2, 117.27 t·hm−2 and 380.98 t·hm−2. And clarified the research contents that need further attention: (1) The current research cannot form a complete Pinus densata forest classification system, supplementary field investigations is needed to complete the relevant understanding of community characteristics; (2) Recent researches on biomass, carbon storage and productivity were concentrated in the arbor layer, and the research areas were concentrated in Linzhi City and Shangri-La County, data of other regions, understory vegetation and underground parts should be supplemented, and the variation laws and driving factors of biomass and productivity along various geographic gradients should be explored; (3) Recent researches on water conservation in mature forests were lacking, and the water-holding capacity of soil layer and moss layer should be focused.

  • 高山松(Pinus densata Mast.)林是我国西南山区特有的森林类型,分布范围大致为北纬28°~33°,东经93°~104°,东起四川岷江流域,西迄西藏朗县,北起四川道孚,南至云南永胜,在川西地区分布海拔为2 000~3 800 m,在滇西北为3 000~3 400 m,藏东南为2 600~3 500 m[1-3]。根据第六次全国森林资源清查(1999~2003年)数据,高山松林面积为180.47 × 104 hm2 [4],其蓄积量、生物量密度和碳密度在中国森林类型中均处于较高水平[5-6],是西南地区重要的碳库,在固碳释氧、固土保肥、水源涵养等方面发挥着重要作用[7-9],具有较高的生态保育价值。

    系统认识高山松林的群落类型与结构、生态系统服务功能等是非常有必要的,对于高山松林的科学保护与管理以及资源的合理开发利用具有重要的理论价值。系统查阅了1980~2020年发表的高山松林相关文献资料,包括文献58篇,专著5本。综述如下方面的研究现状:(1)高山松林群落类型与特征;(2)群落生物量与生产力;(3)生态系统水源涵养能力;并进一步梳理出当前需要聚焦的方向,为高山松林的深入研究提供基础。

    • 以《中国植被志》研编内容与规范[10]为群落分类标准,对已有资料进行梳理,将高山松林划分为6个群系9个群丛组6个群丛(见表1)。除上述群落类型外,高山松还可与丽江云杉(Picea likiangensis (Franch.) E. Pritzel)、川西云杉(Picea likiangensis var. rubescens Rehder & E. H. Wilson)、白桦(Betula platyphylla Suk.)、帽斗栎(Quercus guajavifolia H. Léveillé)等形成混交林,灌木层优势种还包括云南杜鹃(Rhododendron yunnanense Franch.)、大白杜鹃(R. decorum Franch.)、腋花杜鹃(R. racemosum Franch.)、圆锥山蚂蟥(Desmodium elegans Candolle)等,草本层优势种还包括秦岭槲蕨(Drynaria baronii Diels)、金茅(Eulalia speciosa (Debeaux) Kuntze)等[1, 14-16, 19]

      群系群丛组群丛文献来源
      高山松
      Pinus densata Mast.
      高山松高山松陈伟烈等, 1980[11]
      P. densata Mast.P. densata Mast.
      高山松-草本高山松-羊茅郭立群, 1982[12]
      P. densata Mast. - HerbP. densata Mast. - Festuca ovina L.
      高山松-尼泊尔大丁草罗建, 2008[13]
      P. densata Masters - Leibnitzia nepalensis (Kunze) Kitamura
      高山松-灌木高山松-矮高山栎吴征镒, 1987[14]
      P. densata Mast. - ShrubP. densata Masters - Q. monimotricha Handel-Mazzetti
      高山松-川滇高山栎四川植被协作组,1980[15]
      P. densata Masters - Q. aquifolioides Rehder & E. H. Wilson
      高山松-灌木-草本高山松-川滇高山栎-疏穗野青茅四川森林编辑委员会, 1992[16]
      P. densata Mast. - Shrub - HerbP. densata Masters - Q. aquifolioides Rehder & E. H. Wilson - Deyeuxia effusiflora Rendle
      高山松+长穗高山栎高山松+长穗高山栎-灌木-草本郭立群, 1982[12]
      P. densata Mast. + Quercus longispica (Hand.-Mazz.) A. CamusP. densata Mast. + Q. longispica (Hand.-Mazz.) A. Camus - Shrub - Herb
      高山松+川滇高山栎高山松+川滇高山栎-灌木-草本段代祥等, 2010[17]
      P. densata Mast. + Q. aquifolioides Rehder & E. H. WilsonP. densata Mast. + Q. aquifolioides Rehder & E. H. Wilson - Shrub - Herb
      高山松+大果红杉高山松+大果红杉-灌木-草本郭立群, 1982[12]
      P. densata Mast. + Larix potaninii var. australis A. Henry ex Handel-MazzettiP. densata Mast. + L. potaninii var. australis A. Henry ex Handel-Mazzetti - Shrub - Herb
      高山松+华山松高山松+华山松-灌木罗建, 2008[13]
      P. densata Mast. + P. armandii Franch.P. densata Mast. + Pinus armandii Franch. - Shrub
      高山松+山杨高山松+山杨-灌木-草本王雪, 2011[18]
      P. densata Mast. + Populus davidiana DodeP. densata Mast. + Populus davidiana Dode - Shrub - Herb

