TAXONOMIC STATUS OF BARK BEETLES (COLEOPTERA,CURCULIONIDAE)

许多学者将小蠹作为象虫总科Curculionoidea中一个独立的科Scolytidae,而目前国际上普遍认为小蠹为象虫科Curculionidae中的一个亚科,即小蠹亚科Scolytinae.Erichson在1842年即提出了小蠹为象虫科的一个亚科,20世纪以R.A.Crowson为代表的学者对此观点表示支持并进行了较为严谨的科学论证,而以S.L.Wood为代表的另外一些学者则坚持小蠹的科级地位,我国学者蔡邦华、殷惠芬等一直沿用该分类系统.近20年来,国际上许多学者通过对小蠹类昆虫的成虫和幼虫外部特征的深入研究,以及支序系统学和分子生物学等一些最新结果,普遍认为小蠹类昆虫应为象虫科的一个业科,这一观点已被国际上广泛接受.本文全面介绍了小蠹分类研究的历史及新的研究进展,并提出和建议采用“小蠹亚科”这一分类地位,与象虫亚科并列,均隶属于象虫科.     

象甲科线粒体基因组特征及系统发育分析

昆虫线粒体基因组广泛应用于系统发育关系的重新建立、分子进化、谱系地理学及物种诊断等领域。为揭示象甲科昆虫线粒体全基因组序列的主要结构特征,探究其系统发育相关信息,为进化遗传学研究和分子标记选取等提供参考依据,本研究利用比较基因组学和生物信息学方法,对NCBI上已公布的35种象甲科物种线粒体全基因组序列进行了分析。结果显示:(1)象甲科tRNA基因存在排序及数目异常情况,不同物种中蛋白质编码基因和2种rRNAs排列相同,线粒体全基因组具有明显AT偏向;(2)COX1、ATP6、ND5、ND4、ND4L和ND1基因除标准三联密码子外,还存在特殊的起始密码子AAT、TTG和GTG;(3)13种蛋白质编码基因的进化速率顺序为COX3ATP8ND2ND5ND1ND4ND6ND4LND3ATP6CytbCOX1COX2;(4)13个蛋白编码基因和rRNAs基因中,ND5、rrnL、ND4和ND2基因变异位点数较高,可作为备选的分子标记;(5)各亚科的系统发育关系可能为(((小蠹亚科Scolytinae+长小蠹亚科Platypodinae)+(隐喙象亚科Cryptorhynchinae+魔喙象亚科Molytinae+象虫亚科Curculioninae)+((孢喙象亚科Cyclominae+粗喙象亚科Entiminae)+(隐颏象亚科Dryophthorinae+长小蠹亚科))),为象甲科的系统发育分析有提供参考。     

耳喙象亚科的分类简史及其系统地位简述

耳喙象亚科Otiorrhynchinae是象虫科最大的一个亚科,包括短喙象和耳喙象两大类群。据作者初步统计,全世界已记录800余属(亚属),约8500种,我国已记录70属约400种。耳喙象的分类系统地位,一直存在着争议,国内学者尚未进行这方面的研究和讨论。本文就耳喙象亚科的分类简史及其系统地位作一简单介绍和评述。一、分类简史1758年,林奈在《自然系统》第10版中首创了Curculio属,记载了80种象虫,C.palma-run为模式种,其中我国有一种(C.chinensis)。1826年,Schoenherr首次将象虫分为Brach-yerides、Entimides、Pachyrychides、Cleonides,Cyclomide…     

