基于线粒体基因组序列的鞘翅目肉食亚目水生类群系统发育分析

【目的】明确肉食亚目(Adephaga)水生类群线粒体基因组的基本特征,并基于线粒体基因组序列分析肉食亚目水生类群的系统发育关系。【方法】基于Illumina HiSeq X Ten测序技术测定了圆鞘隐盾豉甲Dineutus mellyi和齿缘龙虱Eretes sticticus的线粒体全基因组序列,对其进行了基因注释,并对其tRNA基因二级结构进行了预测分析。加上已公布的鞘翅目(Coleoptera)肉食亚目水生类群17个种的线粒体基因组序列,对该类群共19个种线粒体的蛋白质编码基因(protein-coding genes, PCGs)开展了比较基因组学分析,包括AT含量、密码子偏好性、选择压力等。基于13个PCGs的氨基酸序列和核苷酸序列,利用最大似然法(ML)和贝叶斯法(BI)分别构建鞘翅目肉食亚目水生类群的系统发育关系,并通过FcLM分析进一步评估伪龙虱科(Noteridae)和瀑甲科(Meruidae)的系统发育位置。【结果】圆鞘隐盾豉甲和齿缘龙虱的线粒体基因组全长分别为16 123 bp(GenBank登录号: MN781126)和16 196 bp(GenBank登录号: MN781132),都包含13个PCGs、22个tRNA基因、2个rRNA基因和1个D-loop区(控制区)。19个肉食亚目水生类群线粒体基因组PCGs的碱基组成都呈现A+T偏好性,在密码子使用上也都偏向于使用富含A+T的密码子;在进化过程中13个PCGs的进化模式相同,都受到纯化选择。基于线粒体基因组13个PCGs的氨基酸序列的肉食亚目水生类群的系统发育关系为(豉甲科Gyrinidae+(沼梭甲科Haliplidae+((壁甲科Aspidytidae+(两栖甲科Amphizoidae+龙虱科Dytiscidae))+(水甲科Hygrobiidae+(瀑甲科Meruidae+伪龙虱科Noteridae)))))。【结论】研究结果表明,豉甲科是肉食亚目水生类群的基部类群,接下来是沼梭甲科和龙虱总科;伪龙虱科和瀑甲科互为姐妹群,并一起作为龙虱总科内部的一个分支;两栖甲科与龙虱科具有更近的亲缘关系。     

豉甲科及水生肉食亚目的分子系统发育学分析(昆虫纲：鞘翅目)

利用PAUP和MrBayes软件,对线粒体COⅠ基因序列3个密码子位置的数据模块分别进行了豉甲科(Gyrinidae)和水生肉食亚目(Hydradephaga)在亚科或科水平上的系统发育学分析,结果表明第二密码子数据模块获得了理想的分析结果。由PAUP生成的豉甲科最优树来自第二密码子数据模块的分析,而由MrBayes生成的最优树来自全部密码子数据模块的分析。此外,用对应的氨基酸序列生成的ME和MP树与第二密码子数据模块分析的结果也一致。亚科Orectochilinae和Gyrininae以高的支持率形成了单系。然而,来自亚科Enhydrinae的种Porrorhynchus landaisi landaisi呈现了异常的位置。SH-test检验也支持该异常位置,表明这个种可能代表了一个科。在来自第二密码子数据模块的水生肉食亚目最优ML树中,整个Hydradephaga树呈现单系,豉甲科位于树的基部,表明了该科在水生肉食亚目中是一个早期的分支。在树中还产生了一个单系的Dytiscoidea总科,由Dytiscidae、Hygrobiidae、Noteridae和Amphizoidae 4个科组成,单系的Haliplidae与之成为姐妹群。此外线粒体分子钟的结果表明豉甲科的5对相近种间的分化是一个短时期内发生的(0.01～1.81百万年前),这点可能与它们的特殊地理分布有关。     

长白山北坡静水水体中水甲虫分布与环境关系的典范对应分析

应用典范对应分析（CCA）对长白山北坡静水水体12个样点中28种水甲虫与环境关系的研究表明,长白山5种环境因子中水底腐殖化程度和海拔对水甲虫的分布起主要作用,与排序轴的相关系灵敏分别高达0.8371和0.7206,而水温和植被密度也有较大的影响,在环境因子的影响下不同生境中水甲虫分布的种群不同,深刻斜凹龙虱,端钩切眼龙虱,布朗沟牙甲,沼梭科水甲虫等主要分布在深水区,与水温没有关系。而异毛龙虱和舟型牙甲等与海拔和水泡子的腐殖化程度呈正相关。     

