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 Aedeastria, however, suggests a mono- Gymnotympana, Papuap.saltria, Scottotympana phyletic origin with the group containing Baelu- and Venustria are not clear, as indicated by the ria, Chlorocysta, Glaucopsaltria, Gymnotym- polytomy in fig. 1 . Guineapsaltria and Papuap- pana, Guineapsaltria. Owra, Papuapsaltria. saltria are possibly sister groups, based on a very Scottotympana and Venustria (see fig. 1 ques- small and rounded male operculum in many of tion marks) their species (6). Chlorocysta, Glaucopsaltria The coloured tegmina of Cystosoma and Cys- and Owra share a very broad vertex with the topsaltria which are also found in Mirabilopsal- small ocelli wide apart with most species of triaand Thaumastopsaltria might also indicatea Gymnotympana and all but one species of Gui- relationship between these genera though this neapsaltria; the distance between the lateral character is also found in some genera of ocelli often exceeds three times the width of the Prasiini. frontal ocellus. The fairly short meracanthus of Biogeography the species of Chlorocysta, Glaucopsaltria and Owra and the vague colour pattern found on the Biogeographic patterns of Chlorocystini and a head and thorax of two of the Chlorocysta similarly distributed group of cicadas, the sub- species, might indicate a close relationship with tribe Cosmopsaltriaria, were recently compared Gymnotympana, Scottotympana and Venustria. and analysed (De Boer, 1995c). It appears that The phylogenetic analysis showed that a recon- most of the species of these groups are found on struction with Chlorocysta, Glaucopsaltria and terranes derived from a historic oceanic island Owra as sister group of Baeluria, Gymnotym- arc, which developed since about 40 million pana, Scottotympana and Venustria is slightly years ago as a result of subduction of the former more parsimonious than the option with Gui- Tethys Sea under the Pacific tectonic plate (De neapsaltria and Papuapsaltria as sister group of Boer, 1995c). This arc is known as the Outer that group. On a tree with a total length of 906 Melanesian Arc (Hamilton 1979; Holloway, steps the latter option is only one step longer (De 1979, 1984; Duffels, 1986; Rangin etal., 1990a,

1 a!., 1 99 ). to the continuous Boer, 1995d). 990b; Daly et 1 Due Cystosoma and Cystopsaltria share a strongly northward movement of Australia and the west- furrowed pronotum and a very narrow border ward movement of the Pacific this volcanic along the hind margin of tegmen with Aedeas- island arc broke up and fragments collided with they tria, Mirabilopsaltria and Thaumastopsaltria, a the northern craton of Australia where now strongly bent proximal spine on the fore femur form part of New Guinea (Pigram and Davies,

with Mirabilopsaltria and Thaumastopsaltria 1 987). It is supposed that the Outer Melanesian and an angularly protruding postclypeus with Arc, before breaking up, formed an important Thaumastopsaltria. All these characters are also route of dispersal for south-east Asian biota widely distributed in Prasiini and are presumed invading the Pacific (Duffels 1986, Duffels and plesiomorphic. Furthermore, Cystosoma and De Boer, 1990, De Boer, 1995c). Cystopsaltria have small male opercula which The distribution of Chlorocystini over the arc are regarded plesiomorphic; the opercula of Cys- terranes. the fact that their presumed sister tosoma schultzi and Cystopsaltria immaculata. group (the oriental Prasiini as defined by De

1 in Sulawesi that though relatively small, do slightly extend medi- Jong, 985) is most speciose and these groups combined (the ally of the meracanthus but the operculum of the sister group of Cystosoma saundersii does not. The two species genus Muda) occurs in south-east Asia and the )

94 A. J. DE BOER greater Sunda Islands, suggest that the ancestor Venustha, an endemic Australian genus that of that tribe also used the island arc as route of should possibly be included in Gymnotympana dispersal. In fact, the distribution and phylogeny (see discussion above: De Boer, 1995a; 1995d). of these cicadas reflect the geotectonic history in is also restricted to the Cape York Peninsula. that the main vicariant speciation events in The two groups of endemic Australian genera the cladograms correspond with presumed have a much wider distribution, reaching farther sequences of fragmentation of the island arc southward. These have a strikingly similar dis- (Dc Boer, 1995c). This means that the ancestor tribution in northern and eastern Qld, including of Chlorocystini originates from Asia and that the Cape York Peninsula and north-eastern all its Australian species must have invaded Aus- NSW. tralia by dispersal presumably through New The distribution patterns of the species of Guinea (De Boer. 1995c). endemic and non-endemic Australian genera Two patterns of distribution of Australian and their relative positions in the cladogram

Chlorocystini can be recognized. It appears that suggest different ages. There must have been at species not belonging to endemic genera but least two periods of dispersal from New Guinea belonging to genera found in New Guinea as well to Australia. For the non- Australian genera a (Guineapsalttia Jlava, Gymnotympana rufa, dispersal during Pliocene-Pleistocene low sea-