      Table 1.  The classification of Pinus densata forest

    • 少量研究对群落优势种高山松的胸径、高度和冠幅结构进行了分析。卢杰等[20]对林芝八一镇高山松种群调查发现,其径级结构、高度结构和冠幅结构均呈反“J”形,为增长型种群。在林芝地区高山松天然次生林中,幼龄林的胸径分布呈偏左的近似正态分布,随着林龄的增加,偏度和峰度均变小,中龄林为中间高两边低的近似正态分布[21]。对西藏林芝地区高山松种群格局的分析结果表明,其在幼苗-幼树-立木的发育过程中,空间格局由集群分布转变为随机或均匀分布[22-23]。左政等[24]对香格里拉的高山松林直径结构分析发现,其林分直径结构以迈耶负指数分布函数的拟合效果最好。

    2.   群落生物量与生产力
    • 在林分水平上,少量研究以蓄积量或林龄为自变量,建立了高山松林乔木层生物量回归模型(见表2);还有多个研究以香格里拉地区的高山松林为研究对象,构建了高山松林地上生物量遥感估测模型[28-33],但精度较低,均不超过80%。对高山松林群落生物量已有调查数据进行收集(见表3),结果表明,高山松成熟林群落生物量为81.24~318.79 t·hm−2,其中乔木层为79.39~311.53 t·hm−2,灌木层为0.105~1.49 t·hm−2,草本层为0.035~0.3318 t·hm−2,凋落物层为0.515~90.84 t·hm−2。群落细根生物量的研究仅1个,何永涛等[42]测定了林芝八一镇高山松林0~50 cm土层中细根生物量,为431.2 g·m−2,其中活细根生物量为326.1 g·m−2,死细根为105.1 g·m−2;在垂直分布上,69.6%的活细根生物量集中在0~10 cm,而40~50 cm土层内无细根分布。

      估算模型样本量相关系数文献来源
      B = 0.5168V + 33.237816R = 0.94Fang et al., 2001[25]
      B = 0.5272V1.079319R = 0.9978黄从德等, 2008[26]
      B = 162.21 / (1 + 3.6259e−0.0578a/R2 = 0.966徐冰等, 2010[4]
      B = 0.81V + 11.89/R2 = 0.912Qiu et al., 2020[27]
        注:B为林分生物量密度(t·hm−2),V为林分蓄积量(m3·hm−2),a为林龄(a)。

      Table 2.  Biomass density estimation model of Pinus densata forest

      研究区域林龄/a群落总生物量乔木层灌木层草本层凋落物文献来源
      地上部分地下部分
      西藏林芝20~10831.26~318.79//30.55~311.53///罗天祥, 1996[34]
      /101.84~210.59//////张剑等, 2008[35]
      成熟林////0.16330.33187.3437邹林红等, 2005[36]
      幼龄林/////1.491.79杨阳, 2013[37]
      中龄林/////1.757.81
      成熟林/////1.677.2
      云南香格里拉15////3.79651.6966/王利民等, 2006[38]
      /98.89~234.90//////程鹏飞等, 2011[39]
      //95.7//22.3892.2675.165岳彩荣, 2012[40]
      Yue, 2012
      40294.306264.47329.049293.5220.1050.0410.638吴兆录等, 1994[19]
      100231.495194.32436.5230.8240.1210.0350.515
      四川木里成熟林/419.661//1.490.218/杨劲夫, 2016[41]

      Table 3.  Community biomass of Pinus densata forest

      对高山松林碳储量的研究包括树种含碳率和群落碳密度两个方面。王金亮等[43]测定了香格里拉高山松林不同林龄不同器官的含碳率,树种平均含碳率为51.31%,与张坤[44]基于1978—1994年期间关于中国生物量的文献和全国第三次森林资源清查(1984—1988年)数据的计算结果接近,为50.09%。杨阳[37]测定了林芝地区不同林龄高山松林的草本和凋落物含碳量,幼龄林、中龄林和成熟林草本地上含碳量分别为441.44 g·kg−1,412.84 g·kg−1和433.30 g·kg−1;草本地下含碳量分别为389.12 g·kg−1,370.54 g·kg−1和393.12 g·kg−1;凋落物含碳率分别为500.90 g·kg−1,507.35 g·kg−1和480.37 g·kg−1。对高山松林群落碳储量已有调查数据进行收集,结果表明,高山松林乔木层碳密度为49.543~103.24 t·hm−2,灌木层碳密度为10.964 t·hm−2,草本层碳密度为0.868 t·hm−2,凋落物碳密度为1.43~18.68 t·hm−2,土壤层(0~100 cm)碳密度为216.274 t·hm−2 [39, 45-50]

