扁蚜亚科昆虫虫瘿多样性研究(半翅目,蚜科)

虫瘿是蚜虫诱导植物异速生长的结果,虫瘿作为蚜虫重要的延伸特征,对蚜虫系统分类、系统发育关系、以及起源演化等研究具有非常重要的作用.而且虫瘿的形态结构、着生部位等在蚜虫的物种间存在非常丰富的多样性,是蚜虫重要的生物学特征,也是物种鉴定的重要依据之一.本文在已有标本采集记录和资料的基础上,从结瘿的植物、虫瘿着生部位、形态结构及类型等4个方面对扁蚜亚科虫瘿的多样性进行了系统研究.结果表明该亚科蚜虫大多都在原生寄主上形成虫瘿,个别属及种可在次生寄主上成瘿;虫瘿在类型上有虫瘿和伪虫瘿之别;在着生部位上,有叶片、叶脉、叶柄、小枝、粗枝等;虫瘿的形状也十分多样,有管状、袋状、球状、半球形、刺球状、纺锤形、圆锥形、分支状、香蕉束状等;在结构上既有单室、多室之分,也有开放型、封闭型之别.对于虫瘿多样性的研究,可为虫瘿演化规律的探讨提供重要信息,也是基于虫瘿进行物种鉴定的重要基础.     

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、     

不同寄主植物叶片上蚜虫的形态适应

昆虫对寄主植物之间的形态适应性是两者相互关系的重要组成部分, 本文以植食性蚜虫为研究对象, 研究其对不同种类寄主植物叶片的形态适应。选取寄生于禾本科、 杨柳科、 壳斗科和松科4科7属植物叶片上的26种蚜虫, 基于有翅孤雌蚜和无翅孤雌蚜, 在光学显微镜下观察、 测量并统计比较了37个形态特征及其种内变异。以克隆平均值为基础数据,选取与蚜虫取食和附着于植物表面相关的形态特征（喙末端、跗节Ⅰ、Ⅱ和爪）为变量,分别得出无翅、 有翅孤雌蚜和所有蚜型3个聚类分支图,并将各种蚜虫的寄主植物科、 属分别对应到分支图上。结果表明：体型、腹管和触角的形态在不同蚜虫的科间有一定差别, 喙末端、 跗节Ⅰ、跗节Ⅱ及爪的形态则在不同的寄主植物间存在差异; 在蚜虫种内各形态特征存在一定变异, 变异系数的范围为1.89%~26.08%。3个聚类分析的结果显示,不同种类蚜虫形成的分支基本对应不同科、属的寄主植物;而杨一条角蚜Doraphis populi、三堡瘿绵蚜Epipemphigus imaicus和杨柄叶瘿绵蚜Pemphigus matsumurai则处在较为特殊的分支位置, 那是因为前者为次生寄主型, 后两者均能产生虫瘿。研究结果表明寄生于不同植物相同部位上的蚜虫存在形态特征的明显分异, 对应不同的寄主植物, 蚜虫与寄主植物之间存在着形态适应关系。     

蚜小蜂科一中国新纪录属——原蚜小蜂属记述（膜翅目：小蜂总科）

记述蚜小蜂科1中国新纪录属——原蚜小蜂属Protaphelinus Mackauer,对该属模式种尼氏原蚜小蜂Protaphelinus nikolskajae(Yasnosh)作了重新描述,附有雌、雄形态特征及其蚜虫寄主照片。本研究的尼氏原蚜小蜂采自黑龙江五大连池,寄生火山杨Populus sp.上的远东枝瘿绵蚜Pemphigus borealis Tullgren。目前该属世界仅记载1种,是造瘿蚜虫瘿绵蚜科Pemphyidae瘿绵蚜属Pemphigus的专性寄生蜂。     

