摘要
蝗总科是直翅目中较大的一个类群，全世界有1万余种，分布于热带、温带的草原和沙漠地区。蝗虫是典型的植食性昆虫，由于有些种类的翅长且个体较为强壮，具有迁飞能力，是重要的农林害虫。但由于其营养成分含量高，因此它也是重要的食用资源昆虫。蝗虫的研究不仅具有一定的理论意义，而且在害虫防治和资源利用上具有重要的现实意义。
特有种是指在一个区域分布而其他区域没有分布的种类，其形成受生物地理历史过程和生态环境的双重影响。我国东北地区特有属种资源丰富，但鲜见有相关的研究报道。本论文首先对东北地区特有属种的名录进行了概括总结，并对其分布也进行了记述。由于特有属种中绝大部分种类属于近缘种，在形态上极其相似不易区分。因此，本论文除了从外部形态学方面对东北地区特有属种的34种蝗虫进行研究之外，又从解剖学、细胞学以及分子生物学方面对这34种蝗虫进行了较详尽的研究，以探讨它们之间的系统发育关系。
1．形态学方面
本部分对采自东北地区的34种特有属种蝗虫的外部形态进行了描述并进行传统分类，在采集过程中得到2新种——鳞翅跃度蝗和狭额跃度蝗，其中鳞翅跃度蝗已发表在Entomologica Fennica上。采用了夏凯龄（1985）等以印象初（1982）系统为基础，做了修改的用于编写的中国蝗总科志系统，把蝗总科分为8个科。研究结果如下。
本论文研究的34种蝗虫隶属于3科9属，分别是：
①斑腿蝗科Catantopidae：无翅蝗属Zubovskia Dov.–Zap；安秃蝗属Anapodisma Dovnar-Zapolskii；翘尾蝗属Primnoa Fischer-Waldheim；秃蝗属Podisma Berthold；玛蝗属Miramella Dovnar-Zapolskii；
②网翅蝗科Arcypteridae：跃度蝗属Podismopsis Zubovski；
③剑角蝗科Acrididae：绿洲蝗属Chrysochraon Fisch；迷蝗属Confusacris Yin et Li；鸣蝗属Mongolotettix Rehn。
2.解剖学方面
本部分实验运用生理解剖和扫描电镜技术对25种特有属种蝗虫的消化道外部形态、前胃以及贲门的内部形态进行了研究，并利用蝗虫消化道前肠、中肠、后肠，以及前肠的嗉囊、前胃、胃盲囊及贲门的各部分长度进行了聚类分析。结果如下。
（1）蝗虫消化道可分为前、中、后肠三个部分，不同科、属、种之间的差别主要是消化道各段长度的不同以及前肠的前胃和贲门的内部显微结构的差异。
（2）对于蝗总科高级分类阶元：不同科间、属间消化道的内部与外部形态特征具有稳定的显著的差异，其可以作为不同科、属间的分类依据。
（3）对于蝗总科低级分类阶元：由于本实验选取的蝗虫种类亲缘关系较近，同属不同种的蝗虫在消化道外部以及内部显微结构存在差异，但差异不明显。但即便如此，消化道的形态特征对同属不同种间的鉴定仍然可以起到辅助作用。
（4）蝗虫消化道的形态结构是其对植物长期选择和适应的结果，换句话说，消化道的形态结构与蝗虫的食性具有紧密的相互关系，对这种相互关系的研究，不仅能够为分析不同种类蝗虫的食性提供基础资料，还对蝗害的防治有帮助。
（5）聚类结果与形态学分类结果基本一致，表明聚类分析方法有一定的参考价值。
3.细胞学方面
本部分试验运用染色体常规压片法对2科6属18种蝗虫的染色体核型进行了研究，利用染色体的相对长度进行了聚类分析，同时制作了其核型模式图。
（1）蝗虫的性别决定机制均为XX♀/X0♂，斑腿蝗科染色体数目为2n(♂)=23或21，均为端部着丝粒染色体；网翅蝗科染色体数目为2n(♂)=17，第一至第三对染色体为亚中部着丝粒染色体，其余都为端部着丝粒染色体。
（2）所研究的斑腿蝗科中的无翅蝗属、翘尾蝗属、秃蝗属、玛蝗属以及安秃蝗属均为丧失飞行能力的属，它们的染色体核型表现出一定的相似性，可能与其在长期进化过程中都适应了相似的外部环境有关。
（3）染色体核型在蝗总科昆虫高级分类阶元中具有十分稳定的差异性，可作为鉴定不同科及属间蝗虫的一个分类依据。
（4）由于同属不同种的蝗虫亲缘关系较近，染色体核型存在一定的共性，但种间也存在染色体组式、染色大小及相对长度不同，性染色体类型不同等差异。研究的属内蝗虫种类越多，这种差异表现得就越明显，因此染色体核型也可作为种间的分类指标。
（5）染色体核型的近似程度在一定程度上与形态分类结果相吻合，二者存在正态相关关系，或者说核型进化与形态进化是平行的。
4.分子生物学方面
