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退化红壤植被恢复对土壤节肢动物群落的影响

Influences of Vegetation Restoration on Soil Arthropods in Degraded Red Soil

【作者】 丁程成

【导师】 胡锋;

【作者基本信息】 南京农业大学 , 生态学, 2007, 博士

【摘要】 土壤动物是森林生态系统的重要组成部分,在系统养分循环、能量流动中具有重要的地位。土壤动物群落与植物群落间存在动态的相互影响,从生态系统的角度来看,地上植物群落的变化必然会对其中的土壤动物群落产生影响。土壤动物通过对凋落物的分解、直接取食植物根系以及对土壤的耕耘作用,使土壤理化性质发生改变,对地上植物群落产生直接或间接的作用,从而影响其结构、功能和恢复动态。关于土壤动物群落与植物群落之间相互作用的研究,目前已经成为国际上土壤动物研究领域的热点之一,国内鲜有报道。论文选择退化红壤地区4种主要的植被恢复类型,即旱生性草坡、稀疏针叶林、针叶林、针阔混交林,另设裸地和顶级的常绿阔叶林2种类型作为研究的对照。于2005年秋至2006年夏,对这6个样地中的土壤节肢动物群落进行了详细的四季调查,包括大型和中小型节肢动物。大型节肢动物按凋落物层,0-10cm土层,10-20cm土层;中小型节肢动物按凋落物层,0-5cm土层,5-10cm土层进行取样。同时,对各类型的植被群落和土壤养分进行调查和分析。在全面了解植被恢复对土壤节肢动物的影响的基础上,分析土壤大型土壤节肢动物的种类分布与植被群落的关系,以及中小型节肢动物(螨类和弹尾虫)与土壤养分之间的关系。这有助于深入地认识植被恢复对土壤生物多样性的影响,对于进一步探讨土壤退化的内部机制具有重要的理论意义。主要研究结果如下:1.该地的土壤节肢动物种类丰富,其中大型土壤节肢动物21目107科;螨类和弹尾虫构成该地中小型节肢动物的主体,二者共计60科,A/c值高达7.6。大型节肢动物的优势类群为路舍蚁属,占总数的10.24%;常见类群金龟甲科幼虫(Scarabaeidaelarvae),食虫虻科幼虫(Asilidae larvae)、蚁属(Formica)等20类,二者共占大型动物总数的73.85%。中小型节肢动物中,土壤螨类计4亚目53科,前气门亚目17科,中气门亚目8科,甲螨亚目27科,粉螨亚目1科;弹尾虫计2亚目7科23属。螨类的优势类群有甲螨亚目的矮汉甲螨(Nanhermanniidae)和单翼甲螨科(Haplozetidae);常见类群包括囊螨科(Ascidae)、长须螨科(stigmaeidae)、跗线螨科(Tarsonemidae)、珠足甲螨科(Belbidae)、真卷甲螨科(Archoplophoridae)、菌甲螨科(Scheloribatidae)等16类;其余35类均为稀有类群。弹尾目中的优势类群包括优势类群为符跳属(Folsomia)、类符跳属(Folsomina)、小圆跳属(Sminthurinus)、棘跳属(Onychiurus);常见类群包括长跳属(Entomobrya)、土跳属(Tullbergia)、裔符跳属(Folsomides)等3类;其余16类为稀有类群。2.土壤节肢动物在空间上存在明显的成层现象;在时间上,节肢动物四季的类群数、个体数量、优势类群和多样性都发生明显的变化。在空间分布上,无论类群数还是个体数,均是凋落物层最高,0-10 cm层次之,10-20cm层最低。在时间上,就类群数而言,夏季的类群数最高,达到112类,其次为春季,有92类,秋季与冬季的类群数相同,均为82类。就个体数而言,夏季的个体数量也最多,占全年总数的33.42%;其次是冬季,占全年总数的25.45%;而秋季的个体数量最少,仅占全年总数的17.42%。优势类群也随着季节发生改变,春季的优势类群有路舍蚁属和举腹蚁属,夏季的优势类群仅有路舍蚁属,秋季的优势类群包括路舍蚁属和近扭尉属,到了冬季,蚁属和华扭尉属成了优势类群。