【作者】 许士国；

【导师】 安利国； 付荣恕；

【作者基本信息】 山东师范大学 ， 动物学， 2011， 博士

【摘要】 本文首次深入系统地研究黄河三角洲土壤动物,从多层次探讨各年代土壤动物群落特点及演替过程。本次调查主要依据成土年限、植被类型、土壤性质、人类干扰程度等因素设置采样点,从土壤动物、土壤螨类、土壤甲螨和土壤跳虫4个层面4个角度分析群落结构特点,并从种类组成、数量分布、季节变动、多样性和相似性等方面分析群落间的差别,主要根据成土年限和植被类型对群落结构的影响,探讨土壤动物群落的演替趋势。本次研究结果表明,黄河三角洲地理环境单一,土壤动物无论从数量上还是从种类上均较贫乏,全区平均密度不足1万,在纲或目分类水平上仅17类群。螨类和跳虫在11个样点中均为优势类群,且每个样点中均只有这两个优势类群。A/C值全区全年平均为2.04,明显大于1。4个季度中,冬季和夏季的A/C值较大,说明螨类对低温或干旱等不利环境适应力和抵抗力强于跳虫。该地区土壤动物优势类群明显,群落异质性差,群落的多样性较低。本次调查的11个样点中,多样性指数最大的是样点A(农田),说明该地区土壤环境差,土壤贫瘠,盐碱化,不利于土壤动物生存,相对而言,人类的活动会改善土壤环境,土壤动物的种类和数量均的所增加。在4个季度中,土壤动物群落多样性最高的是秋季,而冬季和夏季均偏低,说明土壤动物对土壤温度和土壤含水量的变化非常敏感。11个样点间土壤动物群落相似性较高,群落间为极相似或相似,尽管土壤形成年代相差久远,植被差别明显。土壤群落群落相似性高,说明对土壤动物影响最为直接和重要的环境因子是“水”和“盐”。本次调查结果显示,隐气门亚目数量最多,前气门亚目数量次之,中气门亚目数量最少。在4个季度中,隐气门亚目和前气门亚目百分比较为稳定,1月份< 4月份< 7月份< 10月份,中气门亚目百分比变化较大。3个年代三角洲内3个亚目所占比例差别不大,均是隐气门亚目>前气门亚目>中气门亚目。6种植被生境内3个亚目所占比例差别明显。中气门亚目在芦苇湿地和芦苇-獐毛荒地生境中比例较低,前气门亚目在农田生境中比例最低,而隐气门亚目在农田生境中比例最高。该地区土壤螨类虽然III层明显少于I层和II层,但表聚性并不强。本次调查获得土壤甲螨38科52属64种,包括47个已知种和17个未知种,其中优势种1个:大洋上罗甲螨;亚优势种6个:威氏多奥甲螨、滑菌甲螨、新小奥甲螨、覆盖头甲螨、细条木单翼甲螨、日本礼服甲螨。结合文献资料对47种甲螨已知种进行区系分析,本区与华中区重叠率最高,其次是东北区和华南区。11个样点土壤甲螨群落的种类组成和数量分布具有极显著差异,样点A、B和E种类和数量最多,该3个样点为农田或宜耕地。3个年代的三角洲无论是种类数还是个体数变动趋势一致,古代三角洲>近代三角洲>>现代三角洲,随三角洲成土年限的增加,土壤甲螨种类增多,密度增大。6种植被生境土壤甲螨群落的种类组成的数量分布均具有极显著的差异。无论是种类数还是个体数均芦苇湿地<<芦苇-獐毛荒地<碱蓬-獐毛荒地<<柽柳-海洲蒿荒地。在4个季节中,土壤甲螨群落的种类组成为秋季>冬季>春季≈夏季;从个体数量上看,秋季最多,其他3个季度的密度差别不大。7个优势属或亚优势属的季节变动规律差别较大,上罗甲螨在4个季节中均为群落的优势属或亚优势属,数量变动最小,礼服甲螨和木单翼甲螨变动趋势一致,夏季数量最少,盖头甲螨则夏季数量最多,菌甲螨秋季数量较少,其他3个季节中数量较多,小奥甲螨则是秋季数量最多,其他3个季节中数量较少,多奥甲螨数量变动最大,在冬季和春季数量最多,而在夏季和秋季数量较少。无论是从种类数上还是从个体数量上分析,黄河三角洲土壤甲螨群落三大类群比例均衡,群落结构均属于O型,对3个年代三角洲、6种不同植被生境及4个季度土壤甲螨群落的测定结果一致,大多数为O型。MGP-分析I和MGP-分析II测定结果有所不同。黄河三角洲地区土壤甲螨群落异质性高,多样性指数较高。各样点的多样性差异明显,大多数样点多样性指数在2.3~3.0之间。3个年代的三角洲土壤甲螨群落异质性高,多样性指数都比较大。6种不同植被类型的生境中,土壤甲螨群落无论是类群数还是多样性指数均差异明显。11个样点间土壤甲螨群落的相似性较低,没有超过0.75的系数,大多数群落之间为不相似或极不相似。3个年代三角洲和6种植被生境土壤甲螨群落间相似性差别较大。古代三角洲与近代三角洲间相似性较大,芦苇湿地与其他植被生境间的相似性系数均较低。本次调查获得土壤跳虫8科22属,其中棘跳、类符跳、球角跳和符跳为优势属,构成土壤跳虫群落的主体。与土壤甲螨群落相比,土壤跳虫出现的属数少,优势属多、稳定而明显。11个采样点土壤跳虫群落的种类组成和数量分布均有显著差异。3个年代的三角洲土壤跳虫无论是种类数还是个体数量均是古代三角洲最多,近代三角洲次之,现代三角洲最少。6种植被生境中,芦苇湿地土壤跳虫种类和数量最少,其他5种生境差异不大。无论从种类组成上还是从个体数量上看,土壤跳虫均为秋季>春季>夏季>冬季。在4个季度中的优势属相同,但优势属的季节变动趋势有着明显不同,棘跳为典型的土内种,数量比较稳定;类符跳随冬—春—夏—秋相对数量逐次增多;球角跳则是冬季最多,秋季较少;符跳在春季仅出现于4个样点,数量较其他3个季节明显减少。11个样点、3个年代的三角洲、6种不同植被类型生境中土壤动物的多样性、均匀性、优势度差异不大,相似性差别较大。本次研究结果显示,在高级分类阶元上分析土壤动物群落组成差别不大,不能充分表现出土壤动物群落演替过程。也就是说在高级分类阶元上分析,土壤动物群落相似性较大,在此水平上研究土壤动物群落演替过程意义不大。但随成土年限或植物群落的演替过程,土壤动物群落间的相似性表现出一定的规律性,即成土年限相距越近,群落相似性越高;随芦苇湿地、芦苇-獐毛荒地、碱蓬-獐毛荒地、柽柳-海洲蒿荒地的演替过程,土壤动物群落间的相似性逐渐降低,演替阶段越接近,土壤动物群落间的相似性就越大。调查结果还表明在纲或目分类阶元上,不同植被类型的生境中,土壤动物群落虽然均达相似,但随植物群落的演替,其组成仍表现出一定的差异,且这种差异远大于土壤形成年代所造成的差异。3个年代三角洲和6种植被生境中土壤螨类的密度差异明显,土壤螨类的数量随三角洲成土年限的增加而增多,且均是隐气门亚目>前气门亚目>中气门亚目,随植被的演替过程而增多。隐气门亚目的比例随植被的演替过程而增加,中气门亚目的比例则随植被的演替过程而减少,前气门亚目的变化没有明显的规律性,因此,研究动物群落的演替过程,隐气门亚目和中气门亚目要比前气门亚目重要的多。本次调查结果显示,3个年代的三角洲内土壤甲螨无论是种类数还是个体数变动趋势一致,古代三角洲>近代三角洲>>现代三角洲,随三角洲成土年限的增加,土壤甲螨种类增多,密度增大。6种植被生境土壤甲螨群落的种类组成的数量分布均具有极显著的差异。无论是种类数还是个体数随植被的演替过程,土壤甲螨群落变化表现出一定的规律性,即芦苇湿地<<芦苇-獐毛荒地<碱蓬-獐毛荒地<<柽柳-海洲蒿荒地。随植物群落的演替过程,土壤甲螨群落的结构变化表现出一定的规律性,即随芦苇湿地→芦苇-獐毛荒地→碱蓬-獐毛荒地→柽柳-海洲蒿荒地的演替过程,土壤甲螨群落的M-群的类群数和相对数量逐渐减少,P-群的类群数和相对数量逐渐增多。3个年代的三角洲土壤甲螨群落异质性高,多样性指数都比较大,但仍能看出土壤甲螨的种类数和多样性均随土壤形成年限的增加而增加。6种不同植被类型的生境中,土壤甲螨群落无论是类群数还是多样性指数均差异明显。随植物群落的演替过程,土壤甲螨群落的类群数逐渐增多,均匀性逐渐增大,多样性逐渐升高。比较3个年代三角洲土壤甲螨群落相似性,可知成土年限相距越远,群落相似性越小。6种植被生境土壤甲螨群落相似性差别较大。芦苇湿地与其他植被生境间的相似性系数均较低。随植物群落的演替过程,土壤甲螨群落间的相似性逐渐降低,演替阶段越接近,群落间的相似性就越大。