枣食芽象甲线粒体基因组全序列测定与系统发育分析

【目的】从线粒体基因组水平上探讨枣食芽象甲Scythropus yasumatsui与近缘种的系统发育关系。【方法】利用Illumina MiSeq测序平台对枣食芽象甲线粒体基因组进行测序,对基因组序列进行拼装、注释和特征分析;利用贝叶斯法和最大似然法构建基于象甲科13个物种的线粒体基因组13个蛋白质编码基因核苷酸序列的系统发育树。【结果】结果表明,枣食芽象甲线粒体基因组全长为16 472 bp (GenBank登录号: MF807224),包含13个蛋白质编码基因、22个tRNA基因、2个rRNA基因和2个非编码控制区,37个基因的排列顺序与祖先昆虫的线粒体基因排列顺序一致。13个蛋白质编码基因的起始密码子为ATN,其中除了cob和nad1基因的完全终止密码子为TAG外,其余11个基因的完全终止密码子为TA(A)。22个tRNA基因中除了trnS1缺少DHU臂,反密码子由GCT变为TCT外,其余均能形成典型的三叶草结构。基于13个蛋白质编码基因序列构建的系统发育树结果显示,象甲科8个亚科系统发育关系为：(((隐喙象亚科(Cryptorhynchinae)+(象虫亚科(Curculioninae)+魔喙象亚科(Molytinae)))+长小蠹亚科(Platypodinae))+(粗喙象亚科(Entiminae)+Cyclominae亚科))+隐颏象亚科(Dryophthorinae)+小蠹亚科(Scolytinae))。【结论】在13种象甲科昆虫物种中,同属于粗喙象亚科的枣食芽象甲与南美果树象甲Naupactus xanthographus在系统发育树中聚为同一分支,表明基于线粒体基因组全序列的分子系统发育结果与传统的形态分类结果是一致的。     

基于mtDNA CO I基因的十种长小蠹分子系统进化研究 (鞘翅目:长小蠹科)

全世界记录的长小蠹(鞘翅目:长小蠹科)有1 500余种,其分类地位一直存在争议.本研究通过分子基因信息探讨长小蠹科昆虫的分子系统进化关系,测定了中对长小蠹Euplatypus parallelus(Fabricius)、希氏长小蠹E. hintzi Schaufuss、散对长小蠹E. solutus Schedl、杯长小蠹Dinoplatypus cupulatus Chapuis、小杯长小蠹D.cupulatulus Schedl、芦苇黄截尾长小蠹D.calamus Blandford、五棘长小蠹Diapus quinquespinatus Chapuis、锥长小蠹Treptoplatypus solidus Walk、栎长小蠹Platypus quercivorus Murayama和东亚长小蠹P.lewosi Blandford等长小蠹科5属10种mtDNA CO I基因部分序列(549 bp);采用MEGA3.1分析了序列组成及遗传距离,应用PAUP4.0分别构建了长小蠹NJ,MP和ML等3种分子系统树,同时结合长小蠹的形态分类,探讨10种长小蠹及其所在属的系统进化.结果表明:长小蠹科昆虫在碱基使用频率上有很大的偏向性;长小蠹科与外群小蠹科松瘤小蠹Orthotomicus erosus Wollaston之间的遗传距离(0.288～0.338)远大于科内种间距离(0～0.226);Diapus属分化最早,Euplatypus属独成一支,Treptohlatypus,Dinoplatypus和Platypus 3属分化为一支.长小蠹科分子系统进化研究结果与Wood(1993)新修订的分类系统基本一致,说明长小蠹科的分类系统更趋于合理.     

盲蝽科昆虫的分类系统概述

依据目前已有的资料 ,概述了盲蝽科昆虫的分类系统。盲蝽科隶属于半翅目异翅亚目臭虫型的盲蝽总科 ,这在半翅目学者中意见一致。但关于盲蝽科亚科及族级水平的分类系统观点不一。最合理的为 6亚科及 8亚科的分类系统。8亚科分类系统是目前最被接受和应用的系统。     

