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梅园节肢动物群落动态及朝鲜球坚蚧与其寄主、天敌间的关系
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摘要
本文系统地总结了国内外节肢动物群落的研究进展,并对果园节肢动物群落、天敌利用以及害虫与植物互作关系等进行了综合分析。运用生态学、统计学、分类学和化学分析测定等知识综合研究了梅园节肢动物群落生态学,对节肢动物群落组成结构与功能、多样性、种群动态、主要天敌对害虫的控制作用、农用化学物质对节肢动物群落的影响以及害虫与植物互作关系等进行了探讨,为梅园主要害虫综合防治提供科学依据。其主要研究结果如下:
     1.对梅园节肢动物群落结构系统调查,共发现节肢动物169种,分属4纲18目90科,按照营养取食关系,划分为4个亚群落,6个功能类群。植食类、瓢虫类、蜘蛛类、寄生类和中性昆虫的物种数分别占总物种数的41.62%、25.80%、21.40%,7.59%和3.59%。植食类是主要害虫类群,瓢虫类和蜘蛛类均为主要捕食性天敌类群,其中朝鲜球坚蚧和黑缘红瓢虫分别是梅园害虫和天敌优势种;节肢动物群落结构较为稳定,天敌自然控害效果比较明显。
     2.应用聚集度指标、Iwao法和Taylor幂法则分别研究了3-9月份朝鲜球坚蚧和黑缘红瓢虫的分布型及其变化动态,结果表明:朝鲜球坚蚧和黑缘红瓢虫在不同方位和冠层的枝条上以及所有密度下均呈聚集分布,以上层和南向冠层的聚集度最高,下层和北向冠层最低,在上、中和下冠层均呈现前期明显的高聚块,后期呈逐渐降低的趋势;提出了该蚧最适抽样数及序贯抽样模型。同时,运用地统计学研究了不同时期朝鲜球坚蚧若虫及黑缘红瓢虫种群的空间格局和空间相关性,结果表明不同时期朝鲜球坚蚧若虫种群和黑缘红瓢虫幼虫种群的半变异函数均为球型,其空间格局为聚集分布,二者间具很强追随关系,说明黑缘红瓢虫是朝鲜球坚蚧的天敌优势种。
     3.对梅园瓢虫亚群落组成及相对丰盛度等研究表明,该亚群落有瓢虫68种,隶属15科48属,黑缘红瓢虫、红点唇瓢虫等为优势种;影响瓢虫亚群落多样性的主要因素前中期为均匀性,后期则为丰富度;优势瓢虫种群在各方位的空间分布型均属于聚集分布。
     4.对中性昆虫亚群落研究表明,中性昆虫占节肢动物总群落比例>10%,与天敌跟随趋势明显且略早于优势种害虫,弥补了天敌自然控害的不足;蚤蝇、水蝇和蚊科中性昆虫同天敌在时间生态位重叠值较大,可作为一个功能团共同起作用;天敌与中性昆虫种群的灰色关联度R(Y,X_j)排序前3位的分别为:黑缘红瓢虫>异色瓢虫>草间小黑蛛。
     5.对杀虫剂和除草剂干扰下的节肢动物群落结构与组成分析表明,长期施用杀虫剂可明显增加植物性害虫种类及其优势度,显著降低群落多样性,减少了蜘蛛目、膜翅目、鞘翅目、双翅目、天敌和中性昆虫功能集团的物种数,削弱了冠层天敌种类和数量上对害虫的制约潜能,诱发了叶部植食害虫的暴发:除草剂对地面节肢动物群落、各亚群落的多样性、均匀度、优势集中性和物种丰富度影响较大,而对冠层节肢动物群落影响较小。
     6.首次采用土施法和OECD(经济合作与发展组织,Organization for EconomicCo-operation and Development,简称OECD)推荐的滤纸接触法系统研究了5种稀土元素对梅园土栖动物群落的影响以及对地下害虫蛴螬的生态毒性。结果表明:5种稀土元素对土栖动物群落主要群落参数影响较为显著;对蛴螬的急性和慢性毒性大小均依次为La>Ce>Pr>Nd>Sm,但毒性响应不同。14 d时所有稀土对蛴螬生长均有刺激作用;28 d后Nd和Sm由刺激作用转为抑制作用,La和Ce刺激作用减弱,而Pr刺激作用增强。稀土能改变土栖动物群落的结构,并对其产生一定的生态毒性。
     7.调查研究了梅园杂草群落结构组成及杂草群落对节肢动物群落影响,结果表明:梅园杂草有28科97种,其种类较为丰富,且各时期杂草优势种突出;对群落相关生态指标的模糊聚类分析得出不同生长时期梅园杂草和昆虫群落的优势种,并认为杂草群落的变动对节肢动物群落有一定程度的影响。
     8.对梅园节肢动物群落的时空动态和生态位研究结果表明,节肢动物群落结构较丰富,且各群落特征参数随时间推移而变化;以植食性的桃蚜和桃小食心虫空间生态位宽度最大,朝鲜球坚蚧时间生态位宽度最大;在捕食性天敌中黑缘红瓢虫时间和空间生态位宽度均最大,异色瓢虫、红点唇瓢虫及寄生蜂次之;黑缘红瓢虫与朝鲜球坚蚧的生态位重叠值在时空二维上均较大,且二者在时间上的同步性和空间上的同域性均优于其它天敌,为天敌优势种群。
     9.对黑缘红瓢虫捕食功能作用研究表明,黑缘红瓢虫捕食功能反应和寻找效应与自身密度关系用HollingⅡ和Hassell方程模拟程度较好,黑缘红瓢虫对朝鲜球坚蚧具有较强的捕食能力:其对猎物的寻找效应和种内干扰随着自身密度的增大而降低;随着空间异质性趋于复杂而捕食作用率减弱。
     10.在室外对梅园主要节肢动物生物学特性和室内对黑缘红瓢虫人工饲料研究表明:黑缘红瓢虫1年1代,以成虫越冬,以高龄幼虫和成虫捕食朝鲜球坚蚧控制作用最强;朝鲜球坚蚧1年l代,以2龄若虫越冬,主要以固定若虫和雌成虫危害;明确了以草鱼肉、鲜猪肝、夜蛾幼虫粉为主的人工饲料各配方基本满足了黑缘红瓢虫成虫的营养需要,其存活率、饲料利用率和转化率均较高,捕食功能反应均符合HollingⅡ模型:但各配方人工饲料饲育的成虫捕食能力随饲育时间延长而减弱,寻找效应随着猎物密度增加而下降。
