黄土丘陵沟壑植被恢复区节肢动物群落特征及时空动态研究
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摘要
本论文以纸房沟流域为研究单元,采用群落生态学的方法,分析不同林分节肢动物群落的结构组成、时空动态、综合指数及稳定性,并分析了优势害虫、天敌之间的生态位及空间格局,探讨群落内物种之间的相互联系、相互制约的内在机制。以便在多元林分生态系统中寻找节肢动物类群综合生物指标,评价植被恢复状况,为黄土高原生态环境建设中的植被恢复提供理论依据。主要研究结果如下:
1.不同林分节肢动物种-面积关系符合Sq=Cqm,得出相应最小调查面积,大小排序为:自然灌木林>自然乔木林>杨树刺槐混交林>柠条沙棘混交林>柠条林>沙棘林>刺槐林>柳树林。建立了不同林分节肢动物种-多度模型,天然恢复林地节肢动物以LN分布模型最为适合,表明群落中个体数量居中的种类最多,稀有种和常见种分不均衡;在混交林和纯林中以LC分布模型最为适合,群落中稀有种和常见种较多。
2.不同林分的节肢动物群落中,共有节肢动物420种,分属15目93科。其中鳞翅目的种类最多,鞘翅目次之。各林分节肢动物以天然恢复林中物种数最多,其次为混交林,纯林的物种数较少。天然恢复林中优势科均为夜蛾科、尺蛾科和金龟科,混交林优势科前两个为夜蛾科和尺蛾科,四种人工纯林优势科差异较大。
3.从特征指数来分析:林分组成结构越单一,节肢动物群落目级水平上多数目的多样性、均匀性及种间相遇几率在降低。天然恢复林节肢动物群落优势度值均低于其它林分,而种间相遇几率值比其它林分均高;不同功能亚类群H、J、PIF值大小为:天然恢复林>混交林>单纯林。优势度以纯林最高,其次为混交林,天然恢复林最低;在时间序列过程中多样性指数以天然恢复林>混交林>纯林分,且各林分在7月下旬至8月下旬节肢动物群落多样性均达到最大值;均匀度以纯林和混交林的节肢动物群落变化幅度较大,天然恢复林变化幅度最小;群落的优势度与种间相遇几率(PIF)、均匀度值的变化呈相反趋势。
4.根据主成分分析原理,建立节肢动物群落多样性评价综合性指数,不同林分节肢动物目级水平、亚群落及时序三种综合指数排序一致表现为天然恢复林>混交林>纯林。目级水平、亚群落综合指数聚类以天然恢复林距离较小,发生情况最接近,混交林之间距离较近,四种纯林之间距离较小。时序聚类以天然恢复林发生情况最接近,混交林与拧条林、沙棘林发生情况最接近,柳树林和刺槐林发生情况最接近。采用最优分割方法,两种天然恢复林的时序划分为相同三个时段,而混交林和单纯林的时间格局相互不一致。
5.天然恢复林节肢动物群落稳定性最高,混交林和纯林稳定性相对较低,其群落稳定性测度综合性指数小,不同林分群落稳定性总体综合指数的排序与多样性综合指数的排序不完全一致,但二者仍表现为天然恢复林>混交林>纯林。
6.生态位分析表明,在柠条林中害虫之间以三种种实害虫三维生态位重叠较高,相互之间竞争激烈。从天敌三维生态位重叠看,豆象盾腹茧蜂与姬小蜂重叠值最高,二者之间竞争激烈。天敌与害虫之间以豆象盾腹茧蜂与拧条豆象生态位重叠值最高,说明豆象盾腹茧蜂对拧条豆象重控制和跟随作用最强。
7.在柳树林中,害虫之间以柳弯角卷叶蛾与柳尖胸沫蝉三维生态位重叠最高,二者在时空资源上竞争激烈。天敌之间以七星瓢虫与异色瓢虫重叠值最高。从害虫与天敌生态位重叠看,七星瓢虫与柳瘤大蚜重叠值最高,说明七星瓢虫对柳瘤大蚜重控制和跟随作用最强。
8.根据柳厚壁叶蜂为害特点,表明柳厚壁叶蜂幼虫均以个体群的形式呈聚集分布,在不同的空间阶元验证其分布型,而且在不同的空间其聚集原因存在一定的差异。根据公式,确定各种密度下的最适抽样数,通过公式以枝条空间m_0=10;叶片上m_0=0.2为防治指标制定了序贯抽样分析表。
The arthropods were investigated from 2006 to 2008 in 8 stands of Zhifanggou area on the loess plateau in China. The species-area and species-abundance relationship of arthropods in various stands were discussed by some mathematical models,Structure of arthropods community, space-time dynamics, diversity comprehensive index and stability in various stands were concluded by the analysis on diversity, principal component and clustering. And the multidimensional niches of the pests and their enemies in Periploca sepium and Caragana mocrophylla and spatial distribution pattern of Pontania dolichura larvae and sampling technique were analyzed as follows:
1.The species-area relationship of arthropods in various stands accorded with the formula Sq=Cqm , the smallest investigation area was computed, and the order was: Natural bush > Natural arbor > Populus davidiana+Robinia pseudoacacia > Hippaphae rhamnoides L+Caragana mocrophylla >Periploca sepium> Hippaphae rhamnoides L > Robinia pseudoacacia > Caragana mocrophylla, the more complex stand type was, the larger the need to investigate minimum area was. On the basis of sampling investigation, species-abundance model of arthropods in various stands was established, lognormal distribution model was the most suitable in natural recovery stand, there were more species of medial individual amount ,and there were fewer abundant species and rare species in community。logcauchy distribution model was the most suitable in mixed stand and pure stand, there were more abundant species and rare species than in natural recovery.
