蛾蛉类昆虫化石研究进展

蛾蛉类昆虫是脉翅目中化石记录最完整的的类群之一,现生类群蛾蛉、美蛉和山蛉统称为蛾蛉科,但是现生类群与化石类群分类标准的不一致性,为蛾蛉类昆虫化石研究带来较大的困难。本文统计了世界已发现的蛾蛉类昆虫化石属种名录,介绍了蛾蛉类昆虫化石研究历史、地质年代及地理分布、系统发育研究进程,并提出了现今有待解决的问题以及对未来研究的展望。     

鞘翅目系统演化关系研究进展

现生鞘翅目类群是昆虫纲中种类最多的一个类群,由4个亚目组成。形态学研究表明,鞘翅目是一个单系,但在加入了分子数据之后,其结果证明鞘翅目为并系类群。最早的甲虫起源于二叠纪,与现生的原鞘亚目具有许多相似特征。大多数学者认为与鞘翅目亲缘关系最近的类群是脉翅类昆虫,但是也有的学者认为鞘翅目与捻翅目互为姐妹群。本文对鞘翅目高级阶元4个亚目的系统关系和有关鞘翅目的起源和演化等研究进行了总结,并对相关问题做了进一步探讨和展望。     

中国长翅目昆虫化石研究现状

长翅目昆虫在地史纪录上可以追溯到早二叠纪,是全变态昆虫中最原始的种类之一.截至目前,29篇关于中国长翅目昆虫化石分类的论著发表,共描述我国长翅目昆虫化石11科28属51种,这些化石分布于从三叠纪到白垩纪的不同地层中.本文通过图表提供了我国已发现的长翅目化石名录并介绍了其分布和年代,回顾了我国长翅目昆虫化石的研究进展,指出了一些分类存在的问题,简要慨述了长翅目中一些科的起源与演化以及长翅目与其它全变态昆虫的关系.我国长翅目化石种类丰富,但在基础分类、系统演化方面还缺乏系统的研究,因而许多工作甚至最基础的分类工作亟待开展.     

昆虫的变态与进化

昆虫的起源与进化问题,在昆虫学界讨论颇多,许多问题已基本取得了一致的意见,如昆虫纲起源于多足纲;昆虫的增节变态是最原始的变态类型;表变态是由增节变态演化而来;有翅亚纲中的无翅类是属于后生无翅等。但仍有许多问题处于争论之中,其中之一是有翅亚纲的几种变态类型：原变态（Ｐｒｏｍｅｔａｂｏｌａ）,不完全变态（Ｈｅｍｉｍｅｔａｂｏｌａ）,完全变态（Ｈｏｌｏｍｅｔａｂｏｌａ）的起源的问题。作者认为：有翅亚纲中的三种变态类型中,不完全变态是最原始的变态形式,原变态类和完全变态类昆虫是起源于祖先的不完全变态类。…     

昆虫的翅

介绍了昆虫翅的起源、基本结构、类型、脉相、连锁方式及其功用。     

中生代脉翅目昆虫的叶状拟态

拟态是一种生物模拟另一种生物或环境中的其它物体从而使自身获得好处的现象,是昆虫在进化过程中特化出的一种极为重要和有效的防御策略。脉翅目昆虫是一类完全变态性昆虫,在其超过2.7亿年的演化历史中,出现了形态多样的翅斑。美翼蛉属Bellinympha Wang,Ren,Liu,Shih&Engel,2010是脉翅目溪蛉科丽翼蛉亚科的一个绝灭属,发现于中国内蒙古道虎沟地区中侏罗世九龙山组地层。该属的发现代表了迄今报道的最古老的叶状拟态现象,也是唯一报道的昆虫模拟裸子植物或蕨类植物叶片。本文简要概述了脉翅目昆虫研究概况,总结了中生代脉翅目翅斑的多样性,对美翼蛉属进行了描述,并对其叶状拟态及其生物学意义进行了探讨。     

"古翅类"系统发育关系研究进展(昆虫纲,有翅类)

古翅类的系统发育问题是六足总纲中有争议的热点问题之一。对现存古翅类(孵蝣目 蜻蜒目)与新翅类之间的系统发育关系有3种主要观点：古翅类(＝蜉蝣目 蜻蜒目) 新翅类,蜉蝣目 (蜻蜒目 新翅类),蜻蜒目 (蜉蝣目 新翅类)。第1种观点得到化石、形态和部分基因证据支持,第2种观点得到较多形态特征支持,支持第3种观点的证据较少。这一问题的解决有赖于更多昆虫种类、化石以及分子证据的发现和研究。     

