摘要
异翅亚目(昆虫纲：半翅目)是昆虫纲中种类较多,世界范围内广泛分布的一个类群。本论文以异翅亚目部分种类作为研究对象,进行了DNA条形码的筛选及应用、谱系地理学和花蝽科(狭义)的分子系统发育三方面的研究。
第一部分,DNA条形码的筛选及应用：在动物界中DNA条形码所面临的主要困难是找到一个理想的基因区段可以区分任何物种。已往对异翅亚目部分种类的研究中发现单独使用C0I序列时存在无法区分物种的情况,因此有必要筛选合适的备选分子标记,以备C0I无法区分时使用。而备选的标记也必须考虑近缘种和地理种群等因素的影响。本研究选择斑红蝽属的四个种：小斑红蝽Physopelta cincticollis Stal,1863,突背斑红蝽Physopelta gutta (Burmeister,1834),浑斑红蝽Physopelta robusta Stal,1863和显斑红蝽Physopelta slanbuschii (Fabricius,1787),依据广域大样本量多基因研究,结合种间-种内遗传距离分析和多种方法建树分析,结果支持Cyt b和COI都适合作为半翅目-异翅亚目DNA条形码的理想分子标记。
将DNA条形码技术应用于分类研究中,可以对以前的分类结果进行新的判断。本研究通过中国普缘蝽属Plinachtus Stal(半翅目：异翅亚目：缘蝽科)已知的4个种：刺肩普缘蝽Plinachtus dissimilis Hsiao,1964,钝肩普缘蝽Plinachtus bicoloripes Scott,1874,黑普缘蝽Plinachtus acicularis (Fabricius,1803)和棕普缘蝽Plinachtus basalis (Westwood,1842)的COI(1338 bp)序列分析,计算其种间／种内的遗传距离,并进行基于距离法、最大简约法和贝叶斯法的分析,结果均支持将刺肩普缘蝽和钝肩普缘蝽合并为1个种。
第二部分,在异翅亚目内部还未见以中国为主要区域的谱系地理学研究报道。对加入来自中国和伊朗的分子数据对稻绿蝽(异翅亚目：蝽科)世界范围谱系地理学的重新分析表明：位于青藏高原在上新世-更新世时期的快速隆起造成了东亚和西亚种群的分隔。同时与以往研究不同的结论有：由于中新世非洲微板块,阿普利亚和阿拉伯半岛与欧洲板块的碰撞后,形成了一系列连续的阻隔物(如：红海,扎格洛斯山脉)造成了稻绿蝽非洲与其它地区种群的隔离。在此类群中,Cyt b也被证明是DNA条形码在异翅亚目中除COI外合适的分子标记。在欧原花蝽群(异翅亚目：花蝽科)内通过重塑种内的系统发育关系、祖先分布地、扩散路径以及种间分化的时间等,表明欧原花蝽群起源于横断山区,目前的分布格局主要受青藏高原近期两次抬升的影响。随着青藏高原的抬升而隆起的一些山脉促进了欧原花蝽群的分化。隔离和扩散在欧原花蝽群的分布区形成上都有非常重要的作用。
第三部分,已往只有基于形态数据的花蝽科内部的系统发育研究。本研究是在狭义花蝽科内首次应用分子数据进行的系统发育研究。选用18S rDNA,16S rDNA和细胞色素氧化酶I (COI)序列作为分子标记,总长度为4 kb。基于ML和Bayesian两种算法的系统发育结果可分为两大支系。支系Ⅰ包括原花蝽族Anthocorini(松花蝽属Elatophilus除外)小花蝽族Oriini,但小花蝽族中的圆花蝽属Bilia与其它属不属于一个支系,沟胸花蝽Dufouriellini的叉胸花蝽属Amphiareus,齿股花蝽族Scolopini的Calliodis属。支系Ⅱ包括点刻花蝽族Almeidini,原花蝽族Anthocorini的松花蝽属Elatophilus,沟胸花蝽族Dufouriellini(叉胸花蝽属Amphiareus除外),仓花蝽族Xylocorini,齿股花蝽族Scolopini的齿股花蝽属Scoloposcelis和刷花蝽族Blaptusterini;其中沟胸花蝽族Dufouriellini的沟胸花蝽属Dufouriellus为一单独分支,Cardiastethus, Dysepicritus和Physopleurella组成另一分支,二者关系亦不近缘。由此可知,已往的原花蝽族Anthocorini、沟胸花蝽族Dufouriellini、小花蝽族Oriini和齿股花蝽族Scolopini均为并系类群,需要设立新族以分别容纳原花蝽族Anthocorini中的松花蝽属Elatophilus和小花蝽族Oriin中的圆花蝽属Bilia。沟胸花蝽族Dufouriellini应拆分成在3个族级单元以分别容纳沟胸花蝽Dufouriellus,((Dysepicritus + Cardiastethus)+Physopleurella)和Amphhiareus。齿股花蝽族Scolopini的两个亚族Scolopina和Calliodina应该提升为族。
Heteroptera is a large group in insect with a world wide distribution. The main contents of this dissertation include three sections:1) selection and application of the proper DNA barcoding markers in Heteroptera; 2) the phylogeographic studies of some taxa in Heteroptera; 3) the molecular phylogenetic analysis of the Anthocoridae (sensu stricto).