2.There were 420 spcies of arthropods in different stands; they belong to 93 families and15 orders. Most of them were Lepidoptera, 25 families and 122 species, followed by Coleoptera, 12 families, 107 species. In various stands, arthropods species in natural recovery stand were the most, followed by mixed stand, and species in pure stand were the least. Most arthropods dominant families in various stands were Lepidoptera and Coleoptera. The proportion of dominant species was the most in pure stand, followed by mixed stand, and dominant species were obvious. The proportion of dominant species was the least in natural recovery stand, and dominant species were not prominent. The result showed that arthropods were stable in natural recovery stand, which had more complex bio-system, followed by mixed stand; pure stand eco-system was single and had the least stability.
3.The arthropod community structure of natural recovery stand was the most complex in different orders, and the species of arthropods were the most, followed by mixed stand, pure stand was the least. Lepidoptera and Coleoptera had the most families and orders, followed by Hemiptera, Hymenoptera, Araneae, Diptera and Homoptera, etc, and other orders were the least. From individual numbers in different orders, the arthropod amounts of natural stand were relative mean in various population, followed by mixed stand, pure stand were the worst. The phytophagous group is common dominance group, and species and individuals were the highest in different function groups. The species and individuals of natural enemies in natural recovery stand were obviously higher than other stand. From season changes of arthropod community in various stands, arthropods chiefly occurred from late April to September. the species number of arthropod were more from late July to mid August, and the most in Natural bush and Natural arbor. the individuals of arthropods was more from mid-June to early August, and the most in Periploca sepium and Caragana mocrophylla, followed by mixed stand, Hippaphae rhamnoides L and Robinia pseudoacacia, the individual change of arthropods in the natural recovery forests were the smallest
4.From features index: arthropods community in different orders had the diversity index(H), the eveness index(J), the dominanc index(D)and probability of interspecific encounter(PIF), which associated with stand complexity. The more complex the stand was, the more orders which had higher indexes in H, J and PIF was, and H, J and PIF increased in majority of orders. On the contrary, the less complex the stand was, the larger dominance was, and dominance index increased. the sequence of H,J and PIF in various function sub-community was natural recovery stand, mixed stand, pure stand. Evenness index of arthropods community changed in the following order: natural stand, mixed stand, and pure stand. D was the highest in pure stand, followed by mixed stand, and the the lowest in natural recovery stand. In time series, diversity index of arthropods community in various stands was: natural recovery stand> mixed stand> pure stand, and diversity index of arthropods community in various stands was the highest from late July to late August. Evenness index of arthropods community changed larder in pure stand and mixed stand, smaller in natural recovery stand. Dominance of community was opposite to PIF and J.