     11.对8种寄主上朝鲜球坚蚧种群动态、寄主选择性及其与枝条物理结构、寄主和蚧体内化学物质间关系进行系统的研究,结果表明:不同生长势寄主对朝鲜球坚蚧的适合度由高至低顺序均是桃、梅、李、杏、苹果、梨、樱和山楂,且分成桃、梅、李、杏最适宜寄主;苹果、梨、樱桃适宜寄主和山楂次适宜寄主三类,均以6月份的寄主适合度最高,且以生长势好的寄主最高,中等的次之,弱的最差:朝鲜球坚蚧更趋向于最适宜寄主上刺吸取食和产卵,且若蚧存活率、卵的孵化率以及羽化率都较高,自然种群增长最快;朝鲜球坚蚧对寄主的取食和产卵选择性分别与寄主间的表皮毛密度、长度、皮层厚度、枝条直径和黄酮、单宁等之间均达极显著负相关,相关系数r分别在-0.9413~-0.8816和-0.9624~-0.8943之间;同样,取食和产卵选择性分别与8种寄主间的可溶性糖、可溶性蛋白质、游离氨基酸、含水量、CarE、AChE、GSTs和蚧体可溶性蛋白质等之间均达极显著正相关,相关系数,r范围分别为0.9025~0.9413和0.8937~0.9201;取食和产卵选择性一致。以取食表皮较为光滑、表皮毛稀疏、长度短、皮层薄,寄主可溶性蛋白质、游离氨基酸和含水量多,黄酮和单宁少的桃、梅、李和杏等寄主的蚧体内CarE、AChE、GSTs和可溶性蛋白质活性较高,对寄主选择性较强,危害严重;取食苹果、梨和樱桃的次之;取食山楂的最低。
     12.运用功能法评价了杀虫剂对捕食性瓢虫集团捕食功能的影响,结果表明:杀虫剂对各捕食性瓢虫功能反应模型基本结构没有影响,均符合HollingⅡ模型,只改变了模型各参数,但药剂处理后捕食性瓢虫的控虫能力、寻找效应都显著降低;杀虫剂造成的各捕食性瓢虫的死亡率(D_t)和捕食功能减退率(FD_t)各不相同,且按乐斯本、阿维菌素和速扑杀顺序减小,表明速扑杀是一种对捕食性瓢虫致死作用和捕食功能影响均较小而安全性较高的药剂。
The advance on arthropod community was systemically reviewed in the thesis.The summary of arthropod community,utilizes of natural enemies and relationship between pests and plants in orchards were summed up.According to the investigation,field and indoor trial in ecology,structure and diversity of arthropod community,population dynamic,control function,effect of chemic substance and relationship between pests and plants in plum orchard,by using knowledge and technique of ecology,statistics,taxonomy and chemic identification and analysis etc,the results were as follows:
     1.The structure and dynamics of the arthropod community in plum orchard was investigated,identified,classified and analyzed in 2006 in plum orchard of Anhui Science and Technology University and horticulture of Fengyang County.169 species belonged to 4 classes,18 orders and 90 families were discovered in the plum orchards.According to nutritional and feeding relationship,the arthropod community was divided into 4 sub-communities and 6 functional groups;the relative abundance of the 4 sub-communities was respectively 41.62%of phytophages,47.20%of predators,7.59%of parasitoids and 18.7%of neutralities of arthropods.Dominant species of phytophages were D.koreanus Borchsenius;of natural enemies were C.rubidus Hope and spiders in plum orchards.By the analysis of number dynamics,ecological dominance and diversity of the functional groups,the results showed that the structure of arthropod communities in the plum orchards was relatively stable.It was obvious that natural enemies were able to control pests effectively.
