中国蝗总科昆虫系统发育和支序生物地理学研究
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摘要
1.支序生物地理学的研究概况和对中国蝗总科昆虫进行支序系统学和支序生物地理
学研究的意义和必要性
支序生物地理学(Cladistic Biogeography)是在支序系统学(Cladistics)理论
的基础上发展起来的一种历史生物地理学(Historical Biogeography)研究方法。由
于该方法是在研究现在生物物种系统发育关系的基础上来分析生物地理分布的演化
过程，重建生物地理历史过程。它是基于严格的逻辑和计算过程，所以越来越为人
们所接受和使用。它与现在的生物地理区划方法的本质区别在于它对生物地理区域
关系的研究是基于生物的系统发育关系，而不是区域间生物分布种类和格局的异同。
从60年代产生支序系统学的思想产生到现在，支序生物地理学的方法随着他的
发展而发展，经过了以下几种方法的进化：①累进原则(The progression rule)、
②隔离分化生物地理学(Vicariance biogeography)Rosen 1976年的方法、③删
减区域支序图法（Reduced area cladogram)Platnick and Nelson 1978年的方法、
④祖先种的地图法(Ancestral species maps)(Wiley的方法1950,1981)⑤组
成成分分析(组元分析)Component analysis Humphries(1956)、⑥Brook的简
约分析方法(Brooks parsimony analysis)、⑦量化的组成成分分析
(Quantification of component analysis）和⑧特有性的简约性分析(Parsimony
analysis of Endemicity)。⑤-⑧为目前几种比较成熟的方法，其中⑤和⑥经常结
合在一起使用，并且最常用到。⑦由于要求的系统发育研究较多，应用较少。⑧适
用于系统发育关系不太清楚的类群。
现在支序生物地理学方法已在动物、植物的一些类群得到了应用，尤其是以昆
虫的研究为最多。并且取得了一定的成果，但相对与数量众多的生物门类和种类来
说还远远不够。从研究的地区来看，一般研究的范围都比较大，划分比较粗。在小
范围的地理区划应用的极少。
我国的支序生物地理学研究工作开展较晚，尹文英(1993)关于原尾自夕蚖科
的生物地理学研究是我国首次应用支序生物地理学的方法对昆虫生物地理的研究，
但除累进原则外，并没有应用支序生物地理学的新的方法。《动物分类学报》1998
年的“中国动物地理学研究”专辑是中国支序生物地理学研究的一个突破。该专辑
收录了支序生物地理学方面的论文9篇，约为总论文数的一半。涉及的动物门类主
要是鱼类(4篇)和昆虫(5篇)。其中昆虫类群有同翅目(1篇)、半翅目(3篇)
和鳞翅目(1篇)。就分析的方法来说有：BPA、假说1和假说2、PAE、删减区域支
序图法和扩散-分替理论。半翅目和鳞翅目方面的研究主要采用的是结合系统发育关
系人工进行地理起源分析。相对与众多的昆虫门类来说，这些研究只能算是沧海一
粟，特别是对于昆虫来说，系统的支序生物地理学研究方法只是在沫蝉和蜡蝉总科
的部分分类单元略有应用。从整个世界来看，还没有直翅目昆虫的支序生物地理学
研究方面的论文发表。
我国的蝗虫己知有860多种，约占亚洲种类的2／3。但我国蝗总科的支序系统
学研究工作开展比较晚也比较零散。1992年龚静对我国网翅蝗科的部分属进行了支
序系统学研究，并根据累进原则对网翅蝗科的起源进行了推测。李保平门994）对
中国斑腿蝗科属级单位进行了支序分类，虽然没有得到斑腿蝗科属级系统发育关系，
但成功地将斑腿蝗科分为7个支（稻蝗支、切翅蝗支、凸额蝗支、长腹蝗支、刺胸
蝗支、裸蝗支和黑蝗支人许升全等曾对我国稻蝗属和竹蝗属的染色体性状进行了支
序分析（许升全郑哲民1997，许升全等1997人许升全等1999年在对蝗总科的雌
性下生殖板进行比较研究的基础上，运用支序系统学的方法研究了蝗总科的系统发
育关系。郑哲民等门）依据蝗总科各科形态性状研究了 8科的系统发育关系，
主要讨论了长腹蝗亚科Leptacrinae的系统地位。这些蝗总科的支序系统学研究工
作相对于，根据蝗总科昆虫的系统发育关系进行支序生物地理学研究的需要远远不
够，因此必须研究蝗总科的系统发育关系。
2本文的研究目的
本文的研究目的就是在用支序系统学方法研究中国蝗总科系统发育关系的基础
1！
上，对中国的动物地理区划进行支序生物地理学方面的研究，以便为动物地理区划
提供依据。
3材料和方法
3．l支序系统学
在对中国的蝗总科昆虫进行支序系统学研究时，分别从属间和种间关系的角度
进行了研究。
在进行属问系统发育关系研究时；考虑到我国的谷个蝗总科分类系级夫于一些
类群的系统地位问题有冲突，这些冲突主要在斑腿蝗科、网翅蝗科、斑翅蝗科、褪
角蝗科和剑角蝗科，而癫蝗科在不同的分类系统中都处于比较原始的地位，锥头蝗
科和瘤锥蝗科有时分开有时合并这样的因素，将中国蝗总科的8个科按照系统发盲
关系分成3个大类群：第一类为癫蝗科，以蚌属为外群研究了全部13个属的属间关
系，第H类为锥头蝗科和瘤锥蝗科，以蠢蝗属和波腿蝗属为外?