Gymnotympana varicolor and Thaumastopsal- level stands has been proposed (De Boer, 1 992b; tria globosa) are restricted to the northern and 1995c). The two groups of endemic Australian eastern parts of the Cape York Peninsula; the genera branch off lower down in the cladogram only exception ( T. globosa) is also recorded from and must therefore be older. Their ancestors Groote Eylandt, NT (Moulds, 1990). This same must have reached Australia earlier, possibly pattern is found in several species that do not following the first collision between Australia belong to Chlorocvstini: Diceropvga subapicalis and a fragment of the Outer Melanesian Arc, (Walker. 1868) (see Duffels, 1977), Lembeja which is dated at about 25 mya (Pigram and paradoxa (Karsch, 1 890) and Lembeja vittieoilis Davies, 1987; De Boer, 1995c). (Ashton, 1912) (see De Jong, 1982) belong to For maps of distribution of the Australian non-endemic genera and are restricted to the species described here see Moulds ( 1 990) and De Cape York Peninsula (Moulds, 1990). Boer (1995c).

Key to males

I Tegmen with variable venation, often differing between left and right tegmen of individuals, forming 9 or more apical areas and several subapical areas between apical and ulnar areas 2 Tegmen venation regular, with 8 or 9 apical areas, but without subapical areas 10

2( 1 Tegmen hyaline, with fairly broad hyaline border along hind mar-

gin and broad costal area. Proximal spine of fore femur erect (fig. 20). Postclypeus not swollen ventrally (figs 4, 45). Pronotum smooth, without distinct medial furrow 3 Tegmen opaque or weakly reddish or greenish tinged, with nar- row border along hind margin and narrow costal area. Proximal spine of fore femur strongly bent, adjacent to femur (fig. 60). Postclypeus distinctly swollen ventrally (fig. 70). Pronotum with distinct medial furrow 7

3(2) Aedeagus Z-curved in apical part, proximal part straight (cf. fig. 12). Tymbal with 9 or more ridges 4 Aedeagus S-curved (figs 31. 42). Tymbal with 6-8 ridges ... 6 4(3) Head and thorax without colour markings. Tymbal with 9-10 ridges. Clasper base forming a low collar around base of anal valves, which is sloping proximally of aedeagus (fig. 6). Aedeagus not incised at apex (figs 11, 12) Chloroeysta vilripenni.s FHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 95

Head and thorax with pattern of dark brown spots and stripes. Tymbal with 11-12 ridges. Clasper base forming a high collar around base of anal valves, which is erect proximally of aedeagus (tigs 14, 25). Aedeagus incised at apex (figs 18, 27) 5 5(4) Distomedial corner of operculum not reaching medially of mer- acanthus (fig. 24). Ventral part of postclypeus darkly suffused. Tymbal with 12 ridges, 4 of which do not reach ventral tymbal margin Chlowcysta suffusa — Distomedial part of operculum almost pointed, elongate, reach-

ing medially of meracanthus (fig. 1 5). Ventral part of postclypeus

not darkly suffused. Tymbal with 1 1 ridges, 4 of which do not reach ventral tymbal margin Chlowcysta fumea 6(3) Body length over 28 mm. Abdomen strongly inflated. Seventh tergite wedge- shaped in lateral view, long dorsally and short at ventral margin; pygofer turned to a ventral position. Distal part of operculum longer than basal part and reaching beyond apex of meracanthus (fig. 44). Wing with 6 apical areas Glaucopsaltria viridis Body length under 18 mm. Abdomen weakly inflated. Seventh tergite not wedge- shaped in lateral view; pygofer directed pos- teriad. Distal part of operculum much shorter than basal part and not reaching to apex of meracanthus (fig. 33). Wing with 5 apical areas Owra insignis 7(2) Tegmina rounded at apex, venation not reticulate. Wings with 6 apical areas. Pronotum not forming a sharply edged lateral crest. Head and pronotum not triangle-shaped; anterior margins of postclypeus and vertex lobes not forming a continuous line with lateral margins of pronotum. Cruciform elevation of mesonotum broader than long across its centre. First and second sternites not adjacent. Clasper straight, with broad domed clasper hollow and with laminiform and sharply pointed medial protrusion. Tymbal with 6 ridges Thaumastopsaltria globosa — Tegmina pointed at apex, venation reticulate in distal half. Wings with more than 6 apical areas. Pronotum forming a sharply edged lateral crest. Head and pronotum triangle-shaped; anterior mar- gins of postclypeus and vertex lobes forming an almost continu- ous line with lateral crest of pronotum. Cruciform elevation of mesonotum narrower than long across its centre. First and second sternites adjacent. Clasper hook-shaped, with narrow clasper hol- low in downwardly directed apical part and without medial pro- trusion. Tymbal with 7 or more ridges 8 8(7) Veins CuA and M not fused at corner of basal area (fig. 72). First radial area of tegmen divided by longitudinal vein. Venation of apical and ulnar areas reticulate. Pygofer opening narrow, V- shaped (fig. 78). Aedeagus with dorsal appendages near its apex (figs 85, 86). Tymbal with 10 ridges .Cystopsahria immaculata Veins CuA and M fused at corner of basal area (fig. 81). First radial area of tegmen not divided. Venation of apical areas reticu- late, ulnar areas normal. Pygofer opening broad, U-shaped (fig 57). Aedeagus without dorsal appendages (figs 62. 77). Tymbal with 7 ridges 9 96 A. J. DE BOER

