湖南会同亚热带次生阔叶林群落特征及种间关系研究
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摘要
湖南会同亚热带次生阔叶林地处于中国-日本森林植物区的核心地带,为中亚热带地带性植被人为破坏后自然恢复的植被类型,具有较强的过渡性。植物区系地理成分中,科属组成以泛热带分布为主,温带成分占有一定比例；种组成以中国特有分布和东亚分布为主,而温带成分占较大比例,并伴有一定的热带成分。研究的次生阔叶林群落内植物种类组成丰富,层次较为复杂,常绿性高于落叶性为研究的次生阔叶林典型代表。
为了研究亚热带次生阔叶林群落特征和种间关系,在代表性地段设置0.96hm2调查样地,样地内胸径大于4cm的林木1532株,分别隶属76种,27科,57属。群落物种丰富度Margalef指数(R)为10.28,Shannon-Wienner多样性指数(H)为5.11,生态优势度指数Simpson多样性指数(λ)值为0.05,Pielou均匀度指数为0.82。样地的植物种类丰富,在群落内的分布较均匀,群落的生态优势度值小,群落中没有占绝对优势的优势种,为多个优势种形成了群落共建种。群落的优势种群为拟赤杨(Alniphyllum fortunei)、杉木(Cunninghamia lanceolata)、南酸束(Choerospondias axillaris)、马尾松(Pinus massoniana)、枫香(Liquidambar formosana)、青冈栎(Cyclobalanopsis glauca)、檵木(Loropetalum chinense)、狭叶润楠(Machilus rehderii)、刨花楠(M. pauhoi)等,这些树种优势度为72.62%,在群落中占有明显优势,决定着森林群落的结构和演替的趋势。
会同亚热带次生阔叶林群落所有林木、主要优势种群中演替后期植物的直径结构为呈“J”型分布,而演替早期种的直径结构呈现多峰或单峰型分布；在垂直方向上,群落及其主要优势种群的树高结构分布呈偏左正态分布,说明该群落存在种内和种间的光资源的竞争。
采用Hegyi的单木竞争指数模型,对调查群落中优势种的地上部分种内与种间竞争强度进行定量分析。群落内13个优势种的竞争强度排序为：马尾松<刨花楠<红皮树<毛豹皮樟<赤杨叶<城口桤叶树<枫香<青冈栎<狭叶润楠<杉木<檵木<南酸枣<黄杞。其中,种内竞争大于种间竞争的优势种群为狭叶润楠、杉木、南酸枣、黄杞；种内竞争小于种间竞争的有马尾松、刨花楠、红皮树、毛豹皮樟、城口桤叶树、枫香、青冈栎；种内和种间竞争相差不大的树种为檵木和赤杨叶。各优势树种与上层树种之间的竞争较下层强烈,树木竞争强度与胸径呈负相关。
地上竞争改变林木的生存环境和林地土壤特性,从而对地下根系产生影响,而林木细根是受环境影响反应最敏感的部分,其形态、构型及分布会随着环境因子的改变而发生变化。选择群落内拟赤杨、枫香和青冈等3个优势树种,探讨亚热带次生阔叶混交林中林木之间的竞争强度以及林木细根分布、形态对竞争的适应状况。3个树种竞争指数CI的大小为：青冈(2.16)>拟赤杨(1.69)>枫香(0.97)。在0-45cm土层中,拟赤杨、枫香和青冈的活细根总生物量分别为104.1 g·m-2、122.4 g·m-2和54.4 g·m-2,3个树种的细根生物量主要分布在0-30cm土层,且均随着土壤深度的增加而明显减少(p<0.05)。拟赤杨和枫香的细根生物量在各土层中的分布规律较为相似,而青冈细根的分布却与其它两个树种的不同。在水平分布上,拟赤杨、枫香和青冈的细根生物量无显著差异(p>0.05),其中拟赤杨和枫香的细根各水平分布较为均匀,而青冈的细根靠树干近的分布较高。
各树种细根根长密度、根表面积密度与生物量垂直分布和水平分布方式相似,随着土壤深度的增加而明显减少,拟赤杨和枫香的细根根长密度和根表面积密度的水平分布较均匀,而青冈表现出靠近树干分布的趋势。在垂直分布上,枫香的比根长(SRL)、比表面积(SRA)和比根尖密度(SRT)在0-15cm的表层土壤中均高于拟赤杨和青冈,并随着土壤深度的增加而逐渐减少。与枫香相反,青冈的SRL、SRA和SRT随着土壤深度增加而增加。在水平分布上,枫香和青冈的SRL、SRA和SRT分别在不同的水平距离处高于其它树种。拟赤杨的SRL、SRA和SRT在各土壤层次和各水平距离处分布均匀。青冈的竞争指数最大,细根适应性较强,在分布和构型(SRL、SRA和SRT)具有较大的可塑性,可与枫香共存,而拟赤杨的可塑性较差,对竞争的响应不明显。
用Ripley's K(t)点格局分析函数研究群落各优势树种的分布格局以及种群间联结关系。群落内所有林木、拟赤杨、马尾松和枫香呈聚集分布格局,南酸枣和青冈栎则呈现局部小尺度上的聚集或随机分布。南酸枣和拟赤杨、拟赤杨和马尾松之间的空间分布呈负相关。植物主要竞争地上生长空间和光资源,南酸枣和马尾松、青冈栎和枫香之间呈现负相关。如果植物具有相似的竞争能力或竞争被某些外部因素削弱,植物可以共存,如拟赤杨、枫香和马尾松3者之间的空间联结关系为正相关的共存关系。除资源利用生态位分化外,群落内因倒木形成林窗,优势树种的共存与生长率、耐荫性、树冠存留时间和植物繁殖能力密切相关,促进了植物更新,如青冈栎与枯死木为正相关。种内聚集可以增加种内竞争事件,使系统向稳定共存条件转化,群落优势树种的种内聚集分布,缓解了种间竞争。随着群落演替,次生阔叶林中落叶树种将被演替后期的常绿阔叶树种所替代。
