Reevaluation of Blapimorpha and Opatrinae: addressing a major phylogeny‐classification gap in darkling beetles (Coleoptera: Tenebrionidae: Blaptinae)

The taxonomic concepts of Blapimorpha and Opatrinae (informal and traditional, morphology‐based groupings among darkling beetles) are tested using molecular phylogenetics and a reassessment of larval and adult morphology to address a major phylogeny‐classification gap in Tenebrionidae. Instead of a holistic approach (family‐level phylogeny), this study uses a bottom‐up strategy (tribal grouping) in order to define larger, monophyletic lineages within Tenebrioninae. Sampling included representatives of 27 tenebrionid tribes: Alleculini, Amarygmini, Amphidorini, Blaptini, Bolitophagini, Branchini, Cerenopini, Coniontini, Caenocrypticini, Dendarini, Eulabini, Helopini, Lagriini, Melanimini, Opatrini, Pedinini, Phaleriini, Physogasterini, Platynotini, Platyscelidini, Praociini, Scaurini, Scotobiini, Tenebrionini, Trachyscelini, Triboliini and Ulomini. Molecular analyses were based on DNA sequence data from four non‐overlapping gene regions: carbamoyl‐phosphate synthetase domain of rudimentary (CAD) (723 bp), wingless (wg) (438 bp) and nuclear ribosomal 28S (1101 bp) and mitochondrial ribosomal 12S (363 bp). Additionally, 15 larval and imaginal characters were scored and subjected to an ancestral state reconstruction analysis. Results revealed that Amphidorini, Blaptini, Dendarini, Pedinini, Platynotini, Platyscelidini and Opatrini form a clade which can be defined by the following morphological features: adults—antennae lacking compound/stellate sensoria; procoxal cavities externally and internally closed, intersternal membrane of abdominal ventrites 3–5 visible; paired abdominal defensive glands present, elongate, not annulated; larvae—prolegs enlarged (adapted for digging); ninth tergite lacking urogomphi. To accommodate this monophyletic grouping (281 genera and ～4000 species), the subfamily Blaptinae sens. nov. is resurrected. Prior to these results, all of the tribes within Blaptinae were classified within the polyphyletic subfamily Tenebrioninae. The non‐monophyletic nature of Terebrioninae has already been postulated by previous authors, yet no taxonomic decisions were made to fix its status. The reinstatement of Blaptinae, which groups ～50% of the former Tenebrioninae, helps to clarify phylogenetic relations among the whole family and is the first step towards a complete higher‐level revision of Tenebrionidae. The Central Asian tribe Dissonomini (two genera, ～30 species) was not included in Blaptinae due to a lack of representatives in the performed phylogenetic analyses; however, based on morphological features, the tribe is listed as a potential addition to the subfamily.     

Higher level molecular phylogeny of darkling beetles (Coleoptera: Tenebrionidae)

Insect diversity represents about 60% of the estimated million‐and‐a‐half described eukaryotic species worldwide, yet comprehensive and well‐resolved intra‐ordinal phylogenies are still lacking for the majority of insect groups. This is the case especially for the most species‐rich insect group, the beetles (Coleoptera), a group for which less than 4% of the known species have had their DNA sequenced. In this study, we reconstruct the first higher level phylogeny based on DNA sequence data for the species‐rich darkling beetles, a family comprising at least 20 000 species. Although amongst all families of beetles Tenebrionidae ranks seventh in terms of species diversity, the lack of knowledge on the phylogeny and systematics of the group is such that its monophyly has been questioned (not to mention those of the subfamilies and tribes contained within it). We investigate the evolutionary history of Tenebrionidae using multiple phylogenetic inference methods (Bayesian inference, maximum likelihood and parsimony) to analyse a dataset consisting of eight gene fragments across 404 taxa (including 250 tenebrionid species). Although the resulting phylogenetic framework only encompasses a fraction of the known tenebrionid diversity, it provides important information on their systematics and evolution. Whatever the methods used, our results provide strong support for the monophyly of the family, and highlight the likely paraphyletic or polyphyletic nature of several important tenebrionid subfamilies and tribes, notably the polyphyletic subfamilies Diaperinae and Tenebrioninae that clearly require substantial revision in the future. Some interesting associations in several groups are also revealed by the phylogenetic analyses, such as the pairing of Aphtora Bates with Phrenapatinae. Furthermore this study advances our knowledge of the evolution of the group, providing novel insights into much‐debated theories, such as the apparent relict distribution of the tribe Elenophorini.     

