瘿绵蚜科虫瘿的多样性研究

是否形成虫瘿及虫瘿的位置、形态等是蚜虫生物学的重要特征.本文在已有标本采集记录和资料的基础上,从形成虫瘿的寄主植物、虫瘿的类型、虫瘿着生的部位和虫瘿的形态结构4方面对瘿绵蚜的虫瘿多样性进行了系统研究.结果表明该科蚜虫的虫瘿在类型上有虫瘿和伪虫瘿之别;在着生部位上,有叶片、侧脉、主脉、叶柄、复叶总轴、嫩枝等不同部位;就虫瘿而言,在形状上有尖椒状、鸡冠状、袋状、球状、黄瓜状、枫叶状和倍花状之分,在结构上有单室和多室两类.同时,基于系统分类及虫瘿多样性研究的结果,初步探讨了瘿绵蚜科虫瘿的演化,为进一步从分子水平深入研究虫瘿演化奠定基础.     

球蚜科分类概述

球蚜科是蚜虫类的一个重要类群,包括多种林业害虫和检疫害虫,本文从形态特征、生活史、寄主植物、地理分布,进化起源等方面对世界及中国球蚜物种进行了综合论述.结果表明,球蚜科具有较多原始的特征,是蚜虫中较为古老的一类;球蚜具有包括异寄主全周期、异寄主不全周期、同寄主不全周期等复杂的生活周期,形成虫瘿是其主要的为害特征;球蚜科物种专性寄生在松科植物上,主要包括云杉属Picea、松属Pinus、落叶松属Larix、冷杉属Abies等属的植物,在蚜虫物种和寄主植物之间存在着平行演化的现象;球蚜的分布与寄主植物的分布相一致,主要是北半球温带和亚热带高山.世界球蚜有8属59种,区系成分主要是新北区、古北区.中国球蚜科有6属20种,分布在黑龙江、云南、四川等地,主要是古北区成分.初步推测球蚜科可能首先在松属植物上起源.     

五节根蚜亚科（半翅目：蚜总科：瘿绵蚜科）基于线粒体基因COⅡ的分子系统学研究

五节根蚜亚科蚜虫大多营异寄主全周期生活,具有明显的原生寄主专一性,且能在原生寄主上形成形态各异的虫瘿,因此成为一个倍受关注的类群.但是亚科内族的划分问题却一直存在争议.基于线粒体基因COⅡ部分序列,用PAUP*4.0、MrBayes3.0等软件分别构建了该亚科的MP树、ML树和贝叶斯树,研究了五节根蚜亚科的系统发育关系.结果显示3种算法所构建的系统树都强烈支持五节根蚜亚科是一个单系;系统树明显分为两大分支,提示该亚科下分两个族.将蚜虫主要形态特征及其原生寄主植物信息对应到系统树上,经分析认为这两大分支分别对应倍蚜族和五节根蚜族.依据DNA分类理论,基于系统树的拓扑结构及序列间遗传距离,初步推测术知种No.16961是肚倍蚜Kaburagia rhusicola,但究竟是哪一个亚种尚不确定.     

福建省蚜虫类昆虫多样性研究

在已有标本采集记录和资料的基础上,从分类阶元、寄主植物、寄生方式和寄生部位等角度探讨了福建省蚜虫类昆虫的物种多样性.福建省蚜虫已知9科80属164种,东洋区种类占明显优势;寄主植物种类繁多,涉及60科158属;寄生方式为3种,不形成虫瘿、形成虫瘿和形成伪虫瘿;寄生部位多样化,以叶片、茎、嫩尖、嫩枝为主要寄生部位.福建省植物物种多样性特点决定了福建省蚜虫类物种多样性特点,很好的反映了蚜虫和其寄主植物的密切关系.     

