东北贫营养泥炭沼泽几种植物的种群生态学研究
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摘要
贫营养泥炭沼泽的形成是气候、地貌、水文等因素复合作用的结果，大多分布在亚寒带与寒温带地区。在我国，贫营养泥炭沼泽主要发育东北地区的大、小兴安岭和长白山地。本研究选择长白山西麓哈泥泥炭沼泽和小兴安岭南坡东段汤北林场泥炭沼泽作为研究区，对不同生活型的植物(主要包括中位泥炭藓、疣壁泥炭藓、白齿泥炭藓、桧叶金发藓、狭叶杜香、甸杜和米典苔草)进行了种群生态学研究。
(1) 六种植物种群构件的年龄结构以非增长型为主。疣壁泥炭藓、桧叶金发藓和甸杜种群分株以及米典苔草根茎构件呈衰退型年龄结构；中位泥炭藓、泥炭沼泽边缘的狭叶杜香以及米典苔草分株和活动芽构件均为稳定型年龄结构；仅泥炭沼泽开阔地的狭叶杜香种群分株呈增长型年龄结构。
(2) 所有五种常绿植物分株生物量均为衰退型年龄结构：米典苔草种群分株呈增长型年龄结构，但根茎呈衰退型年龄结构。
(3) 六种植物种群分株生物量均表现出年龄依赖规律。三种苔藓植物种群分株均重均随龄级增加呈线性增长规律，常绿小灌木种群分株平均生物量随龄级增加呈指数函数增加，狭叶杜香种群分株干物质积累基数大于甸杜(以沼泽边缘为例)但增长率较小；米典苔草种群因2a株生殖株比例高，2a株均重低，1a和3a分株均重相似。
(4) 三种苔藓植物和两种小灌木植物种群分株高度随龄级增加呈线性增加的规律。中位泥炭藓的高度的增长率大于疣壁泥炭藓。有无孢子体生产对于桧叶金发藓种群分株平均高度无显著影响，但孢子体生产限制了高度的生长速率，亦提高了各龄级分株高度的变异程度。
(5) 三种苔藓种群在生物量和高度生长两性状间均表现出等速生长规律。两种小灌木种群表现出异速生长规律，均侧重生物量的生长。
(6) 米典苔草根茎长度和生物量生长上未表现出异速生长规律。种群7、8月全体根茎单位长度生物量无显著差异，但1a根茎表现出将有限的干物质储备用于增加长度、扩展种群空间生态位的作用。
(7) 两种小灌木种群分株在高度和生物量两方面均表现出很高的生
态可塑性，泥炭沼泽边缘和的小灌木种群分株平均高度和平均生物量均显
著大于边缘。
(8)除乔木分布局限于沼泽边缘外，小灌木植物种群均表现出对低
pH值和低养分条件较强的耐受能力，在泥炭沼泽边缘和开阔地两种生境
中均有分布。除金露梅和大果毛篙豆外，小灌木种群盖度均以泥炭沼泽边
缘显著偏大。
(9)哈泥泥炭沼泽泥炭蓦植物以中位泥炭鲜盖度、统壁泥炭醉、偏
叶泥炭鲜为共优势种。尖叶泥炭鲜盖度在边缘和开阔地生境间无差异。中
位泥炭醉和流壁泥炭醉在开阔地中盖度较大，其他泥炭醉和桧叶金发醉则
相反。
(10)所有小灌木均分布于鲜丘上，除甸杜和大果毛篙豆外，不同种
群的在醉丘不同部位的盖度不同。小灌木种群在沼泽开阔地垂直空间生态
位略宽，即使在水深为Ocm处尚有少量分布。
(11)在醉丘斑块微生境中，中位泥炭鲜、毛壁泥炭鲜和疵壁泥炭醉
均可成为优势物种，而在丘间微生境中，白齿泥炭鲜(沼泽边缘)或偏叶
泥炭醉(沼泽开阔地)占优势;其他苔鲜物种则均为伴生种。中位泥炭鲜、
锈色泥炭鲜以及疵壁泥炭鲜两两之间以及三者与白齿泥炭醉、偏叶泥炭醉
间存在明显的垂直空间生态位分离现象。而中位泥炭鲜、锈色泥炭薛以及
庆壁泥炭醉与大泥炭醉、尖叶泥炭鲜和红叶泥炭鲜在垂直空间生态位上既
有分离又有重叠。
(12)三种泥炭鲜种群的最小和最大繁殖年龄均分别为la和4a，平
均生殖年龄均接近3a。桧叶金发鲜种群最小、最大以及平均繁殖年龄分
别为Za、6a和4.22a，狭叶杜香种群则分别为6a、16a和10.74a，甸
杜种群则分别为3a、sa和5.18a。研究中的七种植物种群生殖株年龄结
构均为明显的衰退型。
(13)四种苔鲜植物抱子体生产数量由大到小依次为疵壁泥炭醉>中
位泥炭鲜>白齿泥炭鲜>桧叶金发醉。抱子体均重由大到小依次为桧叶金
发鲜(0 .59 mg)>统壁泥炭醉(0 .34 mg)>中位泥炭鲜(0 .31 mg)>白
齿泥炭醉(0 .25 mg)。桧叶金发鲜抱子体均重分别约为中位泥炭鲜、疵壁
泥炭鲜和白齿泥炭醉的1.9、2.0和2.5倍。狭叶杜香和甸杜种群单株生产
花数平均分别为16.92个和7.15个，花均重分别为0.0089和0.0149。
(14)以现存的抱子体或花的生物量与分株地上部分总生物量比值衡
量生殖分配，除甸杜种群外，狭叶杜香、桧叶金发鲜、中位泥炭醉、白齿
泥炭鲜、统壁泥炭醉的繁殖分配均小于10%。按类群看，桧叶金发醉种
群繁殖分配最大、其次为小灌木植物，泥炭鲜植物最小。
(15)若使用抱子体的生物量与分株当年生长片段的生物量比值衡量
苔醉植物的繁殖分配，四种苔醉植物种群的繁殖分配由大到小依次为桧叶
金发鲜(18.34%)>白齿泥炭薛(2.29%)>中位泥炭鲜(2.11%)>疵