     2.Spatial pattern and time-series dynamics of Didesmococcus koreanus Brochseniusenius and Chilocorus rubidus Hope were analyzed by calculating five indices of aggregation,and parameters of Iwao model and Taylor's power law model form March to September.The results showed that the spatial pattern of D.koreanus Brochsenius in horizontal and upright dimension,at different direction,altitude and population density in plum orchard belonged to an aggregated distribution pattern,and the fundamental component was the group composed of several individual and the individual was aggregated.The Aggregation intensity of D.koreanus Brochsenius was the highest in upper and southern part of crown,and the lowest in its lower and northern part.The aggregation was due to the biological habits of D.koreanus Brochsenius and environmental heterogeneity.No mater D.koreanus Brochsenius was in the upper,middle, or lower crown of plum tree,its aggregation intensity was higher before mid-ten days of April,and then gradually decreased after the last ten days of April because of preying of natural enemies and so on C.rubidus Hope.Based on the data of the spatial distribution, the model of the most suitable sampling quantity is established and the suitable sampling quantity at different density of D.koreanus Brochsenius and error level is presented.The sequential sampling diagram was drawn to investigate the D.koreanus Brochsenius population levels in plum orchard.The pattern of C.rubidus Hope in plum orchard fitted the aggregated distribution pattern,and the aggregation of it was due to environmental effects.Meanwhile,the spatial pattern and correlation of D.koreanus Brochsenius and C. rubidus Hope in different periods were investigated with Geostatistics.The results showed that the semivariogram of D.koreanus Brochsenius and C.rubidus Hope could be described by spherical model,indicating that their spatial pattern were in agreement.The spatial variation was aroused by its spatial autocorrelation in small scales,and the spatial pattern had no apparent changes in different periods.The range varied from 9.56-15.70 m and from 18.83-19.74 m,respectively.The amount and spatial distribution of C.rubidus Hope was closely related to that D.koreanus Brochsenius,which revealed that C.rubidus Hope was the dominant natural enemy of D.koreanus Brochsenius.