池沼、稻田中水生昆虫(成虫)的识别

在昆虫采集中,用水网在池沼、稻田中捞捕水生昆虫,常见的主要是鞘翅目和半翅目少数科(属),且一般可用肉眼观察鉴定其性状和生态特征,使用下列简易检索表不难识别,种名可依其科(属)名称呼也是允许的。 1(8)前翅角质,其内缘在体背中央合成中缝,后翅膜质。(鞘翅目) 2(3)中足和后足特别扁短,整形(图A);常在水面作快速旋转游泳………………………………………豉甲科Gyrinidae (常见为鼓甲属Dineutus) 3(2)中、后足不如上述;无回旋游泳习性。 4(5)触角短,不超过9节,端部3-4节膨大且具绒毛。常潜水中,不时露头呼吸…………水龟虫科(牙(?)科)ydrophilidae(常见为水龟虫属Hydrophilus) 5(4)触角细长,10-11节,端部不膨大且无毛,常栖水底。 6(7)后足粗扁,有长毛如刷(图B);后足基节正常………………龙虱科Dytiscidae (常见为龙虱属Cyhister) 7(6)后足细长,无刷状长缘毛;后足基节成平板状(图C),盖住大部腹部………………………………沼梭科Haliplidae     

《Review of Diving Beetles of the world》书评

<正>拥有一本某一类群系统的世界性分类鉴定手册,对昆虫分类工作者来说十分重要。在水生甲虫中,龙虱科Dytiscidae是最大的类群,然而一直缺乏一本世界性的鉴定手册。MillerBergsten编著的《世界龙虱》的出版为了解和鉴定龙虱提供了方便。该书为320页的精美硬装本,也有可购买的电子版。该书可分为以下几部分:1.导言:介绍了龙虱的生活史和行为学、     

酷甲虫“屎”里逃生

正甲虫遇到蛙,会是什么结果?当然是被吃掉喽!这个答案对,也不对。一般情况下,甲虫遇到蛙,很快被吞下,不久后它会被消化,生命就这样结束啦!然而有一种甲虫与众不同,能凭自己的本事与命运斗争,最终活下来。究竟是哪种甲虫这么厉害?"重生"的甲虫它就是掌握独特逃生秘诀的梭形扁牙甲。梭形扁牙甲是一种水生甲虫,往往和蛙类生活在同一环境中。蛙作为食虫高手,见到这种甲虫当然不会客气。"啊呜!"还没等梭形扁牙甲反应过来,眼疾嘴快的黑斑侧褶蛙就把它吞进肚子里了。     

鞘翅目高级阶元分子系统学： 研究现状及存在的问题

鞘翅目是世界上物种最丰富的类群, 分为原鞘亚目（Archostemata Kolbe, 1908）、 藻食亚目（Myxophaga Crowson, 1955）、 肉食亚目（Adephaga Schellenberg, 1806）和多食亚目（Polyphaga Emery, 1886）。随着分子生物学的发展,分子系统学的技术被广泛应用于鞘翅目系统学研究中。本文综述了鞘翅目高级阶元的分子系统学的研究进展及存在问题。基于分子生物学手段, 分子分类学家提出了关于鞘翅目高级阶元分子系统学很多假说, 分子分析结果支持鞘翅目的4个亚目各为单系, 而亚目间的系统关系还不统一。基于分子手段对于亚目内的系统发育关系的研究也有了一定的进展, 比如： 分子系统学结果支持肉食亚目的水生类群和陆生类群分别为单系, 水生类群为一次起源。目前, 鞘翅目高级阶元分子系统学的研究还不够成熟和完善, 主要表现为： 材料选择有限且不均衡、 基因数目和适合度不理想, 以及一些关键节点研究的欠缺。     