本部分实验提取了蝗总科昆虫26种昆虫基因组总DNA。采用2对特异性引物扩增并测定了DNA序列，获得长度为492bp的Cytb基因序列21条，420bp长度的16SrDNA序列23条；使用DNASTAR、DNAMAN、DNAsp5和MEGA5.05软件对DNA序列的碱基组成、氨基酸组成、碱基替换及遗传距离进行了分析；采用邻接法（NJ）、最小进化法（ME）、最大简约法（MP）和贝叶斯法对分别对Cytb和16SrDNA基因序列进行了蝗总科的系统发育关系重建，在此基础上联合2段基因数据集同样利用4种建树方法构建系统发育树。结果如下。
（1）Cytb（492bp）基因序列中，A、T、G、C的平均含量分别为40.3%、32.4%、16.0%、11.3%。A+T平均含量较高，为72.7%，而G+C的平均含量仅为27.3%。其中密码子第一位点A+T平均含量为75.0%，第二位点A+T平均含量为64.2%，第三位点A+T平均含量最高，为78.6%。
16SrDNA（420bp）基因序列中，A、T、G、C的平均含量分别为33.4%、37.9%、17.5%、11.2%。A+T平均含量较高，为71.3%，而G+C的平均含量仅为28.7%。其中密码子第一位点A+T平均含量为72.1%，第二位点A+T平均含量为68.1%，第三位点A+T平均含量最高，为72.9%。
Cytb基因序列与16SrDNA基因序列中的A+T和G+C含量相似。2种线粒体基因片段碱基组成均表现出明显的A+T组成偏向性。
（2）21条蝗总科昆虫Cytb基因序列中，核苷酸替代的转换数与颠换数的比值（R）的平均值为1.29。转换的平均数高于颠换的平均数。密码子不同位点的转换数与颠换数的比值（R）存在差异。密码子第二位点的R值明显高于第一位点和第三位点的R值，密码子第三位点的转换和颠换的频率都比第一位点的和第二位点的高。
23条蝗总科昆虫16SrDNA基因序列的颠换数略大于转换数，转换数与颠换数的比值（R）的平均值为0.9，与Cytb基因之间存在差异。转换以A-G为主（7），颠换以T-A为主（10），这一点与Cytb基因的相似。
（3）用P距离与R值作图发现P距离与R值之间的关系是存在依赖性的，这种距离依赖性的R值是昆虫线粒体DNA的典型特征，即随着P距离的增大，R值基本呈现下降的趋势。
（4）采用4种方法对Cytb与16SrDNA单个基因及联合2个基因构建NJ、MP、ML、贝叶斯树，比较Cytb与16SrDNA单个基因及联合2个基因所建的树，虽然个别属种的聚类结果存在出入，但各建树结果仍存在一定的相似性，并且联合基因建的树的各分支的支持率较单个基因建的树的各分支的支持率显著提高。
（5）单个基因建树与联合建树的结果均支持斑腿蝗科与网翅蝗科为非单系群的结论，该结论与前人的研究结果相一致。
（6）通过本研究，我们推测网翅蝗科跃度蝗属昆虫中，狭翅跃度蝗Podismopsisangustipennis、长须跃度蝗Podismopsis dolichocerca、土门岭跃度蝗Podismopsistumenlingensis与牡丹江跃度蝗Podismopsis mudanjiangensis是同种异名。
通过以上研究结果可以看出，采用扫描电镜技术、染色体常规压片法和DNA测序等现代分类手段对东北地区特有属种部分种类的研究结果基本一致，虽然个别种类在种属上的分类与形态分类略有出入，但大体上与传统的形态分类结果一致。同时，我们还发现应用现代分类手段有助于区分近缘种类。现代分类学方法以其准确、直观、灵敏等优点弥补了传统分类手段的不足，尤其是对近源种的研究方面。但由于传统分类手段具有直观性等优点，也不可将其摒弃。将传统分类与现代分类手段相结合，是未来解决系统发育关系的一个趋势。
Acridoidea is a bigger superfamily in Caelifera of Orthoptera, with more than10000known species in worldwide. Acridoidea is typical phytophagous insects. Several specieswhose tegmina developed and body robusted have strongly migration capacity, so theybecome important pests in agriculture and forest. However, because of high content ofnutrients, they also become important food resource insects. This study on Acridoidea is notonly have theoretical significance but also practical value in pests control, plant protectionand resources usage.