     3.The structure and relative abundance of ladybugs sub-community of plum orchard were carried out,after primarily classification and authenticating in laboratory.There were 68 species belong to 48 geneses,15 families.The dominant species were Chilocorus rubidus Hope,Chilocorus kuwanae Silvestri,Harmonia axyridis,Coccinella septempunctata Linnaeus,Progylaea japonica and Stethorus punrtillum Weise.Changes of time pattern of the species richness(SR),total individual number(N),index of diversity (H′),and evenness(J) were preliminarily studied by using multiple early and medium-term of growth of stepwise regression analysis.The results showed that in plum tree,where the main factor affecting ladybugs was evenness and in lopper it was the species richness.Calculation the diversity of result of spatial pattern for time,space and three dimensions(time-vertical and level space) showed that the spatial pattern of ladybugs in plum orchard all belonged to aggregated distributions.
     4.The study of neutral insect sub-community was carried out.It was showed that neutral insect whose relative abundance was more than 10%was one of important parts and played an undoubtedly role in the arthropod community of plum orchards of Anhui science and technology university and horticulture of Fengyang county form March to November month in 2007.Meanwhile,the occurrence tendency of the neutral insect which accompanied with the occurrence of natural enemy was somewhat earlier than the dominant pest,which could,to some extent,make up with the defect of natural enemies who take place always behind the pests in the natural situation.The study on time niche among natural enemies and neutral insects showed that encounter opportunity of natural enemies to Chilocorus rubidus Hope,Chilocorus kuwanae Silvestri,Harmonia axyridis, Coccinella septempunctata Linnaeus,Progylaea japonica and Stethorus punrtillum Weise in time sequences were larger.And that is to say,the neutral insect has active effect to maintain the population of natural enemies in the plum orchard.The principal component analysis was used to explore the neural insect's function in the community,a conclusion was drawn that predominant effect of neutral insects was more scattered and they played the role by the functional group.Through the calculation of grey correlation analysis of natural enemy's effect to mixed neutral insect population,it was showed that the sequence of grey relational grade was as follow:Chilocorus rubidus Hope>Harmonia axyridis>Erigonidiurn graminicola.
     5.Disturbed by Chlorpyrifos,Supracid and Abamectins,the composition and structure of arthropod community were systematic investigated and analyzed at representative sampling sites.And the results showed that application Chlorpyrifos and Supracid caused some phytophagous species and dominant index of arthropod community becoming abundant observably in plum orchard.Species diversity of general community was decreased significantly.The numbers of species in community were decreased remarkably.Specially,species in the categories of Araneae,Hymenoptera,Coleoptera and Diptera,or functional groups of predators,parasitoids and neutrals were reduced. Therefore,the potential impacts of the phytophagous natural enemies of predators and parasitoids weakened in category and individual.To some extent,this might lead to outbreak of the phytophagous in the plum leaves.The compared with plot of Abamectins and no being control was not significant difference.So indiscriminate application of Chlorpyrifos and Supracid should be prohibited in integrated pest management(IPM). Consequently,it is advisable that an ecological pest management(EPM) program be implemented with Abamectins insecticides and horticulture measure.The arthropod community was divided into four sub-communities:pests,predatory,parasitoids and neutralities by the standard of nutrition and hunting.The differences of the diversity, abundance,evenness and structures of arthropod community and sub-community in plum orchard among treatments Gramoxone,Simazine and contrast(CK) were analyzed by using a number of community parameters.The results showed that the diversity and evenness index of arthropod community and four sub-communities were low in herbicide plots,while the dominant concentration was high,the arthropod community stability was lower than the contrast,and the pest out break was more possible in herbicide treatments than that in the contrast.
     6.By the method of OECD(Organization for Economic Co-operation and Development,OECD) filter paper contact firstly,the effects of five rare earth elements on soil fauna community structure and their ecological toxicity to Holotrichis parallels in Plum Orchard were studied.The results showed that there were significant differences between treatments and the control in soil fauna species,quantity of main species,diversity index,evenness index,Dominant centralization,ecological dominant index,species richness and function group.Urgent and chronic toxic test showed that the differences between the treatments and the control were significant.Toxicity order of five rare earth elements were La>Ce>Pr>Nd>Sm,but their toxicity response were Different.The growth of Holotrichis parallels and its pupae were stimulated by all the rare earth elements in 14 d,and weight of Holotrichis parallels and its pupae were increased obviously;28 d after the Nd and Sm convert the stimulating effect into inhibition,the stimulating of La and Ce were weakened,and the stimulating of Pr were enhanced.Rare earth elements could change the structure of the soil fauna,and the soil fauna had a certain eco-toxicity.