3个年代的三角洲和6种植被生境中土壤跳虫无论是种类数还是个体数量均有明显差异,且变动趋势一致,古代三角洲>近代三角洲>现代三角洲;芦苇湿地种类和数量最少,其他5种生境差异不大。3个年代的三角洲和6种植被生境中土壤动物的多样性、均匀性、优势度差异不大。3个年代三角洲土壤跳虫群落相似性随三角洲的成土年限不同而有规律的变化,成土年限越接近,相似性越高。相似性系数的最大值发生在碱蓬-獐毛荒地与柽柳-海洲蒿荒地间,碱蓬-獐毛荒地、柽柳-海洲蒿荒地和芦苇-獐毛荒地三者之间均为相似或极相似,相似性系数随植被演替过程而有规律地减小。本文创新之处在于本研究首次深入系统地研究黄河三角洲土壤动物,从多层次探讨各年代土壤动物群落特点及演替过程。研究结果表明,该地区土壤环境差,土壤贫瘠,盐碱化,不利于土壤动物生存,致使土壤动物优势类群明显,群落异质性差,群落的多样性较低。研究结果表明,土壤动物高级分类阶元不能很好地反映群落的演替过程。隐气门亚目和中气门亚目随成土年龄、植被演替而发生有规律的改变,因此,研究土壤动物群落演替应重视对隐气门亚目或中气门亚目的深入研究。特别是土壤甲螨随成土年限或植被的演替过程均表现出相似的规律性:种类增多,密度增大,均匀性逐渐增大,多样性逐渐升高,成土年限或演替阶段相距越远,群落相似性越小。但随植物群落的演替过程,土壤甲螨群落的变化较成土年限的影响更为明显。土壤跳虫更易受其他环境因子(如温度、湿度)的影响,对成土年限的反应不如土壤甲螨规律,降低了其在研究动物群落演替中的重要性。更多还原

【Abstract】 Soil animal community in the Yellow River Delta was studied intensively and systematically for the first time in this study, and its characteristics and succession process were discussed from multi-levels as well. Sampling sites were set on the basis of soil-forming period, vegetation type, soil properties, human disturbance and other factors. Communities structure characteristics were analyzed from 3 levels and 4 perspectives of total soil animals, total soil mites, soil oribatid mites and soil collembola, and the differences among communities were analyzed from the aspects of species composition, quantitative distribution, seasonal changes, diversity and similarity and other aspects. Succession of the soil animal community was explored mainly based on the influence of soil-forming period and vegetation type to the community structure.Results of this study show that, the geographical environment of the Yellow River Delta is simple, both species number and quantity of the soil animals are poor, average density of animal is less than 10 thousand individuals in the entire region, and only 17 classes or orders of the soil animals are obtained in this study. For the soil animals among these 11 sampling sites from three ages, 6 different vegetation habitat, diversity, evenness and dominance show little difference, but the similarity varies greatly.Mites and Collembola are the dominant groups in all the 11 sampling sites, and they are the only two dominant groups at each site. The annual average A / C value of the whole area is 2.04, significantly greater than 1. A / C values in winter and summer are comparatively larger, indicating that adaption and resistance of mites to low temperature or drought is stronger than that of Collembola.Dominant groups of soil animals in this area are significant, community heterogeneity and diversity are low. Among the 11 sampling sites, site A (farmland) has the largest diversity index, indicating that the soil environment is poor, the soil is lean, salinized, and not suitable for soil animals,to survive, on contrast, human activities improved the soil environment, both the species and number of soil animals were increased, Soil animals has the highest diversity in autumn in the four seasons, while the diversities in winter and summer are low, indicating that the soil animals are very sensitive to soil temperature and soil moisture changes.Although the soil formation age and the vegetation are very different,soil animal community similarity among the 11 sampling sites is high, indicating that the most important environmental factors that directly impact on soil animals are "water" and "salt."Survey results show that, the number of Cryptostigmata(Oribatida) is