九连山自然保护区毛红椿天然种群生态学特征
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摘要
毛红椿(Toona ciliata var.pubescen)被列为国家二级保护濒危物种。目前面临资源破坏较严重,濒危局势日趋严峻。在自然保护区内,其分布区域的种群数量同样面临缩减趋势。本文以江西九连山国家自然保护区的天然群落为研究对象,从种群层面研究分析其有性、无性繁殖特性与种群动态变化特征,种子雨、土壤种子库以及林窗对其种群天然更新的影响,寻求制约其种群繁衍更新的关键因子,初步揭示其致濒机理。主要研究成果如下：
(1)通过调查九连山自然保护区现存虾公塘与大丘田2个毛红椿天然群落,分析其组成、结构及多样性特征。结果表明：群落中毛红椿占据优势地位,物种多样性指数大小依次是灌木层、草本层和乔本层,群落组成的植物种类多、结构较复杂,科属组成较为分散；分别共有维管束植物63科、95属和138种(含变种)和62科,94属,128种(含变种)。
(2)以虾公塘观测站毛红椿人工林为参照,通过天然群落结实标准株的野外花期观察和分析表明,毛红椿花期为5月底至7月初,2010年单枝花序开花数200朵以上,但天然林林分落花率超过70%,结实率较低。开花位置是影响毛红椿结实格局的主要因素,蒴果常集中在主花轴的中上部,垂直分布以树冠中、上层为主,水平分布东、南向居多。主成分分析表明,以结实母树数量、结实母树比率、乔冠层林分密度、母树胸径作为单株母树种子产量的测产依据。结果枝的结果数、结实率、单果种子数、种子饱满率均较低,分别在12-44、7.99%-13.19%、23～43、2.9%-40.4%之间,遗传变异系数2%～37%,单果枝结果数的遗传变化最大。
(3)毛红椿天然种群的种子雨和种子库等研究表明,种子雨散布时间为10月下旬至12月下旬,高峰期为11月中下旬。2010年种子雨强度251.6～284.7粒.m-2,以22.6-34.3粒.m-2完好种子供土壤萌发更新。天然种群的种子雨强度空间差异性不明显,随时间变化表现一定的差异性。种子库以3-6m树冠内的地表枯落层空间分布的种子密度最大,不同方位间种子的腐烂量及其腐烂率差异显著,北部居高。土壤种子储量取决于结实量、鸟类取食和种子活力保存等因素,鸟类取食是其种子储量大幅下降的首要因素；种子不耐储藏以及大量腐烂,有效贮藏期不足1个月。12月天然林不同种群结实单株种子库8.8～15.7粒.m-2,平均萌发数≤2株.m-2,次年1月土壤种子库种子量最少,6.7～11.8粒·m-2,平均仅萌发为0.4-0.6株.m-2,与林下实生幼苗分布极少吻合。
(4)林窗对毛红椿天然群落更新研究表明,天然林窗下种子发芽率5.98%,土壤湿度过高是影响种子发芽低的重要因子,光照是其主要影响因子,但不一定与其种子发芽率正相关。林窗内幼苗数量变化经历5-6月的第1次高死亡率,6-8月和越冬期间又出现第2和第3次较高死亡,其余月份间趋于平稳,12月毛红椿实生苗存活率仅2.89%。
(5)毛红椿种群动态及其生命活动的过程分析结果表明,天然种群变化以第Ⅰ龄级有较高的死亡率和致死率,进入Ⅱ龄级及以后,死亡率和致死率曲线总体变化趋于平缓,在Ⅶ龄级、Ⅷ龄级的死亡率和致死率小幅度上升。毛红椿天然种群结构波动性较大,幼龄个体数量少,中老年个体大,属衰退型种群。大丘田毛红椿天然种群以有性生殖为主,种群存活曲线趋于Deevey-Ⅲ型,天然有性生殖能力弱；虾公塘毛红椿天然种群以无性生殖为主,种群存活曲线趋于Deevey-Ⅰ型,具有一定的无性更新能力。
T. ciliata var. pubescens, a national secondary endangered species, is now facing serious resource destruction, and the situation of being endangered is getting worse even if being located in natural protection reserves. In order to explore endangered processes and mechanisms of natural population, and to seek the key factors to its population multiply and regeneration in this paper, materials as natural dominant community and its plantation of T. ciliata var. pubescens in Jiulianshan National Reserve in Jiangxi Province, the population dynamics characteristics being studied, including sexual and clone breeding traits, seed rain, soil seed bank and forest gaps for the effects on natural regeneration at population level. The main results show that:
(1) The observation data from T. ciliata ver pubescens natural communities in between in Xiagongtang and in Daqiutian station, and the comparison with the composition, structure and diversity were analyzed in Jiulianshan nature reserve. The results indicated that T. ciliata var pubescens occupied an advantage position on different communities, and plant species belonged to63families,95genera,138species and62families,94genera,128species, respectively. Their species diversity indices ware both in order layer of shrub, herb and arboraceous layer. The community composition was more plant species, complex structure, more dispersed of the family and genus composed.