     7.The structure of weed communities in plum orchards of Anhui science and Technology University and horticulture of Fengyang County which many arthropods dwelled was studied,and there were 97 species belonged to 28 families.The temporal dynamics of individual number,species abundance and diversity of both weed and arthropod communities were dealt with,and it was also analyzed that the arthropod communities characteristics index were affected by the corresponding index of weed communities,and the change trend of the first and second dominant species both weed and arthropod communities in all of two kinds of plum orchards were similar.The cluster analysis indicated that there existed similarity on the time dynamics of weed and arthropod communities in two plum orchards to some extent.
     8.The arthropods communities in plum orchard were investigated on Space-time dynamic and temporal spatial niche.The results showed that the arthropod community was abundant.The individuals of species,diversity indices,and evenness changes with time. Myzus persicae and Asiaarposina sasokii had the widest spatial niche breadth,while Didesmococus koreauus Borchs had the widest temporal niche breadth.Among the natural enemies Chilocorus rubidus Hope and Didesmococus koreauus Borchs overlapped larger than that of the others,which indicated their synchrony in temporal dimension and their similarity in spatial dimension.As the dominant natural enemies the two populations should be protected and utilized to control plum pest.
     9.The experiment was conducted to study the functional response,density of predator and prey,and their relationship,mutual interference,space heterogeneity and temperature effects on Chilolocorus rubidus Hope to Didesmococcu koreanus Borchs in the laboratory. The results were showed that the functional response,and effects of mutual interference and temperature on predacious efficiency of Chilocorus rubidus Hope at different larvae and the adult ladybugs were determined as HollingⅡand Hassell equation,x~2 test indicated that the theoretical expectation fits the observe values,and the differences were not significantly different(p<0.05).The searching efficiency decreased with the increase of predator density.The mutual interference among individual predators and preys were more obvious with the increase of their density.There were mutual interference among the larves and adults of the ladybugs,and the predatory rate increased as its density decreased. Predacious efficiency of Chilocorus rubidus Hope was dropped with complexity of space heterogeneity.
     10.Using field observations and periodic sampling of Chilocorus rulolus Hope and Didesmococcus koreanus Borchs of plum orchard and studing on diets of Chilocorus rulolus Hope in Anhui Science and Technology University in 2005,studies on the bionomics of them showed that all 1 generation developed annually.The adult of C. rulolus Hope oveiwinters in weeds and seams and it is an important natural enemy of D. koreanus Brochsenius.It mainly occurs in peach and plum trees,and suppresses the population of D.koreoaaus Borchs efficiently.Its larvae and adult were the main preyer of the scale insect.The second nymphal stage of D.koreoaaus Borchs over wintered on brush, emerge in March of the next year,from April to May was the adult's stage and begin to hatch among the first ten days of June and mid-July.The D.koreanus Borchs attack trees mainly in forms of fixed nymsphs and adult females.Meanwhile,natural enemies had high capability of control,so it was suggested to adopt integrated pest management.The feeding results of C.rubidus Hope on the five artificial diet indicate that the mixture of grass carp, pig-liver and powder of Noctidae larvae could met basica nutrition requirement of the adult respectively.The adult fed with the mixture had high a survival rate,but the females had a relatively low egg production.Preying functional response of C.rubidus Hope adults on D. koreanus Borchs was fitted to HollingⅡmodel.Adults fed with No.3 and No.5 diets preyed strongly on D.koreanus Borchs.However,from 25 to 290 days,their preying capacities(1-31 head) decreased with raising duration.Food-seeking efficiency of C. rubidus Hope(0.3802-0.0717) declined with increasing the densities of D.koreauus Borchs(2-10 head).A better efficiency was detected in C.rubidus Hope adults fed with No.3 and No.5 diets.Adding Preying stimuli in the adult diet could significantly increase the survival rate from winter,egg number praying ability and searching efficiency.