The oldest opatrine terrestrial darkling beetle (Coleoptera: Tenebrionidae: Tenebrioninae) from the Paleocene of Menat (France)

The oldest terrestrial darkling beetle of the opatrine lineage, Palaeosclerum pohli gen. et. sp. nov. (subfamily Tenebrioninae) is described from the Middle Paleocene of Menat (France). The complex of characters, combining the peculiar shape of body (robust, adapted to a fossorial lifestyle), structure of fossorial protibia, 5-segmented antennal club demonstrates a considerable similarity of the new genus to the extant representatives of the ‘sclerina’ generic group of the subtribe Opatrina. Palaeosclerum gen. nov. is compared with recent genera Sclerum Dejean, 1834, Platynosum Mulsant and Rey, 1859 and Eurycaulus Fairmaire, 1868.     

Enzyme classification with peptide programs: a comparative study

Background  

Efficient and accurate prediction of protein function from sequence is one of the standing problems in Biology. The generalised use of sequence alignments for inferring function promotes the propagation of errors, and there are limits to its applicability. Several machine learning methods have been applied to predict protein function, but they lose much of the information encoded by protein sequences because they need to transform them to obtain data of fixed length.     

The taxocene of lichen-feeding darkling beetles (Coleoptera,Tenebrionidae: Helopini) in a forest-steppe ecotone

The taxocene of lichen-feeding darkling beetles from the tribe Helopini is studied in the steppes of the south of Rostov Province. The species of lichens grazed on by the representatives of this tribe are identified for the first time and dependence of circadian activity and spatial distribution of adults on temperature and air humidity is shown. Global warming and human activities may have caused degradation of steppe lichen communities that formed the dietary base for the lichen-feeding darkling beetles in natural ecosystems. As a result, 3 out of 4 beetle species completely left steppe biotopes for tree shelterbelts. This caused a change in the trophic relationships (transition from epigeic to epiphytic lichens) and in the spatial distribution of darkling beetles, promoted the establishment of competitive interactions, and increased the duration of circadian and seasonal activity due to the lengthening of the periods with optimum temperature and air humidity for each species.     

Life forms and strategies of lichen-feeding darkling beetles (Coleoptera,Tenebrionidae: Helopini)

Trophic relations of lichen-feeding darkling beetles of the tribe Helopini in Turkey, the Caucasus, and Anti-Atlas are considered. This group is the most diverse in relict Anatolian forests of Cedrus libani, Abies cilicica, Pinus nigra, Juniperus spp., and Quercus cerris, where multitaxon lichenophagous assemblages exist. Three life strategies for attenuation of interspecific competition are distinguished: (1) differentiation of the consumed lichens, (2) separation of the life forms based on the types of the lichens consumed, and (3) segregation of circadian activity in adults. We assume that adaptation of the Helopini larvae to the soil life promoted their wide distribution in different landscapes by elimination of the limiting factor for completing their life cycles, namely the need for ligneous substrate of old rotting trees. The possibility of using lichen-feeding beetles of the tribe Helopini in ecosystem and lichen indication is considered.     

Toward genetics-based virus taxonomy: comparative analysis of a genetics-based classification and the taxonomy of picornaviruses

Virus taxonomy has received little attention from the research community despite its broad relevance. In an accompanying paper (C. Lauber and A. E. Gorbalenya, J. Virol. 86:3890-3904, 2012), we have introduced a quantitative approach to hierarchically classify viruses of a family using pairwise evolutionary distances (PEDs) as a measure of genetic divergence. When applied to the six most conserved proteins of the Picornaviridae, it clustered 1,234 genome sequences in groups at three hierarchical levels (to which we refer as the "GENETIC classification"). In this study, we compare the GENETIC classification with the expert-based picornavirus taxonomy and outline differences in the underlying frameworks regarding the relation of virus groups and genetic diversity that represent, respectively, the structure and content of a classification. To facilitate the analysis, we introduce two novel diagrams. The first connects the genetic diversity of taxa to both the PED distribution and the phylogeny of picornaviruses. The second depicts a classification and the accommodated genetic diversity in a standardized manner. Generally, we found striking agreement between the two classifications on species and genus taxa. A few disagreements concern the species Human rhinovirus A and Human rhinovirus C and the genus Aphthovirus, which were split in the GENETIC classification. Furthermore, we propose a new supergenus level and universal, level-specific PED thresholds, not reached yet by many taxa. Since the species threshold is approached mostly by taxa with large sampling sizes and those infecting multiple hosts, it may represent an upper limit on divergence, beyond which homologous recombination in the six most conserved genes between two picornaviruses might not give viable progeny.     