5.Based on the principal component analysis,the synthetic indices for community diversity evaluation were established.The results of the order of arthropods’synthetic index in order level, sub-community and season showed that natural recovery stand> mixed stand> pure stand. The clustering of order level and sub-community synthetic index showed that Natural bush and Natural arbor had shorter distance, Populus davidiana + Robinia pseudoacacia and Hippaphae rhamnoides L+Caragana mocrophylla had shorter distance , and four pure stands had shorter distance. The clustering of timing synthetic index showed that Natural bush and Natural arbor had shorter distance, Populus davidiana+Robinia pseudoacacia, Hippaphae rhamnoides L+Caragana mocrophylla, Periploca sepium and Hippaphae rhamnoides L had shorter distance, Robinia pseudoacacia and Caragana mocrophylla had shorter distance. Using the optimal segmentation, time pattern of arthropods community was divided, and all kinds of stands were divided to three phases by ordered phase, which could better reflect the community’s variation.
6.Arthropods community had the higher stability in natural recovery stand, and lower stability in mixed stand and pure stand, which had small community stability synthetic index. The sort of community stability synthetic index was not completely consistent with diversity synthetic index in various stands. Community diversity synthetic index was high, and community stability of arthropods was not necessarily high, but they showed that natural recovery stand> mixed stand> pure stand. It illustrated that natural recovery stand bio-system was complex, it increased the relative stability of arthropods community, and provided a theoretical basis for pest ecological control.
7.Niche analyses showed that three kinds of seed pests had higher dimensional niche overlap between pests in Periploca sepium, and they had fierce competition each other, secondly .From dimensional niche overlap of natural enemies, Phanerotomella sp. and Entedon sp. had the highest overlap index and fierce competition. From dimensional niche overlap of natural enemies and pests, Phanerotomella sp., Entedon sp. had higher overlap index with three kinds of seed pests, Phanerotomella sp. and Kytorhinus immixtus Motschulsky had the highest overlap index, which showed that Phanerotomella sp. had the strongest control to Kytorhinus immixtus Motschulsky among them. Among all the natural enemies of aphid, Aphidius sp. had the highest overlap index. Pandemis heparana and Aphrophora costalis had the highest overlap index between pests in Caragana mocrophylla, which showed that they had fierce competition in time and space resources. From niche overlap of natural enemies, Coccinella septempunctata and Harmonia ascyridis had the highest overlap index, which had fierce competition. From niche overlap of pests and enemies, Coccinella septempunctata and Tuberolachnus salignus had the highest overlap, which showed that Coccinella septempunctata had the strongest control and follow function to Tuberolachnus salignus. So niche overlap should be considered in controlling pests, and took measures to control the pests. In pests’ecological regulation, natural enemies’protection measures should be taken to give full play to their roles.