基于mtDNA COⅠ基因的十种长小蠹分子系统进化研究（鞘翅目：长小蠹科）

全世界记录的长小蠹（鞘翅目：长小蠹科）有1 500余种,其分类地位一直存在争议。本研究通过分子基因信息探讨长小蠹科昆虫的分子系统进化关系,测定了中对长小蠹Euplatypus parallelus（Fabricius）、希氏长小蠹E. hintzi Schaufuss、散对长小蠹E. solutus Schedl、杯长小蠹Dinoplatypus cupulatus Chapuis、小杯长小蠹D. cupulatulus Schedl、芦苇黄截尾长小蠹D. calamus Blandford、五棘长小蠹Diapus quinquespinatus Chapuis、锥长小蠹Treptoplatypus solidus Walk、栎长小蠹Platypus quercivorus Murayama和东亚长小蠹P. lewisi Blandford等长小蠹科5属10种mtDNA COⅠ基因部分序列（549 bp）;采用MEGA3.1分析了序列组成及遗传距离,应用PAUP4.0分别构建了长小蠹NJ,MP和ML等3种分子系统树,同时结合长小蠹的形态分类,探讨10 种长小蠹及其所在属的系统进化。结果表明：长小蠹科昆虫在碱基使用频率上有很大的偏向性;长小蠹科与外群小蠹科松瘤小蠹Orthotomicus erosus Wollaston之间的遗传距离（0.288～0.338）远大于科内种间距离（0～0.226）;Diapus属分化最早,Euplatypus属独成一支,Treptohlatypus, Dinoplatypus和Platypus 3属分化为一支。长小蠹科分子系统进化研究结果与Wood（1993）新修订的分类系统基本一致,说明长小蠹科的分类系统更趋于合理。     

叶甲科三亚科十三种叶甲酯酶同工酶的研究(鞘翅目,叶甲总科)

应用聚丙烯酰胺凝胶电泳方法分析研究了叶甲科3亚科13种昆虫的酯酶同工酶.结果显示,其聚类结果与传统分类结果基本相一致,说明以酯酶同工酶作为研究手段来进行叶甲类昆虫亚科以下阶元的分类是可行的,同时也说明了它们酯酶同工酶酶谱的差异和其分类地位是一致的;但跳甲亚科和叶甲亚科先聚为一类,再与萤叶甲亚科聚为一类,与前人的研究有差异,作者认为:酯酶同工酶的编码基因可能是快进化单位,在解决亚科以下阶元的系统关系时是很好的分子标记,而对于研究叶甲科、亚科间的系统关系,就不一定很合适.     

中国根小蠹属记述(小蠹研究之四)

根小蠹属Hylastes Erichson(1836年)是齿小蠹科中一个较小的属,隶属于海林小蠹亚科Hylesininae,根小蠹族Hylastini;Hagedorn(1910年)曾把广义的Hylastes属分为Scierus,Hylastes,Hylurgops和Hylastinus四个亚属;但Reitter(1913)以后学者把上述各亚属都提升为属,现已被各国学者所通用。     

基于内转录间隔区基因片段对叩甲科昆虫的系统发育研究

【目的】采用叩甲科昆虫的内转录间隔区(ITS-2)片段对叩甲科昆虫部分种类进行系统发育分析,明确各亚科类群间的亲缘关系并同传统分类系统进行比较,验证ITS-2片段是否能用于叩甲科昆虫的分子系统学研究。【方法】基于叩甲科昆虫的内转录间隔区(ITS-2)片段,对10个亚科69个种类的叩甲进行了系统发育分析。进行系统发育信号检测,计算各亚科之间的遗传距离,采用邻接法、似然法和简约法三种模型构建系统发育树。【结果】3种方法构建系统发育树结构基本一致,各亚科都能被很好的聚类,且各亚科间遗传距离符合传统分类学观点。同时,分析结果也支持对传统分类系统进行修改:尖鞘叩甲亚科Oxynopterinae和异角叩甲亚科Pityobiinae被聚入齿胸叩甲亚科Denticollinae,建议将尖鞘叩甲亚科和异角叩甲亚科同齿胸叩甲亚科合并;槽缝叩甲亚科Agrypninae和单叶叩甲亚科Conoderinae并入萤叩甲亚科Pyrophorinae;梳爪叩甲类群Melanotinae被聚类到叩甲亚科Elaterinae内,建议将梳爪叩甲亚科并入叩甲亚科。【结论】分析结果显示,ITS-2片段适用于叩甲科昆虫低级分类阶元的系统发育分析,各亚科之间的亲缘关系符合传统的分类学观点,但聚类结果也有同传统分类系统有所不同,可为分类系统的修改提供依据。     