the largest, followed by Prostigmata, and Mesostigmata is the least. Percentages of Cryptostigmata and Prostigmata are stable in the four seasons, with a varity order of January <April <July <October, but that of Mesostigmata changes largely. The percentages of the 3 suborders are not very different in this area during the 3 ages, with a order of Cryptostigmata > Prostigmata >Mesostigmata. The proportion of the three suborders among the 6 vegetations habitats are different significantly. The proportion of Mesostigmata in Phragmites australis wetland and in Phragmites australis - Aeluropus sinensis wasteland habitats is relatively low, and Prostigmata in farmland is the lowest, while Cryptostigmata in the farmland is the highest. Although the soil mites in layer III is significantly less than in layer I and layer II, the surface concentration is not strong.64 species of soil oribatid mites belonging to 52 genera of 38 families were obtained in this investigation, including 47 known species and 17 unknown species. With one dominant species: Epilohmannia pallida pacifica, 6 sub-dominant species: Multioppia wilsoni, Scheloribates laevigatus, Oppiella nova, Tectocepheius velatus,, Xylobates tenuis, and Trhypochthonius japonicus. Fauna analysis was carried out on these 47 known species in combination with the literature data, the overlapping coefficient between this area and central China is the highest, followed by Northeast and South China area.Species composition and quantity distribution of soil oribatid mite communities among these 11 sites are different significantly, but the change is similar. Site A, B and E in the farmland with the largest number of species and quantity. Both the change tendency of species number and the individual number in this area of three ages is coherent, with a sequence of ancient age > recent age> present age The species number and the density of soil oribatid mite increased. with the increase of soil-forming period. Species composition and quantity distribution of the soil oribatid mite communities differ significantly among these 6 vegetation habitats . Both the number of species and the individual have the same change sequence of Phragmites australis wetland < Phragmites australis - Aeluropus sinensis wasteland < Suaeda salsa - Aeluropus sinensis wasteland < Tamarix chinensis - Artemisia fauriei wasteland.Species composition of the soil oribatid mite community change with a sequence of autumn> winter> spring≈summer; individuals number in autumn is the most, and densities do not differ much in the other three seasons. The difference of seasonal fluctuation in the seven dominant genera or subdominant genera is observable, Epilohmannia is the dominant genera or sub-dominant genera in four seasons with the smallest number changes; Trhypochthonius and Xylobates seasonal fluctuation trends are the same, with the least number in summer. Number of Tectocepheius in summer is the largest. The number of Scheloribates in the fall is comparatively low, while much many in the other three seasons,while for Oppiella, with the largest number in autumn but very few in the other three seasons. Multioppia has the largest fluctuation in number, with the largest number in winter and spring, but fewer in summer and autumn.Both the number of species and individuals of the three groups of