Ancient pinnate leaf mimesis in Mesozoic lacewings

拟态是一种生物模拟另一种生物或环境中的其它物体从而使自身获得好处的现象,是昆虫在进化过程中特化出的一种极为重要和有效的防御策略.脉翅目昆虫是一类完全变态性昆虫,在其超过2.7亿年的演化历史中,出现了形态多样的翅斑.美翼蛉属Bellinympha Wang,Ren,Liu,Shih&Engel,2010是脉翅目溪蛉科丽翼蛉亚科的一个绝灭属,发现于中国内蒙古道虎沟地区中侏罗世九龙山组地层.该属的发现代表了迄今报道的最古老的叶状拟态现象,也是唯一报道的昆虫模拟裸子植物或蕨类植物叶片.本文简要概述了脉翅目昆虫研究概况,总结了中生代脉翅目翅斑的多样性,对美翼蛉属进行了描述,并对其叶状拟态及其生物学意义进行了探讨.     

中国襀翅目昆虫系统分类研究进展

襀翅目昆虫广泛分布在多种类型的水域中,对水中的化学物质较为敏感,是一类重要的水质监测指标生物。本文回顾了我国襀翅目分类的研究历史,其中Klapálek、Navás、Banks及胡经甫对我国早期的襀翅目分类研究做出了重要贡献。同时总结了我国襀翅目物种多样性的研究现状,即我国已记录襀翅目10科、66属、657种(包括台湾分布的31种、香港分布的4种),约占世界已记录种类的17.8%;我国学者定名的种类占全国记录种类的74.7%,其中1949年建国后的70年间定名的种类占67.4%。此外,还对我国襀翅目昆虫的地理分布及系统发育研究进行了总结分析,其中我国学者共测序襀翅目昆虫全线粒体基因组60种,占已测序种类的90.9%,为襀翅目的分子系统发育研究积累了重要分子数据;同时利用这些数据对襀翅目的分子系统发育进行研究,取得了较好的研究结果。最后,文章分析了我国襀翅目系统分类研究存在的主要问题,并对未来的研究进行了展望。     

广东南岭国家级自然保护区大东山昆虫名录(Ⅳ)

本文报道广东南岭国家级自然保护区大东山管理站所辖林区广翅目1科2属2种、脉翅目3科8属10种、长翅目1科1属1种、鳞翅目(蛾类)27科213属299种的昆虫名录.     

A mitochondrial genome phylogeny of the Neuropterida (lace-wings, alderflies and snakeflies) and their relationship to the other holometabolous insect orders

We present a mitochondrial (mt) genome phylogeny inferring relationships within Neuropterida (lacewings, alderflies and camel flies) and between Neuropterida and other holometabolous insect orders. Whole mt genomes were sequenced for Sialis hamata (Megaloptera: Sialidae), Ditaxis latistyla (Neuroptera: Mantispidae), Mongoloraphidia harmandi (Raphidioptera: Raphidiidae), Macrogyrus oblongus (Coleoptera: Gyrinidae), Rhopaea magnicornis (Coleoptera: Scarabaeidae), and Mordella atrata (Coleoptera: Mordellidae) and compared against representatives of other holometabolous orders in phylogenetic analyses. Additionally, we test the sensitivity of phylogenetic inferences to four analytical approaches: inclusion vs. exclusion of RNA genes, manual vs. algorithmic alignments, arbitrary vs. algorithmic approaches to excluding variable gene regions and how each approach interacts with phylogenetic inference methods (parsimony vs. Bayesian inference). Of these factors, phylogenetic inference method had the most influence on interordinal relationships. Bayesian analyses inferred topologies largely congruent with morphologically‐based hypotheses of neuropterid relationships, a monophyletic Neuropterida whose sister group is Coleoptera. In contrast, parsimony analyses failed to support a monophyletic Neuropterida as Raphidioptera was the sister group of the entire Holometabola excluding Hymenoptera, and Neuroptera + Megaloptera is the sister group of Diptera, a relationship which has not previously been proposed based on either molecular or morphological data sets. These differences between analytical methods are due to the high among site rate heterogeneity found in insect mt genomes which is properly modelled by Bayesian methods but results in artifactual relationships under parsimony. Properly analysed, the mt genomic data set presented here is among the first molecular data to support traditional, morphology‐based interpretations of relationships between the three neuropterid orders and their grouping with Coleoptera.     