壁泥炭薛(2.08%)。
(16)四种苔辞植物在不同水深的条件下，均表现出不同的出生率、
死亡率和增长率等种群数量特征的变化。总体看来，桧叶金发醉种群对水
深变化的适应幅度最宽，其次为中位泥炭醉和疵壁泥炭鲜种群，二者相似，
白齿泥炭醉种群的适应幅度最窄。
(17)三种泥炭醉种群的分株生物量均与水深间存在线性相关关系，
均随水深的增加而增加。三种泥?
The formation of oligotrophic mires resulted from special complex effects of climate, physiognomy, hydrology etc. They are mainly distributed in sub-frigid zone and frigid-temperate zone. In China, the mountain region of Northeast China, including Daxing'an Mountains, Xiaoxing'an Mountains and Changbai Mountains are concentrated distribution areas of oligotrophic mires. Hani mire in the foot of Changbai Mountains and Tangbei Forestry Centre mire in the east part of the south slope of Xiaoxing'an Mountains were chosen as the study areas. Population ecology of plants, mainly including Sphagnum magellanicum, S. papillosum, S. girgensohnii, Polytrichum juniperum, Ledum palustre var. angustum, Chamaedaphne calyculata and Carex middendorffii were studied.(1) Non-increased age structure is predominant in the six plant populations. The ramets of S. papilloum, P. juniperum and Ch. calyculata and rhizomes of C. middendorffii showed a declined age structure. Ramets of S. magellanicum population and L. pallustre var. angustum population in open area and the ramets and active buds of C. middendorffii population showed stable age structure. Only ramets of I. pallustre var. angustum in the margin of mire showed an increased age structure.(2) Accumulative biomass of ramets in All the five evergreen plant populations showed a declined age structure, while accumulative biomass of ramets and that of rhizomes in C middendorffii population showed an increased and a declined age structure respectively.(3) Biomass of ramets in all the six populations showed age dependence. Mean biomass of ramets in every age class increased with a linear rule as the ramets aged in the three bryophyte populations, while an exponential increased rule was found in L. pallustre var. angustum population. In margin of the oligotrophic mire, base of dry materials accumulation in L. pallustre var. angustum was higher than that in Ch. Calyculata, but with a lower increscent rate. Due to a high proportion of sexual reproductive ramtes in 2 a ramets,
