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Old method not old-fashioned: parallelism between wing venation and wing-pad tracheation of cockroaches and a revision of terminology


The Comstock–Needham system, the foundation of main vein homology of insects, is grounded in the comparison between wing venation and wing-pad tracheation, particularly that of Plecoptera and Blattodea. However, the trachea-based approach has come into great dispute in recent years. Meanwhile, the terminology used to describe the wing venation of cockroaches requires a revision according to homology and the practical need. These motivated us to examine the wings of cockroaches, comparing the tracheation of nymphs with the tracheation and venation of adults. We find a correspondence between the tracheation and the main vein venation; therefore the trachea-based approach is effective in cockroaches. It is supported that Neopteran insects have six homologous main veins, namely, Sc, R, M, Cu, Pcu and V. We interpret the wing morphology of cockroaches in detail, including veins, furrows, folds and apical folding. RA and RP in hindwing, ScP, CuA and CuP in tegmen and hindwing, are recognized. Hindwing R + M is the only normal venal fusion in cockroaches. In addition, we propose several descriptive terms: cmv, V[1], V[s], pseudostem and characteristic posterior branch. The venation of 41 species (representing 41 genera), and the wing-pad tracheation of four of them including Diploptera punctata, which has unique apical folding, are illustrated.

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Most of the materials examined were collected by Dr. Shun-Hua Gui, Dr. Yan Shi, Dr. Jian-Yue Qiu, Mr. Zhi-Wei Qiu, Mr. Lu Qiu, Mr. Hong-Guang Liu and XRL. We thank our colleagues Ms. Li-Li Wang and Mr. Lu Qiu for drafting several figures in Online Resource 1. We also want to thank Mr. Tie-Xiong Zhao and Dr. Gui-Qiang Huang for collecting and donating the specimens of Spelaeoblatta sp. and Catara rugosicollis, respectively. The reviewers’ constructive comments improved the manuscript. Dr. Han-Qiang Wang (Institute of Plant Physiology and Ecology, SIBS, CAS) made one of the references accessible. This study is financially supported by the National Natural Sciences Foundation of China (nos. 31093430 and 31472026).

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Correspondence to Zong-Qing Wang.

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Li, X., Zheng, Y., Wang, C. et al. Old method not old-fashioned: parallelism between wing venation and wing-pad tracheation of cockroaches and a revision of terminology. Zoomorphology 137, 519–533 (2018).

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  • Blattodea
  • Dictyoptera
  • Polyneoptera
  • Trachea
  • Vein
  • Wing folding