四十五种叶蝉的染色体研究（同翅目：叶蝉总科）

研究观察了45种中国雄性叶蝉的减数分裂,其中44种的核型为首次报道,染色体数目变化在2n=12～26之间,性别决定均为XO型。从叶蝉总科的组型图来看,该科染色体数目变化在2n=8～28之间,峰值为2n=18(16+XO),另外几种类型2n=16,20,22也有较高的出现频率。科内染色体数目的进化不具有明显的方向性,2n=22(20+XO)是该科的原始核型,易位导致的不均等互换可能是染色体数目进化的主要机制。从精子发生来看叶蝉总科与角蝉总科的关系较为密切,两者的共同特点是：①精母细胞体积较大,显著不同于沫蝉和蝉科;②减数分裂行为及精子变态过程相似;③染色体数目较少,染色体体积较大;④减数分裂前期具有典型的花束期,没有弥散期,因而不同于蜡蝉。但是由于叶蝉总科的染色体变异范围明显大于角蝉总科,而角蝉总科的核型相对较为保守,从核型上来说角蝉总科是比叶蝉总科较为原始的类群。     

中国19种蜡蝉的核型研究(同翅目：蜡蝉总科)

研究了中国19种蜡蝉的染色体数目、性别决定和染色体行为。2n=28(26+XO)被认为是蜡蝉科的原始核型特征。由于蜡蝉精巢具有被膜,减数分裂前期Ⅰ具有弥散期,因此作者认为蜡蝉与半翅目的关系更为密切。     

沫蝉总科Cercopoidea昆虫

沫蝉总科Cercopoidea隶属于昆虫纲Insecta半翅目Hemiptera头喙亚目Auchenorrhyncha，包括沫蝉科Cercopidae、尖胸沫蝉科Aphrophoridae、巢沫蝉科Machaerotidae及长盾沫蝉科Clastopteridae4个科，我国只有前3个科的种类。沫蝉总科目前全世界记录约3000种，广泛分布于世界各地，中国已知的种类300余种。沫蝉的若虫第七、八腹节具有发达的泡沫腺，能分泌胶质，与呼出的气体相混，形成泡沫状液体盖住身体以作保护，因而得名沫蝉。本期封面照片拍摄于安徽宣城。     

同翅类昆虫的雄性生殖系统及精子发生(昆虫纲:半翅目)

本文比较了同翅类昆虫雄性生殖系统的结构、减数分裂期间染色体的行为和精子尾部的超微结构。研究表明蜡蝉总科和异翅类的精巢具有被膜,而蝉总科、叶蝉总科、沫蝉总科、角蝉总科、木虱总科、蚜总科、粉虱总科和蚧总科的精巢均不具有被膜。也可以根据精巢小叶的形状将精巢分为三类,蝉总科、叶蝉总科、沫蝉总科、角蝉总科、蚜总科和粉虱总科的精巢小叶为球形,蜡蝉总科、木虱总科和蚧总科的精巢小叶为管状,而异翅类的精巢小叶为片层状。减数分裂可以被分为5类:①蝉型(Cicadoidtype);②蜡蝉型(Fulgoroidtype);③木虱型(Psyloidtype);④蚜型(Aphidoidtype);⑤粉虱型(Aleyrodoidtype)和⑥蚧型(Coccoidtype),至少具有四个类群的减数分裂前期I具有弥散期,它们是:木虱总科、蜡蝉总科、蚧总科和异翅类。除粉虱总科和蚧总科的精子尾部退化以外,其余种类的精子鞭毛均具有典型的9 9 2轴丝结构。     