mean biomass of 1 a and 2 a ramets was similar, and 2 a ramets were lighter.(4) Height of ramtes in the three bryophytes and two dwarf shrubs populations linearly increased with the ramtes aging. The increscent rate of S. magellanicum was higher than that of S. papillosum. No significant difference was found in P. juniperum populations no matter with sporophyte production or not, but sporophyte production limited the increscent rate of height and increased the variation level of height of ramets in different age classes.(5) On the two hands of growth of biomass and height, the three bryophytes showed an equal velocity growth. The two dwarf shrubs populations showed allometry, and emphasized particularly on biomass growth.(6) On the two hands of growth of biomass and length in rhizomes of C. middendorffii populations, no allometry was found. There were no significant difference in mean biomass per unit length of rhizomes in C. middendorffii population in July and August, but 1 a rhizomes showed a role to increase length to expand spatial niche by using limited dry material accmulation.(7) The two dwarf shrubs populations showed high ecological plasticity in height and biomass. Mean height and biomass in the margin of the mire was significantly higher.(8) Except that arbor only grew in the margin of mires, dwarf shrubs showed a strong tolerant capacity in low pH and low nutrient conditions, and could be seen both in open area and margin of mires. The coverage of dwarf shrubs, but Dasiphora fruticosa and Oxycoccus quadripetalus, was significantly higher in the margin of mires(9) In Hani mire, no difference between coverage of S. acutifolium in the margin and that in the open area. S. magellanicum, S. papillosum and S subsecundum are dominant Sphagna. The coverage of S. magellanicum and S. papillosum are higher in the open area, while by contraries, other Sphagna and P. juniperum with a higher coverage in the margin of the mire.(10) All the dwarf shr