虫瘿多样性及其与寄主植物和环境间关系

虫瘿是自然界极常见的生物现象,凝聚着昆虫与植物间显著、复杂而密切的协同关系。本文主要阐述了致瘿昆虫的主要类群及其在植物上的致瘿部位、虫瘿外部形态、虫瘿发育过程、虫瘿内部结构、虫瘿寄主植物多样性以及虫瘿空间分布规律等,探讨了致瘿昆虫和寄主植物间相互关系,以及影响虫瘿空间分布的环境因素等。最后对目前虫瘿生物学存在的问题及以后的研究方向进行了讨论,以期为有害虫瘿的控制和有益虫瘿的开发与利用,以及致瘿昆虫与寄主植物间协同演化关系、致瘿昆虫的致瘿机理等研究奠定一定的理论基础。     

基于三个分子标记的粉毛蚜亚科DNA条形码

【目的】粉毛蚜亚科昆虫是重要的林业害虫,但是由于蚜虫体型较小,形态特征趋于简化,可用于物种鉴定的有效特征非常有限,因此一般基于外部形态特征难以对蚜虫物种实现快速准确的鉴定。本研究获取该亚科2属10种的DNA条形码标准序列,解决部分物种的分类问题,同时比较了3种标记对粉毛蚜亚科（Pterocommatinae）物种快速鉴定的效率。【方法】基于蚜虫的线粒体细胞色素氧化酶C亚基I（cytochrome oxidase subunit I, COI）基因、细胞色素b（cytochrome b, Cytb）基因和蚜虫初级内共生菌Buchnera 6-磷酸葡萄糖酸脱氢酶（gluconate-6-phosphate dehydrogenase, gnd）基因,对2属10种共197号样品进行NJ分析、遗传距离的计算以及基于相似性的物种鉴定分析。【结果】与K-2P模型相比,基于p-distance模型计算得到的遗传距离更小,序列差异频次图上种内距离与种间距离的重叠区域也小于前者;COI序列的物种鉴定成功率最高。获取了粉毛蚜亚科近200条DNA条形码标准序列,并建立了基于3个标记的该亚科物种DNA条形码序列库。【结论】在粉毛蚜亚科DNA条形码研究中,p-distance模型要优于K-2P模型;COI序列具有最高的条形码分析效率;增毛卷粉毛蚜Plocamaphis assetacea可能为蜡卷粉毛蚜Plocamaphis flocculosa的同物异名。     

肚倍蚜瘿内种群动态与虫瘿生长发育的关系

【目的】本研究拟对肚倍蚜Kaburagia rhusicola瘿内种群动态与虫瘿生长发育关系进行研究,为深入研究肚倍蚜的生物学特性奠定基础。【方法】在2014年4月27日至7月16日期间,在湖北竹山田间每间隔10 d随机选择长势基本一致的青麸杨,分别采集健康肚倍蚜虫瘿,每次采集样本30个。采用排水法测量虫瘿体积,之后将虫瘿解剖,取出瘿内蚜虫,置于培养皿内,采用四分法统计数量。对虫瘿体积增长与瘿内蚜虫种群变化进行了相关性分析。【结果】虫瘿体积增长与蚜虫种群数量增长之间呈极显著正相关(r=0.960,P0.01)。虫瘿体积与蚜虫种群数量增长分别呈现3个峰,即5月6日至5月16日、5月26日至6月6日和6月26日至7月6日。肚倍蚜瘿内密度变化的主要趋势是先降后升,中间有一个小的起伏。【结论】本研究表明虫瘿大小和瘿内蚜虫种群数量有关,虫瘿体积可用于评价干母的环境适合度。     

瘿绵蚜属五新种和三堡瘿绵蚜新属新种

《蚜虫概论》(1957)中瘿绵蚜属仅有2种。本文记载9种,包括5新种。另记述三堡瘿绵蚜新属新种。这些蚜虫都为害杨树,在枝或叶上形成虫瘿。 中国已知的9种瘿绵蚜和二堡瘿绵蚜(节省篇幅,旧种记述略): 1.早螺瘿绵蚜Pemphigus protospirae Lichtenstein (图1-6,66)寄主:青杨。分布:北京;欧洲。 2.柄脉叶瘿绵蚜 Pcmphigus anobursarius Chang 新种 3.杨柄叶瘿绵蚜Pemphigus populi Courchet(图13—17,65)寄主:小叶杨。分布:北京、黑龙江、     