    • 高山松林生产力的相关研究很少,仅3篇。吴兆录等[51]建立了高山松林乔木器官净第一性生产力的优化回归模型,对香格里拉吉迪林场林龄40年和100年的高山松林生产力进行了估算,结果分别为12.192 t·hm−2·a−1和10.013 t·hm-2·a−1,其中乔木层分别为12.160 t·hm−2·a−1和9.980 t·hm−2·a−1,主要分配在叶和树干中;灌木层分别为0.018 t·hm−2·a−1和0.021 t·hm−2·a−1,草本层分别为0.014 t·hm−2·a−1和0.012 t·hm−2·a−1。唐佳[52]估算出工布自然保护区高山松林年生产力为9.85 t·hm−2·a−1。基于四川、云南和西藏17个样地数据,高山松成熟林全林和乔木层生产力分别为5.48~18.07 t·hm−2·a−1和4.29~14.23 t·hm−2·a-1;在地理分布格局上,包括高山松林在内的温性松林生物生产力随纬度的增加而递减,随经度的增加而递增;而在水热分布格局上,呈现一种双曲面的递增函数,随降水量的增加呈自然对数递增,随温度的增加呈线性递增,随温暖指数和潜在增散量的增加也呈递增趋势[34]

    3.   水源涵养能力
    • 高山松林林冠截留量仅在色季拉山和纳帕海开展过研究,结果表明高山松林林冠截留率为24.32%~28.87%,且林冠截留量与降雨量间以幂函数拟合效果最好[53-55]

    • 高山松林林下苔藓层持水特性研究仅1例,陈甲瑞和王小兰[56]对色季拉山东坡高山松林的调查发现,林下苔藓层平均厚度为1.13 cm,生物量为1.02 t·hm−2,自然含水量为1.83 t·hm−2,最大持水量为8.69 t·hm−2;浸水实验表明,苔藓在前0.25 h内吸水速率最大,可达654.279 g·kg−1·h−1,随后迅速下降,2 h后下降速度减缓,8 h后吸水速率基本稳定并趋向于零,吸水速率与浸泡时间呈幂函数关系。

    • 已有研究对色季拉山成熟林以及纳帕海幼龄林和中龄林凋落物层持水能力和吸水特性进行了测定(见表4表5),结果表明高山松成熟林凋落物层最大持水量为117.27 t·hm−2;浸水实验发现,在第5 min时吸水速率最大,可达6210 g·kg−1·h−1,随后迅速下降,8~10 h后吸水量基本达到最大值,吸水速率与浸泡时间呈幂函数关系[59]

      研究区域林龄/a最大持水量/(t·hm−2最大持水率/%有效拦蓄量/(t·hm−2文献来源
      未分解层半分解层未分解层半分解层未分解层半分解层
      纳帕海1034.9512.82///////陆梅等, 2011[57]
      2566.7718.5///////
      2017.178.22///////
      1922.295.8116.48129.52136.39127.2614.984.3410.64周祥等, 2011[58]
      1935.79//138.48//15.67//石小亮等, 2015[55]
      色季拉山成熟林117.2753.4563.82 /106.54156.93 /30.9447.25喻武等, 2010[59]

      Table 4.  Water-holding capacity, water-holding rate, and effective interception of litter layer in Pinus densata forest