9(8) Body length over 40 mm. Abdomen globularly inflated. Medial margin of operculum not reaching medially of mcracanthus (fig. 68) Cystosoma saundersii — Body length under 30 mm. Abdomen distinctly but not globularly inflated. Medial margin of operculum reaching medially of mcr-

acanthus (fig. 71) Cystosoma schmeltzi

1 0( 1 ) Wing with extremely slender anal fields; 2nd anal field rudimen- tary, not hyaline. Abdomen with reddish ventral band. Tergites sharply folded towards sternites ventrolaterally. Ventral part of pygofer not incurved to its margins; ventral and basal margins of pygofer forming a continuous, sharply edged, ridge 11 — Wing with broad anal fields; 2nd anal field distinct and hyaline. Abdomen without reddish ventral band. Tergites gradually curved towards sternites ventrolaterally. Ventral part of pygofer gradually incurved to its margins; ventral and basal margins of pygofer not forming a sharply edged ridge 12

1 1(10) Tymbal with 5 ridges. Tymbal and lateral part of 2nd tergite sep- arated by broad wedge-shaped cavity. Second tergite strongly curved between auditory capsule and 2nd sternite. Caudodorsal beak erect and conically shaped; its lateral margins curving inwards and fusing. Aedeagus short and almost straight, directed upwards, not reaching beyond ventral margin of pygofer Gymnolympana rufa Tymbal with 4 ridges. Tymbal and lateral part of 2nd tergite sep- arated by narrow gap. Second tergite straight between auditory capsule and 2nd sternite. Caudodorsal beak strongly curved, but not conically shaped, its lateral margins not fusing. Aedeagus long and strongly curved down, reaching beyond ventral margin of pygofer Gymnotympana varicolor 12(10) Body length over 20 mm. Operculum large, covering most of tymbal cavity and reaching medially of meracanthus. Basal part of operculum medially distinctly longer than laterally (fig. 55). Tegmina slightly bronzed, with 9 apical areas. Clasper broad and lobate, with large clasper hollow and strongly curved down, not forming a shaft around aedeagus (fig. 5 1 ). Aedeagus erect, its apex adjacent to base of anal valves. Aedeagus broad, swollen in proxi- mal part and without subapical lobe Venustria superba

— Body length under 1 6 mm. Operculum small, not covering tymbal cavity and not reaching medially of meracanthus. Basal part of operculum oblong. Tegmina clear hyaline, with 8 apical areas. Clasper long, slender and straight, without distinct clasper hol- low, directed posteriad and forming a shaft around aedeagus. Aedeagus directed posteriad between claspers, its apex near apex of clasper. Aedeagus very slender, with distinct subapical lobe. Guineapsaltria jlava

Chlorocysta, Glaucopsaltria and Owra inflated 1.4-2.3 X as long as head and thorax in together, females 0.9- 1 .2 X . Head broad and Description. The species of Chlorocysta and short (fig. 3), 2.4-3.0 X as wide as long and 1.8- Glaucopsaltria are large compared to other 2.2 X as wide as distance between eyes. Postcly- Chlorocystini, body length 20-33 mm; only peus broad and bluntly rounded anteriorly, 2.3- Owra is distinctly smaller, under 16 mm. Body 3.3 X as wide as long. Anterior margin of post- reddish brown or greenish, generally uniformly clypeus continuous with anterior margins of ver- coloured, but abdomen often with reddish seg- tex lobes. Sides of postclypeus with about 8-10 mental hind margins and a row of dark coloured distinct furrows, ending in short rows of short ventrolateral spots. Male abdomen strongly parallel ridges, which form a narrow band along PHYLOGENY AND BIOGEOGRA1MIV OF AUSTRALIAN CICADAS 97