The most ancient bark beetle known: a new tribe,genus and species from Lebanese amber (Coleoptera,Curculionidae, Scolytinae)

Abstract Cylindrobrotus pectinatus gen. et sp.n. , a new scolytine species from Cretaceous Lebanese amber, is described. A new tribe, Cylindrobrotini trib.n. , is proposed for this unique species, which demonstrates an unusual combination of some archaic and many advanced characters. This finding suggests that the Scolytinae became a distinct lineage of Curculionoidea from the Lower Cretaceous. Fossil records are reviewed, and some remarks on the origin and taxonomic position of bark and ambrosia beetles are made. Some comments on the various phylogenetic interpretations of the last 30 years are given, particularly in respect of their correspondence with the fossil record. The early appearance of Scolytinae in the fossil record before other Curculionidae (which appeared in the Upper Cretaceous) can be used as evidence against the hypothesis of bark beetles as offspring of weevils. The question of the taxonomic rank of bark beetles (separate subfamily or family) and their placement among other groups of the superfamily remains unsolved.     

Brief Review of the Associations of Xylobiont Nematodes with Bark Beetles (Coleoptera,Curculionidae: Scolytinae)

Nematode pathogens cause wilt diseases in conifers and deciduous trees. The longhorn beetles (Coleoptera: Cerambycidae) and bark beetles (Coleoptera: Curculionidae: Scolytinae) act as nematode vectors spreading the invasive juvenile stages during their maturation feeding or during oviposition on the plant hosts. There are numerous reviews of nematodes associated with bark beetles on conifers, while little attention has been paid to the nematodes of deciduous trees. The development of Dutch elm disease and ash dieback is mainly caused by fungal pathogens transmitted by bark beetles; the latter act as vectors of not only fungi but also nematodes enclosed in nematangia under their elytra, and also in the tracheae and Malpighian canals. Apart from phytopathogenic nematodes, bark beetles transmit mycophagous and bacterivorous nematodes and own parasites of bark beetles. The ecological groups of nematodes associated with Scolytinae are reviewed; the known records of associations of nematodes with bark beetles are listed for coniferous host plants of Russia and neighboring countries; the world-wide list of these associations for deciduous plant hosts is given.

     

The Banded Elm Bark Beetle, Scolytus schevyrewi Semenov (Coleoptera, Curculionidae, Scolytinae) in North America: a taxonomic review and modifications to the Wood (1982) key to the species of Scolytus Geoffroy in North and Central America

In 2003, an Asian bark beetle, Scolytus schevyrewi Semenov (Coleoptera: Curculionidae: Scolytinae), the banded elm bark beetle, was detected for the first time in North America. This paper modifies the Wood (1982) key to the species of Scolytus Geoffroy to enable identification of Scolytus schevyrewi in North and Central America. Variation of diagnostic characters in Scolytus schevyrewi is discussed.     

Bark and Ambrosia Beetles (Coleoptera,Curculionidae: Scolytinae) of Northwest Russia: History of the Study,Composition and Genesis of the Fauna

Entomological Review - An essay of the fauna of bark beetles (Coleoptera: Curculionidae: Scolytinae) in five federal subjects of Northwest Russia is presented. The species composition of scolytids...     

Relative performance of Lindgren multiple-funnel, Intercept panel, and Colossus pipe traps in catching Cerambycidae and associated species in the southeastern United States