soil oribatid mites in the Yellow River Delta are even. The community structure is O type, the Structure characteristics of the soil oribatid mites communities in this area of three ages, the 6 different vegetation habitats and four seasons are similar, most are O type. Results are different partially with MGP- I and MGP II analysis.The heterogeneity and diversity of soil oribatid mite in the Yellow River Delta are high, The diversity indexes at most sampling sites are between 2.3 to 3.0, and the differences among every site are significant. The heterogeneity of soil oribatid mite of this area of three ages is high, and the diversity index is relatively high. Both the number of groups and the diversity index of the soil oribatid mite are significantly different in the 6 different types of vegetation habitats.None similarity index is higher than 0.75 of the soil oribatid mite communities among these 11 sampling sites, so the similarity is relatively low, and the majority of communities are not similar or extremely not similar. The similarity of the soil oribatid mite communities in this area among 3 ages and 6 vegetation habitats varies greatly. The similarity of this area between ancient and recent age is relatively high, and all the similarity coefficients among Phragmites australis wetland and other vegetation habitats are relatively low.22 genera belonging to 8 families of soil Collembola are obtained in the investigation, of which Onychiurus, Folsomina, Hypogastrura and Folsomia are the dominant genus, constituting the main body of the soil Collembola community. Compared to the soil oribatid mite community, the number of soil Collembola genera is less, with more stable and significant dominant genera. The species composition and quantitative distribution of the soil Collembola community among these 11 sampling sites are significantly different. Both the number of species and individuals of the soil Collembola in this area from the three ages change at the same order: ancient age >recent age>persent age. There are the least number of Collembola in Phragmites australis wetland habitats, and no significant difference for the other 5 habitats.Both the species composition and the number of individuals of soil Collembola change at the same order: fall > spring> summer> winter. The dominant genera in the four seasons is the same, but the seasonal change trend of the dominant genera shows significant difference. Onychiurus is a typical deep-soil-living type with relatively stable number. number of Folsomina successively increases with winter→Spring→Summer→Autumn changing. while Hypogastrura has the largest number in winter. Folsomia were relatively less in Autumn, found at only four sampling sites, and the number decreased significantly compared with the other three seasons.The results of this study show that, the soil animal communities composition is not very different at high-level taxa, which can not fully demonstrate the process of their succession. That is to say, the similarity of soil animal communities is relatively high at high-level taxa, and it is less use to investigate the succession of soil animal communities at this level. However, with the succession of soil formation period or plant community, similarity among soil animal communities shows certain regularity, namely, the nearer the age interval