敦煌莫高窟及周边环境拟步甲虫害调查研究

自2018年以来,每年春夏季在莫高窟窟区及周边荒漠都会出现大规模的拟步甲科昆虫活动,对壁画保存和游客参观造成了影响。为明确拟步甲虫害情况,采用样方法、陷阱法对该区域进行全面普查,以明确拟步甲虫害类型、分布、暴发特点以及对文化遗产的危害。结果表明,在莫高窟暴发的拟步甲科昆虫主要有4种：洛氏脊漠甲Pterocoma loczyi、克氏扁漠甲Sternotrigon kraatzi、三沟胸鳖甲Colposcelis trisulcata和光滑胖漠甲Trigonoscelis sublaevigata,优势种为洛氏脊漠甲。莫高窟约1/3的调查洞窟内有拟步甲活动,对起甲和酥碱壁画威胁较大;其主要取食白刺Nitraria tangutorum,危害荒漠植被。虫害暴发与区域降雨量增加和植物食源丰富相关;亟需采取有效防控措施,以确保文物及其赋存环境安全。本研究为莫高窟虫害的监测防治和文物保护提供了科学依据。     

荒漠景观拟步甲科昆虫多样性及其对生境的指示作用

本研究在宁夏灵武白芨滩国家级自然保护区北部荒漠景观下的磁窑堡地区,选取典型荒漠草原区(DS)、人工柠条林种植区(CCL1,CCL2)及流沙人工封育区(SDE1,SDE2,SDE3)3类生境共6块样地,于2010年4月至8月以巴氏罐诱法采集该地区拟步甲科昆虫,研究该地区拟步甲科昆虫多样性及其对生境的指示作用。结果共获得标本3406头,计12属22种,其中东鳖甲属Anatolica和小鳖甲属Microdera为优势属,波氏东鳖甲Anatolica potanini Reitter、阿小鳖甲Microdera kraatz alashanica Skopin、小丽东鳖甲Anatolica amoenula Reitter和尖尾东鳖甲Anatolic amucronata Reitter为优势种。在流沙人工封育区的SDE1样地物种丰富度指数、均匀度指数和多样性指数最高,SDE3样地个体数量和优势度指数最高;人工柠条林种植区的CCL1样地个体数量最低。聚类分析结果表明流沙人工封育区与荒漠草原区、人工柠条种植区在物种组成及生境相似性方面差异较大。利用指示值法确定不同生境的特征指示种类,结果显示波氏东鳖甲、谢氏宽漠王Mantichorula semenowiReitter、尖尾东鳖甲可作为流沙封育区特征指示种;突角漠甲Trigonocnera pseudopimelia(Reitter)、中华砚甲Cyphogenia chinensis(Faldermann)和纳氏东鳖甲AnatolicanuretiSchusteretReymond可作为荒漠草原特征指示种;蒙古漠王Platyope mongolica Faldermann、多毛宽漠甲Sternoplax setosa setosa(Bates)、克小鳖甲Microdera kraatzi(Reitter)是生境恢复的监测种;小丽东鳖甲Anatolica amoenula Reitter等种类可作为生境退化的监测种。     

基于雌蚊尾器形态的中国阿蚊属分类研究

【目的】明确阿蚊Armigeres雌蚊尾器在分类鉴定中的价值。【方法】依据在云南长期采集的标本, 观察了我国阿蚊属Armigeres正式记录的16种雌蚊尾器构造, 对雌蚊尾器的形态进行了系统描述和分类研究。【结果】编制两个亚属和16种阿蚊的雌蚊尾器分类检索表。【结论】每一种雌蚊尾器与雄蚊尾器一样, 都具有种的固定的特征, 是种的重要分类依据。     

宁夏贺兰山拟步甲科昆虫分布与地形的关系

利用典范对应分析(CCA)及曲线回归拟合,研究了宁夏贺兰山拟步甲科昆虫多样性及分布与地形的关系。结果表明:共调查到14属42种,其中,小圆鳖甲和阿小鳖甲为优势种,其个体数量分别占总捕获个体数的36.09%和19.14%。CCA分析显示,海拔、坡度和平面曲率对拟步甲昆虫的分布格局有显著影响。拟步甲昆虫丰富度和多样性分别与海拔呈显著的线性关系,个体数量与平面曲率呈显著的二次曲线关系。拟步甲昆虫的分布与坡度和坡向没有明显的回归关系,但拟步甲倾向于0°-15°坡度和西北坡向聚集。优势种小圆鳖甲与5个地形因子均不显著相关,因此在贺兰山的分布比较广泛。而优势种阿小鳖甲分布受海拔、坡度和平面曲率影响显著。     