lorum. Postclypeus not swollen vcntrally; part of second tergite hardly longer than lateral anterior margin (lateral view) straight or weakly parts, proximal margin of second tergite weakly convex (fig. 4). Vertex very broad and smooth; convex medially. Lateral parts of 2nd tergite diverging fissures from centre of head to corners weakly swollen at anterior margins and adjacent of postclypeus weakly developed. Vertex lobes to tymbals. Tergite part between auditory cap- sometimes with some weak longitudinal sules and stcrnitc 2 almost straight, forming a wrinkles. Vertex 1.7-2.4 X as wide as long; 1.3- distinct ridge along tymbal cavity. Sternites 1 1.6 as wide X as postclypeus and 1.6-2.4 X as and 2 not adjacent (fig. 1 3 arrow). Auditory cap- wide as eye. Ocelli small and far apart. Distance sules not swollen, hardly elevated relative to between lateral ocelli 2.4-4. 1 X width of frontal connecting bar between abdomen and tymbal. ocellus and 0.8-1.2 X distance between lateral Female abdomen shorter and more robust than ocellus and eye. Pronotum 2.3-2.6 X as wide as that of male, with short and broad pygofer (fig. long and fairly smooth, without medial furrow. 22). Ovipositor sheaths almost reaching to apex Pronotal collar laterally weakly amplified and of sharply pointed caudodorsal beak. Male slightly curving down at anterior margin of lat- pygofer short and rounded, with convex distal eral amplifications. Tcgmina and wings hyaline, and ventral margins. Caudodorsal beak short venation reddish or sometimes ochraceous. Teg- and erect; not curving over basal part of claspers men venation variable with 9-15 apical areas or anal valves and bluntly rounded, truncate, or and a more or less continuous band of subapical rectangular at apex. Lateral lobes of pygofer areas. Costal area hyaline and very distinct, weakly curved inwards with small, weakly widening towards tegmen apex. A distinct hyal- inflated, protuberances. Ventral margins of ine border along hind margin of tegmen (fairly pygofer converging, generally forming a sharp narrow in Owra). Wings with 5, in Glaucopsal- angle at base of pygofer opening. Claspers fairly tria 6, apical areas and a distinctly broader hyal- slender and hook-shaped; sharply curving down ine border than in tegmina. Legs ochraceous and at half-length. Apical part of claspcr with large, unmarked. Fore femur (fig. 20) with row of 3 sharply edged, clasper hollow (in Owra without erect and sharply pointed spines, diminishing in clasper hollow). Claspers weakly diverging length towards tibia. Tymbal with 6-12 parallel towards rounded apices. Basal part of claspcr sclerotized ridges. The most proximal ridges forming a continuous ring-shaped collar around often not reaching ventral margin of tymbal. base of anal valves. Aedcagus S-curvcd (in Short intercalary ridges forming a lateral band at Chlorocysta more Z-curvcd in apical part), with half- width across tymbal. Opercula very small. very narrow lateral crests along proximal part. Basal part of operculum slightly vaulted with Monophyly of the three genera. In the follow- two rounded elevations and weakly wedge- ing discussion figures in parentheses refer to shaped; longest medially and gradually tapering apomorphies in the cladogram (fig. 2). Chloro- towards lateral margin. Basal part with distinct cysta, Glaucopsaltria and Owra form a mono- crest around its distolateral corner, lateral part phyletic group based on the following synapo- of this crest very short and in males often glo- morphies: 1. a continuous band of subapical bularly swollen. Distal part of operculum in areas in tegmen; 2, a wedge shaped basal part of males angularly oblong and longer than basal operculum; and 3, small male auditory capsules part (though extremely short in Owra) and not, (De Boer, 1995d). or only partly covering tymbal cavity in ventral view; often leaving most of folded membrane C. fumea exposed. Lateral margin of male operculum C. suffusa straight or weakly convex, slightly directed mesiad and forming a distinct and obtuse angle C. vitripennis with crest of basal part and with straight distal ' O. insignis lies laterally margin. The straight medial margin 1 2 3 of meracanthus. Meracanthus fairly short, but G. vlridis generally reaching beyond operculum. Female Figure 2. Cladogram of the species of Chlorocysta. operculum shorter than in male and sickle- Glaucopsaltria and Owra. Numbers refer to apomor- shaped. Male abdomen very delicate and dis- phies discussed in the text. tinctly inflated. First tergite in male medially often more than half as long as 2nd tergite (but The tegmen venation tends to be variable and Glaucopsaltria with extremely long 2nd tergite) often differs between left and right tegmen and not hidden under metanolum. Middorsal within individuals. The number of apical areas 98 V J. DE BOER

varies from 9-10 in Owra to 11-13 in Chloro- species sharing a large tymbal with 11-12 ridges cysta and to 13-15 in Glaucopsallria but is (7). Furthermore, these species share a wide always more than 8. the number most common pygofer opening and a distinct colour pattern on in related genera and in most cicadas. Further- head and thorax but the phylogenetic value of more, the tegmina have a more or less continu- these characters is not clear since they also occur ous band sometimes of a double row of subapical in several related genera. areas between apical and ulnar areas. Similarly high numbers of apical areas in the tegmina are Chlorocysta West wood Boer also found in Baeturia inconstans De Cvstusoma (Chlorocvsta) Wcstwood, 1851: 208. — (De Boer, 1994c), Gymnotympana (4 species, De Walker. 1852: 1133. Boer. 1995a), Mirabilopsaltria (2 species, De Chlorocvsia.-Sl&\. 1863: 575. — God ing and Frog-