蚜虫寄主植物与取食部位的多样性

在中国科学院动物研究所已有标本采集记录和国内外文献资料的基础上,分别从蚜虫类Aphidina科级和属级阶元系统研究世界范围内蚜虫寄主植物与取食部位的多样性.蚜虫的寄主植物种类繁多,涉及267科2120属,尤其以菊科、禾本科、豆科、伞形科、唇形科、蔷薇科、茜草科、兰科、壳斗科、杨柳科、胡桃科等植物为主.13科蚜虫的寄主植物差别很大:在科级水平,球蚜科Adelgidae、纩蚜科Mindaridae和平翅绵蚜科Phloeomyzidae的寄主植物类群比较单一,蚜科Aphididae和瘿绵蚜科Pemphigidae的寄主植物范围最为广泛;在属级水平,各属蚜虫间寄主植物也有明显差异,有143属蚜虫的寄主植物多于2科,其中蚜科的属占多数,蚜属Aphis、瘤蚜属Myzus、长管蚜属Macrosiphum、粗额蚜属Aulacorthum和声蚜属Toxoptera的寄主植物最多,各超过100科290属.在不同的寄主植物上寄生着不同种类的蚜虫,其中16科47属寄主植物上寄生的蚜虫多于14属.蚜虫在寄主植物上的取食部位丰富多样,可分为7类,分别为叶片、嫩梢、嫩枝、茎、花、根部、果实等.以叶片、茎、嫩枝和嫩梢为蚜虫主要取食部位.平翅绵蚜科主要取食嫩枝、茎和根部,短痣蚜科Anoeciidae主要取食叶片、嫩梢和根部,球蚜科、群蚜科Thelaxidae和毛管蚜科Greenideidae取食叶片、嫩梢、嫩枝和茎等部位,纩蚜科取食叶片、嫩梢、茎和花等部位,大蚜科Lachnidae和斑蚜科Drepanosiphidae除取食叶片、嫩梢、嫩枝、茎等部位外,前者还在根部取食,后者还寄生在果实上,根瘤蚜科Phylloxeridae在除花以外的其它6个部位取食,蚜科、瘿绵蚜科和毛蚜科Chaitophoridae的取食部位最为多样,它们在上述7个部位均可取食.还初步讨论了在不同寄主植物上蚜虫物种的分化,以及蚜虫与寄主植物之间的对应关系.     

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。     

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。     

角倍蚜虫瘿的组织学结构与功能解析

五倍子是一类具有重要的经济价值的蚜虫虫瘿,在中医、化工、纺织、食品等行业用途广泛。角倍是我国五倍子生产中最重要的一种,是由角倍蚜Schlechtendalia chinensis在盐肤木Rhus chinensis叶轴翅上刺激形成,产量占我国五倍子总产量的75%以上。本研究发现五倍子虫瘿柄部分化出了异常膨大的木质部与叶轴直接相连以稳定支撑虫瘿。虫瘿壁由疏松的薄壁组织构成,有效降低了虫瘿重量并维持虫瘿的足够硬度。柄部除了异常膨大的木质部外,还分布有大量的维管束及裂生道,并从虫瘿柄部一直延伸至虫瘿末端。维管束及裂生道有大量弯曲的细小分枝,形成包裹整个虫瘿的网状结构。从内壁到外壁维管束与裂生道逐渐减小,这既能防止外源微生物入侵,也有效阻止虫瘿内水分向外挥发,保持了虫瘿内蚜虫所需的高湿度生境。内壁大量的分枝有利于虫瘿内蚜虫吸收营养,网状分枝结构在有限倍体中增大了裂生道及维管束数量、体积与表面积,使其能运输更多的营养及提供最大面积的物质交换场所。封闭的虫瘿外壁长有大量绒毛,绒毛中分布有气孔,绒毛有保护功能,可以抵御其它食草动物的攻击;虫瘿内壁粗糙,凹凸不平,有利于蚜虫的附着,同时增加了内壁的表面积;内壁中还分布有大量的缝隙和孔,这些缝隙可以吸收蚜虫排泄的蜜露,使瘿内能保持清洁。本文通过解析角倍蚜虫瘿的结构和功能,从生态学角度阐明蚜虫与植物之间相互适应关系,为研究昆虫与植物协同进化提供新的有力证据。     