Special species refers species which distribution in one region, and at the same timeother regions do not have. These species are impact by biogeographic history of the processand the ecological environment. Special species are resourceful abundant in Northeast Area ofChina, but revelant reports are rare. In this paper, i first list a cataloge of Acridoidea specialspecies and describe their distribution, which distribute in Northeast Area. Because of most ofthem are closely related species, and are difficult to distinguish by morphological, then icollected34species of Acridoidea which belong to3families,10genera, decribed theirconfigurations separately, and researched them with the methods of anatomy, cytology andmolecular. Their purpose is to discuss the phylogenetic relationships.
1．Morphology
In this part, the configurations of34special genera and species of Acridoidea weredescribed using the traditional classification methods. During the collection process idiscovered two new species (P. angustifronsensis sp. nov. and P. squamopennis Lu, Wang etRen). The classification system that Xia brought up in1985, which devided Acridoidea intoeight families was used. The results are as follows.
The34species of Acridoidea belong to3families,9genera are:
①Family Catantopidae:Genus Zubovskia Dov.–Zap;Genus Anapodisma Dovnar-Zapolskii;Genus Primnoa Fischer-Waldheim;Genus Podisma Berthold;Genus Miramella Dovnar-Zapolskii;
②Family Arcypteridae:Genus Podismopsis Zubovski;
③FamilyAcrididae:Genus Chrysochraon Fisch; Genus Confusacris Yin et Li;Genus Mongolotettix Rehn.
2．Anatomy
In this part, the alimentary canals of25species of Acridoidea were analyzed using themethod of the external anatomy and scanning electron microscope. The relation betweenmorphological characters of alimentary canals was briefly discussed, and the clusteringanalysis of these25species was studied based on their lengths of foregur, midgut, hindgut,crop, proventriculus, gastric cecum and cardia. The results are as follows.
(1) The alimentary canals of grasshoppers include three parts: foregut, midgut andhindgut. The significant differences between different genera exist in the proventriculus andcardia.
(2) For Acridoidea high level: there are some obviously differences in the morphologicalcharacters of alimentary canals between different families and different genera. It can be usedin classification of different families and different genera.
(3) For Acridoidea low level: the selected species which used in this experiment areclosely related species. There some differences in the morphological characters of alimentarycanals among them, but this difference is not obvious. Even though, the morphologicalcharacteristics still can play a supporting role in the classification of the different species ofthe same genus.
(4) The morphological characters of alimentary canals of grasshoppers are the result oflong-term selection and adaptation with plants. In another word, there is closely relationshipbetween morphological characters of alimentary canals and feeding habits. These interactionrelationship not only to provide basic information to analysis the feeding habits of differentkinds of grasshoppers, but also practical value in pests control.
(5) The results of clustering analysis are consistent with those of morphologicalclassification．It showed that clustering analysis method has some reference value.
3．Cytology
In this part, the chromosomal conventional karyotypes of18species of grasshopperswere studied, which belong to2families,6genera. At the same time the clustering analysis ofthese species was studied based on their chromosomal relative lengths, and the karyotypespattern graph was made. The results are as follows.
(1) All the species of grasshoppers have the basic sex determining mechanism XX♀/X0♂. The family Catantopidae has the chromosome numbers of2n=23or21, chromosomeare all telocentric chromosomes; the family Arcypteridae has the chromosome numbers of2n=17, the first to the third paris of chromosome are sumbmetacentric chromosome, othersare telocentric chromosomes.