中国斑蚜科物种多样性及地理分布格局

从分类群的多样性、寄主植物多样性和地理分布多样性3个方面系统研究了中国斑蚜科蚜虫的物种多样性。分类群的多样性从亚科、属和种3个不同阶元进行了描述。寄主植物的多样性体现在：不同蚜虫类群寄主植物的丰富度,其中角斑蚜亚科的寄主最为丰富,包括14个科的植物;同类寄主植物上取食蚜虫的多样性,其中桦木科植物上有15个属的蚜虫取食。在地理分布上,斑蚜科在中国主要分布在古北界的4个区和东洋界的3个区,以古北界的成分占优势;而且华北区种类最为丰富,其次为东北区、华中区、华南区和蒙新区。属的分布类型可分为12种,分别为中国特有分布型、东亚特有分布型、古北界特有分布型、东洋界特有分布型、全北界分布型、全北界和澳洲界共有分布型、古北界和东洋界共有分布型、全北界和东洋界共有分布型、全北界和非洲界共有分布型、古北界、东洋界和非洲界共有分布型、东洋界和澳洲界共有分布型及东洋界、非洲界和澳洲界共有分布型,其中以东亚特有分布型为最丰富,有10个属;其次为全北界分布型,有7个属,同时中国的特有成分也相当高,涉及6个属。纵观整个中国斑蚜科属级阶元的分布类型,中国的斑蚜科以全北界成分为主,其次为东洋界成分。斑蚜科在中国的地理分布,由东向西,种类越来越贫乏,特别是青藏高原,种类很少;从南北向来看,以华北区为中心,向两边扩散,东北区、华南区和华中区种类都有所减少。     

两种毛蚜线粒体基因组比较分析

【目的】线粒体基因组分析已被应用于昆虫系统发育研究。本研究以蚜科Aphididae重要类群毛蚜亚科物种为代表,测定并比较分析了该类蚜虫的线粒体基因组特征,探讨了基于线粒体基因组信息的蚜虫系统发育关系重建。【方法】以毛蚜亚科三角枫多态毛蚜Periphyllus acerihabitans Zhang和针茅小毛蚜Chaetosiphella stipae Hille Ris Lambers,1947为研究对象,利用长短PCR相结合的方法测定线粒体基因组的序列,分析了基因组的基本特征;基于在线t RNAscan-SE Search Server搜索方法预测了t RNA的二级结构;基于12个物种(本研究获得的2个物种和10个Gen Bank上下载的物种数据)的蛋白编码基因(PCGs)序列,利用最大似然法和贝叶斯法重建了蚜科的系统发育关系。【结果】两种毛蚜均获得了约94%的线粒体基因组数据,P.acerihabitans获得了14 908 bp,控制区为1 205 bp;C.stipae获得了13 893 bp,控制区为609 bp。两种毛蚜同时获得33个基因,包含接近完整的13个蛋白编码基因(PCGs)(nad5不完整),18个tRNA,2个rRNA基因;ka/ks值表明,C.stipae的进化速率更快。从基因组组成、基因排列顺序、核苷酸组成分析、密码子使用情况、t RNA二级结构等特征来分析,两种蚜虫线粒体基因组基本特征相似。系统发育重建结果表明毛蚜亚科、蚜亚科的单系性得到了支持,毛蚜亚科位于蚜科的基部位置。【结论】两种毛蚜线粒体基因组的基本特征相似,符合蚜虫线粒体基因组的一般特征,两种线粒体基因组的长度差异主要来自控制区长度的不同;系统发育重建支持毛蚜亚科与蚜亚科的单系性,毛蚜亚科位于蚜科较为基部的位置。研究结果为蚜虫类系统发育重建提供了参考。     

Social parasitism and behavioral interactions between two gall-forming social aphids