Phylogenetic significance of the wing-base of the Holometabola (Insecta)

The present knowledge of the wing-base morphology of the holometabolous insects is summarized, and the value of these structures for phylogenetic analysis is demonstrated. An autapomorphy of the Holometabola is a locking mechanism composed of a knob on the basalare and a corresponding cavity on the ventral wing-base. Two synapomorphic hindwing-base characters support a sister-group relationship of Coleoptera and Neuropterida. Only few data are available on the wing-base of the Hymenoptera. An autapomorphy of the taxon is a modification of the wing locking mechanism with reduced size of the basalare and its knob. It is demonstrated that wing-base characters are helpful for the analysis of the relationships between strepsipteran families. However, characters of the wing-base support neither a relationship of Strepsiptera and Coleoptera nor of Strepsiptera and Antliophora.     

The First Mitochondrial Genome for the Fishfly Subfamily Chauliodinae and Implications for the Higher Phylogeny of Megaloptera

Megaloptera are a basal holometabolous insect order with larvae exclusively predacious and aquatic. The evolutionary history of Megaloptera attracts great interest because of its antiquity and important systematic status in Holometabola. However, due to the difficulties identifying morphological apomorphies for the group, controversial hypotheses on the monophyly and higher phylogeny of Megaloptera have been proposed. Herein, we describe the complete mitochondrial (mt) genome of a fishfly species, Neochauliodes punctatolosus Liu & Yang, 2006, representing the first mt genome of the subfamily Chauliodinae. A phylogenomic analysis was carried out based on the mt genomic sequences of 13 mt protein-coding genes (PCGs) and two rRNA genes of nine Neuropterida species, comprising all three orders of Neuropterida and all families and subfamilies of Megaloptera. Both maximum likelihood and Bayesian inference analyses highly support the monophyly of Megaloptera, which was recovered as the sister of Neuroptera. Within Megaloptera, the sister relationship between Corydalinae and Chauliodinae was corroborated. The divergence time estimation suggests that stem lineage of Neuropterida and Coleoptera separated in the Early Permian. The interordinal divergence within Neuropterida might have occurred in the Late Permian.     

9-Genes Reinforce the Phylogeny of Holometabola and Yield Alternate Views on the Phylogenetic Placement of Strepsiptera

Background

The extraordinary morphology, reproductive and developmental biology, and behavioral ecology of twisted wing parasites (order Strepsiptera) have puzzled biologists for centuries. Even today, the phylogenetic position of these enigmatic “insects from outer space” [1] remains uncertain and contentious. Recent authors have argued for the placement of Strepsiptera within or as a close relative of beetles (order Coleoptera), as sister group of flies (order Diptera), or even outside of Holometabola.

Methodology/Principal Findings

Here, we combine data from several recent studies with new data (for a total of 9 nuclear genes and ∼13 kb of aligned data for 34 taxa), to help clarify the phylogenetic placement of Strepsiptera. Our results unequivocally support the monophyly of Neuropteroidea ( = Neuropterida + Coleoptera) + Strepsiptera, but recover Strepsiptera either derived from within polyphagan beetles (order Coleoptera), or in a position sister to Neuropterida. All other supra-ordinal- and ordinal-level relationships recovered with strong nodal support were consistent with most other recent studies.

Conclusions/Significance

These results, coupled with the recent proposed placement of Strepsiptera sister to Coleoptera, suggest that while the phylogenetic neighborhood of Strepsiptera has been identified, unequivocal placement to a specific branch within Neuropteroidea will require additional study.     

