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Download Full Article 2.0MB .Pdf File Memoirs of the Museum ol'Victoria 1 — 56< I );9 1 23 (1997) 28 February 1997 https://doi.org/10.24199/j.mmv.1997.56.05 PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN GENERA OF CHLOROCYSTINI (INSECTA: HOMOPTERA: TIBICINIDAE) A.J. De Boer lnstituut voor Systematiek en Populatie Biologic (Zoologisch Museum). University of Amsterdam. PO Box 94766, 1090 GT Amsterdam. The Netherlands Abstract Boer. A.J. de. 1997. Phylogeny and biogeography of Australian genera of Chlorocystini' (Insecta: Homoptera: Tibicinidae). Memoirs of Museum vf Victoria 56(1): 91-123. Six cicada genera belonging to the Chlorocystini are endemic to Australia. These (Chlor- ocysta Westwood, 1851; Cystopsaltha Goding and Froggatt, 1904; Cyslosoma Westwood. 1 842; Glaueopsallria Goding and Froggatt, 1 904; Owra Ashton, 1912; and Venustria Goding and Froggatt, 1904) can be divided into three monophyletic groups. The phylogeny and biogeography of these groups is discussed, descriptions are given of the groups and all species concerned. A key to males is presented for all species of Chlorocystini in Australia. Introduction relationships of and between Australian genera are discussed separately. A computer analysis of The "Baeluria and related genera complex" was the distribution of shared characters of all 148 denned as a supposedly monophyletic group for species of the Chlorocystini (sensu stricto) has which aedeagal characters are regarded synapo- shown that the Australian genera can be subdiv- morphic (De Boer, 1 990). Recently this complex ided into three groups: was identified as the tribe Chlorocystini (sensu 1. Cystopsaltria and Cyslosoma. as mono- stricto) (De Boer, 1995d) comprising about 150 phyletic group; species attributed to 14 genera. Most species 2. Chlowcysta, Glaueopsallria and Owra, as occur in New Guinea but the distribution of the monophyletic group: and tribe includes Maluku and Timor, the Bismarck 3. Venustria. Archipelago, Solomon Islands, Vanuatu, The latter takes a somewhat isolated position, Samoa, Tonga and parts of northern and eastern but is presumably closely related to Gymnotym- Australia. pana Stal, 1861 and shares several characters Of the 13 species in Australia two (Thaumas- with the Australian species (G. rufa and G. var- topsaltria globosa Distant, 1897 and Guineap- icolor) (De Boer, 1 995a; 1 995d). saltria Jlava (Goding and Froggatt, 1904)) also The results of phylogenetic analysis (De Boer, occur in New Guinea. All others are endemic to 1 995d) are summarised and relationships within Australia and, apart from Gymnotympana rufa and between the Australian groups is treated in (Ashton, 1914) and Gymnotympana varicolor more detail. The groups are diagnosed and all (Distant, 1907), belong to endemic Australian species are described. A key to males of all genera. Australian species of the Chlorocystini is The present publication forms part of a phy- presented. logenetic and biogeographic study of the Chlor- ocystini stricto) and deals with the (sensu Methods endemic Australian genera: Chlowcysta West- is wood, 1 85 1 ; Cystopsaltria Goding and Froggatt, The material examined for this study pre- 1 904; Cyslosoma Westwood, 1 842; Glaueopsal- served in the following collections: BMNH, lria Goding and Froggatt, 1904; Owra Ashton, Natural History Museum (formerly: British 1912; and Venustria Goding and Froggatt, 1 904. Museum (Natural History)). London; BPBM, Cyslosoma has two species, Chlowcysta three Bernice P. Bishop Museum. Honolulu; CSIRO, and the others are monotypic. Commonwealth Scientific and Industrial All nine species were recently discussed Research Organisation, Canberra; DEI,, Entomologisches Institut, Eber- (Moulds, 1 990) but descriptions and drawings of Deutsches male genitalia are given for the first time here. swalde; IZW. Polska Akademia Nauk. Instytut The redescriptions recount the characters used Zoologii, Warszavva; KBIN, Koninklijk Belgisch in a phylogenetic reconstruction of the Chloro- lnstituut voor Natuurwetenschappen, Brussels; cystini as a whole (De Boer, 1995d). Here, the Moul, personal collection Mr M.S. Moulds, 91 42 A. .1. HE BOER medial fis- Sydney; RMNH. Nationaal Natuurhistorisch vaulted pronotum without distinct border along the Museum (formerly Rijksmuseum van Natuur- sure (4) and a distinct hyaline Other genera have a lijke Historie). Leiden; SEM, Snow Entomologi- hind margin of tegmen (5). pronotum with a cal Museum, Lawrence, Kansas; SMD, Staat- somewhat wrinkled head and fissure on the pronotum liehes Museum fiir Tierkunde, Dresden; SMF, fairly distinct medial Natur Museum und Forschungs Institut "Scn- and a very narrow border along the hind margin kenberg", Frankfurt am Main; SMN, Staatliches of tegmen; these character states also occur in Prasiini, the presumed