Boer. 1 996) and all but one species of Thaumas- gatt, 1904: 566, 596, 658. — Distant, 1906: 153, 159. topsaltria (De Boer, 1992b). A continuous band — Boulard. 1979: 35. — Duffels and Van der Laan, Boer, of subapical areas is regarded apomorphic for 1985: 249. — Moulds. 1990: 185-186. — De 1990: 64. — De Boer, 1991: 2-3. — De Boer, 1992a: Chlorocysta, Glaucopsahria and Owra (1 in fig. 164. — De Boer, 1993a: 16-17. — De Boer, 1993b: 2). Although in the tegmina of the species of 142. — De Boer. I 994a: 3. — De Boer, 1 995a: 4. 8, 24. Gymnotympana, Mirabilopsaltria and Thau- — De Boer. 1 995b: 6. mastopsaltria noted above some subapical areas Glaucocvsta Coding and Froggatt. 1904: 566. occur, these never form a continuous band. Mardalana Distant, 1905: 213, 215. — Distant, Subapical areas do not occur in Baeturia. 1906: 154, 159. — Mctcalf, I 963: 257. — Duffels and The basal part of the male operculum is Van der Laan. 1985: 249. slightly wedge-shaped, longest medially and Mardarana fsic] Kato, 1932: 185. — Kato, 1956: slightly tapering to its lateral margin. The oper- 70. 46. culum base is generally oblong in species of Mardalena [sic] Boulard, 1979: Chlorocystini but in most species of Gymnotym- Type species. Cvstosoma vitripennis Westwood, pana the operculum base has its greatest length 1851. laterally (De Boer, 1995a). Only in Gymnotym- pana ru fa, G. varicolor and Venustria superba the Diagnosis. Green body. Basal part of operculum strongly inflated. basal part of operculum is longest medially but wedge shaped. Male abdomen inflated. narrows more abruptly at about one third of its Male auditory capsules weakly Teg- width. The weakly wedge-shaped operculum mina with more than eight apical areas and con- base is regarded apomorphic for Chlorocysta, tinuous band of subapical areas. Tymbal with 9-12 ridges. Glaucopsallria and Owra (2). Aedeagus in lateral view Z-shaped Chlorocysta. Glaucopsahria and Owra have near apex. small and weakly protruding auditory capsules. Remarks. Chlorocysta, originally described as a Although similarly unswollen capsules are pres- subgenus of Cystosoma, is the type genus of ent in Thaumaslopsaltria (De Boer, 1992b) they Chlorocystini Distant, 1 905. The genus contains presumably represent a parallel development three species but several undescribed species and can be regarded as for apomorphic this have been distinguished (Moulds, pcrs. coram.). group (3). The peculiarly Z-curved aedeagus is a supposed Phylogenetic relationships. Chlorocysta and apomorphy for Chlorocysta. Owra share wings with 5 apical areas synapo- morphic for these genera (4). Wings with 5 apical Chlorocysta vitripennis (Westwood) areas occur in several genera of cicadas but only sporadically in other genera of the Chlorocys- Figures 3-13 tini. Cvstosoma (Chlorocvsia) vitripennis Westwood. The three species of Chlorocysta share a some- 1851: 208. what Z-curved aedeagus (a modification of the Chlorocysta vitripennis. — Coding and Froggatt,

S-curved aedeagus, an apomorphy of the Chlor- 1 904: 566, 659, pi. xix fig. 6. — Distant, 1 906: 1 59. — ocystini) and a high number (9-12) of tymbal Burns, 1 957: 643. — Metcalf, 1 963: 256-257. — Duf- fels and Van der Laan, 1985: 249. ridges (5, 6) as apomorphies. Owra insignis — Moulds, 1990: 1 88- 189, pi. 22 figs 2, 2a-b. — De Boer. 1995a: 16,77. shares an incised aedeagal apex with C. fumea — De Boer. 1995b: 5. and C suffusa but similarly incised aedeagal Cicada congrua Walker, 1862: 303-304. — Coding apices occur in several related genera (e.g., Aed- and Froggatt. 1904: 611, 657. eastria, Guineapsaltria and Papuapsaltria). C. Chlorocvsta maenda Stal, 1 863: 575. — Godingand fumea and C sujfusa arc supposed to be sister Froggatt. 1904: 566. 659, 660. PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 99

Figures 3-13. Chlovocysia vitripennis Westwood, 1851: 3, male head in dorsal view, Mt Tamborine; 4. male postclypeus in lateral view, Mt Tamborine; 5, pygofer in lateral view, Mt Tamborine; 6, claspers and aedeagus, Mt Tamborine; 7. pygofer in oblique view, Mt Tamborine; 8, male caudodorsal beak in dorsal view, Mt Tam- borine; 9, female operculum, Dorrigo; 10, female caudodorsal beak in dorsal view. Dorrigo; 1 1, aedeagus in lateral view. Mt Tamborine; 1 2, aedeagus in lateral view, ex. coll. v Voixem; 1 3, male operculum, Mt Tamborine, arrow indicating gap between sternites 1 and 2. bp = basal part of operculum; c = crest around distolateral corner of basal part of operculum; cb = caudodorsal beak; dm = distal margin of operculum; di = distal margin of pygofer; do = dorsal margin of pygofer; dp = distal part of operculum; ho = clasper hollow; la = lateral margin of operculum, m = medial margin of operculum; me = mcracanthus; mc = medial corner of operculum; pr= protuberance on lateral lobe of pygofer; ve = ventral margin of pygofer. 102 A. J. DE BOER