(2) The reference to T. ciliata var pubescens plantation in Xiagongtang observatory, flowering of the standard seed setting trees were observed and analyzed by comparison with them. The results indicated flowering of T. ciliata var pubescens begins at end of May to early July.. Flowers per flowering branch were normally above200flowers in2010. However the flower dropping rate is above70percent in natural forest, which indicated that the seed setting rate was much low. The flowering position is the main spatial pattern of seed setting in different population, so fruiting branches were often concentrated in the middle and upper positions of main floral axis. The seed setting was mainly concentrated on vertical distribution of middle, upper canopy, and of east, south for horizontal distribution. Seed tree, seed yield and forest conditions were large difference between in different populations. The analysis of mainly factors indicated that the seed yield per seed tree mainly depended on the basis of predicted indexes such.as the number of seed trees, percent of seed trees, tree density on canopy layer, stem diameter of seed tree. The number of cones per branch and of seeds per cone, setting rate, percentage of full-developed seeds in difference forests were12～ 44,23～ 43,7.99%～ 13.19%,2.9%～ 40.4%, respectively. Their genetic variation coefficients were within2%～ 37%, of which, genetic variation in the number of cones per branch is the largest.
(3) The dynamics of seed rain and seed bank, seed germination and seedling status in T. ciliata var. pubescens natural populations were investigated and evaluated. Results showed the seed rain of T. ciliata var. pubescens was dispersed from late September to the end of November. Seed densities of different natural populations in2010decreased in order of:Xiagongtang protection station (284.7±24.2) seeds-m"2and Daqiutian protection station (251.6±24.7) seeds·m-2, and supplied intact seeds of34.3seeds·m-2, and22.6seeds·m-2into soil for seed germination update, respectively. The spatial differences of seed rain in natural populations were little significant, but a certain differences within time. The seed density of seed bank occupied largest over litter layer under3～6m canopy, the rotting rate and number of seeds rotted showed significantly difference in different populations, for north was the most. Seed bank reserves were affected by such factors as strong volume, bird feeding, seed viability, etc. of which birds feeding was the prime factor for a substantial drop of seed bank. A large number of seeds rotted with seed intolerance so that seeds could hardly be effectively stored beyond the length of one month. Soil seed reserves in December were8.8～15.7seeds m-2in natural populations, and its germination seedlings were less than2stands·m-2, the following year being the minimum of soil seed reserves (6.7～11.8seeds m-2) in January, and seedling of0.4～ 0.6stands·m-2, which was consistent with rare distribution of natural seedlings of T. ciliata var. pubescens, natural populations.
(4) Results from forest gaps for T. ciliata var pubescens regeneration in the natural community showed that its everage germination rate in forest gaps was extremely low to5.98%. High soil moisture is an important factor to low seed germination. Light is a main impact factor for seed germination, but was positively related uncentainly. Seedling dynamics passed the first time of high mortality rate within May-Jun., then the second of a low high mortality rate of within Jul.-Aug., and the same of the third for during winter, other times tended a stable change. On December there total lived38stands, survival rate of only2.89%.
(5) The analysis from the population dynamics and life process of T. ciliata var pubescens in Jiulianshan nature reserve showed a higher level of mortality rate and fatality rate at the I age class, they tended to level off totally entering and after the II age class, but a slight increase at ⅩⅡ, Ⅷ age class. Their population structures existed in a large fluctuation, of which, short of young seedlings, and the proportion of elderly individuals increasing, indicated that population of the species inclined to the type of decline. The breeding system of natural population in Xiagongtang is mainly sexual reproduction, but whose sexual reproduction was weak. Its survival curve of the natural population appeared to be a type of Deevey-Ⅰ. While clonal reproduction for population in Daqiutian, it showed a certain renewal ability to clone reproduction. And the survival curve of natural population appeared to be a type of Deevey-Ⅲ.