Mitochondrial phylogenomics illuminates the evolutionary history of Neuropterida

Neuroptera (lacewings) and allied orders Megaloptera (dobsonflies, alderflies) and Raphidioptera (snakeflies) are predatory insects and together make up the clade Neuropterida. The higher‐level relationships within Neuropterida have historically been widely disputed with multiple competing hypotheses. Moreover, the evolution of important biological innovations among various Neuropterida families, such as the origin, timing and direction of transitions between aquatic and terrestrial habitats of larvae, remains poorly understood. To investigate the origin and diversification of lacewings and their allies, we undertook phylogenetic analyses of mitochondrial genomes of all families of Neuropterida using Bayesian inference, maximum likelihood and maximum parsimony methods. We present a robust, fully resolved phylogeny and divergence time estimation for Neuropterida with strong statistical support for almost all nodes. Mitochondrial sequence data are typified by significant compositional heterogeneity across lineages, and parsimony and models assuming homogeneous rates did not recover Neuroptera as monophyletic. Only a model accounting for compositional heterogeneity (i.e. CAT‐GTR) recovered all orders of Neuropterida as monophyletic. Significant findings of the mitogenomic phylogeny include recovering Raphidioptera as sister to Megaloptera plus Neuroptera. The sister family of all other lacewings are the dusty‐wings (Coniopterygidae), rather than Nevrorthidae. Nevrorthidae are instead returned to their traditional position as the sister group of the spongilla‐flies (Sisyridae) and closely related to Osmylidae. Our divergence time analysis indicates that the Mesozoic was indeed a ‘golden age’ for lacewings, with most families of Neuropterida diverging during the Triassic and Jurassic and all extant families present by the Early Cretaceous. Based on ancestral character state reconstructions of larval habitat we evaluate competing hypotheses regarding the life style of early neuropteridan larvae as either aquatic or terrestrial.     

Morphological and molecular evidence converge upon a robust phylogeny of the megadiverse Holometabola

We present the largest morphological character set ever compiled for Holometabola. This was made possible through an optimized acquisition of data. Based on our analyses and recently published hypotheses based on molecular data, we discuss higher‐level phylogeny and evolutionary changes. We comment on the information content of different character systems and discuss the role of morphology in the age of phylogenomics. Microcomputer tomography in combination with other techniques proved highly efficient for acquiring and documenting morphological data. Detailed anatomical information (356 characters) is now available for 30 representatives of all holometabolan orders. A combination of traditional and novel techniques complemented each other and rapidly provided reliable data. In addition, our approach facilitates documenting the anatomy of model organisms. Our results show little congruence with studies based on rRNA, but confirm most clades retrieved in a recent study based on nuclear genes: Holometabola excluding Hymenoptera, Coleopterida (= Strepsiptera + Coleoptera), Neuropterida excl. Neuroptera, and Mecoptera. Mecopterida (= Antliophora + Amphiesmenoptera) was retrieved only in Bayesian analyses. All orders except Megaloptera are monophyletic. Problems in the analyses are caused by taxa with numerous autapomorphies and/or inapplicable character states due to the loss of major structures (such as wings). Different factors have contributed to the evolutionary success of various holometabolan lineages. It is likely that good flying performance, the ability to occupy different habitats as larvae and adults, parasitism, liquid feeding, and co‐evolution with flowering plants have played important roles. We argue that even in the “age of phylogenomics”, comparative morphology will still play a vital role. In addition, morphology is essential for reconstructing major evolutionary transformations at the phenotypic level, for testing evolutionary scenarios, and for placing fossil taxa.
© The Willi Hennig Society 2010.     

The morphology and evolution of the female postabdomen of Holometabola (Insecta)

In the present article homology issues, character evolution and phylogenetic implications related to the female postabdomen of the holometabolan insects are discussed, based on an earlier analysis of a comprehensive morphological data set. Hymenoptera, the sistergroup of the remaining Holometabola, are the only group where the females have retained a fully developed primary ovipositor of the lepismatid type. There are no characters of the female abdomen supporting a clade Coleopterida + Neuropterida. The invagination of the terminal segments is an autapomorphy of Coleoptera. The ovipositor is substantially modified in Raphidioptera and distinctly reduced in Megaloptera and Neuroptera. The entire female abdomen is extremely simplified in Strepsiptera. The postabdomen is tapering posteriorly in Mecopterida and retractile in a telescopic manner (oviscapt). The paired ventral sclerites of segments VIII and IX are preserved, but valvifers and valvulae are not distinguishable. In Amphiesmenoptera sclerotizations derived from the ventral appendages VIII are fused ventromedially, forming a solid plate, and the appendages IX are reduced. The terminal segments are fused and form a terminal unit which bears the genital opening subapically. The presence of two pairs of apophyses and the related protraction of the terminal unit by muscle force are additional autapomorphies, as is the fusion of the rectum with the posterior part of the genital chamber (cloaca). Antliophora are supported by the presence of a transverse muscle between the ventral sclerites of segment VIII. Secondary egg laying tubes have evolved independently within Boreidae (absent in Caurinus) and in Tipulomorpha. The loss of two muscle associated with the genital chamber are likely autapomorphies of Diptera. The secondary loss of the telescopic retractability of the postabdomen is one of many autapomorphies of Siphonaptera.     

