长白山典型树种径向生长与气候因子的关系研究
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摘要
20世纪以来,全球气候变化对生态系统产生了深远的影响。大量研究表明物种对气候变化的响应差异会影响植物的群落结构与组成,最终导致一个地区植被类型的改变。因此,利用树木年轮学方法研究不同物种生长及其与气候因子的关系有利于定量研究气候变化对植物群落结构及组成的影响。本文以长白山地带性植被阔叶红松林的建群树种红松(Pinus koraiensis)和先锋树种山杨(Populus davidiana)以及暗针叶林建群树种鱼鳞云杉(Picea jezoensis var. komarovii)作为研究对象。设立6个海拔梯度获取树轮资料,分别为海拔600米(红松分布的下限,下文简称DX和XF),海拔800米(红松分布的中海拔地区,简称GM和DZ),海拔1000米(红松分布的中海拔地区,简称CBM1),海拔1200米(红松分布上限,鱼鳞云杉分布的下限,简称CBM2),海拔1400米(红松分布的生理极限,鱼鳞云杉分布的中心地带,简称CBM3)海拔1600米(鱼鳞云杉分布上限,简称CBM4)。从海拔,物种特性,性别和火干扰等方面分析树种径向生长特征及其与气候因子(上年5月到当年9月的月平均、最高和最低温度以及月降水量)的关系,并尝试模拟自然条件下树木径向生长与气候因子的关系,以期探讨气候变化对长白山典型树种生长的影响。主要结果如下：
(1)红松径级在海拔梯度上的分布存在差异,呈逐渐增加趋势,鱼鳞云杉径级则呈现递减趋势；各典型树种胸径与年龄的关系也有所不同,红松径级和年龄之间相关性较差,鱼鳞云杉和山杨的胸径则与年龄存在显著正相关关系(p<0.05)。在不同样地内的红松生长情况有差异,而鱼鳞云杉基本一致；同时相同采样点内不同树种径向生长量也存在一定差别,红松的生长量的年际变化均较大,山杨次之,鱼鳞云杉生长量变化幅度最小。由各典型树种宽度年表序列主分量的时间序列图可知共有的气候和环境条件及树种本身的特性对它们的径向生长年间变化具有一定影响,且主要集中在1971～1976年,1982～1994年和1999年～2006年间,但影响程度不同,因此有必要对典型树种与气候因子的响应关系进行系统研究。
(2)不同海拔梯度上,红松树木生长与气候因子的关系存在一定差异性,而鱼鳞云杉则较为一致。在红松分布的低海拔地区,红松的径向生长主要受到降水因子的影响,与当年7月降水正相关；随着海拔高度上升,温度对红松径向生长的作用凸显,与降水的相关性也发生了改变,在中海拔林区,红松径向生长均与当年4月的月平均温度和月最低温度显著正相关, GM采样点红松径向生长与上年8月的降水显著负相关, CBM1采样点红松径向生长则与上年8月的降水显著正相关；高海拔地区红松径向生长主要受到生长季温度的制约,与当年7月的月平均温度和月平均最低温度以及8月的月平均温度呈正相关关系。在极端气候条件下,影响红松径向生长的主要气候因子与径向生长对逐月气候因子的响应结果相比有所不同。尤其是低海拔地区,春季降水成为主要制约因子。与红松不同,在各海拔梯度上,鱼鳞云杉径向生长均与上一年7月的温度和当年5月的降水显著负相关,而与当年5月的月平均温度和月平均最低温度呈显著正相关。在极端高温的基础条件下,鱼鳞云杉径向生长主要受春季温度的影响,且均成正比。
(3)火干扰促进了红松的生长并使其对气候变化更为敏感。在火烧初期和1949-1981年间,火烧红松的径向生长量要显著高于未过火红松；火烧红松主要受到当年生长季温度和上一年生长季降水的影响,而未过火红松则主要受到上年生长季温度和当年降水的影响；同时在近5年内样地空间格局未发生变化的情况下,火烧红松更易受到相邻树木的影响。
(4)相同立地条件下不同树种对气候因子的响应关系有差异。阔叶红松林内,红松径向生长与上年11月、当年4月的月平均最低温度以及当年7月的降水量显著正相关,而与上一年7月和8月的降水负相关。山杨与当年1月的月平均温度,上一年6月和当年9月的降水显著负相关。在阔叶红松林和暗针叶林过渡带内,红松径向生长与当年7月平均温度显著正相关。鱼鳞云杉则与当年5月份的月平均温度显著正相关,而与该月降水显著负相关。
(5)不同性别间山杨植株生长量存在差异。近5年和10年内,山杨雄株的生长量显著高于雌株。山杨雌株与雄株的径向生长与降水因子的关系基本一致,均与生长季末(当年9月)的降水负相关；但与温度因子的关系则有所不同,山杨雌株分别受到冬季(上年10月和当年2月)的温度和春季降水(当年3月)的负而影响,而春季(当年3月)温度升高将促进雄株径向生长。
(6)除鱼鳞云杉外,红松和山杨雌雄植株的径向生长与气候因子的关系均能够被很好地模拟,尤其是火烧红松。结合长白山地区温度上升和降水将少的变化格局,可以预测出处于低海拔和火烧后遗留的红松生长将受到抑制；中海拔和高海拔的红松径向生长将得到促进；山杨雌株生长将降低,而山杨雄株生长在一定程度上将提高。
Since the 20th century, the effect of global climate change on ecological system had been more profound. Much research indicated that different response of plant to climate factor would influence the structure and composition of community and then lead the variation of vegetation pattern. Thus, study on radial growth dynamic of different plant and its' relationship with climate factors using dendroecological method will be in favor of quantitating the effect of climate change on the structure and composition of plant community. Pinus koraiensis, Populus davidiana and