of the soil formation period, the higher the community similarity; With the succession process of Phragmites australis wetland, Phragmites australis - Aeluropus sinensis wasteland, Suaeda salsa - Aeluropus sinensis wasteland and Tamarix chinensis - Artemisia fauriei wasteland, the similarity of soil f animal communities decreased gradually, the closer succession stage, the greater the similarity among the soil animal communities. We also show that, on the taxa of class or order classification level, for habitats of different vegetation types, although the soil animal communities are similar, but with the succession of plant communities, their composition still showe some differences, and this difference is much greater than the difference caused by the soil formation age.The density differences of the soil mites in this area among 3 ages and 6 different vegetation habitats are significant, soil mites number increased with the increase of soil formation period, and all with a sequence of Cryptostigmata > Prostigmata >Mesostigmata, and increase with the succession process of vegetation. The proportion of Cryptostigmata increases while Mesostigmata decreases with the succession of vegetation, and the change of Prostigmata shows no apparent regularity, and therefore, for the study of the succession of soil animal communities, Cryptostigmata and Mesostigmata are much more important than Prostigmata.Results of this investigation show that, both the change tendency of species and individual number of soil oribatid mite in this area from three ages are the same, with a sequence of ancient age> recent age> present age, species number and density of soil Oribatida increased with the increasing of soil-forming period. Both the species composition and quantitative distribution of soil oribatid mite communities in the 6 different vegetations habitants show extremely significant differences. Both the numbers of species and individuals of soil oribatid mite community show a certain regularity with the succession process of vegetation, that is, with a order of Phragmites australis wetland << Phragmites australis - Aeluropus sinensis wasteland < Suaeda salsa - Aeluropus sinensis wasteland << Tamarix chinensis - Artemisia fauriei wasteland.With the succession of plant communities, the structure of soil oribatid mite community changes at a certain regularity, the species number and relative number of M-group of soil oribatid mite community decreased gradually, while the P-group increased gradually with the succession of Phragmites australis wetland→Phragmites australis - Aeluropus sinensis wasteland→Suaeda salsa - Aeluropus sinensis wasteland→Tamarix chinensis - Artemisia fauriei wasteland.The heterogeneity of soil oribatid mite communities in this area of three ages is high, the diversity indexes are relatively large, and both the species number and diversity of the soil oribatid mite communities increase with the soil formation period. Both the number of species and the diversity index of the soil oribatid mite communities are significantly different among the 6 different types of vegetation habitats. The species number, evenness and diversity of the oribatid communities increased gradually with the succession of the plant communities.By comparison of the similarity of soil oribatid mite communities in this area from