Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola)
Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola)
复制标题
DOI：
10.1111/j.1365-3113.2007.00396.x
复制
发表时间：
2008-01-01
影响因子：
4.8
通讯作者：
Aspock, Horst
中科院分区：
文献类型：
--
作者：
Aspock, Ulrike;Aspock, Horst
雄性针翅目第9节（segment 9）包括terterpium、sternite、gonocoxites、gonostyli和gonapophyses，是衡量针翅目、广翅目和脉翅目3个脉翅目的雄性和雌性末端同源性的基准。与生殖器相关的节段在男性中为9、10和11，在女性中为7、8和9。脉翅目全形态学和最近的分子分支分析结果支持姐妹群关系：（1）针翅目和分支Megaloptera +脉翅目，和（2）亚目Nevrorthiformia和所有其他脉翅目。这些研究结果之间的主要差异是在分子分析中Hemerobiiformia亚目的非单系性。广翅目的单系性（这一直受到质疑）进一步证实了迄今为止被忽视的地面模式autapomorphy：在Corydalidae和Sialidae的融合gonocoxites 11复杂的可翻转囊的存在。最近在日本尾龟属（Nevronorthus）中发现的成对生殖器10（参数）复合体表明，日本尾龟属和南方尾龟属的胸骨9的奇怪尖端是生殖器10的骨片与胸骨9的合并，解释为突触形态。在分子生物学研究中，Nevrorthidae，Sisyridae和Osmylidae分支在连续的分裂事件，这一结果是支持的男性生殖器片的分析报告。非凡的平行apomorphies（如过度扩大和修改gonocoxites 10结束在一个线程一样的“penisfilum”）在派生的代表Coniopturidae，Berothidae，Rhachiberothidae和Mantispidae证实dilarid分支的形态分析，并导致我们假设姐妹组的关系Coniopturidae与dilarid分支。重新解释的tignum作为gonocoxites 11，这意味着以前称为gonarcus的结构代表融合gonocoxites 9。在Hemerobiidae中，相应的骨片因此也被同源为融合生殖器9。在这两个家庭的生殖器的侧翼扩大被解释为一个synapomorphy。过度扩大的gonostyli 11在Polystoechotid分支和Myrandiontiformia支持这两个分支的姐妹群关系。雌性生殖器结构（生殖器和生殖突）的某些系列同源物的出现，例如某些Myraburontidae的第8节中的指状突和扁平的附件，或某些Berothidae的第7节中的疣状突和扁平的圆形片，以及某些Nemopteridae和Coniopteridae中的生殖器8作为假胸骨的存在，可能是特征反转。在Rhachiberothidae和Austroberothella（Berothidae）中的terternost9（pseudogonocoxites）的指状突起要么是独立发育的具有产卵功能的收购物，要么是同源的片骨，可能是epipleurite起源。
Segment 9 of male Raphidioptera, comprising tergite, sternite, gonocoxites, gonostyli and gonapophyses, is a benchmark for homologies in the male and female terminalia of the three Neuropterida orders Raphidioptera, Megaloptera and Neuroptera. The segments relating to genitalia are 9, 10 and 11 in males and 7, 8 and 9 in females. Results from holomorphological and recent molecular cladistic analyses of Neuropterida agree in supporting the sister-group relationships between: (1) the Raphidioptera and the clade Megaloptera + Neuroptera, and (2) the suborder Nevrorthiformia and all other Neuroptera. The main discrepancy between the results of these studies is the nonmonophyly of the suborder Hemerobiiformia in the molecular analysis. The monophyly of the Megaloptera (which has been repeatedly questioned) is further corroborated by a hitherto overlooked ground pattern autapomorphy: the presence of eversible sacs within the complex of the fused gonocoxites 11 in Corydalidae and Sialidae. The recently discovered paired complex of gonocoxites 10 (parameres) in Nipponeurorthus (Nevrorthidae) indicates that the curious apex of sternite 9 of Nevrorthus and Austroneurorthus is the amalgamation of the sclerites of gonocoxites 10 with sternite 9, interpreted as synapomorphic. In the molecular study, the Nevrorthidae, Sisyridae and Osmylidae branch off in consecutive splitting events, a result that is supported by the analysis of male genital sclerites reported here. Extraordinary parallel apomorphies (e.g. excessive enlargement and modification of gonocoxites 10 ending in a thread-like 'penisfilum') in derived representatives of Coniopterygidae, Berothidae, Rhachiberothidae and Mantispidae corroborate the dilarid clade of the morphological analysis and leads us to hypothesize a sister-group relationship of the Coniopterygidae with the dilarid clade. A re-interpretation of the tignum of Chrysopidae as gonocoxites 11 means that the structure previously called the gonarcus represents the fused gonocoxites 9. In Hemerobiidae, the corresponding sclerite is consequently also homologized as fused gonocoxites 9. The enlargement of the lateral wings of the gonocoxites in both families is interpreted as a synapomorphy. Excessive enlargement of gonostyli 11 in the Polystoechotid clade and Myrmeleontiformia supports a sister-group relationship of these two clades. The occurrence of certain serial homologues of female genitalia structures (gonocoxites and gonapophyses), such as the digitiform processus together with the flat appendices in segment 8 of certain Myrmeleontidae, or the wart-like processus together with the flat circular sclerites in segment 7 of certain Berothidae, as well as the presence of gonocoxites 8 as pseudosternites in certain Nemopteridae and Coniopterygidae, are probably character reversals. The digitiform processus of tergite 9 (pseudogonocoxites) in Rhachiberothidae and Austroberothella (Berothidae) are either independently developed acquisitions with a function in oviposition, or are homologous sclerites, possibly of epipleurite origin.