Picea jezoensis var. komarovii were selected in this paper as a typical species, pioneer tree in a zonal vegetation, broad-leaved Korean pine forest and a typical species in dark forest respectively. Tree-ring chronologies were developed at six altitudes:600 m (lower altitudinal border of Korean pine forest, later referred to as site DX and XF),800 m (middle altitudinal of Korean pine forest, site GM and DZ),1000 m (middle altitudinal of Korean pine forest, site CBM1),1200 m (high altitudinal border of Korean pine forest and lower altitudinal border of dard forest, site CBM2),1400m (ecological boundary of Pinus koraiensis and middle altitudinal border of Picea jezoensis var. komarovii, site CBM3), and 1600 m (high altitudinal border of Picea jezoensis var. komarovii, site CBM4). Radial growth dynamics of Pinus koraiensis, Populus davidiana and Picea jezoensis var. komarovii, and their relation with climate factors were analyzed from four aspects, elevation, fire disturbance species characteristics, and the gender. Climatic variables were monthly mean, maximum and minimum temperature and monthly total precipitation over the period from previous year May through current year September. We also try to establish the growth model in nature condition to discuss the effect of climate change on typical trees'growth. The main conclusions were as follows:
(1) The distributions of diameter at breast height (dbh) of Pinus koraiensis were different along the gradient of elevation. The dbh size increased with the elevation. There was a poor relationship between the dbh and age for Pinus koraiensis. The dbh of Picea jezoensis var. komarovii decreased with the elevation. The relation between the dbh and age was different from each site. There were positive relations between the dbh and age of Picea jezoensis var. komarovii and Populous davidiana (p<0.05). Tendency of Pinus koraiensis's increment varied among different sites. The increments of Picea jezoensis var. komarovii were accord with each site. Under the same environment condition, interannual variation of increment was different among three typical species, with the sharpest one was Pinus koraiensis, then Populus davidiana and the slowest one was Picea jezoensis var. komarovii. Through time series analysis of principle component of different species, we also found the interannual variations were influenced by common signals (climate, environmental condition and species characteristics). Though the impact was different, it concentrated on the period of 1971-1976,1982-1994 and 1999-2006. Therefore, it was necessary to analyze the relationship between the typical species and climate factors systemly.