中国新纪录属--小裂绵蚜属(同翅目,瘿绵蚜科)

研究了中国瘿绵蚜科Pemphigidae 1新纪录属--小裂绵蚜属Schizoneurella Hille Ris Lambers,1973;记述1新纪录种,印度小裂绵蚜Schizoneurella indica Hille Ris Lambers,1973.标本采自云南省昭通市,该种在苹果Malus pumila根部取食,蚜虫是重要的苹果害虫,可造成较大经济损失.文中提供了各型检索表、形态记述、寄主植物及地理分布,配有形态特征图和野外生态照片.研究标本保存在中国科学院动物研究所动物标本馆和英国自然历史博物馆.     

蚜虫对杨属植物不同寄生部位的形态适应(英文)

为了明确取食相同植物不同部位和营不同虫瘿的蚜虫是否存在形态适应,论文以取食杨属Populus植物的10种蚜虫为研究对象,基于蚜虫喙末端、各足跗节和爪的形态测量数据,对122个克隆的形态变异通过一般的统计描述、典型变量分析(canonical variates analysis,CVA)以及聚类分析等方法进行了研究。结果显示,不同的蚜虫克隆间形态特征存在分异,与不同的虫瘿类型和取食部位相关。3个典型变量分析明显地区分了形成虫瘿和不形成虫瘿的蚜虫克隆,在叶片上形成真虫瘿和伪虫瘿的克隆以及不产生虫瘿的克隆,并形成明显的克隆簇。而用于分析的形态学特征,如喙末端、跗节和爪的测量值,其一般统计描述的差异也支持这些区分。蚜虫自然种群不同克隆簇的区分很好地对应了不同的取食部位和不同的虫瘿类型,且各簇的形态特征体现了各自的特点,表明了蚜虫对杨属植物已经形成了良好的形态适应。同时,也初步讨论了不同蚜虫克隆簇形态适应产生的原因。并建议在基于形态特征探讨昆虫的系统发育关系和进行传统分类时,必须考虑昆虫形态特征的适应性;在深入研究昆虫与寄主植物相互之间关系时,需要将各种形态特征综合考虑并关注其它的影响因素。     

A galling aphid furnishes its home with a built-in pipeline to the host food supply

Galling insects modify the developmental pathways of their host plants and create a protected and favourable microhabitat for their offspring. Galling aphids reproduce in their galls and the resulting clones often comprise hundreds of individuals. We followed the histological changes in the host Pistacia palaestina (Anacardiaceae) induced by the galling aphid Geoica wertheimae (Aphidoidea, Pemphigidae: Fordinae). We showed that the leaf tissues are altered in the gall in a way that gives the aphids easy access to the vascular system from which they obtain their nourishment. Specifically, the cuticle that lines the normal leaf epidermal cells is absent on the surface of the inner gall cavity, and the normal palisade cells are replaced by parenchymatous tissue with numerous wide latex ducts. Associated with these ducts, many new sieve tubes (phloem elements) are formed just a few cell layers below the inner gall surface. This arrangement enables the hundreds of aphids to feed simultaneously and continuously in the gall throughout the summer.     

The evolution of host plant manipulation by insects: molecular and ecological evidence from gall-forming aphids on Pistacia