介形纲(甲壳亚门)系统分类中核基因18S rDNA的应用研究

本研究采用贝叶斯法(Bayesian-inference,BI)、最大似然法(Maximum-likelihood,ML)和最大简约法(Maximumparsimony,MP),基于18SrDNA序列对介形纲4目12超科间以及其它相关泛甲壳动物(Pancrusacea)间的系统分类关系进行了分析,其目的是探讨单独的核基因18SrDNA序列研究介形纲各阶元分类时的稳定性,及分析介形纲两大主要类群内的系统分类关系。结果发现,应用单独的核基因18SrDNA序列对介形类进行研究时,所获的三种系统树的拓扑结构在介形纲的目、亚目和超科阶元均表现稳定,表明单独的核基因18SrDNA序列在该3个阶元的分类研究中可发挥重要作用。基于此,进一步对介形类内两主要类群的系统分类研究,结果支持Martin和Davis(2001)对介形纲中壮肢亚纲(又称丽足亚纲,Myodocopa)在目、亚目和超科阶元上的分类;但尾肢亚纲(又称速足亚纲,Podocopa)中各主要类群间的系统发育关系却与Martin和Davis(2001)的分类略有不同,其中Punciidae游离于尾肢亚纲主要类群尾肢目(又称速足目,Podocopida)之外,而该结果却与Liebau(2005)对Punciidae归类一致;此外尾肢目中5个主要类群的分类关系呈现[浪花介亚目Cytherocopina+[金星介亚目Cypridocopina+简肢亚目Platycopina(小花介科Cytherellidae)+[达尔文介亚目Darwinulocopina+巴氏介亚目Bairdiocopina]]]。     

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。     

豉甲科分类研究历史

本文对豉甲科分类研究的重要历史事件进行了系统总结回顾,重点介绍了三位著名的国外豉甲分类学家R&#233;gimbart M.,Ochs G和Brinck P.的工作,国内从事豉甲分类工作的专家有周明(羊)先生等.他们都为豉甲分类事业作出了重大贡献.本文将有助于区系分类工作的进一步开展,可为今后的应用研究作准备.     

Phylogeny of hydradephagan water beetles inferred from 18S rRNA sequences

Several families in the beetle suborder Adephaga have an aquatic life style and are commonly grouped in the "Hydradephaga," but their monophyly is contentious and relationships between and within these families are poorly understood. Here we present full-length 18S rRNA sequence for 84 species of Hydradephaga, including representatives of most major groups down to the tribal level, and a total of 68 species of the largest family, Dytiscidae. Using a direct optimization method for the alignment of length-variable regions, the preferred tree topology was obtained when the cost of gaps and the cost of nucleotide changes were equal, and three hypervariable regions of 18S rRNA were downweighted by a factor of five. Confirming recent molecular studies, the Hydradephaga were found to be monophyletic, indicating a single colonization of the aquatic medium. The most basal group within Hydradephaga is Gyrinidae, followed in a comb-like arrangement by families Haliplidae, Noteridae, Amphizoidae, and Hygrobiidae plus Dytiscidae. Under most alignment parameters, Hygrobiidae is placed amid Dytiscidae in an unstable position, suggesting a possible data artifact. Basal relationships within Dytiscidae are not well established, nor is the monophyly of subfamilies Hydroporinae and Colymbetinae. In contrast, relationships at the genus level appear generally well supported. Despite the great differences in the rates of change and the significant incongruence of the phylogenetic signal in conserved vs hypervariable regions of the 18S rRNA gene, both contribute to establish relationships at all taxonomic levels.     

The morphological evolution of the Adephaga (Coleoptera)

The evolution of the coleopteran suborder Adephaga is discussed based on a robust phylogenetic background. Analyses of morphological characters yield results nearly identical to recent molecular phylogenies, with the highly specialized Gyrinidae placed as sister to the remaining families, which form two large, reciprocally monophyletic subunits, the aquatic Haliplidae + Dytiscoidea (Meruidae, Noteridae, Aspidytidae, Amphizoidae, Hygrobiidae, Dytiscidae) on one hand, and the terrestrial Geadephaga (Trachypachidae + Carabidae) on the other. The ancestral habitat of Adephaga, either terrestrial or aquatic, remains ambiguous. The former option would imply two or three independent invasions of aquatic habitats, with very different structural adaptations in larvae of Gyrinidae, Haliplidae and Dytiscoidea.     

On the systematic position of the family Gyrinidae (Coleoptera: Adephaga)