sister Museum fiir Naturkunde, Stuttgart; ZIM, Zool- most species of the ogisches Institut und Zoologisches Museum, group of the tribe. Hamburg; ZMA, Instituut voor Systematiek Baeturia en Populatie Biologic (Zoologisch Museum), Scottotympana Amsterdam; ZMB, Institut fiir Spezielle Zoo- logie und Zoologisches Museum der Humboldt- Gymnotympana Universitiit, Berlin; and ZMH, Zoological Mu- Venustria seum of the University of Helsinki, Helsinki. Some of the terms used in the descriptions are Chlorocysta 3 4 5 Glaucopsaltrla explained in figs 5 and 13. To examine the male Owra genitalia the pygofer was pulled out after over- Guineapsaltria night softening, with a sharp needle inserted between the pygofer and the 8th abdominal seg- c Papuapsaltria acdeagus was pulled out at the same ment. The Aedeastria time by inserting the needle between the Thaumastopsaltria claspers. Descriptions were made from dried museum material. This drying often affects r^ Mirabilopsaltria colour. Bright green becomes yellowish brown, ' Cystopsaltria but colour marks like blackish stripes and spots Cystosoma remain intact. Measurements arc based on all , available specimens. Only some of the most • Prasiini important and the most recent systematic litera- Figure 1. Tentative cladogram of the Chlorocystini. species is cited. ture concerning the genera and Numbers refer to apomorphies discussed in the text. For more complete lists of literature is referred to the catalogues of Metcalf (1963) and Duffels Another subdivision can be made based on male and Van der Laan (1985). operculum size. Venustria forms a monophyletic group with Baeturia, Gymnotympana and Scot- Phylogeny totympana. These genera share a fairly large In the following discussion figures in parenth- operculum as apomorphy (7) in which the eses refer to apomorphies in the cladogram (fig. medial margin lies medial to the meracanthus. 1 ). The Chlorocystini (sensu stricto) form a In other genera of Chlorocystini, in Prasiini and monophylctic group for which an S- curved aed- in Muda the medial margin of the operculum lies eagus with winged lateral crests is the supposed generally lateral to the meracanthus. apomorphy (1). The phylogenetic relationships Several characters of Venustria are also found between the 148 described species of this tribe in Scottotympana and many species of Gymno- were analysed with the aid of the program PAUP tympana and indicate a close relationship with (Swofford, 1993) using a data matrix of 154 these two genera: characters and 409 character states (Dc Boer, 1. tegmina with reddish venation and a broad 1 995d). The tribe Prasiini was used as the sister hyaline border along the hind margin; group and the genus Muda as outgroup. The 2. a long proximal spine on the fore femur, often result of this analysis showed that on a generic longer than the distance to middle spine; level the support of some of the proposed phy- and logenetic relationships is very weak. 3. a very short meracanthus. Two major subdivisions can be made. Chlor- However, Baeturia and Scottotympana presum- ocysta, Glaucopsaitria. Owra and Venustria can ably form a monophyletic group; claspers which be grouped with Baeturia, Guineapsaltria, Gym- are not fused at the base arc a presumed synap- notympana, Papuapsaltria and Scottotympana omorphy for these two genera (8). V. superba based on the following synapomorphics: proxi- shares a very similarly shaped basal part of the mal spine of fore femur erect (3), a smoothly operculum with Gymnotympana rufa and PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 93 G. vahcolor while the female of V. superba shares of Cystosoma and most species of Thaumastop- a similar thorn-shaped protuberance on the saltria and Mirabilopsaltria have a narrow tym- pygofer with G. rufa. (De Boer, 1995a: 1995d). bal cavity; the tergite part between the auditory The last two characters suggest that Venustha capsule and the 2nd sternitc is very short. This should be included in and be synonymiscd with character is presumably apomorphic for Cystos- Gymnotympana but no unambiguous synapo- oma, Cystopsaltria, Mirabilopsaltria and Thau- morphy for Venustria and Gymnotympana mastopsaltria together but then lost in some of together has been found. The phylogenetic the species (2). It is not clear whether or not Aed- analysis showed that Venustria as the sister eastria should also be included in this group; group of Gymnotympana is the most parsimoni- though Aedeastria has a wide tymbal cavity it ous solution (De Boer, 1995d). shares distinct diverging fissures on the vertex as The phylogenetic relationships between a possible synapomorphy with Mirabilopsaltria Chlorocysta, Glaucopsaltria and Owra and the and Thaumastopsaltria. The erect fore femoral group formed by Baeluria, Guineapsaltria, spine of
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