(1)通过调查九连山自然保护区现存虾公塘与大丘田2个毛红椿天然群落,分析其组成、结构及多样性特征。结果表明：群落中毛红椿占据优势地位,物种多样性指数大小依次是灌木层、草本层和乔本层,群落组成的植物种类多、结构较复杂,科属组成较为分散；分别共有维管束植物63科、95属和138种(含变种)和62科,94属,128种(含变种)。
(2)以虾公塘观测站毛红椿人工林为参照,通过天然群落结实标准株的野外花期观察和分析表明,毛红椿花期为5月底至7月初,2010年单枝花序开花数200朵以上,但天然林林分落花率超过70%,结实率较低。开花位置是影响毛红椿结实格局的主要因素,蒴果常集中在主花轴的中上部,垂直分布以树冠中、上层为主,水平分布东、南向居多。主成分分析表明,以结实母树数量、结实母树比率、乔冠层林分密度、母树胸径作为单株母树种子产量的测产依据。结果枝的结果数、结实率、单果种子数、种子饱满率均较低,分别在12-44、7.99%-13.19%、23～43、2.9%-40.4%之间,遗传变异系数2%～37%,单果枝结果数的遗传变化最大。
(3)毛红椿天然种群的种子雨和种子库等研究表明,种子雨散布时间为10月下旬至12月下旬,高峰期为11月中下旬。2010年种子雨强度251.6～284.7粒.m-2,以22.6-34.3粒.m-2完好种子供土壤萌发更新。天然种群的种子雨强度空间差异性不明显,随时间变化表现一定的差异性。种子库以3-6m树冠内的地表枯落层空间分布的种子密度最大,不同方位间种子的腐烂量及其腐烂率差异显著,北部居高。土壤种子储量取决于结实量、鸟类取食和种子活力保存等因素,鸟类取食是其种子储量大幅下降的首要因素；种子不耐储藏以及大量腐烂,有效贮藏期不足1个月。12月天然林不同种群结实单株种子库8.8～15.7粒.m-2,平均萌发数≤2株.m-2,次年1月土壤种子库种子量最少,6.7～11.8粒·m-2,平均仅萌发为0.4-0.6株.m-2,与林下实生幼苗分布极少吻合。
(4)林窗对毛红椿天然群落更新研究表明,天然林窗下种子发芽率5.98%,土壤湿度过高是影响种子发芽低的重要因子,光照是其主要影响因子,但不一定与其种子发芽率正相关。林窗内幼苗数量变化经历5-6月的第1次高死亡率,6-8月和越冬期间又出现第2和第3次较高死亡,其余月份间趋于平稳,12月毛红椿实生苗存活率仅2.89%。
(5)毛红椿种群动态及其生命活动的过程分析结果表明,天然种群变化以第Ⅰ龄级有较高的死亡率和致死率,进入Ⅱ龄级及以后,死亡率和致死率曲线总体变化趋于平缓,在Ⅶ龄级、Ⅷ龄级的死亡率和致死率小幅度上升。毛红椿天然种群结构波动性较大,幼龄个体数量少,中老年个体大,属衰退型种群。大丘田毛红椿天然种群以有性生殖为主,种群存活曲线趋于Deevey-Ⅲ型,天然有性生殖能力弱；虾公塘毛红椿天然种群以无性生殖为主,种群存活曲线趋于Deevey-Ⅰ型,具有一定的无性更新能力。
T. ciliata var. pubescens, a national secondary endangered species, is now facing serious resource destruction, and the situation of being endangered is getting worse even if being located in natural protection reserves. In order to explore endangered processes and mechanisms of natural population, and to seek the key factors to its population multiply and regeneration in this paper, materials as natural dominant community and its plantation of T. ciliata var. pubescens in Jiulianshan National Reserve in Jiangxi Province, the population dynamics characteristics being studied, including sexual and clone breeding traits, seed rain, soil seed bank and forest gaps for the effects on natural regeneration at population level. The main results show that:
(1) The observation data from T. ciliata ver pubescens natural communities in between in Xiagongtang and in Daqiutian station, and the comparison with the composition, structure and diversity were analyzed in Jiulianshan nature reserve. The results indicated that T. ciliata var pubescens occupied an advantage position on different communities, and plant species belonged to63families,95genera,138species and62families,94genera,128species, respectively. Their species diversity indices ware both in order layer of shrub, herb and arboraceous layer. The community composition was more plant species, complex structure, more dispersed of the family and genus composed.