     11.By focusing on 8 host plant species including peach,plum,cherry plum,apricot, apple,pear,cherry and hawthorn,we investigated the feeding,oviposition,population dynamics of Didesmococcus koreanus Broth in the field,and the relationship among the selectivity of D.koreanus Borchs,branch physical structure and chemical substance in the hosts and D.koreanus Borchs in laboratory.The fitness of eight host plants to D.koreanus was quantitatively measured with selective index(SI),damage index(DI),and preference index(PI).Moreover,index of population trend(I) was used as an important estimate for the influence of host plants on the natural population of D.koreanus.The results indicated that,under all three types of plant growing conditions,the order of fitness was peach,plum,cherry plum,apricot,apple,pear,cherry and hawthorn.The fitness of 8 host plants,presenting as a normal distribution,were consistently the best in June with a good growing condition~inferior medium growing condition~weak condition pattern.D.koreanus preferred to feed and oviposit on peach,plum,cherry plum and apricot,where they featured higher survival,eclosion and hatched rate and more rapid nymphae increase,and also higher values for other four indexes(SI value>0.875, PI value>5.469,I value>5.75,DI value>1.049).According to the fitness indexes,the host plants of D.koreanus may be divided into three categories,with peach,plum,cherry plum and apricot as the most suitable hosts,apple,pear and cherry the suitable,and hawthorn the least suitable category.With the best linear regression equation was PI=-0.3389+DI, there was an obvious linear relation between PI,SI and DI(r_(0.01)=0.937).Measuring and scanning showed that there was a significant negative correlated(r_(0.01)=-0.976) between fitness and the epicuticle hair numbers,hair length,epidermis thickness and branch diameter.The feeding and oviposition have been controlled by the physical structure of branches.Compared to the most suitable hosts,the epicuticle hair numbers, hair length,epidermis thickness and branch diameter of hawthorn were,respectively,2.73, 3.27,2.77 and 3.47 times higher.Feeding-selectivity and oviposistion-selectivity of D. koreanus Borchs with the eight hosts of the hairs density,length,epidermis thickness, branches diameter,flavones and tannin,and all reached a very significant negative correlation,and their correlated coefficients r were -0.9413 to -0.8816 and -0.9624 to -0.8943 respectively.The same reasoning,feeding-selectivity and oviposistion-selectivity of D.koreanus Borchs with the eight host plants of branch's soluble sugar,host plant's soluble protein,free amino acids,water content,CarE,ACHE,GSTs and the insect's soluble protein content,and all reached a very significant positive correlation,and their correlated coefficients r were 0.9025 to 0.9413 and 0.8937 to 0.9201 respectively. Feeding-selectivity and oviposistion-selectivity of D.koreanus Borchs is consistent on the eight hosts.Activity of CarE,ACHE,GSTs and the soluble protein content of D.koreanus Borchs that feeding peach,plum,cherry plum and apricot which branches relatively smooth surface,low hair density,length short,thin epidermis,more content of host's soluble proteins,free amino acids,water content,and low content flavones and tannins were relatively high.The D.koreanus Borchs of the host's selectivity strong is serious harm to the eight hosts,these of apple,pear and cherry were inferior,and these of hawthorn were the worst.Above results may provide clues for resistance breeding,forecast and optimal control strategies.
     12.The effects of Chlorpyrifos,Supraeid and Abameetins on the predation function of predatory ladies group were evaluated with function method.The result showed that all predating functional responses of predatory ladies were in accordance with the HollingⅡmodel by insecticides treatments,but their model parameters were changed.It was obviously lower than controls that in abilities of controlling pest and the searching efficiency of the predators.The mortalities(D_t) and decreasing rates of predation function(FD_t) of surviving predatory ladies were different,and their order was Chlorpyrifos,Abamectins and Supracid for big to small.Susceptibilities of predatory ladies to Chlorpyrifos were relatively higher(mean of D_t was 74.64%),on the contrary with the Supracid treatment(mean of D_t was 30%),and Abamectins treatment was medium. Supracid was a relatively more securities pesticide to predatory ladies and its affects were small to lethality and predation function.
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