In 2004, we evaluated the relative performance of 8-unit Lindgren multiple-funnel (funnel), Intercept panel (panel), and Colossus pipe (pipe) traps, baited with ethanol and ac-pinene lures, in catching saproxylic beetles (Coleoptera) in pine stands in northern Florida and western South Carolina. Panel traps were as good as, if not better than, funnel and pipe traps for catching Cerambycidae. In particular, more Monochamus titillator (F.) were captured in panel traps than in pipe and funnel traps. Of three species of Buprestidae captured in our study, most Buprestis lineata F. were caught in panel traps, whereas most Acmaeodera tubulus (F.) were caught in funnel traps. Catches of Chalcophora virginiensis Drury and the root-feeding weevils Hylobius pales Herbst an dPachylobius picivorus LeConte (Curculionidae) were unaffected by trap type. Among bark beetles (Curculionidae: Scolytinae), catches of Ips grandicollis (Eichhoff) were unaffected by trap type, whereas most Dendroctonus terebrans (Olivier) were caught in panel traps, most Hylastes salebrosus Eichhoff were caught in panel and pipe traps, and most Hylastes tenuis Eichhoff were caught in funnel traps. Among ambrosia beetles (Curculionidae: Scolytinae), panel traps caught the most Xyleborinus saxesenii (Ratzeburg), whereas pipe traps caught the most Xyleborus Eichhoff spp. More Xylosandrus crassiusculus (Motschulsky) and Dryoxylon onoharaensis (Murayama) were caught in panel and funnel traps than in pipe traps. Among bark beetle predators, more Platysoma Leach spp. (Histeridae) were caught in pipe and panel traps than in funnel traps, whereas most Lasconotus Erichson spp. (Zopheridae) were caught in funnel traps. Variation among trap performance for various species suggests that managers should consider more than one type of trap in their detection programs.     

Exploring associations between international trade and environmental factors with establishment patterns of exotic Scolytinae

Although invasion of exotic ambrosia beetles (fungus feeders) and bark beetles (phloem feeders) (Coleoptera: Curculionidae: Scolytinae) is considered a major threat to forest health worldwide, no studies have quantitatively investigated the anthropogenic and environmental factors shaping the biogeographical patterns of invasion by these insects across large spatial scales. The primary aim of this study was to assess the relative importance of international trade and several environmental variables of the recipient region on species richness of established exotic Scolytinae. As a reference, we also evaluated the relationships between the same environmental variables and species richness of native Scolytinae. Using an information-theoretic framework for model selection and hierarchical partitioning, we evaluated the relative importance of the potential drivers of species richness of native and exotic Scolytinae in 20 European countries and the 48 contiguous continental US states. Analyses were conducted separately for ambrosia and bark beetle species. Value of imports was a strong predictor of the number of exotic Scolytinae species in both regions. In addition, in the USA, warmer and wetter climate was positively linked to increased numbers of both native and exotic ambrosia beetles. Forest heterogeneity and climatic heterogeneity and secondarily forest area were key drivers in explaining patterns of species richness for native bark beetles but not for exotic species in both regions. Our findings suggest that if current infestation levels continue on imported plants and wood packaging material, increasing international trade will likely lead to more establishments of exotic Scolytinae with concomitant negative effects on forest health in both Europe and the USA. Compared to Europe the risk of invasion appears higher in the USA, especially for ambrosia beetles in the southeastern USA where the climate appears highly suitable for exotic establishment.     

Classification of weevils as a data-driven science: leaving opinion behind

Data and explicit taxonomic ranking criteria, which minimize taxonomic change, provide a scientific approach to modern taxonomy and classification. However, traditional practices of opinion-based taxonomy (i.e., mid-20^th century evolutionary systematics), which lack explicit ranking and naming criteria, are still in practice despite phylogenetic evidence. This paper discusses a recent proposed reclassification of weevils that elevates bark and ambrosia beetles (Scolytinae and Platypodinae) to the ranks of Family. We demonstrate that the proposed reclassification 1) is not supported by an evolutionary systematic justification because the apparently unique morphology of bark and ambrosia beetles is shared with other unrelated wood-boring weevil taxa; 2) introduces obvious paraphyly in weevil classification and hence violates good practices on maintaining an economy of taxonomic change; 3) is not supported by other taxonomic naming criteria, such as time banding. We recommend the abandonment of traditional practices of an opinion-based taxonomy, especially in light of available data and resulting phylogenies.     