(1) 六种植物种群构件的年龄结构以非增长型为主。疣壁泥炭藓、桧叶金发藓和甸杜种群分株以及米典苔草根茎构件呈衰退型年龄结构；中位泥炭藓、泥炭沼泽边缘的狭叶杜香以及米典苔草分株和活动芽构件均为稳定型年龄结构；仅泥炭沼泽开阔地的狭叶杜香种群分株呈增长型年龄结构。
(2) 所有五种常绿植物分株生物量均为衰退型年龄结构：米典苔草种群分株呈增长型年龄结构，但根茎呈衰退型年龄结构。
(3) 六种植物种群分株生物量均表现出年龄依赖规律。三种苔藓植物种群分株均重均随龄级增加呈线性增长规律，常绿小灌木种群分株平均生物量随龄级增加呈指数函数增加，狭叶杜香种群分株干物质积累基数大于甸杜(以沼泽边缘为例)但增长率较小；米典苔草种群因2a株生殖株比例高，2a株均重低，1a和3a分株均重相似。
(4) 三种苔藓植物和两种小灌木植物种群分株高度随龄级增加呈线性增加的规律。中位泥炭藓的高度的增长率大于疣壁泥炭藓。有无孢子体生产对于桧叶金发藓种群分株平均高度无显著影响，但孢子体生产限制了高度的生长速率，亦提高了各龄级分株高度的变异程度。
(5) 三种苔藓种群在生物量和高度生长两性状间均表现出等速生长规律。两种小灌木种群表现出异速生长规律，均侧重生物量的生长。
(6) 米典苔草根茎长度和生物量生长上未表现出异速生长规律。种群7、8月全体根茎单位长度生物量无显著差异，但1a根茎表现出将有限的干物质储备用于增加长度、扩展种群空间生态位的作用。
(7) 两种小灌木种群分株在高度和生物量两方面均表现出很高的生
态可塑性，泥炭沼泽边缘和的小灌木种群分株平均高度和平均生物量均显
著大于边缘。
(8)除乔木分布局限于沼泽边缘外，小灌木植物种群均表现出对低
pH值和低养分条件较强的耐受能力，在泥炭沼泽边缘和开阔地两种生境
中均有分布。除金露梅和大果毛篙豆外，小灌木种群盖度均以泥炭沼泽边
缘显著偏大。
(9)哈泥泥炭沼泽泥炭蓦植物以中位泥炭鲜盖度、统壁泥炭醉、偏
叶泥炭鲜为共优势种。尖叶泥炭鲜盖度在边缘和开阔地生境间无差异。中
位泥炭醉和流壁泥炭醉在开阔地中盖度较大，其他泥炭醉和桧叶金发醉则
相反。
(10)所有小灌木均分布于鲜丘上，除甸杜和大果毛篙豆外，不同种
群的在醉丘不同部位的盖度不同。小灌木种群在沼泽开阔地垂直空间生态
位略宽，即使在水深为Ocm处尚有少量分布。
(11)在醉丘斑块微生境中，中位泥炭鲜、毛壁泥炭鲜和疵壁泥炭醉
均可成为优势物种，而在丘间微生境中，白齿泥炭鲜(沼泽边缘)或偏叶
泥炭醉(沼泽开阔地)占优势;其他苔鲜物种则均为伴生种。中位泥炭鲜、
锈色泥炭鲜以及疵壁泥炭鲜两两之间以及三者与白齿泥炭醉、偏叶泥炭醉
间存在明显的垂直空间生态位分离现象。而中位泥炭鲜、锈色泥炭薛以及
庆壁泥炭醉与大泥炭醉、尖叶泥炭鲜和红叶泥炭鲜在垂直空间生态位上既
有分离又有重叠。
(12)三种泥炭鲜种群的最小和最大繁殖年龄均分别为la和4a，平
均生殖年龄均接近3a。桧叶金发鲜种群最小、最大以及平均繁殖年龄分
别为Za、6a和4.22a，狭叶杜香种群则分别为6a、16a和10.74a，甸
杜种群则分别为3a、sa和5.18a。研究中的七种植物种群生殖株年龄结
构均为明显的衰退型。
(13)四种苔鲜植物抱子体生产数量由大到小依次为疵壁泥炭醉>中
位泥炭鲜>白齿泥炭鲜>桧叶金发醉。抱子体均重由大到小依次为桧叶金
发鲜(0 .59 mg)>统壁泥炭醉(0 .34 mg)>中位泥炭鲜(0 .31 mg)>白
齿泥炭醉(0 .25 mg)。桧叶金发鲜抱子体均重分别约为中位泥炭鲜、疵壁
泥炭鲜和白齿泥炭醉的1.9、2.0和2.5倍。狭叶杜香和甸杜种群单株生产
花数平均分别为16.92个和7.15个，花均重分别为0.0089和0.0149。
(14)以现存的抱子体或花的生物量与分株地上部分总生物量比值衡
量生殖分配，除甸杜种群外，狭叶杜香、桧叶金发鲜、中位泥炭醉、白齿
泥炭鲜、统壁泥炭醉的繁殖分配均小于10%。按类群看，桧叶金发醉种
群繁殖分配最大、其次为小灌木植物，泥炭鲜植物最小。
(15)若使用抱子体的生物量与分株当年生长片段的生物量比值衡量
苔醉植物的繁殖分配，四种苔醉植物种群的繁殖分配由大到小依次为桧叶
金发鲜(18.34%)>白齿泥炭薛(2.29%)>中位泥炭鲜(2.11%)>疵
壁泥炭薛(2.08%)。
(16)四种苔辞植物在不同水深的条件下，均表现出不同的出生率、
死亡率和增长率等种群数量特征的变化。总体看来，桧叶金发醉种群对水
深变化的适应幅度最宽，其次为中位泥炭醉和疵壁泥炭鲜种群，二者相似，
白齿泥炭醉种群的适应幅度最窄。
(17)三种泥炭醉种群的分株生物量均与水深间存在线性相关关系，
均随水深的增加而增加。三种泥?