The main difficulty of DNA barcoding is to find an ideal gene that discriminates any species in the animal kingdom. The previous studies indicated that COI alone would simply not be sufficent to identify problematic taxa in Heteroptera. In many cases, when a single gene region did not assure complete taxonomic resolution, it could still be resolved fully with additional genetic or other data. When considering complementary barcoding markers, the combination of factors of geographic variation and closely related species needs to be taken into account. We approached this issue by studying four species of the genus Physopelta Amyot & Serville,1843 in first section:Ph. cincticollis Stal,1863, Ph. gutta (Burmeister,1834), Ph. guadriguttata Bergroth,1894, Ph. robusta Stal,1863 and Ph. slanbuschii (Fabricius, 1787). We used the sequence of CO, Cyt b and 16S rDNA from dozen provinces of China and multiplicate algorithms to analyze the intra- and interspecific relationships between these closely related species in the same genus, the results indicate that both COI and Cyt b are the proper molecular markers for DNA barcoding of Heteroptera..
The DNA barcoding technique can improve the identification of species and to critically examine the precision of morphological traits commonly used in taxonomy. In the first sention, the sequences of COI gene fragments (1338 bp) of 4 species in the genus Plinachtus Stal,1859 (Heteroptera:Coreidae) from China,Pl.dissimilis Hsiao, 1964, Pl. bicoloripes Scott,1874, Pl. acicularis (Fabricius,1803) and Pl. basalis (Westwood,1842) were amplified. Both the results of the genetic distances of inter-and intraspecies, and phylogenetic analysis based on the COI gene sequences by the methods of NJ, MP and Bayesian, support that Pl. dissimilis and Pl. bicoloripes are same species.
Up to now, there are no phylogeographical studies of Heteroptera in China and related area. The results of the phylogeographic reanalysis of the southern green stink bug Nezara viridula (Linnaeus) (Heteroptera:Pentatomidae) in the world by adding the data from China and Iran shows that the acute rising events in the Qinghai-Tibet plateau in the Pliocene-Pleistocene epoch which may have formed the barriers observed separation of the eastern and western Asia clades. The results also support that the basal splitting of African and non-African subgroup was caused by vicariance events related to the collision of the African microplates Apulia and Arabia with Eurasia, and the resultant arise of successive dispersal barriers (e.g. the Red Sea, the Zagros Mountains). The result of the collision of the Arabian Plate with Eurasia during the Miocene probably isolated the African and Eurasian populations. The combined approachs based on phylogenetic analyses, ancestor distritubtions, estimation of node dates, and dispersal-vicariance analyses indicated that the phylogeographical structuring of Anthocoris nemorum group (Heteroptera: Anthocoridae) has been primarily shaped by the two main periods of the Qinghai-Tibet plateau uplift. The DIVA optimal reconstructions suggest that the A. nemorum group diversified from the Miocene to the Pleistocene from a center of origin in the Hengduan Mountains. The rapid uplift of Mountain ranges associated with the uplift of the entire Qinghai-Tibet plateau may have promoted rapid divergence in the A. nemorum group. Vicariance and dispersal were both essential in shaping the present distribution patterns of the A. nemorum group.
The third section deals with the molecular phylogenetic analysis of the Anthocoridae (sensu stricto). The family Anthocoridae (sensu stricto) is known as an important group of predators for pests, some of them have been introduced as biological control agents. The previous studies on the phylogenetic relationships within Anthocoridae were based on morphological characters without molecular support. In this study, the molecular markers of 16S rDNA,18S rDNA and cytochrome oxidase subunit I (COI) gene with a total length about 4 kb were employed to analyze the phylogenetic relationships of Anthocoridae (sensu stricto). The phylogenetic trees analyzed by two algorithms (ML and Bayesian) showed that Anthocoridae could be divided into two main lineages. Lineage I contains Anthocorini (except Elatophilus), Oriini, Dufouriellini (only Amphiareus) and Scolopini (only Calliodis). Almeidini, Anthocorini (only Elatophilus), Dufouriellini (except Amphiareus), Xylocorini, Scolopini (only Scoloposcelis) and Blaptusterini were included in lineage II. However, the genus Bilia does not in the same clade with other genera of Oriini, Dufouriellu forms a single calde, and without the close relateiongship with another clade formed by the genera of Cardiastethus, Dysepicritus and Physopleurella. The results indicated that Anthocorini, Dufouriellini, Oriini and Scolopini are paraphyletic groups. We need to establish new tribes to include the genera of Elatophilus and Bilia respectively. Dufouriellini should be divided into three tribes to include Dufouriellus, ((Dysepicritus+Cardiastethus)+ Physopleurella) and Amphhiareus respectively. Two sbutribes of Scolopini:Scolopina and Calliodina should be rised to tribe level.
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