密度-类群DG指数综合了各方面的因素,反映出土壤大型节肢动物群落多样性的四季变化:夏季>春季>秋季>冬季。土壤小型节肢动物(螨类、弹尾虫)群落多样性的四季变化均为:夏季<春季<秋季<冬季。3.植被恢复对土壤大型节肢动物的组成产生影响,各植被类型的优势类群存在差异。在较高级的分类等级目上看,除了膜翅目是所有类型共有的优势类群,各类型还有自己的优势类群。从低级的科属分类等级上看,不同植被类型的优势类群各不相同。4.植被的恢复对土壤大型节肢动物群落产生影响。顶级常绿阔叶林群落中土壤动物的类群数和个体数均最高,针阔混交林、旱生性草坡、稀疏针叶林的类群数和个体数相近,针叶林的个体数和类群数均显著地低于这些林分,裸地最低。密度-类群DG指数综合考虑了诸多因素,得到不同恢复类型土壤节肢动物群落多样性的变化为:常绿阔叶林>针阔混交林>稀疏针叶林>旱生性草坡>针叶林>裸地。5.植被恢复不同类型的土壤大型节肢动物群落可作如下聚类:裸地、常绿阔叶林群落各为一类、其它4个类型聚为一类。聚类分析的结果表明:裸地中的土壤大型节肢动物群落与林地间存在着本质的区别,常绿阔叶林群落中的土壤动物可以认为是该地的顶级群落。而从灌草丛向常绿阔叶林恢复的过程中,土壤节肢动物群落并没产生显著的变化。6.植被恢复对土壤螨类的密度、类群数、多样性都有显著影响;季节变化对土壤螨类的密度的影响达到显著性水平,但对类群数、多样性的影响未达到显著性水平。应用MGP方法对甲螨群落进行分析,在MGP分析Ⅰ中,各类型的甲螨群落均属于M型;MGP分析Ⅱ中,裸地群落属于MP型,旱生性草坡群落属P型,稀疏针叶林群落、针叶林群落、针阔混交林群落均属于M型,而顶级的常绿阔叶林群落的甲螨群落属于O型。应用MⅠ指数对中气门螨类群落进行分析,表明,各类型的MⅠ指数无明显的变化。应用DCA分析对土壤螨类群落进行排序,将6类型分为3类:裸地、常绿阔叶林群落各为一类,其它4类型归为一大类,显示了土壤螨类群落对植被恢复的响应。7.应用个体密度、类群数及多样性指数等指标,研究植被类型对弹尾虫群落特征的影响。结果表明,各项指标以常绿阔叶林为最高,裸地处于最低水平,基本没有弹尾虫的存在。旱生型草坡、稀疏针叶林、针叶林和针阔混交林等4种植被恢复类型的土壤弹尾虫群落得到了一定的恢复,但各类型之间土壤弹尾虫群落没有明显差异。Bray-Curtis指数显示侵蚀裸地与顶级常绿阔叶林的差异最大(0.99),各植被恢复类型与顶级常绿阔叶林的差异也较明显,但各恢复类型间弹尾虫群落间差异较小。8.应用CCA分析大型节肢动物种类、样地分布与植被群落关系,结果表明,乔木多样性是影响土壤大型动物群落结构和种类组成的最重要的因子,草本优势度也是影响动物种类的重要因子。CCA排序图很好地揭示了土壤动物种类分布对植物群落的适应,狂蚁属、窒蟷科、大蚊科幼虫、大尉属等顶级常绿阔叶林的常见类群,它们分布的环境具有灌木多样性、均匀度高,乔木丰富度高,草本植物的种类、多样性低的特点。近扭尉属、蚁属、幺蚣科等旱生性草坡的常见种类,它们分布的环境具有草本植物盖度高,灌木种类稀少的特点。复翅蠊科、多刺蚁属等是针叶林、针阔混交林的常见类群,它们分布的环境的特征是草本植物的种类、多样性高,灌木种类丰富,但乔木种类单一。9.在退化红壤恢复的各类型,土壤的小型节肢动物组成发生改变,同时对采集的土壤样品性质进行分析。以4类主要中小型动物与主要的土壤性质的9项指标为研究对象,采用典范相关分析,研究影响土壤小型动物的主要因子。结果表明,蔗糖酶、全氮、含水量、有机质的载荷量较高,4类土壤动物中弹尾纲、前气门亚目载荷量较高。在二者作为整体的组成中,蔗糖酶、全氮、含水量对弹尾纲、前气门亚目影响较大,而有机质和全氮对甲螨亚目和中气门亚目影响较大,其它土壤因子与小动物之间的相关性较弱。土壤动物类群组的变化被土壤性质第Ⅰ变量和第Ⅱ变量解释的比例为13.75%和32.71%,仍有53.54%以上变化不能得到解释。