关于停止使用"同翅目Homoptera" 目名的建议

长期以来,在我国昆虫学界,“同翅目Homoptera”和半翅目Hemiptera一直被作为2个并列的昆虫目被广泛使用。传统的“同翅目”被分为3亚目10总科,即鞘喙亚目Coleorrhyncha(包括膜翅蝽总科Peloridioidea)、胸喙亚目Stemorrhyncha(包括木虱总科Psylloidea、粉虱总科Aleyrodoidea、蚧总科Coccoidea和蚜总科Aphidoidea)和头喙亚目Auchenorrhyncha[包括蜡蝉子亚目Fulgoromorpha(包括蜡蝉总科Fulgoroidea)和蝉子亚目Cicadomorpha(包括蝉总科Cicadoidea、沫蝉总科Cercopoidea、叶蝉总科Cicadelloidea和角蝉总科Membracoidea)]。近年来,形态学及分子学特征数据的支序分析研究表明,木虱总科、粉虱总科、蚧总科、蚜总科、蜡蝉总科、蝉总科、沫蝉总科、角蝉总科都是单系群;鞘喙亚目、胸喙亚目、蝉子亚目及蜡蝉子亚目也都是单系群,其相互之间的系统发育关系为：胸喙亚目 (蝉子亚目 (蜡蝉子亚目 (鞘喙亚目 异翅亚目(蝽类)))),它们共同组成了单系的半翅目Hemiptera。系统发育分析表明,在半翅目中,鞘喙亚目与异翅亚目具有最近的亲缘关系,蜡蝉子亚目与鞘喙亚目 异翅亚目是姊妹群,蝉子亚目是蜡蝉子亚目 (鞘喙亚目 异翅亚目)的姐妹群,胸喙亚目是半翅目中最早和最原始的一个分枝。因此传统的“同翅目”并不是一个自然的单系类群,而是一个人为的并系类群。目前,在国际昆虫学界,“同翅目”作为一个人为的并系类群已得到公认和普遍接受,并已不再作为昆虫纲的一个有效目被使用。然而,“同翅目”作为昆虫纲的一个有效目在国内一直被广泛使用,为此,作者建议我国的昆虫学工作者今后应停止使用“同翅目”这一人为的并系目名而使用单系的半翅目目名,即将长期以来一直置于“同翅目”的木虱、粉虱、蚧虫、蚜虫、蝉、沫蝉、叶蝉、角蝉及蜡蝉类昆虫与蝽类昆虫一起作为半翅目的成员对待。     

蝉次目昆虫“足-消化道-马氏管”的形态/功能分化及其在蝉次目昆虫系统演化中的意义

对蝉次目3个总科(蝉总科,沫蝉总科,角蝉总科)代表类群(12种成虫)的足、消化道和马氏管的形态结构及其分化进行了比较形态学研究,提出蝉次目昆虫消化道及马氏管整体结构可被划分为两种类型的观点(蝉总科和沫蝉总科消化道及马氏管整体结构为同一种类型,角蝉总科消化道及马氏管整体结构为另一种类型),并分别绘制了两种类型的模式结构图;基于成虫的马氏管比较形态学研究,提出蝉次目昆虫成虫马氏管可被划分为6部分的观点,并分别绘制了角蝉科、叶蝉科、蝉科和沫蝉科成虫的马氏管区域分化模式结构图。研究结果表明,蝉次目昆虫的足、消化道、马氏管的形态、功能分化与其生物学、行为生态学特性等密切相关,其形态学特征既为蝉次目各类群的单系性提供了重要支持证据,也为进一步从行为学、生态学、生物学等方面探讨蝉次目的系统演化历史提供了新的信息;研究结果支持蝉总科和沫蝉总科为姊妹群的观点,即蝉次目3个总科的系统关系为角蝉总科+(蝉总科+沫蝉总科)。     

蜡蝉总科(昆虫纲:半翅目)系统分类研究进展

蜡蝉总科隶属于半翅目头喙亚目,大致可分为18～21个科。中国记载16科。蜡蝉总科的科级系统发育关系一直存在着广泛争议,至今尚无一个为大多数学者所接受的分类系统。因此,研究蜡蝉总科的系统发育具有重要的理论价值。本文从种及种下阶元的分类鉴定、种上阶元的系统发育、生物地理学等方面对蜡蝉总科的分类研究进行综述。     

半翅目沫蝉总科化石研究进展

回顾半翅目沫蝉总科化石研究简史,列出已发表的化石名录、地理分布及地质年代,共6科、117属、252种;概述了沫蝉总科的地质历史,总结了世界沫蝉总科化石的组成,提出该类群昆虫化石研究中存在的问题,并对今后的研究前景进行展望.     

透明疏广蜡蝉精子超微结构

【目的】昆虫精子的超微结构在昆虫种类鉴定和亲缘关系探讨方面具有重要意义。研究蜡蝉总科(Fulgoroidea)昆虫的精子超微结构,可以为目前仍存较大争议的该类群发育关系分析提供更多证据。【方法】采用超薄切片法并结合光学电子显微镜和透射电子显微镜,观察透明疏广蜡蝉Euricania clara Kato精子形态和超微结构。【结果】透明疏广蜡蝉的成熟精子聚集成束,单根精子无多态性,由头部、颈区和长鞭毛组成。头部包括双层顶体复合体和精子细胞核;颈区可见中心粒和中心粒侧体;长鞭毛主要由一对D形的线粒体衍生物、一对鱼钩状副体和典型的9+9+2微管型轴丝结构组成。【结论】透明疏广蜡蝉的鱼钩状副体与已报道的其他蜡蝉类群的该结构大体一致,但与头喙亚目其他类群的副体结构有显著差异;此外,透明疏广蜡蝉精子线粒体衍生物的数量、大小及横切形状与组成与其他昆虫类群有较大差异,而在头喙亚目内表现出一定的一致性,但也存在明显差异。本研究可以为蜡蝉总科昆虫系统发育分析提供科学资料。     