(2) Along the altitudinal gradient, the relation between the trees'growth of Pinus koraiensis and climate factors was differential, but no difference happened for Picea jezoensis var. komarovii. At low altitude, Pinus koraiensis was positively controlled by precipitation, especially the precipitation in current July. Along the increasing of altitude, the importance of temperature increased and the correlation with precipitation changed. At middle altitude, Pinus koraiensis were positively driven by the mean and maximum temperature in current April. In other hand, the radial growth of Pinus koraiensis in GM site was negative related with the precipitation in previous August, but it turned into positive in CBM1 site. At high altitude, the radial growth of Pinus koraiensis were mainly positively influenced by growing season temperature, mean and minimum temperature in current July, mean temperature in current August. Different from above results, the analysis of climate characteristics in extreme years show that at low altitude, trees were strongly correlated with spring precipitation. Radial growth of Picea jezoensis van komarovii were in accord among different elevation, which negatively related with temperature in previous July and precipitation in current May, but positively significantly related with mean and minimum temperature in current May. Spring temperature had strong and positive impact on its growth in extreme year.
(3) Fire disturbance improve the trees'growth of Pinus koraiensis and make it to be more sensitive with climate change. At the early year of fire occurred and the period between the year of 1949 and 1981, the increment of fire-damaged Pinus koraiensis was significantly higher than the undamaged one. The radial growths of fire-damaged trees were driven by the growing season temperature and precipitation in previous growing season. Oppositely, the undamaged trees were mainly influenced by previous growing season temperature and current growing season precipitation. In addition, the fire-damaged trees were also significantly related with the neighbor competition under the assumption that the spatial pattern didn't changed in the lasted five years.
(4) Different response of differential species to climate factors occurred in the same environmental condition. It also added evidence that growth-climate relationship was species-characteristic. In broad-leaved Koran pine forest, the radial growth of Pinus koraiensis was positively controlled by minimum temperature in previous November and current April, and precipitation in current July, but negatively related with the precipitation in previous July and August. Whereas, Populus davidiana was significantly negative with mean temperature in current January, and precipitation in previous June and current September. In the transition zone of broad-leaved Korean pine forest and dark forest, the radial growth of Pinus koraiensis was significantly and positively related with mean temperature in current July, but Picea jezoensis var. komarovii was significantly positive with the mean temperature in current May, but negative with the precipitation in this month.
(5) Growth of the males of Populus davidiana was significantly higher than that of the females in the lasted 5 and 10 years. The females and males were significantly negative with the precipitation at the end of current growing season (current September). The females showed more sensitive with the climate change. The females were mainly negatively controlled by temperature in previous winter (previous October and current February) and spring precipitation (current March). The males were significantly positive with the spring temperature (current March).
(6) Except for Picea jezoensis var. komarovii, the relationship between the radial growth of Pinus koraiensis and Populus davidiana, and weather variables could be simulated, especially for fire-damaged Pinus koraiensis. Combining the climate variation pattern in Changbai Mountain with temperature increasing and precipitation decreasing, growth of Pinus koraiensis growing at low altitude and disturbed by fire would be inhibited. Growth of Pinus koraiensis at middle and high altitude would be benefit from the climate change. The growth of females of Populus davidiana would be decreased, but that of the males would be improved.
(1)红松径级在海拔梯度上的分布存在差异,呈逐渐增加趋势,鱼鳞云杉径级则呈现递减趋势；各典型树种胸径与年龄的关系也有所不同,红松径级和年龄之间相关性较差,鱼鳞云杉和山杨的胸径则与年龄存在显著正相关关系(p<0.05)。在不同样地内的红松生长情况有差异,而鱼鳞云杉基本一致；同时相同采样点内不同树种径向生长量也存在一定差别,红松的生长量的年际变化均较大,山杨次之,鱼鳞云杉生长量变化幅度最小。由各典型树种宽度年表序列主分量的时间序列图可知共有的气候和环境条件及树种本身的特性对它们的径向生长年间变化具有一定影响,且主要集中在1971～1976年,1982～1994年和1999年～2006年间,但影响程度不同,因此有必要对典型树种与气候因子的响应关系进行系统研究。