【Abstract】 Soil arthropods are important components of forest ecosystems,and they play a particularly significant role in the process of nutrient cycling and energy flowing.From the view of ecosystem,it is the fact that the changes of plant affect the soil arthropods.Soil arthropods alter the availability of nutrients for plant as well as structure and function of plant directly or indirectly,by their effect on the process of decomposition,and their moving ability.The interactions between soil fauna and plant has been one of the hottest topics in soil faunal ecology abroad,however,few has been mentioned in China.Soil arthropods was studied quarterly in degraded red soil,including 5 restored plantatins, i.e.a bare land(Ⅰ),a xeric mesophilous herbosa(Ⅱ),a sparse coniferous woodland(Ⅲ),a coniferous woodland(Ⅳ),a coniferous-broadleaf mixed woodland(Ⅴ) and an evergreen broadleaf forest(Ⅵ)(ⅠandⅥwere used as CK).Soil fauna,including macroarthropods and microarthropods was sampled in the above vegetations from the autumn of 2005 to the summer of 2006.Soil macroarthropods were taken in the litter,0-10cm and 10-20cm,while, soil microarthropods were taken in the litter,0-5cm and 5-10cm.In the meantime,nutrients contents in the soil and species diversity of vegetation at different restored plantations were studied too.Based on a comprehensive knowledge of the soil fauna in the degraded red soil, the effects of vegetation restored on the soil fauna were analyzed.And the relevance between plant and soil macroarthropods as well as the relevance between soil property and soil microarthropods was also studied.It will contribute to,a deep understanding of the effects of vegetation restoration on soil biodiversity and it is theoretically of great significance to further exploration of internal dynamics of soil degradation.The main findings are as follows:1.Soil macroarthropods in the degraded red soil were abundant,a total of 107 families was observed,falling into 21 orders.The dominant group was Tetramorium,accounting for 10.24%of the total;the common groups were Scarabaeidae larvae,Asilidae larvae and Formica and others,altogether making up 63.61%of the total.The main groups of microarthropods were Acarina and Collembolan,A/C value was as high as 7.6.A total of 53 families of Acarina was observed,fairing into 4 suborders(17 Prostigmata,8 Mesostigmata,27 Oribatida and 1 Astigmata),dominant families were Nanhermanniidae and Haplozetidae;common families were Ascidae,Stigmaeidae,Tarsonemidae and others.A total of 23 genus of collembolan was observed,falling into 2 suborder and 7 families,respectively,dominant genus were Folsomia,Folsomina,Sminthurinus, Onychiurus;common genus were Entomobrya,Tullbergia,Folsomides.2.Both the numbers of groups and individuals in the litter were much richer than in 0-10cm and 10-20cm.both the number of soil macroarthropods and individuals varied widely in four seasons.The number of soil macroarthropods was highest in summer (112),the second highest in spring(92) and lowest in autumn and winter(82). Individuals was highest in summer,accounting for 33.42%of the annual total,lowest in autumn,accounting for 17.42%of the annual total The dominant groups changed along with seasons.Tetramorium and Crematogaster were dominant groups in spring, Tetramorium was the only dominant group in summer.By density-groups index(DG), the seasonal fluctuations of diversity of soil macroarthropods can be represented as: summer>spring>autumn>winter,however,the seasonal fluctuations of diversity of soil microarthropods can be represented as:summer<spring<autumn<winter.3.With the restoration of vegetations,both the number of groups and individuals have increased,which tended to fluctuate in the different restored vegetations.In the hierarchy of order,the dominant groups changed in the different vegetations,excepting for Hymenoptera,which was dominant group in all of these vegetations.However,in the hierarchy of family or genus,dominant groups in different vegetations were different.4.Both the number of groups and individuals were highest inⅥ,they were second highest, inⅡ,ⅢandⅤ,and they were lowest inⅠ.By density-groups index(DG),the diversity of soil macroarthropods in different restored vegetations can be represented as. follows:Ⅵ>Ⅴ>Ⅲ>Ⅱ>Ⅳ>Ⅰ.5.Based on Jaccard index and Bray-Curtis index,Soil macroarthropods in different restored vegetations can be clustered as follows:ⅠandⅥwas