We investigated ecological and behavioral aspects of the interactions between two social aphids, Pseudoregma bambucicola and Astegopteryx bambucifoliae (Hormaphidinae, Cerataphidini), both of which produce second-instar, sterile soldiers in galls formed on Styrax suberifolius, in Taiwan. By censusing their galls, either species was found to invade galls of the other species. Twenty-eight (58%) out of 48 A. bambucifoliae galls contained P. bambucicola, while four (6%) of 69 P. bambucicola galls contained A. bambucifoliae. Furthermore, P. bambucicola behaved like social parasites in galls of A. bambucifoliae. Colonies of P. bambucicola produced much fewer or even no soldiers compared with those in natal galls. Our experiments also revealed that individual aphids of P. bambucicola more successfully intruded into galls of A. bambucifoliae than into conspecific galls, and that guarding soldiers of P. bambucicola effectively prevented aphids from invading their galls and permitted only conspecific soldiers to join their colonies. These differences in behavior provide good explanations for the differences in the frequency of invaded galls found between the two species. Received 4 September 2007; revised 7 January 2008; accepted 17 January 2008.     

The evolution of gall parasitism accompanied by a host shift in the gall aphid, Eriosoma yangi (Homoptera: Aphidoidea)

Eriosoma aphids form leaf-roll galls on various elm species. Eriosoma yangi occurring on the Japanese elm is an obligatory "cuckoo" parasite, invading and usurping galls of other Eriosoma species. But another host race of E. yangi induces its own galls on the Chinese elm. Multivariate morphometries show that the two host races differ significantly in quantitative characters. Overwintering eggs of the parasitic form were experimentally transferred to the Chinese elm. The resultant larvae still exhibited a parasitic habit and developed into adults, which multivariate morphometries classified as the parasitic form. This form is considered different enough from the gall maker to be given a species rank. Eriosoma aphids facultatively invade other Eriosoma galls, where they sometimes deposit offspring. It is hypothesized that the parasitic form on the Japanese elm originated from a founder population which had migrated from the Chinese elm. On this new host, only those fundatrices that invade other species' galls will be able to leave offspring, because aphid fundatrices cannot gall plants other than their primary hosts. It is assumed that the parasitic form was established under intense selective pressure following a population bottleneck. This hypothesis was corroborated by field investigations at sites where the two forms occur sympatrically.     

Clonal mixing in the soldier-producing aphid Pemphigus spyrothecae (Hemiptera: Aphididae)

Illuminating the genetic relationships within soldier-producing aphid colonies is an essential element of any attempt to explain the evolution of the altruistic soldier caste. Pemphigus spyrothecae is a soldier-producing aphid that induces galls on the leaf petioles of its host (trees of the genus Populus). At least a quarter of the aphids within the clonally produced gall population are morphologically and behaviourally distinct first-instar soldiers that defend the gall population from predation. Using field trapping and microsatellites, we investigated the degree of clonal mixing within natural gall populations. Field trapping in the UK showed that all the migrants of P. spyrothecae and of two other Pemphigus species were wingless first-instar soldiers. The average degree of mixing estimated from trapping P. spyrothecae migrants was 0.68% (range = 0-15%). Microsatellite genotyping of 277 aphids from 13 galls collected in Italy revealed an average mixing level of 10.4% (range = 0-59%). Six galls contained more than one clone (range = 2-5 clones). Non-kin aphids were not restricted to the soldier caste but were evenly distributed across instars. An additional gall, from which 527 occupants were genotyped, contained 12 non-kin aphids distributed among nine clones, showing that clonal diversity can be high even when mixing is very low. These observations suggest that although soldiers migrate regularly and can moult and reproduce within foreign galls, clonal mixing in this species is generally low and is unlikely to provide a barrier to the evolution of investment by the aphid clones in an altruistic soldier caste.     