Secondary subtropical broadleaved forest in Huitong County, southwest of Hunan Province, southern China, is at the central of Sino-Japan bryoflora. The flora in secondary subtropical broadleaved forest is characterized with abundant plant species containing many ancient and relict plants, complex and wide connected geographical elements, and abundant endemic species. The floristic elements at the family and genus levels are mainly pan tropical with a certain proportion of temperate elements, whereas at the species level, species endemic to China and species of East Asia are the dominants with larger proportion of temperate elements and certain proportion tropical elements. Plant species diversity in the forest area is richer than that in other secondary forests in the same region, which reveals that the community is disturbed moderately before. Plant species in Huitong subtropical secondary forest are high.Tree species in overstory was dominated by Cunninghamia lanceolata, Pinus massoniana, Alniphyllum fortunei, Litsea coreana var. lanuginose, Machilus rehderii, M. pauhoi, Myrica rubra, Choerospondias axillaris, Cyclobalanopsis glauca, and C. myrsinaefolia. Shrub plant in understory was mainly dominated by Camellia oleifera, Loropetalum chinense, Photinia parvifolia, Rhododendron ovatum, Rh. Mariesii, Eurya loquiana, and Quercus serrata var. brevipetiolata.
A 9600 m2 rectangular plot was established to map the location of all trees greater than 4 cm in diameter at breast height (DBH), identify plant species and measure DBH, and total tree height (H). For the forest community, Margalef species richness index, Shannon-Wiener diversity index and Pielou evenness index was 10.28,5.11 and 0.82, respectively, indicating that secondary forest in Huitong had rich and evenness floristic composition. According to dominance value (72.62%), A fortunei, C. lanceolata, C.s axillaris, P. massoniana, Liquidambar formosana, C. glauca, L. chinense, Machilus rehderii, and M. pauhoi were the main dominant tree species for the secondary forest in Huitong.
A reverse-J shaped DBH classes distribution was observed for all stems and trees of later serial species while trees of earlier successional species distributed irregularly. The structures of the height distribution were normal-left distribution for the stand and it's main dominant populations of the Huitong subtropical secondary forest community, and indirectly indicated that both intra- and inter- specific competitions were exist for light resources.