8.The spatial distribution pattern of Pontania dolichum larvae was analyzed with Taylor's power law, Iwao's distribution function, and six aggregation indexes.The results showed that the spatial distribution pattern of P. dolichura larvae was of aggregated, and the basic component of the distribution was individual colony, with the aggregation intensity increased with density. On branches, the aggregation was caused by the adult behavior of laying eggs and the spatial position of leaves, while on leaves, the aggregation was caused by the spatial position of news leaves in s pring when m < 2.37 ,and by the spatial position of news leaves in spring and the behavior o f eclosion and laying eggs when m >2.37.By using the parametersαandβin Iwan's,m*-m regression equation, the optimal and sequential sampling numbers were determined
1.不同林分节肢动物种-面积关系符合Sq=Cqm,得出相应最小调查面积,大小排序为:自然灌木林>自然乔木林>杨树刺槐混交林>柠条沙棘混交林>柠条林>沙棘林>刺槐林>柳树林。建立了不同林分节肢动物种-多度模型,天然恢复林地节肢动物以LN分布模型最为适合,表明群落中个体数量居中的种类最多,稀有种和常见种分不均衡;在混交林和纯林中以LC分布模型最为适合,群落中稀有种和常见种较多。
2.不同林分的节肢动物群落中,共有节肢动物420种,分属15目93科。其中鳞翅目的种类最多,鞘翅目次之。各林分节肢动物以天然恢复林中物种数最多,其次为混交林,纯林的物种数较少。天然恢复林中优势科均为夜蛾科、尺蛾科和金龟科,混交林优势科前两个为夜蛾科和尺蛾科,四种人工纯林优势科差异较大。
3.从特征指数来分析:林分组成结构越单一,节肢动物群落目级水平上多数目的多样性、均匀性及种间相遇几率在降低。天然恢复林节肢动物群落优势度值均低于其它林分,而种间相遇几率值比其它林分均高;不同功能亚类群H、J、PIF值大小为:天然恢复林>混交林>单纯林。优势度以纯林最高,其次为混交林,天然恢复林最低;在时间序列过程中多样性指数以天然恢复林>混交林>纯林分,且各林分在7月下旬至8月下旬节肢动物群落多样性均达到最大值;均匀度以纯林和混交林的节肢动物群落变化幅度较大,天然恢复林变化幅度最小;群落的优势度与种间相遇几率(PIF)、均匀度值的变化呈相反趋势。
4.根据主成分分析原理,建立节肢动物群落多样性评价综合性指数,不同林分节肢动物目级水平、亚群落及时序三种综合指数排序一致表现为天然恢复林>混交林>纯林。目级水平、亚群落综合指数聚类以天然恢复林距离较小,发生情况最接近,混交林之间距离较近,四种纯林之间距离较小。时序聚类以天然恢复林发生情况最接近,混交林与拧条林、沙棘林发生情况最接近,柳树林和刺槐林发生情况最接近。采用最优分割方法,两种天然恢复林的时序划分为相同三个时段,而混交林和单纯林的时间格局相互不一致。
5.天然恢复林节肢动物群落稳定性最高,混交林和纯林稳定性相对较低,其群落稳定性测度综合性指数小,不同林分群落稳定性总体综合指数的排序与多样性综合指数的排序不完全一致,但二者仍表现为天然恢复林>混交林>纯林。
6.生态位分析表明,在柠条林中害虫之间以三种种实害虫三维生态位重叠较高,相互之间竞争激烈。从天敌三维生态位重叠看,豆象盾腹茧蜂与姬小蜂重叠值最高,二者之间竞争激烈。天敌与害虫之间以豆象盾腹茧蜂与拧条豆象生态位重叠值最高,说明豆象盾腹茧蜂对拧条豆象重控制和跟随作用最强。
7.在柳树林中,害虫之间以柳弯角卷叶蛾与柳尖胸沫蝉三维生态位重叠最高,二者在时空资源上竞争激烈。天敌之间以七星瓢虫与异色瓢虫重叠值最高。从害虫与天敌生态位重叠看,七星瓢虫与柳瘤大蚜重叠值最高,说明七星瓢虫对柳瘤大蚜重控制和跟随作用最强。
8.根据柳厚壁叶蜂为害特点,表明柳厚壁叶蜂幼虫均以个体群的形式呈聚集分布,在不同的空间阶元验证其分布型,而且在不同的空间其聚集原因存在一定的差异。根据公式,确定各种密度下的最适抽样数,通过公式以枝条空间m_0=10;叶片上m_0=0.2为防治指标制定了序贯抽样分析表。
The arthropods were investigated from 2006 to 2008 in 8 stands of Zhifanggou area on the loess plateau in China. The species-area and species-abundance relationship of arthropods in various stands were discussed by some mathematical models,Structure of arthropods community, space-time dynamics, diversity comprehensive index and stability in various stands were concluded by the analysis on diversity, principal component and clustering. And the multidimensional niches of the pests and their enemies in Periploca sepium and Caragana mocrophylla and spatial distribution pattern of Pontania dolichura larvae and sampling technique were analyzed as follows:
1.The species-area relationship of arthropods in various stands accorded with the formula Sq=Cqm , the smallest investigation area was computed, and the order was: Natural bush > Natural arbor > Populus davidiana+Robinia pseudoacacia > Hippaphae rhamnoides L+Caragana mocrophylla >Periploca sepium> Hippaphae rhamnoides L > Robinia pseudoacacia > Caragana mocrophylla, the more complex stand type was, the larger the need to investigate minimum area was. On the basis of sampling investigation, species-abundance model of arthropods in various stands was established, lognormal distribution model was the most suitable in natural recovery stand, there were more species of medial individual amount ,and there were fewer abundant species and rare species in community。logcauchy distribution model was the most suitable in mixed stand and pure stand, there were more abundant species and rare species than in natural recovery.