Mutualism in a community context: the positive feedback between an ant–aphid mutualism and a gall-making midge

Although mutualisms are widespread and often described in natural history accounts, their ecological influences on other community members remain largely unexplored. Many of these influences are likely a result of indirect effects. In this field study, we investigated the indirect effects of an ant–aphid mutualism on the abundance, survival rates and parasitism rates of a co-occurring herbivore. Rabdophaga salicisbrassicoides (Diptera: Cecidomyiidae) induces rosette galls on the developing shoots of Salix exigua trees, and populations can reach outbreak densities (up to 1,000 galls/stem) in central Washington State (USA). Ant-tended aphids feed on these same stems and often feed on gall tissue. In this study we used a combination of manipulative experiments and observational surveys to test the hypothesis that the abundances of aphids, ants, and galls have positive and reciprocal effects on one another, in a manner that would create a positive feedback loop in population growth. In addition, we examined whether the combined presence of ants and aphids reduces parasitism rates for the gallers. In support of the positive feedback loop hypothesis, aphids enjoyed higher population growth rates in the presence of ants and galls, the presence of ants and aphids resulted in increased abundance of galls, and the abundances of ants, aphids and galls were all positively correlated with one another. However, the mechanism underlying the positive effect of ants and aphids on galler density remains unknown, as the mutualism did not affect parasitism rates. More broadly, this study demonstrates that mutualisms can have significant and complex indirect effects on community and population ecology.     

A galling aphid with extra life-cycle complexity: Population ecology and evolutionary considerations

In most gall-forming aphids, only the fundatrix is able to induce a gall on the host plant. In Smynthurodes betae Westw. (and a few other species), F₂ descendants emerge from the mother gall and induce their own, morphologically different galls. This constitutes an added complexity to the already very complex life cycle of gall-forming aphids. We investigated the ecology of S. betae on marked trees and shoots at four sites in Israel. Gall initiation, gall distribution and density, and temporal changes in clone size within the galls were investigated during two consecutive years. We discuss the possibility that the two-gall life cycle evolved from the typical one-gall system of most gall aphids, and the possible selective advantage of this added complexity in the life-history strategy of gall aphids. Although the total reproductive output of S. betae is not higher than in related species with a single gall per life cycle, there seems to be an advantage in the subdivision of each aphid clone into several galls, thus reducing the risk of the accidental extinction of the clone (genotype) by environmental factors, including parasites and predators.     

The cost of conflict in aphid societies

Most social aphids are found within plant galls, inside of which clonally‐derived family groups feed, and specialized larval castes forego reproduction and perform various cooperative tasks, including group defence. When unrelated aphids move between clones, conditions are ripe for conflict because galls and cooperative defence are shared resources that are vulnerable to exploitation. A key unknown is whether conflict is costly in aphid social groups. We show that diversity within groups is negatively correlated with performance in the North American social aphid, Pemphigus obesinymphae. A substantial fraction of productivity is invested into drifting. However, drifting aphids tend to mature and depart non‐natal galls prior to the seasonal peak in fecundity. These results suggest that when unrelated individuals move between groups, social aphids may experience conditions consistent with a tragedy of the commons. These results also emphasize the strongly convergent properties associated with conflict across the spectrum of animal and microbial sociality.     

Do aphid galls provide good nutrients for the aphids?: Comparisons of amino acid concentrations in galls among Tetraneura species (Aphididae: Eriosomatinae)

Some aphid species induce leaf galls, in which the fundatrix parthenogenetically produces many nymphs. In order to ensure high performance, galls have to provide the aphids with sufficient nutrients, in particular, amino acids as a nitrogen source. We tested this hypothesis using six Tetraneura aphid species that induce closed galls. We extracted free amino acids from the whole gall tissues of unit weight and quantified the concentration of each amino acid. There were large differences in the total amino acid concentrations among galls of the Tetraneura species. Tetraneura species in which higher concentrations of total amino acids were found in the gall tended to produce larger numbers of offspring. Of the amino acids found, asparagine was predominant in the gall. The asparagine concentration in T. yezoensis galls was several hundred times as high as in control leaves. We discussed why such a high level of asparagine accumulates in aphid galls.