Hegyi's competition index model was used to investigate intraspecific and interspecific competition intensity for dominant species trees. The results showed that competition intensity of the top 13 dominant plant species was ranked in the order of P. massoniana< M. pauhoi< Styrax suberifolius< L. coreana var. lanuginose< A. fortunei< Clethra fargesii< L. formosana< C. glauca< M. rehderii< C. lanceolata < L. chinense< C. axillaries< Engelhardtia roxburghiana. The tree species whose intraspecific competition was more intense than that of interspecific competition were M. rehderii, C. lanceolata, C. axillaris and E. roxburghiana. On the contrary, the tree species with lower intraspecific competition than that of interspecific competition were P. massoniana, M.s pauho, S. suberifolius, L. coreana var. lanuginosa, C. fargesii, L. formosana, C. glauca. The tree species that had an approximate intraspecific and interspecific competition index were L. chinense and A. fortuneii. The dominant and higher trees showed a strong competition intense compared to the lower trees. Relationship between the competition index and the diameter at breast height of objective tree could be expressed as power function and competition intensity appeared to reduce as the diameter at breast height of objective tree increased.
Competition is common natural phenomena. Generally, competition between tree species could be divided into aboveground competition for light and belowground competition for water and nutrient resources in soil. Environmental factors in soil and competition would shift fine root distribution, morphology and architecture. A. fortunei, L. formosan and C. glauca were selected as target to investigate root distribution and morphology in response to abovegroud competition. Competition index (CI) of A. fortunei, L. acakycina and C. glauca was 1.69,0.97 and 2.16, respectively. Total fine root biomass of A. fortunei, L. formosan and C. glauca within 0-45cm soil depth accouted for 104.1g·m-2,122.4 g·m-2 and 54.4 g·m-2, most of which distributed at top 0-30cm soil depth. Fine root biomas of A. fortunei and L. formosan were even distributed in soil depth, but there were soil depth distribution differences between C. glauca and A. fortunei, L. formosana. For horizontal distribution, there were no significant differences among fine root biomass of A. fortunei, L. formosan and C. glauca. (p>0.05) Fine roots of C. glauca generally occupied the places nearby trees, while fine roots of A. fortunei and L. acakycina were not.
Vertical and horizontal distribution of root length density (RLD) and specific root area (SRA) of three tree species was similar to that of fine root biomass. RLD and SRA were significantly decreased with increasing soil depth. RLD and SRA of A. fortunei and L. formosan were similar in different horizontal distance, while RLD and SRA of C. glauca were higher closed to tree species. In terms of vertical distribution, specific root length (SRL), specific root area (SRA) and specific root tips (SRT) of L. formosan which decreased significantly with increased soil depth were higher than that of A. fortunei and C. glauca in surface 0-15cm. In contrast, SRL, SRA and SRT increased with increasing soil depth. In terms of horizontal distribution, SRL, SRA and SRT of L. formosan and C. glauca were higher than other tree species in different horizontal distances, while those of A. fortunei were not predominant in each soil layers and each horizontal distance. Although C. glauca was subjected to strong competition and competitive index was high, its fine roots had great adaptability to changed environment (differential distribution, higher SRL, SRA and SRT), and it can coexist with L. formosan for a long time. On the contrary, suffered from competition from aboveground and belowground, A. fortunei is disadvantageous when competed with L. formosan and C. glauca.
To test the following two hypotheses whether the spatial distribution of dominant tree species is aggregated in subtropical secondary forests, and whether this spatial pattern can be used to infer causal factors for the coexistence of diverse tree species and predict forest succession trends, we employed Ripley's K(t) function analyze spatial patterns and associations. All trees within the forest showed a significant aggregated spatial pattern as did trees of each dominant species. Negative spatial association was detected for trees of C. axillaris versus A. fortunei, L. formosan or Pinus massoniana, and trees of Cyclobalanopsis glauca against A. fortunei, L. formosan, C. axillaris or P. massoniana. The apparent aggregation of spatial distribution for trees of each dominant species is indicative for mosaic patches that partition resources and maintain diverse species coexistence. Repulsive or independent spatial associations between trees of later and earlier serial species demonstrate the gap regeneration process in forest dynamics. Pioneer deciduous species will eventually be replaced by late serial evergreen species within the secondary forest.