2.There were 420 spcies of arthropods in different stands; they belong to 93 families and15 orders. Most of them were Lepidoptera, 25 families and 122 species, followed by Coleoptera, 12 families, 107 species. In various stands, arthropods species in natural recovery stand were the most, followed by mixed stand, and species in pure stand were the least. Most arthropods dominant families in various stands were Lepidoptera and Coleoptera. The proportion of dominant species was the most in pure stand, followed by mixed stand, and dominant species were obvious. The proportion of dominant species was the least in natural recovery stand, and dominant species were not prominent. The result showed that arthropods were stable in natural recovery stand, which had more complex bio-system, followed by mixed stand; pure stand eco-system was single and had the least stability.
3.The arthropod community structure of natural recovery stand was the most complex in different orders, and the species of arthropods were the most, followed by mixed stand, pure stand was the least. Lepidoptera and Coleoptera had the most families and orders, followed by Hemiptera, Hymenoptera, Araneae, Diptera and Homoptera, etc, and other orders were the least. From individual numbers in different orders, the arthropod amounts of natural stand were relative mean in various population, followed by mixed stand, pure stand were the worst. The phytophagous group is common dominance group, and species and individuals were the highest in different function groups. The species and individuals of natural enemies in natural recovery stand were obviously higher than other stand. From season changes of arthropod community in various stands, arthropods chiefly occurred from late April to September. the species number of arthropod were more from late July to mid August, and the most in Natural bush and Natural arbor. the individuals of arthropods was more from mid-June to early August, and the most in Periploca sepium and Caragana mocrophylla, followed by mixed stand, Hippaphae rhamnoides L and Robinia pseudoacacia, the individual change of arthropods in the natural recovery forests were the smallest
4.From features index: arthropods community in different orders had the diversity index(H), the eveness index(J), the dominanc index(D)and probability of interspecific encounter(PIF), which associated with stand complexity. The more complex the stand was, the more orders which had higher indexes in H, J and PIF was, and H, J and PIF increased in majority of orders. On the contrary, the less complex the stand was, the larger dominance was, and dominance index increased. the sequence of H,J and PIF in various function sub-community was natural recovery stand, mixed stand, pure stand. Evenness index of arthropods community changed in the following order: natural stand, mixed stand, and pure stand. D was the highest in pure stand, followed by mixed stand, and the the lowest in natural recovery stand. In time series, diversity index of arthropods community in various stands was: natural recovery stand> mixed stand> pure stand, and diversity index of arthropods community in various stands was the highest from late July to late August. Evenness index of arthropods community changed larder in pure stand and mixed stand, smaller in natural recovery stand. Dominance of community was opposite to PIF and J.