Figures 14-24. 14-19, Ch/orocysta fumea Ashton, 1914: 14, pygofer in lateral view; 15, operculum; 16, cau- dodorsal beak in dorsal view: 17, aedeagus in lateral view; 18, aedeagal apex; 19, claspers. 20-24 Chlorocysta sitffiisa Distant. 1907: 20. male fore femur in lateral view. Cairns; 21, female operculum, Cooktown; 22, female genital segment in lateral view, Cooktown; 23, female caudodorsal beak in dorsal view, Cooktown; 24, male operculum. Julatten. PHYLOGENY AND BIOGEOGRAPFIY OF AUSTRALIAN CICADAS 103 an almost right angle with slightly longer distal sharing a similar colour pattern and incised aed- margin. Distal margin straight, but weakly con- eagal apex. Only one male was available for vex near angular distomedial corner. Medial study. margin straight. Distal part of operculum elongate at distomedial corner, reaching mesiad Chlowcysta suffusa (Distant) beyond apex of meracanthus. Mcracanthus short, not reaching distal margin of oper- Figures 20-29 culum. Mardalana suffusa Distant, 1907: 418. — Burns. Abdomen. Strongly inflated, almost uni- 1957: 644. — Metcalf. 1 963: 258. — Duffels and Van formly brown coloured, but with darkened dcrLaan, 1985: 249. segmental hind margins and a lateral row of Chhwevsta suffusa. — Moulds, 1990: 186-187, pi. darkened spots. 22 figs 7, 7a.

Genitalia. Pygofer in lateral view (fig. 14) Material examined. Australia. S. Qld, Naukler. coll. short and globularly swollen. Dorsal margin Dodd, tf, ZMH. Cairns 9 km W, ii-iv.1985, d. ZMA. weakly concave, convexly bending into erect Cairns. A. P. Dodd d Dct. Chlorocvsta vilripennis G+F. caudodorsal beak. Distal margin slightly convex BPBM. Cairns. 1920, J.F. Illingworth. < 9 Del. Chlor- between beak and lateral protuberance. Ventral oevsta vitripewiisG+F. BPBM. Clohcsi River. S.F. SW

margin angularly convex. Ventral margins con- ofKuranda. 1 8.i. 1 984, M.S. and B.J. Moulds,

1 .2 x. Tegmina of males 1 . 1-1.2 x body length, of Distribution. Endemic to the Cape York Penin- females 1.3-1.4 x. sula, Qld. Moulds (1990) recorded the species Head. With similar colour pattern as in only known from along the old Leo Creek track C. fumea. though strongly varying in intensity. at the southern end of the Mcllwraith Range, at Vertex light brown, darker brown around lateral an altitude of around 300 m. ocelli and sometimes around frontal ocellus and bordering Remarks. C. filmed is the largest species of this with dark brown spots lateral corners genus and is easily recognized by its distome- of postclypeus. Vertex lobe with 2 dark brown dially elongated male operculum. C. fumea is spots between lateral ocellus and eye and a dark closely related to C. suffusa., described next, spot bordering proximomedial corner of eye. 104 A. J. DE BOER

Figures 25-37. 25-29, Chlorocysta suffusa Distant, 1907: 25, pygofer in lateral view, Julatten; 26, pygofer in oblique view, Julatten; 27, claspers and aedeagus, Julatten; 28, male caudodorsal beak in dorsal view, Julatten; 29, aedeagus in lateral view, Julatten.

30-37 Owra insignis Ashton, 1 9 1 2: 30, pygofer in lateral view; 3 1 , aedeagus in lateral view; 32, male caudodorsal beak in dorsal view; 33, male operculum; 34, female caudodorsal beak in dorsal view; 35, pygofer in oblique view; 36, clasper; 37. female operculum. 3