为了研究亚热带次生阔叶林群落特征和种间关系,在代表性地段设置0.96hm2调查样地,样地内胸径大于4cm的林木1532株,分别隶属76种,27科,57属。群落物种丰富度Margalef指数(R)为10.28,Shannon-Wienner多样性指数(H)为5.11,生态优势度指数Simpson多样性指数(λ)值为0.05,Pielou均匀度指数为0.82。样地的植物种类丰富,在群落内的分布较均匀,群落的生态优势度值小,群落中没有占绝对优势的优势种,为多个优势种形成了群落共建种。群落的优势种群为拟赤杨(Alniphyllum fortunei)、杉木(Cunninghamia lanceolata)、南酸束(Choerospondias axillaris)、马尾松(Pinus massoniana)、枫香(Liquidambar formosana)、青冈栎(Cyclobalanopsis glauca)、檵木(Loropetalum chinense)、狭叶润楠(Machilus rehderii)、刨花楠(M. pauhoi)等,这些树种优势度为72.62%,在群落中占有明显优势,决定着森林群落的结构和演替的趋势。
会同亚热带次生阔叶林群落所有林木、主要优势种群中演替后期植物的直径结构为呈“J”型分布,而演替早期种的直径结构呈现多峰或单峰型分布；在垂直方向上,群落及其主要优势种群的树高结构分布呈偏左正态分布,说明该群落存在种内和种间的光资源的竞争。
采用Hegyi的单木竞争指数模型,对调查群落中优势种的地上部分种内与种间竞争强度进行定量分析。群落内13个优势种的竞争强度排序为：马尾松<刨花楠<红皮树<毛豹皮樟<赤杨叶<城口桤叶树<枫香<青冈栎<狭叶润楠<杉木<檵木<南酸枣<黄杞。其中,种内竞争大于种间竞争的优势种群为狭叶润楠、杉木、南酸枣、黄杞；种内竞争小于种间竞争的有马尾松、刨花楠、红皮树、毛豹皮樟、城口桤叶树、枫香、青冈栎；种内和种间竞争相差不大的树种为檵木和赤杨叶。各优势树种与上层树种之间的竞争较下层强烈,树木竞争强度与胸径呈负相关。
地上竞争改变林木的生存环境和林地土壤特性,从而对地下根系产生影响,而林木细根是受环境影响反应最敏感的部分,其形态、构型及分布会随着环境因子的改变而发生变化。选择群落内拟赤杨、枫香和青冈等3个优势树种,探讨亚热带次生阔叶混交林中林木之间的竞争强度以及林木细根分布、形态对竞争的适应状况。3个树种竞争指数CI的大小为：青冈(2.16)>拟赤杨(1.69)>枫香(0.97)。在0-45cm土层中,拟赤杨、枫香和青冈的活细根总生物量分别为104.1 g·m-2、122.4 g·m-2和54.4 g·m-2,3个树种的细根生物量主要分布在0-30cm土层,且均随着土壤深度的增加而明显减少(p<0.05)。拟赤杨和枫香的细根生物量在各土层中的分布规律较为相似,而青冈细根的分布却与其它两个树种的不同。在水平分布上,拟赤杨、枫香和青冈的细根生物量无显著差异(p>0.05),其中拟赤杨和枫香的细根各水平分布较为均匀,而青冈的细根靠树干近的分布较高。
各树种细根根长密度、根表面积密度与生物量垂直分布和水平分布方式相似,随着土壤深度的增加而明显减少,拟赤杨和枫香的细根根长密度和根表面积密度的水平分布较均匀,而青冈表现出靠近树干分布的趋势。在垂直分布上,枫香的比根长(SRL)、比表面积(SRA)和比根尖密度(SRT)在0-15cm的表层土壤中均高于拟赤杨和青冈,并随着土壤深度的增加而逐渐减少。与枫香相反,青冈的SRL、SRA和SRT随着土壤深度增加而增加。在水平分布上,枫香和青冈的SRL、SRA和SRT分别在不同的水平距离处高于其它树种。拟赤杨的SRL、SRA和SRT在各土壤层次和各水平距离处分布均匀。青冈的竞争指数最大,细根适应性较强,在分布和构型(SRL、SRA和SRT)具有较大的可塑性,可与枫香共存,而拟赤杨的可塑性较差,对竞争的响应不明显。
用Ripley's K(t)点格局分析函数研究群落各优势树种的分布格局以及种群间联结关系。群落内所有林木、拟赤杨、马尾松和枫香呈聚集分布格局,南酸枣和青冈栎则呈现局部小尺度上的聚集或随机分布。南酸枣和拟赤杨、拟赤杨和马尾松之间的空间分布呈负相关。植物主要竞争地上生长空间和光资源,南酸枣和马尾松、青冈栎和枫香之间呈现负相关。如果植物具有相似的竞争能力或竞争被某些外部因素削弱,植物可以共存,如拟赤杨、枫香和马尾松3者之间的空间联结关系为正相关的共存关系。除资源利用生态位分化外,群落内因倒木形成林窗,优势树种的共存与生长率、耐荫性、树冠存留时间和植物繁殖能力密切相关,促进了植物更新,如青冈栎与枯死木为正相关。种内聚集可以增加种内竞争事件,使系统向稳定共存条件转化,群落优势树种的种内聚集分布,缓解了种间竞争。随着群落演替,次生阔叶林中落叶树种将被演替后期的常绿阔叶树种所替代。