Various characters of adult and larval members of Adephaga and Cupedidae were analyzed, and suggest that Gyrinidae are the sister-group of the remaining Adephaga, and are not closely related to the remaining aquatic Adephaga. The aquatic families Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae seem to form a well founded monophyletic unit. The following characters are considered as synapomorphies of Adephaga excluding Gyrinidae: bifurcate condition of the muscle (= M.) tentoriopraementalis inferior, reduction of hypopharynx, strongly developed prosternal process, reduction in size and specialized modification of the ventral sclerite of the mesothorax, strongly developed mesofurcal arms, a high mesopleural ridge, globular mesocoxae restricted to rotatory movements, invaginated sternum VIII (coxostemum), the strongly curved base of the median lobe of the aedeagus, which articulates with the parameres, the rotated position of the aedeagus in repose, fusion of the larval clypeolabrum with the frons and reduction of the larval lacinia. Mesal shifting of M. episterno-coxalis prothoracis, and the fusion of the apical portions of the malpighian tubules of either side are considered as synapomorphies of Adephaga excluding Rhysodidae and Gyrinidae. Lateral reduction of the meta “sternal” transverse ridge and the presence of the subcubital setal binding patch of the hind wing are considered as synapomorphic characters of Trachypachidae, Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae. We postulate that the metacoxal fusion occurred independently in gyrmids and the common ancestor of Trachypachidae, Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae. Consequently we consider this character state as another synapomorphy of Trachypachidae and Hydradephaga excluding Haliplidae and Gyrinidae. The following characters are considered as synapomorphies of Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae: Loss of tactile setae on the head capsule, metafurcal origin on the intercoxal wall, expansion of the intercoxal wall, elongation of the subcubital setal binding patch, loss of Mm. furca-coxale anterior and posterior, reduction of the larval abdominal segments IX and X, and the shifting of the uropmphi onto the ventral side of segment VIII. Presence of M. tentorio-mandibularis and M. stipitopalpalis intemus are certainly primitive features of adult gyrinids but the distribution of these character states among most members of Adephaga is yet unclear. Chemical defence gland constituents point towards a very isolated position of Gyrinidae. The old age of the group, documented by a larva found in upper Permian deposits, may support the hypothesis of a sister-group relation-ship between Gyrinidae and the remainder of Adephaga.     

Phylogenomic analysis of the beetle suborder Adephaga with comparison of tailored and generalized ultraconserved element probe performance

Adephaga is the second largest suborder of beetles (Coleoptera) and they serve as important arthropod predators in both aquatic and terrestrial ecosystems. The suborder is divided into Geadephaga comprising terrestrial families and Hydradephaga for aquatic lineages. Despite numerous studies, phylogenetic relationships among the adephagan families and monophyly of the Hydradephaga itself remain in question. Here we conduct a comprehensive phylogenomic analysis of the suborder using ultraconserved elements (UCEs). This study presents the first in vitro test of a newly developed UCE probe set customized for use within Adephaga that includes both probes tailored specifically for the suborder, alongside generalized Coleoptera probes previously found to work in adephagan taxa. We assess the utility of the entire probe set, as well as comparing the tailored and generalized probes alone for reconstructing evolutionary relationships. Our analyses recovered strong support for the paraphyly of Hydradephaga with whirligig beetles (Gyrinidae) placed as sister to all other adephagan families. Geadephaga was strongly supported as monophyletic and placed sister to a clade composed of Haliplidae + Dytiscoidea. Monophyly of Dytiscoidea was strongly supported with relationships among the dytiscoid families resolved and strongly supported. Relationships among the subfamilies of Dytiscidae were strongly supported but largely incongruent with prior phylogenetic estimates for the family. The results of our UCE probe comparison showed that tailored probes alone outperformed generalized probes alone, as well as the full combined probe set (containing both types of probes), under decreased taxon sampling. When taxon sampling was increased, the full combined probe set outperformed both tailored probes and generalized probes alone. This study provides further evidence that UCE probe sets customized for a focal group result in a greater number of recovered loci and substantially improve phylogenomic analysis.     

Ultraconserved elements show utility in phylogenetic inference of Adephaga (Coleoptera) and suggest paraphyly of ‘Hydradephaga’

The beetle suborder Adephaga has been the subject of many phylogenetic reconstructions utilizing a variety of data sources and inference methods. However, no strong consensus has yet emerged on the relationships among major adephagan lineages. Ultraconserved elements (UCEs) have proved useful for inferring difficult or unresolved phylogenies at varying timescales in vertebrates, arachnids and Hymenoptera. Recently, a UCE bait set was developed for Coleoptera using polyphagan genomes and a member of the order Strepsiptera as an outgroup. Here, we examine the utility of UCEs for reconstructing the phylogeny of adephagan families, in the first in vitro application a UCE bait set in Coleoptera. Our final dataset included 305 UCE loci for 18 representatives of all adephagan families except Aspidytidae, and two polyphagan outgroups, with a total concatenated length of 83 547 bp. We inferred trees using maximum likelihood analyses of the concatenated UCE alignment and coalescent species tree methods (astral ii , ASTRID, svdquartets ). Although the coalescent species tree methods had poor resolution and weak support, concatenated analyses produced well‐resolved, highly supported trees. Hydradephaga was recovered as paraphyletic, with Gyrinidae sister to Geadephaga and all other adephagans. Haliplidae was recovered as sister to Dytiscoidea, with Hygrobiidae and Amphizoidae successive sisters to Dytiscidae. Finally, Noteridae was recovered as monophyletic and sister to Meruidae. Given the success of UCE data for resolving phylogenetic relationships within Adephaga, we suggest the potential for further resolution of relationships within Adephaga using UCEs with improved taxon sampling, and by developing Adephaga‐specific probes.     