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 105

Vertex lobes slightly darkened on anterior parts. mental hind margins often darker coloured. Postclypeus in dorsal view dark brown at lateral Ventrolateral row of slightly darkened spots on corners and along anterior margin. Ventral part segments 3-7. Female abdomen with pattern of of postclypeus with large dark medial spot. Lat- broad longitudinal lateral streaks similar as in eral surface of postclypeus with 9-10 irregular Gymnotympana strepitans (De Boer, 1 995a) and rows of short parallel ridges continuing into ten a distinct dark brown ventromedial band. parallel streaks running towards central Female caudodorsal beak reaching to apex of fissure. ovipositor sheaths (fig. 22) and triangular in dor- Thorax. Pronotum ochraceous brown, with a sal view, somewhat swollen to its base and broad lighter ochraceous coloured medial band sharply pointed at apex (fig. 23). that is not, or only slightly, dilating towards dis- Male genitalia. Pygofer in lateral view (fig. 25) tal pronotal margin. Proximal end of this band short and globularly rounded. Dorsal margin marked by a pair of small dark brown parame- slightly concave, continuous with straight and dian spots at pronotal collar. Pronotum light slender caudodorsal beak. Distal margin slightly brown along oblique fissures and towards convex between beak and lateral protuberance. medial band. Mesonotum greyish brown, only Ventral margin straight, but forming a rounded slightly darkened in paramedian spots at prono- corner under lateral protuberance. Ventral mar- tal margin and in vague reticulate pattern of gins converging and forming a sharp angle at angular brown spots, forming lateral streaks base of pygofer opening (fig. 26). Caudodorsal from pronotal margin to corners of cruciform beak in dorsal view (fig. 28) short, triangular and elevation. Cruciform elevation greenish pointed or narrowly rounded at apex. Lateral ochraceous, with brown medial band. 2 black lobe of pygofer curving inwards, forming a tri- spots in front of elevation. angularly swollen protuberance, which does not Tegmina and wings: Hyaline. Tegmen ven- extend beyond distal margin of pygofer. ation variable, differing even between left and Claspers (fig. 27) very similar to C. fumea. right tegmen of individuals. Tegmina with 12- Clasper base fused to a high and continuous col- 13 apical areas and a, at some places double, lar around base of anal valves. Dorsal margin of band of 6-8 subapical areas. Wings with 5 apical clasper ending in right angle on this collar and areas. not forming a crest on lateral surface of clasper Tymbals. 12 dark brown parallel sclerotized base. Clasper base forming collar around base of ridges; 8 ridges spanning the tymbal from dorsal anal valves, which abruptly, but slightly, curves to ventral margin, the 4 most proximal ridges do inwards between claspers. Claspers forming not reach ventral margin and successively short- diverging ridges from dorsal margins to base of ening, with most proximal ridge shortest. 12 anal valves. Distal corner of clasper bending short and lighter coloured intercalary ridges mesiad around aedeagus, supporting aedeagus seem to form a lateral band across tymbal. in upright position. Claspers curving down and of Operculum. Male operculum (fig. 24) closely strongly diverging towards apices. Apical part resembling that of C. fumea in size, but with a clasper with sharply edged hollow at its inwardly more rectangular shaped distal part, not directed side. Aedeagus apically rounded in lat- Z-curved in apical elongated at distomedial corner. Basal part of eral view (fig. 29), slender and almost operculum slighly vaulted and wedge-shaped; third. Basal two-thirds of aedeagus lateral crests and longest medially and tapering towards its lateral straight, with very slender of lateral margin. Basal part forming a distinct, though slightly swollen dorsally, at distal ends apex with dorsoventral very short crest around rectangular distolateral crests. Aedeagal ending in two small, rounded, corner. Distal part of operculum almost rec- incision; aedeagus tangular, adjacent to body and not covering tym- lateral lobes (fig. 27). (mean ± sd): Body length cf; bal cavity in ventral view, though, as in C. Measurements ± 1.1), 9: 20.0-21.5 fumea, covering most of folded membrane. The 24.2-27.9 mm (26.2 mm (20.5 ± 0.6); tegmen length d: 28.2- straight and almost parallel lateral and medial mm mm 26.3-28.3 margins both making an almost right angle with 31.2 mm (30.0 mm ± 1.1), 9: mm ± 0.7); head length d: 2.1-2. 5 mm(2. straight distal margin. Meracanthus short, not (27.5 mm 1.9-2.5 (2.2 mm): pronotum length reaching to distal margin of operculum. Female mm), 9: mm and d: 2.5-2.8 mm (2.7 mm), 9: 2.7-2.8 mm; meson- operculum (fig. 21) short, sickle-shaped otum length d: 5.4-6.4 mm (5.8 mm), 9: 4.9-5.6 erect. Abdomen. Male abdomen strongly inflated, mm (5.3 mm); head width cf: 6.0-6.7 mm (6.4 9: 6.0-6.6 mm (6.3 mm); width of pronotal almost uniformly brown to green coloured, seg- mm ), 106 A. J. DF. BOER