One of the most striking characteristics of gall-forming insects is the variability in gall position, morphology, and complexity. Our knowledge of the driving forces behind the evolutionary divergence of gall types is limited. Natural enemies, competition, and behavioral constraints might be involved. We present a cladogram, based on sequences of COI and COII (1952bp), of mitochondrial DNA for the evolution of 14 species of gall-forming aphids (Fordinae). These insects induce five gall types with remarkable morphological variation on Pistacia spp. hosts. The parsimony cladogram divides the Fordinae into three lineages, Fordini and Baizongiini, and a third (new) sister group including the previously Fordini member, Smynthurodes betae (West). We then use ecological data to trace and explain the evolution of gall morphology. The aphids seem to have evolved gradually towards better ability to manipulate their host plant, induce stronger sinks, and gain higher reproductive success. We suggest that the ancestral gall type was a simple, open, "pea"-sized gall located on the leaflet midvein. Some Fordini and S. betae evolved a two-gall life cycle, inducing a new gall type on the leaflet margin. The Baizongiini improved the manipulation of their host by inducing larger galls near the midvein, with stronger sinks supporting thousands of aphids. Similar gall types are induced at similar sites on different Pistacia hosts suggesting control of the aphids on gall morphology and frequent host shifts. Thus, even extreme specialization (specific gall and host) is flexible.     

The effects of host-plant properties on gall density,gall weight and clone size in the aphidAploneura lentisci (Pass.) (Aphididae: Fordinae) in Israel

Summary Samples of shoots ofPistacia lentiscus carrying galls of the aphid,Aploneura lentisci, were collected at three localities in Israel. Shoots growing near pruning scars carried more galls than elsewhere on the plant, but these galls weighed less and contained fewer aphids (smaller clones). The proportion of empty galls increased with gall density. Crowding of galls at such sites may be due to the early burst of buds at the time of aphid emergence from the overwintering eggs, and not to active search for preferred sites. Shoots bearing larger numbers of leaves carried heavier galls, which contained larger aphid clones. The position of the galled leaf on the shoot had no effect on gall weight nor on clone size. The physiological condition of the plant may be an important environmental (ecological) factor affecting the variation in clone-size and in aphid morphology among galls.     

肚倍蚜对高单宁环境的抗氧化反应研究

肚倍蚜Kaburagia rhusicola Takagi是我国的一种重要的资源昆虫,其瘿内生活环境较特殊(虫瘿单宁含量为70%)。为了解肚倍蚜适应高单宁环境生化机制,本研究以性母为对照,测定了瘿内不同时期肚倍蚜体内总抗氧化能力(T-AOC)以及超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)和谷胱甘肽S-转移酶(GST)的活性,分析了肚倍蚜体内的抗氧化系统对高单宁环境胁迫所作出的相关反应。研究表明,与对照相比,瘿内时期肚倍蚜体内的总抗氧化能力、CAT和POD活性均显著升高;SOD和GST活性除在有翅蚜阶段与对照无差异,在其他阶段的活性均显著高于对照。本研究验证了高单宁环境对肚倍蚜造成氧化胁迫作用,认为肚倍蚜体内的抗氧化能力和抗氧化酶活性提高是其对高单宁环境的一种适应。     

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.     

Histological study of galls induced by aphids on leaves of Ulmus minor: Tetraneura ulmi induces globose galls and Eriosoma ulmi induces pseudogalls

The histological study of galls may provide information on the evolution of the organisms that induce them. The walls of two aphid-induced galls on leaves of Ulmus minor were therefore studied histologically: a globose gall induced by Tetraneura ulmi and a pseudogall induced by Eriosoma ulmi. Galls are regarded as extended phenotypes of aphids, and therefore, they can be used as tools for phylogenetic studies. The walls of the galls induced by T. ulmi are not reminiscent of the ungalled leaf structure of U. minor in any area, showing both cellular hypertrophy and hyperplasia. Galls induced by E. ulmi resemble the leaf structure of U. minor in certain aspects, but in most aspects they do not, showing only cellular hypertrophy. Processes of growth and differentiation are observed in both galls. Based on the findings of this study and other recent publications, we propose to categorize the galls induced by aphids into four types: (1) pseudogalls; (2) closed galls with a “door”; (3) closed galls determined by active processes of growth and differentiation of the lamina of the leaf; and (4) closed galls determined by active processes of growth and differentiation of the midvein.     