5.Based on the principal component analysis,the synthetic indices for community diversity evaluation were established.The results of the order of arthropods’synthetic index in order level, sub-community and season showed that natural recovery stand> mixed stand> pure stand. The clustering of order level and sub-community synthetic index showed that Natural bush and Natural arbor had shorter distance, Populus davidiana + Robinia pseudoacacia and Hippaphae rhamnoides L+Caragana mocrophylla had shorter distance , and four pure stands had shorter distance. The clustering of timing synthetic index showed that Natural bush and Natural arbor had shorter distance, Populus davidiana+Robinia pseudoacacia, Hippaphae rhamnoides L+Caragana mocrophylla, Periploca sepium and Hippaphae rhamnoides L had shorter distance, Robinia pseudoacacia and Caragana mocrophylla had shorter distance. Using the optimal segmentation, time pattern of arthropods community was divided, and all kinds of stands were divided to three phases by ordered phase, which could better reflect the community’s variation.
6.Arthropods community had the higher stability in natural recovery stand, and lower stability in mixed stand and pure stand, which had small community stability synthetic index. The sort of community stability synthetic index was not completely consistent with diversity synthetic index in various stands. Community diversity synthetic index was high, and community stability of arthropods was not necessarily high, but they showed that natural recovery stand> mixed stand> pure stand. It illustrated that natural recovery stand bio-system was complex, it increased the relative stability of arthropods community, and provided a theoretical basis for pest ecological control.
7.Niche analyses showed that three kinds of seed pests had higher dimensional niche overlap between pests in Periploca sepium, and they had fierce competition each other, secondly .From dimensional niche overlap of natural enemies, Phanerotomella sp. and Entedon sp. had the highest overlap index and fierce competition. From dimensional niche overlap of natural enemies and pests, Phanerotomella sp., Entedon sp. had higher overlap index with three kinds of seed pests, Phanerotomella sp. and Kytorhinus immixtus Motschulsky had the highest overlap index, which showed that Phanerotomella sp. had the strongest control to Kytorhinus immixtus Motschulsky among them. Among all the natural enemies of aphid, Aphidius sp. had the highest overlap index. Pandemis heparana and Aphrophora costalis had the highest overlap index between pests in Caragana mocrophylla, which showed that they had fierce competition in time and space resources. From niche overlap of natural enemies, Coccinella septempunctata and Harmonia ascyridis had the highest overlap index, which had fierce competition. From niche overlap of pests and enemies, Coccinella septempunctata and Tuberolachnus salignus had the highest overlap, which showed that Coccinella septempunctata had the strongest control and follow function to Tuberolachnus salignus. So niche overlap should be considered in controlling pests, and took measures to control the pests. In pests’ecological regulation, natural enemies’protection measures should be taken to give full play to their roles.
8.The spatial distribution pattern of Pontania dolichum larvae was analyzed with Taylor's power law, Iwao's distribution function, and six aggregation indexes.The results showed that the spatial distribution pattern of P. dolichura larvae was of aggregated, and the basic component of the distribution was individual colony, with the aggregation intensity increased with density. On branches, the aggregation was caused by the adult behavior of laying eggs and the spatial position of leaves, while on leaves, the aggregation was caused by the spatial position of news leaves in s pring when m < 2.37 ,and by the spatial position of news leaves in spring and the behavior o f eclosion and laying eggs when m >2.37.By using the parametersαandβin Iwan's,m*-m regression equation, the optimal and sequential sampling numbers were determined
引文
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