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An Appraisal of the Higher Classification of Cicadas (Hemiptera: Cicadoidea) with Special Reference to the Australian Fauna

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An Appraisal of the Higher Classification of Cicadas (Hemiptera: Cicadoidea) with Special Reference to the Australian Fauna © Copyright Australian Museum, 2005 Records of the Australian Museum (2005) Vol. 57: 375–446. ISSN 0067-1975 An Appraisal of the Higher Classification of Cicadas (Hemiptera: Cicadoidea) with Special Reference to the Australian Fauna M.S. MOULDS Australian Museum, 6 College Street, Sydney NSW 2010, Australia [email protected] ABSTRACT. The history of cicada family classification is reviewed and the current status of all previously proposed families and subfamilies summarized. All tribal rankings associated with the Australian fauna are similarly documented. A cladistic analysis of generic relationships has been used to test the validity of currently held views on family and subfamily groupings. The analysis has been based upon an exhaustive study of nymphal and adult morphology, including both external and internal adult structures, and the first comparative study of male and female internal reproductive systems is included. Only two families are justified, the Tettigarctidae and Cicadidae. The latter are here considered to comprise three subfamilies, the Cicadinae, Cicadettinae n.stat. (= Tibicininae auct.) and the Tettigadinae (encompassing the Tibicinini, Platypediidae and Tettigadidae). Of particular note is the transfer of Tibicina Amyot, the type genus of the subfamily Tibicininae, to the subfamily Tettigadinae. The subfamily Plautillinae (containing only the genus Plautilla) is now placed at tribal rank within the Cicadinae. The subtribe Ydiellaria is raised to tribal rank. The American genus Magicicada Davis, previously of the tribe Tibicinini, now falls within the Taphurini. Three new tribes are recognized within the Australian fauna, the Tamasini n.tribe to accommodate Tamasa Distant and Parnkalla Distant, Jassopsaltriini n.tribe to accommodate Jassopsaltria Ashton and Burbungini n.tribe to accommodate Burbunga Distant. Additional tribal changes occur within the Australian fauna: Anapsaltoda Ashton, Arenopsaltria Ashton, Henicopsaltria Stål, Neopsaltoda Distant and Psaltoda Stål are transferred from Cyclochilini to Cryptotympanini leaving Cyclochilini monotypic; Chrysocicada Boulard is transferred from Parnisini to Taphurini; Diemeniana Distant, Gudanga Distant and Quintilia infans (Walker) (representing a new genus) from Parnisini to Cicadettini; and Marteena Moulds and Abricta borealis (Goding & Froggatt) (the latter also representing a new genus) from Taphurini to Cicadettini. A new subtribe is proposed in the tribe Taphurini, Tryellina n.subtribe. Diagnostic descriptions are provided for all families, subfamilies, tribes and subtribes treated in this work. Keys to families and subfamilies are presented together with a key to the tribes of Australian Cicadoidea. Terminologies for external and internal morphology of the Cicadoidea are summarized in 30 detailed figures. New structures and features of significant taxonomic importance are identified and named. A stridulatory mechanism found in the Australian genus Cyclochila is described for the first time. MOULDS, M.S., 2005. An appraisal of the higher classification of cicadas (Hemiptera: Cicadoidea) with special reference to the Australian fauna. Records of the Australian Museum 57(3): 375–446. www.amonline.net.au/pdf/publications/1447_complete.pdf 376 Records of the Australian Museum (2005) Vol. 57 Contents Abstract ......................................................................................................... 375 Introduction ......................................................................................................... 377 Terminology ......................................................................................................... 378 Historical reviews .................................................................................................. 384 Historical review of family classification ........................................................ 384 Historical review of tribal classification .......................................................... 386 Overview of family-group concepts, past and present .......................................... 387 Family and subfamily groupings ..................................................................... 387 Cicadidae/Cicadinae Latreille, 1802 .......................................................... 387 Moaninae Boulard, 1976 ............................................................................ 387 Platypediidae/Platyediinae Kato, 1932 ...................................................... 387 Platypleurinae Schmidt, 1918 .................................................................... 388 Plautillidae/Plautillinae Distant, 1905........................................................ 388 Tettigadidae/Tettigadinae Distant, 1905..................................................... 388 Tettigarctidae Distant, 1905 ....................................................................... 388 Tibiceninae Van Duzee, 1916 ..................................................................... 389 Tibicinidae/Tibicininae Distant, 1905 ........................................................ 389 Ydiellinae Boulard, 1973 ........................................................................... 389 Tribal groupings for Australian cicadas ........................................................... 389 Chlorocystini Distant, 1905........................................................................ 390 Cicadettini Buckton, 1889 .......................................................................... 390 Cicadini Latreille, 1802 .............................................................................. 390 Cryptotympanini Handlirsch, 1925 ............................................................ 390 Cyclochilini Distant, 1904.......................................................................... 391 Dundubiini Atkinson, 1886 ........................................................................ 391 Dundubiini, subtribe Cosmopsaltriina Kato, 1932 .................................... 391 Gymnotympanini Boulard, 1979................................................................ 392 Hemidictyini Distant, 1905 ........................................................................ 392 Lyristini Gomez-Menor, 1957 .................................................................... 392 Parnisini Distant, 1905 ............................................................................... 392 Platypleurini Schmidt, 1918 ....................................................................... 392 Prasiini Matsumura, 1917 .......................................................................... 393 Taphurini Distant, 1905 .............................................................................. 393 Thophini Distant, 1904............................................................................... 393 Tibicenini Van Duzee, 1916 ....................................................................... 393 Cicadoidea phylogeny ...................................................................................... 394 Phylogenetic analysis............................................................................................. 394 Taxon representation: groundplan or exemplar? ............................................. 395 Representation of genera.................................................................................. 395 Representation of tribes and higher taxa ......................................................... 396 Choice of outgroup .......................................................................................... 397 Tree searching .................................................................................................. 397 Characters ......................................................................................................... 397 Results and discussion ..................................................................................... 412 Consistency indices .................................................................................... 412 Robustness of clades................................................................................... 412 Some notable character trends ................................................................... 412 Implications for extant families, subfamilies and tribes .................................. 415 On the monophyly of Tettigarctidae .......................................................... 415 On the monophyly and sister group of Cicadidae ..................................... 415 On the monophyly of the subfamily Cicadinae ......................................... 416 On the monophyly of the subfamily Tibicininae ....................................... 416 On the sister group Tettigadidae/Tettigadinae and Platypediinae ............. 416 On Plautilla and the Plautillinae ................................................................ 421 On the monophyly of the Chlorocystini..................................................... 422 On the monophyly of the Cicadettini ......................................................... 422 On the monophyly of the Cryptotympanini, Platypleurini and Thophini . 423 On the monophyly of the Cyclochilini....................................................... 423 On the monophyly of the Dundubiini ........................................................ 423 On the monophyly and sister group of the Parnisini ................................. 424 On the monophyly of the Prasiini .............................................................
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