Secondary subtropical broadleaved forest in Huitong County, southwest of Hunan Province, southern China, is at the central of Sino-Japan bryoflora. The flora in secondary subtropical broadleaved forest is characterized with abundant plant species containing many ancient and relict plants, complex and wide connected geographical elements, and abundant endemic species. The floristic elements at the family and genus levels are mainly pan tropical with a certain proportion of temperate elements, whereas at the species level, species endemic to China and species of East Asia are the dominants with larger proportion of temperate elements and certain proportion tropical elements. Plant species diversity in the forest area is richer than that in other secondary forests in the same region, which reveals that the community is disturbed moderately before. Plant species in Huitong subtropical secondary forest are high.Tree species in overstory was dominated by Cunninghamia lanceolata, Pinus massoniana, Alniphyllum fortunei, Litsea coreana var. lanuginose, Machilus rehderii, M. pauhoi, Myrica rubra, Choerospondias axillaris, Cyclobalanopsis glauca, and C. myrsinaefolia. Shrub plant in understory was mainly dominated by Camellia oleifera, Loropetalum chinense, Photinia parvifolia, Rhododendron ovatum, Rh. Mariesii, Eurya loquiana, and Quercus serrata var. brevipetiolata.
A 9600 m2 rectangular plot was established to map the location of all trees greater than 4 cm in diameter at breast height (DBH), identify plant species and measure DBH, and total tree height (H). For the forest community, Margalef species richness index, Shannon-Wiener diversity index and Pielou evenness index was 10.28,5.11 and 0.82, respectively, indicating that secondary forest in Huitong had rich and evenness floristic composition. According to dominance value (72.62%), A fortunei, C. lanceolata, C.s axillaris, P. massoniana, Liquidambar formosana, C. glauca, L. chinense, Machilus rehderii, and M. pauhoi were the main dominant tree species for the secondary forest in Huitong.
A reverse-J shaped DBH classes distribution was observed for all stems and trees of later serial species while trees of earlier successional species distributed irregularly. The structures of the height distribution were normal-left distribution for the stand and it's main dominant populations of the Huitong subtropical secondary forest community, and indirectly indicated that both intra- and inter- specific competitions were exist for light resources.