three ages, we can see that, the further apart from the soil formation period, the smaller the community similarity. The similarity difference of soil oribatid mite communities for the 6 different vegetations habitats varied greatly. The similarity coefficient among Phragmites australis wetland and other vegetation habitats were relatively low. The similarity of oribatid mite communities decreased with the succession of plant communities, the closer the succession stage, the greater the similarity among communities.Both the change tendency of species and individual number of soil Collembola in the yellow river Delta of three ages and the 6 vegetation habitats show significant difference, but the changing tendency is the same, with a sequence of ancient age> recent age>> present age; the species number and quantity in Phragmites australis wetland is the least , and the differences among the other 5 habitants are not great.For the soil Collembola in the yellow river Delta of three ages and the 6 vegetations habitats, the diversity, evenness and dominance show no great difference.The soil Collembola communities similarity in this area of three ages changed regularly with the different soil formation periods, the closer the soil formation period, the higher the similarity. The similarity coefficient of the soil Collembola communities regularly decreased with the vegetation succession process. The maximum similarity coefficient occurred between Suaeda salsa - Aeluropus sinensis wasteland and Tamarix chinensis - Artemisia fauriei wasteland, among Suaeda salsa - Aeluropus sinensis wasteland, Tamarix chinensis - Artemisia fauriei wasteland and Phragmites australis - Aeluropus sinensis wasteland, it is similar or very similar. We studied the soil animal community in the Yellow River Delta intensively and systematically for the first time, and discussed the characteristics and succession process of the soil animal community from multi-levels.Results show that, the soil environment of this region is poor, and the soil is lean and salinized, and not suitable for soil animals to survive, resulting in significant dominant groups of soil animals, poor community heterogeneity, and low community diversity.It is also shown that, high-level taxa of soil animals can not fully reflect the succession process of community. Cryptostigmata and Mesostigmata change regularly with the succession of soil formation period and vegetation, and therefore more attention should be paid to Cryptostigmata or Mesostigmata in the study of soil animal community succession. Especially, with the succession of soil formation period or vegetation, the soil oribatid mite have shown similar regularity: with increase in species and density, gradual increase in homogeneity and diversity, the further apart from the succession of soil formation period and vegetation, the smaller the community similarity. However, with the succession of plant communities, the impact of changes in soil oribatid mite communities is more evident than that of soil formation period. Soil Collembola are more vulnerable to other environmental factors (such as temperature, humidity), and the response regulation of soil Collembola to soil formation period is not comparable to that of soil oribatid mite, reducing its importance in the study of soil animal community succession.更多还原