(2)不同海拔梯度上,红松树木生长与气候因子的关系存在一定差异性,而鱼鳞云杉则较为一致。在红松分布的低海拔地区,红松的径向生长主要受到降水因子的影响,与当年7月降水正相关；随着海拔高度上升,温度对红松径向生长的作用凸显,与降水的相关性也发生了改变,在中海拔林区,红松径向生长均与当年4月的月平均温度和月最低温度显著正相关, GM采样点红松径向生长与上年8月的降水显著负相关, CBM1采样点红松径向生长则与上年8月的降水显著正相关；高海拔地区红松径向生长主要受到生长季温度的制约,与当年7月的月平均温度和月平均最低温度以及8月的月平均温度呈正相关关系。在极端气候条件下,影响红松径向生长的主要气候因子与径向生长对逐月气候因子的响应结果相比有所不同。尤其是低海拔地区,春季降水成为主要制约因子。与红松不同,在各海拔梯度上,鱼鳞云杉径向生长均与上一年7月的温度和当年5月的降水显著负相关,而与当年5月的月平均温度和月平均最低温度呈显著正相关。在极端高温的基础条件下,鱼鳞云杉径向生长主要受春季温度的影响,且均成正比。
(3)火干扰促进了红松的生长并使其对气候变化更为敏感。在火烧初期和1949-1981年间,火烧红松的径向生长量要显著高于未过火红松；火烧红松主要受到当年生长季温度和上一年生长季降水的影响,而未过火红松则主要受到上年生长季温度和当年降水的影响；同时在近5年内样地空间格局未发生变化的情况下,火烧红松更易受到相邻树木的影响。
(4)相同立地条件下不同树种对气候因子的响应关系有差异。阔叶红松林内,红松径向生长与上年11月、当年4月的月平均最低温度以及当年7月的降水量显著正相关,而与上一年7月和8月的降水负相关。山杨与当年1月的月平均温度,上一年6月和当年9月的降水显著负相关。在阔叶红松林和暗针叶林过渡带内,红松径向生长与当年7月平均温度显著正相关。鱼鳞云杉则与当年5月份的月平均温度显著正相关,而与该月降水显著负相关。
(5)不同性别间山杨植株生长量存在差异。近5年和10年内,山杨雄株的生长量显著高于雌株。山杨雌株与雄株的径向生长与降水因子的关系基本一致,均与生长季末(当年9月)的降水负相关；但与温度因子的关系则有所不同,山杨雌株分别受到冬季(上年10月和当年2月)的温度和春季降水(当年3月)的负而影响,而春季(当年3月)温度升高将促进雄株径向生长。
(6)除鱼鳞云杉外,红松和山杨雌雄植株的径向生长与气候因子的关系均能够被很好地模拟,尤其是火烧红松。结合长白山地区温度上升和降水将少的变化格局,可以预测出处于低海拔和火烧后遗留的红松生长将受到抑制；中海拔和高海拔的红松径向生长将得到促进；山杨雌株生长将降低,而山杨雄株生长在一定程度上将提高。
Since the 20th century, the effect of global climate change on ecological system had been more profound. Much research indicated that different response of plant to climate factor would influence the structure and composition of community and then lead the variation of vegetation pattern. Thus, study on radial growth dynamic of different plant and its' relationship with climate factors using dendroecological method will be in favor of quantitating the effect of climate change on the structure and composition of plant community. Pinus koraiensis, Populus davidiana and Picea jezoensis var. komarovii were selected in this paper as a typical species, pioneer tree in a zonal vegetation, broad-leaved Korean pine forest and a typical species in dark forest respectively. Tree-ring chronologies were developed at six altitudes:600 m (lower altitudinal border of Korean pine forest, later referred to as site DX and XF),800 m (middle altitudinal of Korean pine forest, site GM and DZ),1000 m (middle altitudinal of Korean pine forest, site CBM1),1200 m (high altitudinal border of Korean pine forest and lower altitudinal border of dard forest, site CBM2),1400m (ecological boundary of Pinus koraiensis and middle altitudinal border of Picea jezoensis var. komarovii, site CBM3), and 1600 m (high altitudinal border of Picea jezoensis var. komarovii, site CBM4). Radial growth dynamics of Pinus koraiensis, Populus davidiana and Picea jezoensis var. komarovii, and their relation with climate factors were analyzed from four aspects, elevation, fire disturbance species characteristics, and the gender. Climatic variables were monthly mean, maximum and minimum temperature and monthly total precipitation over the period from previous year May through current year September. We also try to establish the growth model in nature condition to discuss the effect of climate change on typical trees'growth. The main conclusions were as follows:
(1) The distributions of diameter at breast height (dbh) of Pinus koraiensis were different along the gradient of elevation. The dbh size increased with the elevation. There was a poor relationship between the dbh and age for Pinus koraiensis. The dbh of Picea jezoensis var. komarovii decreased with the elevation. The relation between the dbh and age was different from each site. There were positive relations between the dbh and age of Picea jezoensis var. komarovii and Populous davidiana (p<0.05). Tendency of Pinus koraiensis's increment varied among different sites. The increments of Picea jezoensis var. komarovii were accord with each site. Under the same environment condition, interannual variation of increment was different among three typical species, with the sharpest one was Pinus koraiensis, then Populus davidiana and the slowest one was Picea jezoensis var. komarovii. Through time series analysis of principle component of different species, we also found the interannual variations were influenced by common signals (climate, environmental condition and species characteristics). Though the impact was different, it concentrated on the period of 1971-1976,1982-1994 and 1999-2006. Therefore, it was necessary to analyze the relationship between the typical species and climate factors systemly.