歧脊沫蝉属二新种(同翅目:尖胸沫蝉科)

本文记述采自我国西藏、陕西的尖胸沫蝉科Aphrophoridae歧脊沫蝉属Jembrana二新种。模式标本保存于西北农业大学昆虫博物馆。长茎歧脊沫蝉Jembranalongipensis,新种(图1—3) 头顶褐黄色,长约前胸背板的1/3;前缘钝圆,唇基端横向极狭长。触角檐短且厚,褐黄色;额脊、唇基端沟缺无。颜面短宽,棕褐色,二侧横脊浅簿。喙伸出后足转节之外,除东端一点黑褐色外褐黄色。     

Higher‐level phylogeny of the insect order Hemiptera: is Auchenorrhyncha really paraphyletic?

The higher‐level phylogeny of the order Hemiptera remains a contentious topic in insect systematics. The controversy is chiefly centred on the unresolved question of whether or not the hemipteran suborder Auchenorrhyncha (including the extant superfamilies Fulgoroidea, Membracoidea, Cicadoidea and Cercopoidea) is a monophyletic lineage. Presented here are the results of a multilocus molecular phylogenetic investigation of relationships among the major hemipteran lineages, designed specifically to address the question of Auchenorrhyncha monophyly in the context of broad taxonomic sampling across Hemiptera. Phylogenetic analyses (maximum parsimony, maximum likelihood and Bayesian inference) were based on DNA nucleotide sequence data from seven gene regions (18S rDNA, 28S rDNA, histone H3, histone 2A, wingless, cytochrome c oxidase I and NADH dehydrogenase subunit 4) generated from 86 in‐group exemplars representing all major lineages of Hemiptera (plus seven out‐group taxa). All combined analyses of these data recover the monophyly of Auchenorrhyncha, and also support the monophyly of each of the following lineages: Hemiptera, Sternorrhyncha, Heteropterodea, Heteroptera, Fulgoroidea, Cicadomorpha, Membracoidea, Cercopoidea and Cicadoidea. Also presented is a review of the major lines of morphological and molecular evidence for and against the monophyly of Auchenorrhyncha.     

Phylogenetic relationships among superfamilies of Cicadomorpha (Hemiptera: Auchenorrhyncha) inferred from the wing base structure

The infraorder Cicadomorpha is a monophyletic group of the order Hemiptera, suborder Auchenorrhyncha, and is composed of three superfamilies: Cercopoidea (spittle bugs), Cicadoidea (cicadas) and Membracoidea (leafhoppers and treehoppers). Phylogenetic relationships among the superfamilies have been highly controversial morphologically and molecularly, but recent molecular phylogenetic analyses provided support for Cercopoidea + Cicadoidea. In this study, we examined morphology of the wing base structure in Cicadomorpha and tested the previous phylogenetic hypotheses using the characters selected from the wing base. As a result, a sister‐group relationship between Cicadoidea and Cercopoidea was supported by three synapomorphies (presence of a projection posterior to the anterior notal wing process, presence of a novel notal process anterior to the posterior notal wing process, presence of a novel sclerite between the distal median plate and the base of anal vein). The present study provides the first unambiguous and prominent morphological support for Cicadoidea + Cercopoidea.     

Molecular phylogeny of Cicadomorpha (Insecta: Hemiptera: Cicadoidea, Cercopoidea and Membracoidea): adding evidence to the controversy

Abstract. The hemipteran infraorder Cicadomorpha comprises the superfamilies Cicadoidea (cicadas), Cercopoidea (spittlebugs or froghoppers) and Membracoidea (leafhoppers and treehoppers). Earlier attempts to determine relationships among these three monophyletic lineages using either morphological or molecular data suffered from insufficient sampling (taxonomic and data) and problematic tree rooting, leading to discordant results. Presented here are phylogenetic reconstructions within Cicadomorpha based on DNA nucleotide sequence data from multiple genetic markers (18S rDNA, 28S rDNA, and histone 3) sequenced from representative taxa of Cicadidae, Tettigarctidae, Cercopidae, Aphrophoridae, Clastopteridae, Machaerotidae, Epipygidae, Cicadellidae, Membracidae, Myerslopiidae and Aetalionidae. To test the robustness of the phylogenetic signal, these sequence data were analysed separately and in combination under various alignment parameters using both manual alignment (of both attenuated and full sequences) and alignment via clustal x . The results demonstrate clearly that, despite the alignment method used, basing a phylogeny on a single gene region is often misleading. Analyses of the combination of datasets support the major relationships within Cicadomorpha as (Membracoidea (Cicadoidea, Cercopoidea)). Internal relationships recovered within each superfamily shows evidence for: (1) the placement of Myerslopiidae as the sister group of the remaining Membracoidea; (2) the paraphyly of Cicadellidae; (3) the sister-group relationship between Machaerotidae and Clastopteridae; (4) the monophyly of Cercopidae; (5) the diversification of Epipygidae from within the possibly paraphyletic Aphrophoridae.     

