RECONSTRUCTION OF HEXAPOD PHYLOGENY BASED ON THE EVOLUTIONARY TRANSITION OF EMBRYONIC MEMBRANES: EMBRYOLOGICAL STUDY ON DIPLURA

六足动物系统发育中最具争议的问题之一是双足动物。在这个研究项目中，我根据我已经发展和证明的“六足动物和相关节肢动物胚胎膜的进化变化和胚胎固有膜和胚胎膜之间的特化”计划，扩展了涉及双翅动物的系统发育讨论。我批判性地研究了多胚胚胎发生，特别提到了胚胎膜的发育和相关现象。结果表明：1)双翅虫的表皮卵包膜来源于胚体和浆膜，属于胚皮角质层；2)羊膜同源物在双翅目发育，羊膜被认为是胸颌目的胚胎自胚性；3)在双胎中，尽管产生了羊膜，但没有形成任何与羊膜褶相当的结构。这些研究结果可以在“六足动物及其亲缘节肢动物胚胎膜的进化变化和胚胎固有膜与胚胎膜之间的特化”的背景下解读为：1)在六足动物中，双足动物胚皮角质层的产生可以被认为是多形的；2)羊膜的产生可以被认为是双颌和外颌的一种突触形态；(3)羊膜褶皱的形成以及羊膜褶皱与羊膜褶皱的形成之间的联系，并不是羊膜的产生，而是羊膜褶皱的形成，而羊膜褶皱的形成与羊膜褶皱的形成之间的联系，恰恰是异齿龙的自异型。因此，提出了一个系统发育公式为：六足纲=弹尾纲+（双足纲+外齿纲）（=始祖纲+双齿纲（= Zygentoma +翼门））)。这需要对目前公认的六足动物系统发育进行重大修订，即：六足动物=舌颌目（=弹尾目+双翅目）+外颚目（=始祖动物+双齿目动物（= Zygentoma +翼目动物）），拒绝舌颌目的单系地位。这也证实了我所争论的关于“六足动物及其亲缘节肢动物的胚胎膜的进化变化和胚胎固有膜与胚胎膜之间的特化”的方案不需要修改，即使甲壳类或多足动物与六足动物表现出姐妹关系。少

One of the most controversial problems in hexapod phylogeny concerns the Diplura. In this research project, I extend a phylogenetic discussion involving Diplura, in the light of the scheme 'the evolutionary changes of embryonic membranes and the specialization between the embryo proper and embryonic membranes in hexapods and allied arthropods' that I have developed and demonstrated.I critically examined the dipluran embryogenesis with special reference to the development of embryonic membranes and related phenomena. The following results were deduced : 1) the cuticular egg envelope of Diplura is derived both from the embryo proper and serosa, to be categorized in the blastoderm cuticle ; 2) a homologue of the amnion, which has been regarded as an embryological autapomorphy of Ectognatha, develops in Diplura ; and 3) any structures comparable to the amnioserosal fold are not formed, in spite of the production of amnion, in Diplura.These results obtained can be read, in the context of 't … More he evolutionary changes of embryonic membranes and the specialization between the embryo proper and embryonic membranes in hexapods and allied arthropods', as follows : 1) the production of blastoderm cuticle in Diplura is to be recognized as being plesiomorphic within hexapods ; 2) the production of amnion can be designated as a synapomorphy of Diplura and Ectognatha ; and 3) it is not the production of amnion but the formation of amnioserosal folds as well as the linkage of amniotic production to the formation of fold, that should be nominated as the autapomorphy of Ectognatha, were precisely Dicondylia. Consequently, proposed is a phylogeny formulated as : Hexapoda = Collembola + (Diplura + Ectognatha (= Archaeognatha + Dicondylia (= Zygentoma + Pterygota))). This requires a substantial revision to the currently accepted hexapod phylogeny formulated as : Hexapoda = Entognatha (= Collembola + Diplura) + Ectognatha (= Archaeognatha + Dicondylia (= Zygentoma + Pterygota)), rejecting a monophyletic status of Entognatha. It is verified as well that the scheme I have argued concerning 'the evolutionary changes of embryonic membranes and the specialization between the embryo proper and embryonic membranes in hexapods and allied arthropods' does not need to be revised, even if either Crustacea or Myriapoda showing the sister relationship to Hexapoda. Less