      研究区域林龄/a拟合方程适用范围/h相关系数文献来源
      纳帕海10Q= 2.056ln(t) + 15.4100~24R= 0.981陆梅等, 2011[57]
      QL= 1.637ln(t) + 7.0740~24R= 0.960
      V= 15.102t−0.8750~24/
      VL= 6.709t−0.7960~24/
      25Q= 4.129ln(t) + 36.0330~24R= 0.997
      QL= 2.283ln(t) + 10.7200~24R= 0.972
      V= 35.360t−0.8890~24/
      VL= 10.22t−0.8080~24/
      20Q= 0.773ln(t) + 6.4960~24R= 0.983
      QL= 0.983ln(t) + 5.0670~24R= 0.977
      V= 6.389t−0.8880~24/
      VL= 4.886t−0.8320~24/
      19Q= 3.341ln(t) + 13.220~24R2= 0.964周祥等, 2011[58]
      QL= 0.698ln(t) + 3.1950~24R2= 0.963
      QF= 2.643ln(t) + 10.0300~24R2= 0.929
      V= 1.127t−0.690~24R2= 0.953
      VL= 0.300t−0.780~24R2= 0.997
      VF= 0.798t−0.630~24R2= 0.893
      色季拉山成熟林QL= 108.28ln(t) + 37.480~24R2= 0.944喻武等, 2010[59]
      QF= 209.56ln(t) + 25.6320~24R2= 0.987
      VL= 14904t0~1/12R2= 0.987
      VL= 3406.5e(−0.4004t1/12~24R2= 0.987
      VF= 27648t0~1/12R2= 0.989
      VF= 5150.7e(−0.3843t1/12~24R2= 0.989
      Q= 13.143ln(t) + 34.9520~24R= 0.9856李菊和卢杰, 2014[60]
      Y= 68.453ln(t) + 182.040~24R= 0.9856
      V= 10264t-0.1.6420~24R= 0.9914
        注:Q为总持水量(t·hm−2),QL为未分解层持水量(t·hm−2,QF为半分解层持水量(t·hm−2),V为总吸水速率(t·hm−2·h−1),VL为未分解层吸水速率(t·hm−2·h−1),VF为半分解层吸水速率(t·hm−2·h−1),Y为持水率(%),t为浸泡时间(h)。

      Table 5.  Fitting equation of water-holding capacity, water absorption rate and water-holding rate of litter layer in Pinus densata forest

    • 土壤层持水特性的研究集中在林芝地区和纳帕海(见表6),结果表明高山松成熟林0~30 cm土层的土壤最大持水量为380.98 t·hm−2 [61]

      研究区域林龄/a土层/cm持水量/t·hm−2持水率%蓄水量/t·hm−2文献来源
      最大毛管非毛管最大毛管
      巴宜区幼龄林0~1076.5362.88/101.9883.83/刘永春, 1985[61]
      10~2080.869.65/86.4974.48/
      20~3080.3673.26/76.1369.42/
      30~4074.1864.48/68.8760.04/
      40~5077.9765.53/77.3964.93/
      成熟林0~1071.2861.28/157.15135.59/
      10~2080.4367.79/93.0178.19/
      20~3078.9267.47/89.4276.41/
      30~4069.9162.1/81.3569.32/
      40~5080.4472.29/82.4574.14/
      /0~130///47.6/5980王景升等, 2005[62]
      朗县/0~20868.81792.65///59张鹏等, 2019[63]
      香格里拉250~301827.6/406.8///周祥, 2011[53]
      30~501037.4/229.2///
      190~60//637.5//610石小亮等, 2015[55]

      Table 6.  Water-holding capacity, water-holding rate, and water storage capacity of soil layer in Pinus densata forest

    4.   结论与展望
    • 样方调查数据是认识群落类型和结构的基础资料,现有调查资料多形成于上世纪50—70年代,近期的调查集中在林芝市、香格里拉县等高山松林集中分布区;调查对象主要针对高山松纯林,对混交林的关注较少;研究空白点较多,基础数据严重缺乏。在群落类型的划分上,当前研究多基于群落外貌对群落类型进行划分,仅罗建[13]采用了数量分类方法,分类和命名标准不统一,无法形成完整的高山松林群落分类系统。在物种组成方面,已有研究仅列出各层片主要组成物种,缺乏对物种科属统计、生活型和地理区系成分特征的分析。在群落结构上,仅在胸径结构方面有少量研究,垂直结构[20]仅见1个报道。因此,当前对高山松林群落类型、生境条件、物种组成、群落结构等的认识是不全面的,后续需要对高山松林开展野外调查,明确其分布范围和地理分布格局,形成完整的群落分类系统,补全群落特征的相关认识。

    • 森林生物量、碳储量和生产力是研究森林生态系统结构和功能的重要基础数据,目前针对高山松林生物量、碳储量和生产力的研究集中在乔木层,且调查数据集中在林芝地区和香格里拉市,目前仅有1个研究[19]对高山松林各层片的生物量数据均进行了测算。针对高山松林这一群落类型的生物量和生产力空间分布格局和影响因素未见报道。因此,后续应对生物量数据缺失的区域开展补充调查,重点关注林下植被和地下部分的生物量测定;阐明高山松林生物量和生产力在各地理梯度上的变化规律及其驱动因子。

    • 涵养水源是森林生态系统的重要功能之一,当前高山松林水源涵养的研究集中在林芝地区和香格里拉市;研究对象以幼龄林和中龄林居多,成熟林水源涵养能力的数据严重缺乏;层次上以林冠层、凋落物层和土壤层的调查为主,苔藓层相关研究仅1例[56],已有研究对土壤层的调查深度标准不一。因此,后续应对成熟林水源涵养能力开展补充调查,重点关注土壤层和苔藓层的持水能力特性。

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