Cladistic Biogeography is the methods derived from Cladistics to deal with the historical biogeography problems. Under the methods, the biota is analysised with consideraton to the phylogenitic relationships among the organism on the regions. This method is quite different from the general biota methods currently used, not only by the theory but also by the way it works. There are 3 matural methods (assumption 0,1,2) widely accepted, and 2 methods newly setup (quantification of component analysis and PAE).
This paper consisted of two parts: the Cladistics study on the phylogeny of grasshoppers from China, and the Cladistic biogeography study on the grasshoppers from China.
To do the Cladistics study, 8 grasshopper families were treated as 3 groups to study the phylogenv relationships among genera: First group, Pamphagidae (all 13 genera) ; Second group, Chrotogonidae and Prygomorphidae (all 10 genera); third group, Catantopidae, Arcypteridae, Oedipodidae, Gomphoceridae and Acrididae (selected 51 genera). 12 genera are chosen to study the phylogenetic relationships of species, they are: Atractmopha Sauss; Oxya Serv.; Caiyanda Stal; Sinopodisma Chang; Traulict Stal;Bryodema Fieber; Angaracris B.-Bienko; Oedaleus Fieber; Sphingonotus Fieber; Ceracris Walker;Chorthippus Fieber, Acrida L., A new method is adopted to deal with the character states. All the multistate charactors are setted as nonadditative.
To do the Cladistic biogeography study, The Cladogram of the Cladistics studies and distributions of organism were edited into Nexus Format File, and treated by COMPONENT 2.0.
From The Cladistics and Cladistic biogeography studies, we can drawn follow-in Conclusion:
1. Pamphagidae From China is consisted of 2 Cluster: the ancestral cluster includes 4 Genera (Asiotmethis Uv.,Haplotropis Sauss., Pseudottnethis B.-Bienko and Paratmethis Zheng) , the evolving cluster is consisted of 9 genera: (Thrinchus F.-W, Mongolotmethis B.-Bienko, Beybienkia
飞’111
Tzyplenkov,Sinotmethis B.-Bienko, Eoeotmethis Zheng, Eotmethis B.-Bienko, Rhinotmethis Sjostedt, Filchnerella Kamy, Kanotmethis Yin).
2. Chrotogonidae and Piygomorphidae should be merged into one family.
3. The evolutionaiy direction in the grasshopper families: Catantopidae, Arcypteridae, Oedipodidae., Gomphoceridaz and Acrididae is Catantopidae→Arcyptericlae→oedipodidae → Acrididae→Gomphoceridaz.
