New results concerning the morphology of the most ancient dragonflies (Insecta: Odonatoptera) from the Namurian of Hagen-Vorhalle (Germany)

New results concerning the morphology of the most ancient dragonflies (Insecta: Odonatoptera) from the Namurian of Hagen-Vorhalle (Germany)
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DOI:
10.1046/j.1439-0469.2001.00165.x
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发表时间:
2001-12-01
影响因子:
1.9
通讯作者:
Gröning, E
Gröning, E
中科院分区:
生物学2区
文献类型:
--
作者:
Bechly, G;Brauckmann, C;Gröning, E

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“原捐赠型”瓦氏画眉的正模标本(Brauckmann in Brauckmann et al.,1985)和产于德国哈根-沃尔哈勒上石炭统(纳穆里阶B)的巨型“原齿型”Namurotypus sippeli的副模式标本K-13,以及产于德国哈根-沃尔哈勒上石炭统(纳穆里阶B德国)的Erasipteroides cf. Valentini描述。这些新的证据被用于改进蜻蜓目的地面重建和蜻蜓类的系统发育重建。前胸小翼Erasipteroides和没有一个archaeodictyon记录。此外,从Erasipteroides valentini的雌性正模标本中描述了一个非常长且硬化的产卵器,具有生殖角,并提出它不是用于内生而是用于内基质产卵。早期,odonatoids中的前胸小翼的记录,以及它们在化石Palaeodictyoptera和'protorthopteres'中的存在,表明Pterygota的地面平面确实包括三对翅膀。系统发育分析表明,古生代巨型Meganisoptera和“高等”odonatoids(包括。冠群蜻蜓目(Euodonatoptera)是一个单系类群。本文修订了对西氏纳穆罗蒂普斯雄性初级生殖器结构的描述,并提出了新的解释。包括关于其功能的新假设。纳穆罗蒂普斯的雄性有一对阴茎和一对侧旁突,以及一对叶状但仍然分节的生殖足。分节的腿状雄性生殖足被认为是昆虫的一个平面特征,而成对的阴茎被认为是古网翅目、蜉蝣目和蜻蜓目昆虫的一个假定的突触形态。据推测,Namurotypus不直接交配,但保留了外部精囊的古老沉积,就像最初的无翅昆虫一样,S形的雄性尾索可能被放置在雌性头部后面,并用于将雌性拖过精囊,这与一些现存的蜘蛛纲动物(例如Amblypygi)的交配行为非常相似。三个假设的情况下,进化的次级交配在现代蜻蜓目提出。
The holotype specimen of the 'protodonate' Erasipteroides valentini (Brauckmann in Brauckmann et al., 1985) and the paratype specimen K-13 of the giant 'protodonate' Namurotypus sippeli Brauckmann and Zessin, 1989 from the Upper Carboniferous (Namurian B) of Hagen-Vorhalle (Germany) are redescribed, and a new specimen of Erasipteroides cf. valentini is described. The new evidence is used to refine the groundplan reconstruction of Odonatoptera and the reconstruction of odonatoid phylogeny. Prothoracic winglets for Erasipteroides and the absence of an archaedictyon are documented. Furthermore, a very long and sclerotized ovipositor with gonangulum is described from the female holotype specimen of Erasipteroides valentini, and it is proposed that it was not used for endophytic but for endosubstratic oviposition, The record of prothoracic winglets in early, odonatoids, and their presence in fossil Palaeodictyoptera and 'protorthopteres', indicates that the groundplan of Pterygota indeed included three pairs of wings. A phylogenetic analysis suggests that the Palaeozoic giant Meganisoptera and "higher" odonatoids (incl. crowngroup Odonata) together form a monophyletic group which is here named Euodonatoptera. Erasipteroides and the other 'Erasipteridae' are shown to be more closely related to Euodonatoptera than to Eugeropteridae, The description of the male primary genital structures of Namurotypus sippeli is emended and a new interpretation is proposed. including new hypotheses concerning their function. The males of Namurotypus had a paired penis with a pair of lateral parameres, and a pair of leaf-like, but still segmented, gonopods. Segmented leg-like male gonopods are considered as a groundplan character of insects, while a paired penis is regarded as a putative synapomorphy of the palaeopterous insect orders Palaeodictyopteroida, Ephemeroptera, and Odonatoptera. It is proposed that Namurotypus did not mate by direct copulation but retained the archaic deposition of external spermatophores, just like the primarily wingless insects, The sigmoidal male cerci may have been placed behind the female head and used to drag the female over the spermatophore, which is remotely similar to the mating behaviour of some extant arachnids (e.g. Amblypygi). Three hypothetical scenarios regarding the evolution of secondary copulation in modern Odonata are proposed.