Molecular phylogeny of Fordini (Hemiptera: Aphididae: Pemphiginae) inferred from nuclear gene EF-1 alpha and mitochondrial gene COI

The tribe Fordini is a fascinating group because of its complicated life history, primary host specificity and gall-forming characteristic. Different species produce galls with different morphology on different parts of the host plants. The EF-1alpha-based, COI-based and combined sequences-based phylogenetic trees with three algorithms MP, ML and Bayes all strongly suggest that Fordini is a monophyletic group with two clades corresponding to two subtribes, Fordina and Melaphidina, each also monophyletic. Some important morphological characters and primary host plants of aphids were mapped onto the phylogenetic tree to analyse the division of subtribes and to uncover at which level the aphids correspond to their primary hosts, Pistacia and Rhus. Results suggest that the division of subtribes in Fordini is closely related to host selection of aphids. The evolution of gall morphology and the probable driving force behind it in this tribe were also discussed. The Fordini aphids seem to have evolved towards a better ability to manipulate their host plant, induce strong sinks and gain high reproductive success. Galls in this tribe evolved mainly along two directions to attain this goal: (i) by enlarging the gall from small bag to spherical, even big cauliflower-like, and changing the galls' location or forming two galls in their life cycle (Fordina); (ii) by moving the gall position from midrib, petiole of the leaflet, and eventually to the common petiole of the compound leaf (Melaphidina).     

Manipulation of food resources by a gall-forming aphid: the physiology of sink-source interactions

Summary We examined the capacity of the galling aphid, Pemphigus betae, to manipulate the sink-source translocation patterns of its host, narrowleaf cottonwood (Populus angustifolia). A series of ¹⁴C-labeling experiments and a biomass allocation experiment showed that P. betae galls functioned as physiologic sinks, drawing in resources from surrounding plant sources. Early gall development was dependent on aphid sinks increasing allocation from storage reserves of the stem, and later development of the progeny within the gall was dependent on resources from the galled leaf blade and from neighboring leaves. Regardless of gall position within a leaf, aphids intercepted ¹⁴C exported from the galled leaf (a non-mobilized source). However, only aphid galls at the most basal site of the leaf were strong sinks for ¹⁴C fixed in neighboring leaves (a mobilized source). Drawing resources from neighboring leaves represents active herbivore manipulation of normal host transport patterns. Neighboring leaves supplied 29% of the ¹⁴C accumulating in aphids in basal galls, while only supplying 7% to aphids in distal galls. This additional resource available to aphids in basal galls can account for the 65% increase in progeny produced in basal galls compared to galls located more distally on the leaf and limited to the galled leaf as a food resource. Developing furits also act as skins and compete with aphid-induced sinks for food supply. Aphid success in producing galls was increased 31% when surrounding female catkins were removed.     

Factors affecting components of fitness in a gall-making wasp (Cynips divisa Hartig)

Summary The galls of the agamic generation of the cynipid gall-wasp Cynips divisa were studied; these galls are found only on the main veins of the leaf. Galls are concentrated on the middle veins of leaves, and generally occur closer to the midrib than to the edge of the leaf. Survival of the gall-maker and fecundity of the energing wasp depend on wasp size and position of the gall. There is a striking pattern in gall size according to the number of galls on the vein. The number of galls on a vein appears therefore to have a strong effect on potential fecundity.     

Fungal diversity in galls of baldcypress trees

The baldcypress midge (Taxodiomyia cupressi and Taxodiomyia cupressiananassa) forms a gall that originates from leaf tissue. Female insects may inoculate galls with fungi during oviposition, or endophytes from the leaf tissue may grow into the gall interior. We investigated fungal diversity inside of baldcypress galls, comparing the gall communities to leaves and comparing fungal communities in galls that had successful emergence versus no emergence of midges or parasitoids. Galls of midges that successfully emerged were associated with diverse gall fungal communities, some of which were the same as the fungi found in surrounding leaves. Galls with no insect emergence were characterized by relatively low fungal diversity.