4. Oxya Serv. from China should be devided into 2 groups; 0. intricata 0. flavefe~4zura and 0. termacingula belong to the ancestral group, 0. intricata is the most ancerstral species in this group and should be considered as a speices, not as a subspeices of 0. liyla. 0. velox, 0. chinensis and 0. adentata forms a prarphyly, and are the evolving species in another group.
5. Sphingonotus Fieber from China should be devided into 5 groups: Group A is consisted of, S. kirgisicus, S. b~fasciatu~ S. halophilus,S. tsaidarnicus and S. Kweideensis. Group B is consisted of turcmenu.s, S. amplofemurus,S. nig~femoratus, S. yenchiensis and S. Rubescens. Group C is consisted of S. Petikocus, S. maculatus, S. halconemi, S. elegans, S. ningsianus, S. coerulipes, S. bey-bienko, S. carinatus and S. yunnaneu. Group D is consisted of S. qinghaiensis, S. eurasius eurasius, S. salinus, S. tenuipennis and S. longipennis. Group E is consisted of S. tsinhingen.sis, S. octofasciatu.c, S. mongolicus, S. savignyi, S. nebulosus nebulosus, S. zadaensis and S. obsurarus iatiJsim us. The evolutionary direction is A-)~B-~ C-) D-*E.
6. The study on 40 speices of Chorthippus shows that, except some speices of Altichorthippus, the result support that Chorthz~ppus from China be devided into 4 subgenus: Megaulacobothrus, Chorthippus, Glyptobothrus and Altichorthippus, The evolutionary direction is Megaulacobothrus
>Glyptobothrus-*Altichorthippus-)Chorthippus.
7. According to the phylogenetic relationship of grasshopper Genera, the Fanua of China can be devided into Palaearctic Real
学研究的意义和必要性
支序生物地理学(Cladistic Biogeography)是在支序系统学(Cladistics)理论
的基础上发展起来的一种历史生物地理学(Historical Biogeography)研究方法。由
于该方法是在研究现在生物物种系统发育关系的基础上来分析生物地理分布的演化
过程，重建生物地理历史过程。它是基于严格的逻辑和计算过程，所以越来越为人
们所接受和使用。它与现在的生物地理区划方法的本质区别在于它对生物地理区域
关系的研究是基于生物的系统发育关系，而不是区域间生物分布种类和格局的异同。
从60年代产生支序系统学的思想产生到现在，支序生物地理学的方法随着他的
发展而发展，经过了以下几种方法的进化：①累进原则(The progression rule)、
②隔离分化生物地理学(Vicariance biogeography)Rosen 1976年的方法、③删
减区域支序图法（Reduced area cladogram)Platnick and Nelson 1978年的方法、
④祖先种的地图法(Ancestral species maps)(Wiley的方法1950,1981)⑤组
成成分分析(组元分析)Component analysis Humphries(1956)、⑥Brook的简
约分析方法(Brooks parsimony analysis)、⑦量化的组成成分分析
(Quantification of component analysis）和⑧特有性的简约性分析(Parsimony
analysis of Endemicity)。⑤-⑧为目前几种比较成熟的方法，其中⑤和⑥经常结
合在一起使用，并且最常用到。⑦由于要求的系统发育研究较多，应用较少。⑧适
用于系统发育关系不太清楚的类群。
现在支序生物地理学方法已在动物、植物的一些类群得到了应用，尤其是以昆
虫的研究为最多。并且取得了一定的成果，但相对与数量众多的生物门类和种类来
说还远远不够。从研究的地区来看，一般研究的范围都比较大，划分比较粗。在小
范围的地理区划应用的极少。
我国的支序生物地理学研究工作开展较晚，尹文英(1993)关于原尾自夕蚖科
的生物地理学研究是我国首次应用支序生物地理学的方法对昆虫生物地理的研究，