Hegyi's competition index model was used to investigate intraspecific and interspecific competition intensity for dominant species trees. The results showed that competition intensity of the top 13 dominant plant species was ranked in the order of P. massoniana< M. pauhoi< Styrax suberifolius< L. coreana var. lanuginose< A. fortunei< Clethra fargesii< L. formosana< C. glauca< M. rehderii< C. lanceolata < L. chinense< C. axillaries< Engelhardtia roxburghiana. The tree species whose intraspecific competition was more intense than that of interspecific competition were M. rehderii, C. lanceolata, C. axillaris and E. roxburghiana. On the contrary, the tree species with lower intraspecific competition than that of interspecific competition were P. massoniana, M.s pauho, S. suberifolius, L. coreana var. lanuginosa, C. fargesii, L. formosana, C. glauca. The tree species that had an approximate intraspecific and interspecific competition index were L. chinense and A. fortuneii. The dominant and higher trees showed a strong competition intense compared to the lower trees. Relationship between the competition index and the diameter at breast height of objective tree could be expressed as power function and competition intensity appeared to reduce as the diameter at breast height of objective tree increased.
Competition is common natural phenomena. Generally, competition between tree species could be divided into aboveground competition for light and belowground competition for water and nutrient resources in soil. Environmental factors in soil and competition would shift fine root distribution, morphology and architecture. A. fortunei, L. formosan and C. glauca were selected as target to investigate root distribution and morphology in response to abovegroud competition. Competition index (CI) of A. fortunei, L. acakycina and C. glauca was 1.69,0.97 and 2.16, respectively. Total fine root biomass of A. fortunei, L. formosan and C. glauca within 0-45cm soil depth accouted for 104.1g·m-2,122.4 g·m-2 and 54.4 g·m-2, most of which distributed at top 0-30cm soil depth. Fine root biomas of A. fortunei and L. formosan were even distributed in soil depth, but there were soil depth distribution differences between C. glauca and A. fortunei, L. formosana. For horizontal distribution, there were no significant differences among fine root biomass of A. fortunei, L. formosan and C. glauca. (p>0.05) Fine roots of C. glauca generally occupied the places nearby trees, while fine roots of A. fortunei and L. acakycina were not.
Vertical and horizontal distribution of root length density (RLD) and specific root area (SRA) of three tree species was similar to that of fine root biomass. RLD and SRA were significantly decreased with increasing soil depth. RLD and SRA of A. fortunei and L. formosan were similar in different horizontal distance, while RLD and SRA of C. glauca were higher closed to tree species. In terms of vertical distribution, specific root length (SRL), specific root area (SRA) and specific root tips (SRT) of L. formosan which decreased significantly with increased soil depth were higher than that of A. fortunei and C. glauca in surface 0-15cm. In contrast, SRL, SRA and SRT increased with increasing soil depth. In terms of horizontal distribution, SRL, SRA and SRT of L. formosan and C. glauca were higher than other tree species in different horizontal distances, while those of A. fortunei were not predominant in each soil layers and each horizontal distance. Although C. glauca was subjected to strong competition and competitive index was high, its fine roots had great adaptability to changed environment (differential distribution, higher SRL, SRA and SRT), and it can coexist with L. formosan for a long time. On the contrary, suffered from competition from aboveground and belowground, A. fortunei is disadvantageous when competed with L. formosan and C. glauca.
To test the following two hypotheses whether the spatial distribution of dominant tree species is aggregated in subtropical secondary forests, and whether this spatial pattern can be used to infer causal factors for the coexistence of diverse tree species and predict forest succession trends, we employed Ripley's K(t) function analyze spatial patterns and associations. All trees within the forest showed a significant aggregated spatial pattern as did trees of each dominant species. Negative spatial association was detected for trees of C. axillaris versus A. fortunei, L. formosan or Pinus massoniana, and trees of Cyclobalanopsis glauca against A. fortunei, L. formosan, C. axillaris or P. massoniana. The apparent aggregation of spatial distribution for trees of each dominant species is indicative for mosaic patches that partition resources and maintain diverse species coexistence. Repulsive or independent spatial associations between trees of later and earlier serial species demonstrate the gap regeneration process in forest dynamics. Pioneer deciduous species will eventually be replaced by late serial evergreen species within the secondary forest.
引文
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