isolated respectively,all others were classed into one group,which showed the response of soil macroarthropods to these different restored plantations in degraded red soil.The result showed that the soil macroarthropods in the bare land was essentially distinct from those in the forested land,and the soil macroarthropods in the natural forest markedly different from those in the other forested land.In the 4 different restored plantations(Ⅴ,Ⅲ,ⅡandⅣ),there were no remarkable changes in the soil macroarthropod communities.6.The characteristics of soil mite community structure was analyzed by using individual density,number of taxon,diversity,DG index,abundance,and evenness as well as MGP analysis and MI index.The highest density was observed inⅥandⅢ,the lowest density was observed inⅠandⅣ.The same pattern was observed by using other indices.MGPⅠanalysis indicated the soil oribatida communities of all these ecosystems match the M Pattern,however,MGPⅡanalysis identified different patterns for them,ie,MP pattern forⅠ,P pattern forⅡ,O pattern forⅥ,M pattern for others. No difference was observed by MI index among these different restored plantations. These 6 communities were classed into 3 groups by using DCA analysis,ie,ⅠandⅥwas isolated respectively,all others were classed into one group,which showed the response of soil mite to these different restored plantations in degraded red soil.7.The characteristics of soil collembolan community were analyzed by using individual density,number of taxon,diversity,abundance as well as evenness index.The lowest species richness was found in the bare land and highest was found in the evergreen broadleaved forest,difference amongⅡ,Ⅲ,Ⅳ,Ⅴwas not significant.The same pattern was observed by using other indices.The Bray-Curtis between the 5 restored plantations and the evergreen broadleaf forest were high,reaching its maximum(0.99) whenⅠandⅥwere compared.This survey indicated that planting is beneficial to restoration of collembolan community in degraded red soil,although these degraded ecosystems were still in the early stages of the restoration.8.Soil macroarthropods and plant diversity were studied under different restored plantations in degraded red soil,and CCA was used to explore the distributional relationship between soil macroarthropods and plant diversity.The genus(or sites)-enviroument biplots of CCA were automatically mapped using CANOCO 4.5, and the relationships between the distribution of the species and communities with the plant diversity were clearly revealed on these biplots.The results also showed that shannon diversity of tree was the most important factor influencing the distribution of soil macroarthropods,Simpson index of herbage was major factor affecting soil macroarthropods.Common genus of evergreen broadleaf forest such as Paratrechina, Macrotermes,closely linked to high shannon diversity of herbage and high richness of tree.Common genus of xere-mesophilous herbosa such as Pericapritermes, Formica,closely linked to low Shannon diversity of shrubbery and high cover of herbage.Ubiquitous genus such as Polyrhachis,Blattella Caudel,closely linked to low Shannon diversity of tree and high Shannon diversity of herbage.CCA can be a useful tool to understand the distribution of soil macroarthropods in degraded red soil.9.The relationship between soil microarthropods and soil properties was analysed by canonical correlation analysis based on 4 groups of soil microarthropos and 9 indexes of soil properties in the paper,which respectively include Prostigmata,Mesostigmata, Oribatida,Collembola and soil organic matter,pH,moisture content,total nitrogen, available kalium Urease,Sucrase,AP,Protease aimed to find major soil factor variables affecting soil microarthropods in different restored plantations in degraded red soil.The results indicated that CoUembola,Prostigmata were affected firstly by the contents of Sucrase,total nitrogen and moisture content in soil,and Mesostigmata,Oribatida were affected by the contents of organic matters and total nitrogen in soil,while other factors had not much relation with that pattern of distribution.It was found that 13.75%and 32.71%of the variance of soil microarthropods in degraded red soil was respectively explained by the first and the second canonical variable of soil factor variable,the rest was not interpreted yet.

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