但除累进原则外，并没有应用支序生物地理学的新的方法。《动物分类学报》1998
年的“中国动物地理学研究”专辑是中国支序生物地理学研究的一个突破。该专辑
收录了支序生物地理学方面的论文9篇，约为总论文数的一半。涉及的动物门类主
要是鱼类(4篇)和昆虫(5篇)。其中昆虫类群有同翅目(1篇)、半翅目(3篇)
和鳞翅目(1篇)。就分析的方法来说有：BPA、假说1和假说2、PAE、删减区域支
序图法和扩散-分替理论。半翅目和鳞翅目方面的研究主要采用的是结合系统发育关
系人工进行地理起源分析。相对与众多的昆虫门类来说，这些研究只能算是沧海一
粟，特别是对于昆虫来说，系统的支序生物地理学研究方法只是在沫蝉和蜡蝉总科
的部分分类单元略有应用。从整个世界来看，还没有直翅目昆虫的支序生物地理学
研究方面的论文发表。
我国的蝗虫己知有860多种，约占亚洲种类的2／3。但我国蝗总科的支序系统
学研究工作开展比较晚也比较零散。1992年龚静对我国网翅蝗科的部分属进行了支
序系统学研究，并根据累进原则对网翅蝗科的起源进行了推测。李保平门994）对
中国斑腿蝗科属级单位进行了支序分类，虽然没有得到斑腿蝗科属级系统发育关系，
但成功地将斑腿蝗科分为7个支（稻蝗支、切翅蝗支、凸额蝗支、长腹蝗支、刺胸
蝗支、裸蝗支和黑蝗支人许升全等曾对我国稻蝗属和竹蝗属的染色体性状进行了支
序分析（许升全郑哲民1997，许升全等1997人许升全等1999年在对蝗总科的雌
性下生殖板进行比较研究的基础上，运用支序系统学的方法研究了蝗总科的系统发
育关系。郑哲民等门）依据蝗总科各科形态性状研究了 8科的系统发育关系，
主要讨论了长腹蝗亚科Leptacrinae的系统地位。这些蝗总科的支序系统学研究工
作相对于，根据蝗总科昆虫的系统发育关系进行支序生物地理学研究的需要远远不
够，因此必须研究蝗总科的系统发育关系。
2本文的研究目的
本文的研究目的就是在用支序系统学方法研究中国蝗总科系统发育关系的基础
1！
上，对中国的动物地理区划进行支序生物地理学方面的研究，以便为动物地理区划
提供依据。
3材料和方法
3．l支序系统学
在对中国的蝗总科昆虫进行支序系统学研究时，分别从属间和种间关系的角度
进行了研究。
在进行属问系统发育关系研究时；考虑到我国的谷个蝗总科分类系级夫于一些
类群的系统地位问题有冲突，这些冲突主要在斑腿蝗科、网翅蝗科、斑翅蝗科、褪
角蝗科和剑角蝗科，而癫蝗科在不同的分类系统中都处于比较原始的地位，锥头蝗
科和瘤锥蝗科有时分开有时合并这样的因素，将中国蝗总科的8个科按照系统发盲
关系分成3个大类群：第一类为癫蝗科，以蚌属为外群研究了全部13个属的属间关
系，第H类为锥头蝗科和瘤锥蝗科，以蠢蝗属和波腿蝗属为外?
Cladistic Biogeography is the methods derived from Cladistics to deal with the historical biogeography problems. Under the methods, the biota is analysised with consideraton to the phylogenitic relationships among the organism on the regions. This method is quite different from the general biota methods currently used, not only by the theory but also by the way it works. There are 3 matural methods (assumption 0,1,2) widely accepted, and 2 methods newly setup (quantification of component analysis and PAE).
This paper consisted of two parts: the Cladistics study on the phylogeny of grasshoppers from China, and the Cladistic biogeography study on the grasshoppers from China.
To do the Cladistics study, 8 grasshopper families were treated as 3 groups to study the phylogenv relationships among genera: First group, Pamphagidae (all 13 genera) ; Second group, Chrotogonidae and Prygomorphidae (all 10 genera); third group, Catantopidae, Arcypteridae, Oedipodidae, Gomphoceridae and Acrididae (selected 51 genera). 12 genera are chosen to study the phylogenetic relationships of species, they are: Atractmopha Sauss; Oxya Serv.; Caiyanda Stal; Sinopodisma Chang; Traulict Stal;Bryodema Fieber; Angaracris B.-Bienko; Oedaleus Fieber; Sphingonotus Fieber; Ceracris Walker;Chorthippus Fieber, Acrida L., A new method is adopted to deal with the character states. All the multistate charactors are setted as nonadditative.