Two ancient bacterial endosymbionts have coevolved with the planthoppers (Insecta: Hemiptera: Fulgoroidea)

ABSTRACT: BACKGROUND: Members of the hemipteran suborder Auchenorrhyncha (commonly known as planthoppers, tree- and leafhoppers, spittlebugs, and cicadas) are unusual among insects known to harbor endosymbiotic bacteria in that they are associated with diverse assemblages of bacterial endosymbionts. Early light microscopic surveys of species representing the two major lineages of Auchenorrhyncha (the planthopper superfamily Fulgoroidea; and Cicadomorpha, comprising Membracoidea [tree- and leafhoppers], Cercopoidea [spittlebugs], and Cicadoidea [cicadas]), found that most examined species harbored at least two morphologically distinct bacterial endosymbionts, and some harbored as many as six. Recent investigations using molecular techniques have identified multiple obligate bacterial endosymbionts in Cicadomorpha; however, much less is known about endosymbionts of Fulgoroidea. In this study, we present the initial findings of an ongoing PCR-based survey (sequencing 16S rDNA) of planthopper-associated bacteria to document endosymbionts with a long-term history of codiversification with their fulgoroid hosts. RESULTS: Results of PCR surveys and phylogenetic analyses of 16S rDNA recovered a monophyletic clade of Betaproteobacteria associated with planthoppers; this clade included Vidania fulgoroideae, a recently described bacterium identified in exemplars of the planthopper family Cixiidae. We surveyed 77 planthopper species representing 18 fulgoroid families, and detected Vidania in 40 species (representing 13 families). Further, we detected the Sulcia endosymbiont (identified as an obligate endosymbiont of Auchenorrhyncha in previous studies) in 30 of the 40 species harboring Vidania. Concordance of the Vidania phylogeny with the phylogeny of the planthopper hosts (reconstructed based on sequence data from five genes generated from the same insect specimens from which the bacterial sequences were obtained) was supported by statistical tests of codiversification. Codiversification tests also supported concordance of the Sulcia phylogeny with the phylogeny of the planthopper hosts, as well as concordance of planthopper-associated Vidania and Sulcia phylogenies. CONCLUSIONS: Our results indicate that the Betaproteobacterium Vidania is an ancient endosymbiont that infected the common ancestor of Fulgoroidea at least 130 million years ago. Comparison of our findings with the early light-microscopic surveys conducted by Muller suggests that Vidania is Muller's x-symbiont, which he hypothesized to have codiversified with most lineages of planthoppers and with the Sulcia endosymbiont.     

Molecular phylogeny of the homoptera: a paraphyletic taxon

Homoptera and Heteroptera comprise a large insect assemblage, the Hemiptera. Many of the plant sap-sucking Homoptera possess unusual and complex life histories and depend on maternally inherited, intracellular bacteria to supplement their nutritionally deficient diets. Presumably in connection with their diet and lifestyles, the morphology of many Homoptera has become greatly reduced, leading to major controversies regarding the phylogenetic affiliations of homopteran superfamilies. The most fundamental question concerns whether the Homoptera as a whole are monophyletic. Recent studies based on morphology have argued that the Homoptera Sternorrhyncha (Aphidoidea, Coccoidea, Psylloidea, Aleyrodoidea) is a sister group to a group comprising the Homoptera Auchenorrhyncha (Fulgoroidea, Cicadoidea, Cercopoidea, Cicadelloidea) and the Heteroptera, making the Homoptera paraphyletic. We sequenced the 5 580-680 base pairs of small-subunit (18S) ribosomal DNA from a selection of Homoptera, Hemiptera, and their putative outgroups, the Thysanoptera and Psocoptera, to apply molecular characters to the problem of Homoptera phylogeny. Parsimony, distance, maximum-likelihood, and bootstrap methods were used to construct trees from sequence data and assess support for the topologies produced. Molecular data corroborate current views of relationships within the Sternorrhyncha and Auchenorrhyncha based on morphology and strongly support the hypothesis of homopteran paraphyly as stated above. In addition, it was found that Homoptera Sternorrhyncha have extra, GC-rich sequence concentrated in a variable region of the 18S rDNA, which indicates that some unique evolutionary processes are occurring in this lineage.Correspondence to: C.D. von Dohlen