(2) Along the altitudinal gradient, the relation between the trees'growth of Pinus koraiensis and climate factors was differential, but no difference happened for Picea jezoensis var. komarovii. At low altitude, Pinus koraiensis was positively controlled by precipitation, especially the precipitation in current July. Along the increasing of altitude, the importance of temperature increased and the correlation with precipitation changed. At middle altitude, Pinus koraiensis were positively driven by the mean and maximum temperature in current April. In other hand, the radial growth of Pinus koraiensis in GM site was negative related with the precipitation in previous August, but it turned into positive in CBM1 site. At high altitude, the radial growth of Pinus koraiensis were mainly positively influenced by growing season temperature, mean and minimum temperature in current July, mean temperature in current August. Different from above results, the analysis of climate characteristics in extreme years show that at low altitude, trees were strongly correlated with spring precipitation. Radial growth of Picea jezoensis van komarovii were in accord among different elevation, which negatively related with temperature in previous July and precipitation in current May, but positively significantly related with mean and minimum temperature in current May. Spring temperature had strong and positive impact on its growth in extreme year.
(3) Fire disturbance improve the trees'growth of Pinus koraiensis and make it to be more sensitive with climate change. At the early year of fire occurred and the period between the year of 1949 and 1981, the increment of fire-damaged Pinus koraiensis was significantly higher than the undamaged one. The radial growths of fire-damaged trees were driven by the growing season temperature and precipitation in previous growing season. Oppositely, the undamaged trees were mainly influenced by previous growing season temperature and current growing season precipitation. In addition, the fire-damaged trees were also significantly related with the neighbor competition under the assumption that the spatial pattern didn't changed in the lasted five years.
(4) Different response of differential species to climate factors occurred in the same environmental condition. It also added evidence that growth-climate relationship was species-characteristic. In broad-leaved Koran pine forest, the radial growth of Pinus koraiensis was positively controlled by minimum temperature in previous November and current April, and precipitation in current July, but negatively related with the precipitation in previous July and August. Whereas, Populus davidiana was significantly negative with mean temperature in current January, and precipitation in previous June and current September. In the transition zone of broad-leaved Korean pine forest and dark forest, the radial growth of Pinus koraiensis was significantly and positively related with mean temperature in current July, but Picea jezoensis var. komarovii was significantly positive with the mean temperature in current May, but negative with the precipitation in this month.
(5) Growth of the males of Populus davidiana was significantly higher than that of the females in the lasted 5 and 10 years. The females and males were significantly negative with the precipitation at the end of current growing season (current September). The females showed more sensitive with the climate change. The females were mainly negatively controlled by temperature in previous winter (previous October and current February) and spring precipitation (current March). The males were significantly positive with the spring temperature (current March).
(6) Except for Picea jezoensis var. komarovii, the relationship between the radial growth of Pinus koraiensis and Populus davidiana, and weather variables could be simulated, especially for fire-damaged Pinus koraiensis. Combining the climate variation pattern in Changbai Mountain with temperature increasing and precipitation decreasing, growth of Pinus koraiensis growing at low altitude and disturbed by fire would be inhibited. Growth of Pinus koraiensis at middle and high altitude would be benefit from the climate change. The growth of females of Populus davidiana would be decreased, but that of the males would be improved.
引文
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