The formation of oligotrophic mires resulted from special complex effects of climate, physiognomy, hydrology etc. They are mainly distributed in sub-frigid zone and frigid-temperate zone. In China, the mountain region of Northeast China, including Daxing'an Mountains, Xiaoxing'an Mountains and Changbai Mountains are concentrated distribution areas of oligotrophic mires. Hani mire in the foot of Changbai Mountains and Tangbei Forestry Centre mire in the east part of the south slope of Xiaoxing'an Mountains were chosen as the study areas. Population ecology of plants, mainly including Sphagnum magellanicum, S. papillosum, S. girgensohnii, Polytrichum juniperum, Ledum palustre var. angustum, Chamaedaphne calyculata and Carex middendorffii were studied.(1) Non-increased age structure is predominant in the six plant populations. The ramets of S. papilloum, P. juniperum and Ch. calyculata and rhizomes of C. middendorffii showed a declined age structure. Ramets of S. magellanicum population and L. pallustre var. angustum population in open area and the ramets and active buds of C. middendorffii population showed stable age structure. Only ramets of I. pallustre var. angustum in the margin of mire showed an increased age structure.(2) Accumulative biomass of ramets in All the five evergreen plant populations showed a declined age structure, while accumulative biomass of ramets and that of rhizomes in C middendorffii population showed an increased and a declined age structure respectively.(3) Biomass of ramets in all the six populations showed age dependence. Mean biomass of ramets in every age class increased with a linear rule as the ramets aged in the three bryophyte populations, while an exponential increased rule was found in L. pallustre var. angustum population. In margin of the oligotrophic mire, base of dry materials accumulation in L. pallustre var. angustum was higher than that in Ch. Calyculata, but with a lower increscent rate. Due to a high proportion of sexual reproductive ramtes in 2 a ramets,
mean biomass of 1 a and 2 a ramets was similar, and 2 a ramets were lighter.(4) Height of ramtes in the three bryophytes and two dwarf shrubs populations linearly increased with the ramtes aging. The increscent rate of S. magellanicum was higher than that of S. papillosum. No significant difference was found in P. juniperum populations no matter with sporophyte production or not, but sporophyte production limited the increscent rate of height and increased the variation level of height of ramets in different age classes.(5) On the two hands of growth of biomass and height, the three bryophytes showed an equal velocity growth. The two dwarf shrubs populations showed allometry, and emphasized particularly on biomass growth.(6) On the two hands of growth of biomass and length in rhizomes of C. middendorffii populations, no allometry was found. There were no significant difference in mean biomass per unit length of rhizomes in C. middendorffii population in July and August, but 1 a rhizomes showed a role to increase length to expand spatial niche by using limited dry material accmulation.(7) The two dwarf shrubs populations showed high ecological plasticity in height and biomass. Mean height and biomass in the margin of the mire was significantly higher.(8) Except that arbor only grew in the margin of mires, dwarf shrubs showed a strong tolerant capacity in low pH and low nutrient conditions, and could be seen both in open area and margin of mires. The coverage of dwarf shrubs, but Dasiphora fruticosa and Oxycoccus quadripetalus, was significantly higher in the margin of mires(9) In Hani mire, no difference between coverage of S. acutifolium in the margin and that in the open area. S. magellanicum, S. papillosum and S subsecundum are dominant Sphagna. The coverage of S. magellanicum and S. papillosum are higher in the open area, while by contraries, other Sphagna and P. juniperum with a higher coverage in the margin of the mire.(10) All the dwarf shr
引文
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