Trophic associations and specialization of phytophagous beetles (Coleoptera: Chrysomeloidea,Curculionoidea) in the east of the Russian Plain

Data on the trophic associations of beetles with plants in the east of the Russian Plain are summarized and comparative analysis of host specialization of different groups of phytophagous beetles is performed. In terms of the width of the regional trophic spectrum, monophages and narrow oligophages prevail among the Curculionoidea as a whole and in the families Curculionidae and Apionidae in particular, while moderate and broad oligophages prevail in the Chrysomeloidea and in the family Chrysomelidae. Two-thirds of the regional fauna (66%) of Curculionoidea are closely associated with plants of one genus; by contrast, in Chrysomeloidea almost 40% of the species can develop on plants from different genera of one family, the fraction of the narrowly specialized forms comprising only 43%. The higher level of trophic specialization of weevils (Curculionidae, Apionidae) and seed beetles (Bruchidae), as compared to leaf beetles (Chrysomelidae), is probably due to the larval endophagy of most species of these families. Analysis of the distribution of beetles over host plants has shown that the specialized forms are associated with plants of 65 families (about 60% of the regional flora in the east of the Russian Plain). Distribution of beetles over plant families is very non-uniform. Most of the specialized forms (78%) are associated with plants of 15 families, three of which (Asteraceae, Fabaceae, and Brassicaceae) include hosts of more than onethird of the beetle species (37%). Monophages and narrow oligophages are recorded on 201 genera of plants from 59 families. Polyphagous species are recorded on plants of 58 families. The specific features of the distribution of phytophagous beetles over host plants (as compared to other insects) is a high fraction of species developing on coenophobes (particularly those of the family Brassicaceae) typical of the pioneer stages of successions with sparse herbaceous cover, and a small number of species associated with grasses and sedges. These features are most conspicuous in the fauna of Curculionidae.     

Acoustic communication in the pine engraver bark beetle: do signals vary between behavioural contexts?

Acoustic communication is taxonomically widespread in bark beetles and is proposed to play an important role in a variety of social and defensive behavioural contexts. Yet our understanding of how signals vary between contexts is currently limited. The present study tests the hypothesis that acoustic signals vary between behavioural contexts in the female pine engraver beetle Ips pini (Say) (Coleoptera: Curculionidae: Scolytinae). Female Ips pini produce acoustic chirps using a vertex‐pronotal stridulatory organ. Randomly sampled chirps generated under three contexts (i.e. distress, predation and premating) are compared for their duration, number of pulses, interpulse intervals, pulse rate and amplitude envelope shapes. The results obtained show that, during premating events, chirps are significantly longer in duration and tend to have a higher proportion of descending amplitude envelopes than chirps occurring during distress and predation events. Chirps produced during distress and predation conditions are indistinguishable from one another. By contrast to the results from previous bark beetle studies, no support is found for categorizing chirps as ‘interrupted’ or ‘uninterrupted’ types based on temporal patterns. The functional significance of context‐dependent variation in chirp characteristics is discussed. Previous studies on acoustic communication in bark beetles are limited as a result of a general lack of objective sampling and measurement criteria for characterizing signals. Recommendations are outlined for future studies on the functions and evolution of acoustic communication in bark beetles.     

Mouthparts of the bark beetle (Ips acuminatus) as a possible carrier of pathogenic microorganisms

The bark beetle Ips acuminatus Gyllenhal (Coleoptera: Curculionidae: Scolytinae) has been recently reported as one of the most serious secondary pests of pine trees. Since the adult beetles chew the sapwood to create tunnels, they have developed effective drilling mouthparts enough to make galleries directly into the heartwood of the tree. The mouthparts of this bark beetle exhibit typical morphology of mycophagous coleopteran beetles and have those characteristics of chewing mouthparts. Both maxillary and labial palpi have the functions of directing the food to the mouth and holding it while the mandibles chew the food. Although this bark beetle did not have prothoracic mycangial cavities, yeast-like spores were concentrated at the invaginated surface of mouthparts where cuticular hairs are densely packed. In particular, the cuticular surface around the base segments of these palpi has sufficient spaces to accommodate microorganisms during the series of drilling or feeding processes. Therefore, this paper reports detailed observation of the cuticular structure of the mouthpart using the field emission scanning electron microscopy (FESEM) for the purpose of demonstrating its possible implication to act as external carriers of pathogenic microorganisms.