A Unique Box in 28S rRNA Is Shared by the Enigmatic Insect Order Zoraptera and Dictyoptera

The position of the Zoraptera remains one of the most challenging and uncertain concerns in ordinal-level phylogenies of the insects. Zoraptera have been viewed as having a close relationship with five different groups of Polyneoptera, or as being allied to the Paraneoptera or even Holometabola. Although rDNAs have been widely used in phylogenetic studies of insects, the application of the complete 28S rDNA are still scattered in only a few orders. In this study, a secondary structure model of the complete 28S rRNAs of insects was reconstructed based on all orders of Insecta. It was found that one length-variable region, D3-4, is particularly distinctive. The length and/or sequence of D3-4 is conservative within each order of Polyneoptera, but it can be divided into two types between the different orders of the supercohort, of which the enigmatic order Zoraptera and Dictyoptera share one type, while the remaining orders of Polyneoptera share the other. Additionally, independent evidence from phylogenetic results support the clade (Zoraptera+Dictyoptera) as well. Thus, the similarity of D3-4 between Zoraptera and Dictyoptera can serve as potentially valuable autapomorphy or synapomorphy in phylogeny reconstruction. The clades of (Plecoptera+Dermaptera) and ((Grylloblattodea+Mantophasmatodea)+(Embiodea+Phasmatodea)) were also recovered in the phylogenetic study. In addition, considering the other studies based on rDNAs, this study reached the highest congruence with previous phylogenetic studies of Holometabola based on nuclear protein coding genes or morphology characters. Future comparative studies of secondary structures across deep divergences and additional taxa are likely to reveal conserved patterns, structures and motifs that can provide support for major phylogenetic lineages.     

Phylogenetic relationships among insect orders based on three nuclear protein-coding gene sequences

Many attempts to resolve the phylogenetic relationships of higher groups of insects have been made based on both morphological and molecular evidence; nonetheless, most of the interordinal relationships of insects remain unclear or are controversial. As a new approach, in this study we sequenced three nuclear genes encoding the catalytic subunit of DNA polymerase delta and the two largest subunits of RNA polymerase II from all insect orders. The predicted amino acid sequences (In total, approx. 3500 amino acid sites) of these proteins were subjected to phylogenetic analyses based on the maximum likelihood and Bayesian analysis methods with various models. The resulting trees strongly support the monophyly of Palaeoptera, Neoptera, Polyneoptera, and Holometabola, while within Polyneoptera, the groupings of Isoptera/"Blattaria"/Mantodea (Superorder Dictyoptera), Dictyoptera/Zoraptera, Dermaptera/Plecoptera, Mantophasmatodea/Grylloblattodea, and Embioptera/Phasmatodea are supported. Although Paraneoptera is not supported as a monophyletic group, the grouping of Phthiraptera/Psocoptera is robustly supported. The interordinal relationships within Holometabola are well resolved and strongly supported that the order Hymenoptera is the sister lineage to all other holometabolous insects. The other orders of Holometabola are separated into two large groups, and the interordinal relationships of each group are (((Siphonaptera, Mecoptera), Diptera), (Trichoptera, Lepidoptera)) and ((Coleoptera, Strepsiptera), (Neuroptera, Raphidioptera, Megaloptera)). The sister relationship between Strepsiptera and Diptera are significantly rejected by all the statistical tests (AU, KH and wSH), while the affinity between Hymenoptera and Mecopterida are significantly rejected only by AU and KH tests. Our results show that the use of amino acid sequences of these three nuclear genes is an effective approach for resolving the relationships of higher groups of insects.     

Mitochondrial genomes in the Hymenoptera and their utility as phylogenetic markers

Abstract.  In this study, we assessed the ability of mitochondrial genome sequences to recover a test phylogeny of five hymenopteran taxa from which phylogenetic relationships are well accepted. Our analyses indicated that the test phylogeny was well recovered in all nucleotide Bayesian analyses when all the available holometabolan (i.e. outgroup) taxa were included, but only in Bayesian analyses excluding third codon positions when only the hymenopteran representatives and a single outgroup were included. This result suggests that taxon sampling of the outgroup might be as important as taxon sampling of the ingroup when recovering hymenopteran phylogenetic relationships using whole mitochondrial genomes. Parsimony analyses were more sensitive to both taxon sampling and the analytical model than Bayesian analyses, and analyses using the protein dataset did not recover the test phylogeny. In general, mitochondrial genomes did not resolve the position of the Hymenoptera within the Holometabola with confidence, suggesting that an increased taxon sampling, both within the Holometabola and among outgroups, is necessary.