Toward a natural classification: phylogeny of acontiate sea anemones (Cnidaria,Anthozoa, Actiniaria)

Acontia—nematocyst‐dense, thread‐like extensions of the mesenterial filaments―are the characteristic feature of the actiniarian group Acontiaria. Phylogenetic analyses have shown that acontiate taxa form a clade that also includes some taxa without acontia. We analyse five molecular markers from 85 actiniarians to explore the phylogenetic relationships among families in Acontiaria, including acontiate species assigned to other higher taxa and species without acontia that have been allied to Acontiaria. Based on our results, we redefine the group to accommodate those lineages that have lost acontia, and formalize it as superfamily Metridioidea. Based on stable and well supported clades, we resurrect Phelliidae and Amphianthidae, redefine Kadosactinidae and Actinoscyphiidae, and move two species to new genera: that previously termed Sagartiogeton erythraios belongs in Jasonactis gen. nov.; and that previously termed Anthosactis pearseae belongs in Ostiactis gen. nov., type genus of Ostiactinidae fam. nov. We also synonymized Halcampoididae and Halcampidae (as Halcampidae) and Andvakiidae and Isophelliidae (as Andvakiidae). The results of our phylogenetic analyses indicate that the diagnostic morphological characters used in the family‐level taxonomy of acontiate actiniarians such as the nematocysts of the acontia, the marginal sphincter muscle, and mesenteries divisible into macro‐ and micro‐cnemes, have to be revisited, as these features are highly homoplasious.     

Spatial structure of community of darkling beetles (Coleoptera: Tenebrionidae) in the Negev Highlands, Israel

The distribution of niches in resource space of darkling beetles of Negev desert, Israel were studied using discriminant analysis Seventeen environmental variables (substrate structure and vegetation) were measured at 375 plots within 15 grids The first three discriminant axes account for 77% of the variance and reflect a soil structure gradient from sand to clay (the first axis), a gradient of increasing productivity (the second axis), and a type of vegetation architecture (the third axis) Abundance and biomass of tenebrionids increased linearly along the first, the second and the third axes Species richness and diversity had a slanting parabola shape along the substrate axis and increased along the productivity axis There were two distinct spatial guilds of tenebrionid species One guild was composed by species that preferred sandy-gravel soils, and another guild consisted of species that preferred clay soils Niche overlapping of the "sandy-gravel" species was lower than that of the "clay" species Niches of the "sandy-gravel" species were randomly distributed in the resource space Those of the "clay" species formed a tight cluster with most niches placed in the center of the space, representing the centrifugal type of organization The conformity of the guild structure with the different types of organization was supported by compansons between empirical data and stochastic models The border between these two guilds matches with proposed earlier biogeographical boundary (Ramon-Zohar line)     

Toward a classification of lotic habitats

Summary A world-wide system for classifying brooks, streams, and small rivers is proposed, using the following criteria: width, flow, current speed, substrate, summer temperatures, winter temperatures, turbidity, total dissolved organic matter, total dissolved inorganic matter, water hardness, dissolved oxygen, rooted aquatic plants, streamside vegetation. Some of these criteria have much greater biological implications than others. It is contended that widely separated lotic habitats that are similar in the above features have biotas consisting of ecologically similar and parallel clusters of species. There are, nevertheless, many kinds of atypical polluted and unpolluted lotic habitats that cannot be classified with respect to these criteria.Contribution No. 63, Limnology Laboratory, University of Colorado.     

The interaction between bird predation and plant cover in determining habitat occupancy of darkling beetles

Tenebrionid beetles in the Negev Desert exhibit size-related habitat segregation, with larger species found in denser cover. Size-dependent predation by birds has been suggested as the mechanism behind this habitat segregation. Two predictions of this hypothesis were tested: (1) plant cover reduces the predation efficiency of birds upon large tenebrionids, and (2) birds prefer larger species. Both predictions were supported: plant cover reduced predation rate by the most common spring and summer predatory birds: white storks ( Ciconia ciconia ) and stone curlews ( Burhinus oedicnemus ), in cage experiments. Results from preference experiments suggest that tenebrionid species can be divided according to their profitability as prey. Large species are the most profitable, medium-sized species are less profitable but still acceptable and small species are unprofitable and therefore ignored. Field observations demonstrated that the well-vegetated wadi habitats are dominated by large and small species whereas acceptable , medium-sized species are under-represented in this habitat. The results of the cage experiments indicate possible apparent competition between the large profitable and the medium acceptable tenebrionid species in the wadis. Aggregative response of predators in the profitable habitat is suggested as the mechanism leading to truncated distribution of prey species. Large profitable species are refuge-dependent, medium-sized acceptable species use enemy free space and small species are predator independent.     