(2) The reference to T. ciliata var pubescens plantation in Xiagongtang observatory, flowering of the standard seed setting trees were observed and analyzed by comparison with them. The results indicated flowering of T. ciliata var pubescens begins at end of May to early July.. Flowers per flowering branch were normally above200flowers in2010. However the flower dropping rate is above70percent in natural forest, which indicated that the seed setting rate was much low. The flowering position is the main spatial pattern of seed setting in different population, so fruiting branches were often concentrated in the middle and upper positions of main floral axis. The seed setting was mainly concentrated on vertical distribution of middle, upper canopy, and of east, south for horizontal distribution. Seed tree, seed yield and forest conditions were large difference between in different populations. The analysis of mainly factors indicated that the seed yield per seed tree mainly depended on the basis of predicted indexes such.as the number of seed trees, percent of seed trees, tree density on canopy layer, stem diameter of seed tree. The number of cones per branch and of seeds per cone, setting rate, percentage of full-developed seeds in difference forests were12～ 44,23～ 43,7.99%～ 13.19%,2.9%～ 40.4%, respectively. Their genetic variation coefficients were within2%～ 37%, of which, genetic variation in the number of cones per branch is the largest.
(3) The dynamics of seed rain and seed bank, seed germination and seedling status in T. ciliata var. pubescens natural populations were investigated and evaluated. Results showed the seed rain of T. ciliata var. pubescens was dispersed from late September to the end of November. Seed densities of different natural populations in2010decreased in order of:Xiagongtang protection station (284.7±24.2) seeds-m"2and Daqiutian protection station (251.6±24.7) seeds·m-2, and supplied intact seeds of34.3seeds·m-2, and22.6seeds·m-2into soil for seed germination update, respectively. The spatial differences of seed rain in natural populations were little significant, but a certain differences within time. The seed density of seed bank occupied largest over litter layer under3～6m canopy, the rotting rate and number of seeds rotted showed significantly difference in different populations, for north was the most. Seed bank reserves were affected by such factors as strong volume, bird feeding, seed viability, etc. of which birds feeding was the prime factor for a substantial drop of seed bank. A large number of seeds rotted with seed intolerance so that seeds could hardly be effectively stored beyond the length of one month. Soil seed reserves in December were8.8～15.7seeds m-2in natural populations, and its germination seedlings were less than2stands·m-2, the following year being the minimum of soil seed reserves (6.7～11.8seeds m-2) in January, and seedling of0.4～ 0.6stands·m-2, which was consistent with rare distribution of natural seedlings of T. ciliata var. pubescens, natural populations.
(4) Results from forest gaps for T. ciliata var pubescens regeneration in the natural community showed that its everage germination rate in forest gaps was extremely low to5.98%. High soil moisture is an important factor to low seed germination. Light is a main impact factor for seed germination, but was positively related uncentainly. Seedling dynamics passed the first time of high mortality rate within May-Jun., then the second of a low high mortality rate of within Jul.-Aug., and the same of the third for during winter, other times tended a stable change. On December there total lived38stands, survival rate of only2.89%.
(5) The analysis from the population dynamics and life process of T. ciliata var pubescens in Jiulianshan nature reserve showed a higher level of mortality rate and fatality rate at the I age class, they tended to level off totally entering and after the II age class, but a slight increase at ⅩⅡ, Ⅷ age class. Their population structures existed in a large fluctuation, of which, short of young seedlings, and the proportion of elderly individuals increasing, indicated that population of the species inclined to the type of decline. The breeding system of natural population in Xiagongtang is mainly sexual reproduction, but whose sexual reproduction was weak. Its survival curve of the natural population appeared to be a type of Deevey-Ⅰ. While clonal reproduction for population in Daqiutian, it showed a certain renewal ability to clone reproduction. And the survival curve of natural population appeared to be a type of Deevey-Ⅲ.
引文
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