(2) Zubovskia Dov.–Zap, Primnoa Fischer-Waldheim, Podisma Berthold, Miramella Dovnar-Zapolskii and Anapodisma Dovnar-Zapolskii of Catantopidae are withoutflying ability. Their cytology characters are similar. This may relevant that they acclimatizedthemselves to similar environmental conditions during perennial evolution.
(3) There are some obviously differences about the cytological characters betweendifferent families and different genera. It can be used in classification of different families anddifferent genera.
(4) Because of the same genus of different species have closely relationship, theircytological characters have some commonalities. But there are also have some differences ingenome formula, the size of chromosomes and their chromosomal relative lengths, the type ofsex chromosomes. The more species in the same genus which we studies, the moredifferences between species are obviously. Therefore, the chromosomal karyotype could usein different species.
(5) The results of the clustering analysis are consistent with that of anatomy and theseresults are in line with the result of morphological classification. In another word, thekaryotype evolution of grasshoppers is parallel with that of morphology evolution.
4．Molecular Biology
In this part, total genomic DNA was extracted from26species,8genera,3families.21sequences of Cytb (492bp) and23sequences of16SrDNA (420bp) have been acquired byPCR amplification and sequenced using specific two pair primers. The base frequency, aminoacid frequency, base substitution and genetic distance of DNA sequence were analyzed usingsome biosoftware such as DNAMAN, DNAsp5and MEGA5.05software. The phylogeneticrelationships were reconstructed from the Cytb and16SrDNA, and the data was combinedusing PAUP4.0(NJ, MP), PHYMLv2.4.4(ML) and MrBayes3.1.2(BI). The results are asfollows.
(1) In Cytb gene sequences (492bp): the average content of A, T, G, C of Cytb genesequence are40.3%、32.4%、16.0%and11.3%respectively, and the A+T average content is72.7%, the G+C average content is27.3%. The A+T average content of the first site, thesecond site and the third site is75.0%,64.2%and78.6%respectively.
In16SrDNA gene sequences (420bp): The average content of A, T, G, C of Cytb genesequence are33.4%、37.9%、17.5%and11.2%respectively, and the A+T average content is71.3%, the G+C average content is28.7%. The A+T average content of the first site, thesecond site and the third site is72.1%,68.1%and72.9%respectively.
The content about A+T and G+C are similar in Cytb gene sequences and16SrDNA genesequences. They all show a strong A+T bias.
(2) In21sequences of Cytb, the average transition and transversion ratio (R) is1.29.Average transition is highly more than average transversion. The average transition andtransversion ratio (R) are different in different sites of codon. The transition and transversion ratio (R) of the second site is highly more than that of the first and the third site. Thetransition and transversion of the third site are all highly more than that of the first and thesecond site.
In23sequences of16SrDNA, the transversion is highly more than transition. Theaverage transition and transversion ratio (R) is0.9different from that of Cytb. Transitionfrequently occurs between A and G, and transversion usually occurs between T and A. It isthe same as Cytb.
(3) The relationship between P distance and R value are dependence. This dependence ofR is a typical characteristic of the insect mitochondrial DNA. As the P distance increases, theR value showed downward trend basically.
(4) The phylogenetic trees were inferred from the Cytb,16SrDNA single gene andcombined gene sequences using4methods to construct NJ、MP、ML and BI. Although theclustering results of individual species exist difference, there is still a certain degree. And thesupport rates of the various branches of the trees of combined gene sequences significantlyhigher than that of a single gene
(5) The phylogenetic trees were inferred from the Cytb,16SrDNA single gene andcombined gene sequences are all supported that family Catantopidae and Arcypteridae arenon-monophyly group. The conclusion is the same as the former.
(6) I assumed that Podismopsis angustipennsis, Podismopsis dolichocerca, Podismopsistumenlingensis and Podismopsis mudanjiangensis are the same species of the genus ofPodismopsis.
In the above results we can see that, the results of the above three parts are accordant andconsistent with those of morphological classification. Although individual species clusteringresult are difference, the result is the same as that of traditional classification. At the sametime the modern classification methods are absolutely used in identification of closely relatedspecies. The modern classification bear the accuracy, impersonality and delicacy and play as aremedy for the traditional classification, especially for closely related species. Because of thecharacteridtic of intuition, traditional classification is not given up. The results ofphylogenetics analyses based on traditional classification and modern classification are moreconvinced compared with the sole analyses.
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