四种天牛的核型比较研究

为弄清天牛的核型特征, 补充我国天牛染色体分类特征和天牛细胞分类学的基础研究空缺, 本文以4种沟胫天牛亚科天牛为研究对象, 选取天牛的分裂旺盛组织精巢（卵巢）和中肠, 在不同的条件下进行天牛染色体玻片的制备和观察。结果显示： 4种天牛染色体数目均为 2n = 20, 性别决定机制为Xy_p。其中, 云斑白条天牛Batocera lineolata的核型公式： 4L+5M+Xy_p, 大型染色体4对, 中型染色体5对, 性染色体为小型; 榉白背粉天牛Olenecamptus cretaceus marginatus、 中华八星粉天牛Olenecamptus octopustulatus chinensis（粉天牛属）和桑天牛Apriona germari的核型公式均为： 5L+4M+Xy_p, 大型染色体5对, 中型染色体4对, 性染色体为中型。粉天牛属的榉白背粉天牛和中华八星粉天牛的核型指数非常相近; 桑天牛的染色体公式虽然与粉天牛属相同, 但长度和着丝粒位置明显不同。     

台湾和日本拟步甲在纬度分布上的比较研究

台湾和日本处在不同的地理纬度上,同属岛屿海洋性气候,前者地域面积远小于后者,拟步甲的物种多样性却大于后者。为弄清楚这些科学问题,作者采用G-F指数对从台湾到日本不同纬度梯度上的拟步甲多样性分布格局进行了比较分析,得到如下初步结论：(1) G-F指数从大到小依次是：台湾(21°N-25°N)(0．826)>日本(24°N-45°N)(0．824)>日本Ⅱ纬度区(30°N-35°N)(0．792)>日本Ⅰ纬度区(24°N-30°N)(0．765)>日本Ⅲ纬度区(35°N-40°N)(0．761)>日本Ⅳ纬度区(40°N-45°N)(0．603);(2)台湾拟步甲属的多样性(DG)、族的多样性(DF)和G-F指数(DG-F)均最高,分别是4．263、24．464和0．826;(3)各纬度上拟步甲的物种分布情况：台湾(21°N-25°N)(541种)>日本(24°N-45°N)(489种)>日本Ⅰ纬度区(24°N-30°N)(257种)>日本Ⅱ纬度区(30° N-35° N)(231种)>日本Ⅲ纬度区(35° N-40° N)(172种)>日本Ⅳ纬度区(40° N-45° N)(60种)。研究数据显示, G-F指数能较好地反映台湾和日本各地拟步甲族、属的多样性。其物种多样性在纬度上的分布表现为从南向北递减的趋势,并对其基本原因进行分析。作者首次基于台湾和日本两个岛屿拟步甲物种多样性的比较分析,对现有岛屿生物多样性的有关理论提出个人看法,认为岛屿生物地理学的“物种-面积关系理论”中的“岛屿面积越大,物种数量就越多”可能存在一定的局限性,不一定能客观地反映种类众多的现生岛屿昆虫物种多样性的实际情况。     

水松自然种群和人工种群遗传多样性比较

采用ISSR分子标记技术分析水松不同起源种群的遗传多样性.结果表明:10条引物共检测出95个扩增位点,多态位点数占39.0%.与其他濒危裸子植物相比,水松的遗传多样性较低,遗传分化系数Gst为0.3982,基因流Nm仅0.3778,种群间存在一定程度的遗传分化,但种群内变异占主导地位;遗传距离与地理距离呈正相关关系.自然种群的多态位点百分率(P)、Nei的条带多样度(He)和Shannon信息指数(Ⅰ)平均值(39.3%、0.1499和0.2202)分别高于人工种群(30.7%、0.1265和0.1759).自然种群的遗传分化系数(Gst0.4513)和平均遗传距离(D=0.0301)也高于人工种群(Gst=0.3025,D=0.0192).