To do the Cladistic biogeography study, The Cladogram of the Cladistics studies and distributions of organism were edited into Nexus Format File, and treated by COMPONENT 2.0.
From The Cladistics and Cladistic biogeography studies, we can drawn follow-in Conclusion:
1. Pamphagidae From China is consisted of 2 Cluster: the ancestral cluster includes 4 Genera (Asiotmethis Uv.,Haplotropis Sauss., Pseudottnethis B.-Bienko and Paratmethis Zheng) , the evolving cluster is consisted of 9 genera: (Thrinchus F.-W, Mongolotmethis B.-Bienko, Beybienkia
飞’111
Tzyplenkov,Sinotmethis B.-Bienko, Eoeotmethis Zheng, Eotmethis B.-Bienko, Rhinotmethis Sjostedt, Filchnerella Kamy, Kanotmethis Yin).
2. Chrotogonidae and Piygomorphidae should be merged into one family.
3. The evolutionaiy direction in the grasshopper families: Catantopidae, Arcypteridae, Oedipodidae., Gomphoceridaz and Acrididae is Catantopidae→Arcyptericlae→oedipodidae → Acrididae→Gomphoceridaz.
4. Oxya Serv. from China should be devided into 2 groups; 0. intricata 0. flavefe~4zura and 0. termacingula belong to the ancestral group, 0. intricata is the most ancerstral species in this group and should be considered as a speices, not as a subspeices of 0. liyla. 0. velox, 0. chinensis and 0. adentata forms a prarphyly, and are the evolving species in another group.
5. Sphingonotus Fieber from China should be devided into 5 groups: Group A is consisted of, S. kirgisicus, S. b~fasciatu~ S. halophilus,S. tsaidarnicus and S. Kweideensis. Group B is consisted of turcmenu.s, S. amplofemurus,S. nig~femoratus, S. yenchiensis and S. Rubescens. Group C is consisted of S. Petikocus, S. maculatus, S. halconemi, S. elegans, S. ningsianus, S. coerulipes, S. bey-bienko, S. carinatus and S. yunnaneu. Group D is consisted of S. qinghaiensis, S. eurasius eurasius, S. salinus, S. tenuipennis and S. longipennis. Group E is consisted of S. tsinhingen.sis, S. octofasciatu.c, S. mongolicus, S. savignyi, S. nebulosus nebulosus, S. zadaensis and S. obsurarus iatiJsim us. The evolutionary direction is A-)~B-~ C-) D-*E.
6. The study on 40 speices of Chorthippus shows that, except some speices of Altichorthippus, the result support that Chorthz~ppus from China be devided into 4 subgenus: Megaulacobothrus, Chorthippus, Glyptobothrus and Altichorthippus, The evolutionary direction is Megaulacobothrus
>Glyptobothrus-*Altichorthippus-)Chorthippus.
7. According to the phylogenetic relationship of grasshopper Genera, the Fanua of China can be devided into Palaearctic Real
引文
马世骏1959中国昆虫生态地理概述．北京：科学出版社
马恩波，1993斑腿蝗科10属NOR定位及其细胞分类学意义，动物学研究，14(3)：271-276
尹文英，谢容栋，张骏，1994夕蚖宗生物地理学的研究(原尾目，夕蚖科)，昆虫学研究集于小VoL11，p73-80．
方三阳1993中国森林害虫生态地理分布哈尔滨：东北林业大学出版社
牛瑶，郑哲民1994竹蝗属部分种类的染色体分类研究昆虫学研究，陕西师范大学出版社出版。P136-141
王保海，袁维红等1992西藏昆虫区系及演化．郑州：河南科学技术出版社
甘肃省草原总站1997甘肃草地蝗虫．甘肃科学技术出版社．
甘肃省蝗虫调查协作组1985甘肃蝗虫图志．甘肃科学出版社．
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