The number that counts. Phylogenetic implications of the number of nurse cells in ovarian follicles of Coleoptera-Adephaga

The structure of polytrophic ovarioles in both Geadephaga and Hydradephaga is reviewed. Particular attention is paid to the significance of various morphological features and their bearing on the phylogeny of Adephaga. It is concluded that the number of nurse cells in the egg chambers is a valuable character in clarifying the phylogenetic relationships. The comparative analysis of the organization of egg chambers in aquatic and terrestrial adephagans, combined with other data suggests that a relatively small and constant number of nurse cells is a plesiomorphic condition for the Adephaga. This conclusion supports the notion that Haliplidae and Gyrinidae stand close to the ancestral stock of adephagous beetles.     

The systematic position of Aspidytidae, the diversification of Dytiscoidea (Coleoptera, Adephaga) and the phylogenetic signal of third codon positions

Characters of the newly discovered larvae of the South African Cliff Water Beetle Aspidytes niobe were examined and integrated into a data matrix including all families of Dytiscoidea as well as Haliplidae. Fifty-three morphological characters of adults and larvae were analysed separately and combined with molecular data from six nuclear and mitochondrial genes. The phylogeny of the group is reconstructed for the study of the evolution of swimming behaviour and larval feeding habits, as well as the shift in diversification rates leading to the two most speciose lineages. The parsimony analysis of all equally weighted morphological and molecular characters combined resulted in a single well supported tree with the topology (Noteridae (Hygrobiidae ((Aspidytidae, Amphizoidae) Dytiscidae))), in agreement with the molecular data alone, but in contradiction to the morphological data, which favoured a topology in which Hygrobiidae is sister to Dytiscidae. The exclusion of third codon positions of the three protein coding genes resulted in a topology identical to that obtained with the morphological data alone, but the use of Bayesian probabilities or the amino acid sequence resulted in the same topology as that of the tree obtained with parsimony using all equally weighted characters. We concluded that interactions of third codon positions with the other data are complex, and their removal is not justified. There was a significant increase in the diversification rate at the base of the richest families (Noteridae and Dytiscidae), which could be associated with the development of simultaneous stroke and higher swimming performance, although data on the swimming behaviour of some basal groups of Noteridae are incomplete. The presence of larval mandibular sucking channels may have contributed to the diversification of Dytiscidae and the species-rich noterid genera Hydrocanthus and Canthydrus .     

Diversity of aquatic and riparian Coleoptera of the Philippines: checklist,state of knowledge,priorities for future research and conservation

The diversity of the aquatic and strictly riparian beetles of the Philippines is reviewed. A checklist (with 317 species and subspecies) of the aquatic and riparian Coleoptera of the Philippines is provided for the first time herein. This checklist covers the following families (number of species and subspecies in parentheses): Hydroscaphidae (1), Gyrinidae (15), Dytiscidae (65), Noteridae (4), Haliplidae (2), Hydraenidae (15), Hydrochidae (2), Spercheidae (1), Hydrophilidae (72), Scirtidae (49), Elmidae (23), Dryopidae (2), Psephenidae (16), Eulichadidae (1), Limnichidae (20), Heteroceridae (5), Malachiidae (5), Lampyridae (15), Nitidulidae (4). A total of 201 species and subspecies (63%) are considered to be endemic to the Philippines. Aquatic/riparian beetle genera and families that are still unrecorded but likely to occur in this country are briefly discussed. Based on estimates at the genus level, it is concluded that about 850 aquatic and riparian species can be expected to occur in the Philippines. The implication of the Philippine biogeographic history for this diversity is discussed. The potential of aquatic/riparian beetles as bioindicators is briefly examined. The dilemma of aquatic invertebrate conservation is analysed and priorities for future research are outlined.