part of operculum about 4 x as collar <5; 6.7-7.5 mm (7.0 mm), 9: 6.6-7.3 mm exposed. Basal with dis- (6.9 mm). long as distal part, slightly vaulted and tinct crest around rectangular distolateral cor- Distribution. Along the eastern coast of the Cape ner. Basal part wedge-shaped; medially dis- York Peninsula, Qkl. Moulds (1990) recorded tinctly longer than laterally. Distal part of male the species from Iron Range, the Mcllwraith operculum very short, sickle-shaped and erect, Range near Coen and Cooktown south to the hardly more than a continuation of the crest Paluma Range. around distolateral corner of basal part. Mera- Remarks. C, sitffusa closely resembles C. fumea canthus reaching well beyond operculum. can be in tymbal shape and colour pattern but Female operculum (fig. 37) very similar to that easily separated from that species by a dark of male, with extremely short and sickle-shaped brown suffused spot on the ventral side of the distal part. postclypeus. Males of C. suffusa are easily sep- Abdomen. Male abdomen slender and weakly arated from C. fumea by their rectangular oper- inflated, almost unicoloured ochraceous, but culum. segmental hind margins orange-brown. Second Owra Ashton tergite (fig. 33) weakly curved along tymbal cav- — — ity. Female abdomen light brown with reddish Oma Ashton. 1912: 224. > Metcalf, 1963: 252. segmental hind margins and ventrolateral row of Duffels and Van dor Laan, 1985: 248. — Moulds. brown spots on segments 3-8. Ovipositor 1990: 184-185. — Dc Boer. 1992b: 18, 19, 20. — De sheaths not reaching to apex of caudodorsal Boer. 1993a: 16-17. — De Boer, 1993b: 142. — De beak. Female caudodorsal beak in dorsal view Boer. 1995a: 8. — Dc Boer, 1995b: 6. (fig. 34) broad and bluntly rounded at apex. Type species. Owra insignis Ashton, 1912 Male genitalia. Pygofer in lateral view as in fig. (monotypic), 30. Dorsal margin of pygofer straight, strongly Owra insignh Ashton concave to base and angularly bending into straight, slender and erect caudodorsal beak. Figures 30-37 Distal margin slightly convex, angularly bending Qwm insijtnh Ashton, 1912: 224, pi. LI rigs 6-6a. — into margin of beak. Lateral lobe of pygofer Burns. 1957, 642. — Metcalf, 1963: 252. — Duffels strongly curving inwards towards end of distal and Van dor Laan, 1985: 248. — Moulds, 1990: 185, margin and forming an angularly rounded and pi. 22 figs 6. 6a. swollen lateral protrusion. Ventral margin of \liitfiial examined. Australia. Lake Barine, 530 m. pygofer strongly convex near this protrusion, 1- 3 1. 1. ii. 1 964. J. Sedlacek. y. BPBM: ('aims, A. P. but almost straight towards base of pygofer. Dodd. rf Stal. BI'BM. Det. Chlorocyslct nUcrula Ventral margins converging to short and straight Description. Body reddish brown but according basal margin at base of pygofer opening (fig. 35). to Moulds (1990) olive green when alive (for Caudodorsal beak in dorsal view (fig. 32) short photographs see Moulds, 1 990: pi. 22 figs 6, 6a). and broad, truncate at apex. Claspers very dif-

Abdomen in male 1. 5 x head and thorax ferent from those of related species, missing a together, in female l.O x. Tegmina of male l.l x claspcr hollow. Claspcr (fig. 36) broad, globu- body length, of female 1.4 x. larly swollen near base, strongly curved down to Tegmina and wings. Hyaline, venation red- very slender and elongate, almost laminiform brown and costa bright red. Tegmina with 10 apical part. Claspers strongly diverging towards (according to Moulds, 1990, sometimes 9) apical their truncate apices. Claspcr base forming a low areas and a regular band of 3 or 4 subapical collar around base of anal valves. Acdcagus (fig. areas. Costal area very broad. Tegmen with very 31) strongly upcurved at half-length and with narrow hyaline border along hind margin. Wing slender lateral crests. Aedeagal apex incised, as with 5 apical areas and broad hyaline border in C. fumea and C. suffusa, but ending in two along hind margin. more sharply pointed lateral lobes (fig. 35). Tymbals. 5 parallel sclerotizcd ridges span- ning the tymbal from dorsal to ventral margin Measurements. Body length d; 15.1 mm, 9: 13.3 and a 6th, most proximal ridge, almost reaching mm; tegmen length d; 17.0 mm, 9: 19.0 mm: ventral margin. 5 distinct intercalary ridges head length d; 1,6 mm. 9: 1.7 mm; pronotum seem to form a lateral band across tymbal. length d; 1.6 mm, 9: 1 .8 mm; mesonotum length

Opercula. Male operculum (fig. 33) extremely d; 3.0 mm. 9: 3.3 mm; head width d; 4.1 mm, 9: small, not covering tymbal cavity in ventral view 4.6 mm: width of pronotal collar d; 4.0 mm, 9: and leaving folded membrane completely 4.6 mm. PHYLOGENY AND BIOGEOGRAPHY OP AUSTRALIAN CICADAS 107

Distribution. Only known from the south-east- Tymbals. 6 transverse sclerotized ridges span- ern corner of the Cape York Peninsula, north- ning the tymbal from dorsal to ventral margin eastern Qld. Moulds ( 1 990) recorded the species and a 7th, most proximal, ridge almost reaching from Thornton Peak range north of Daintree to ventral margin. 7 short intercalary ridges seem Mission Beach and widespread on the Atherton to form a lateral band across tymbal. Tableland. Operculum (fig. 44). Basal part of operculum slighly vaulted, wedge-shaped; longest medially Remarks. Owra insignis is a very small and tapering towards lateral margin. Basal part ochraceous brown species, at first sight closely forming an irregularly protruding crest around resembling Guineapsaltria Jlava and Gymno- distolateral corner. Distal part of operculum tympana ntfa. Owra, however, is closely related longer than basal part and reaching beyond apex to Chiorocysta and Glaucopsaltria, sharing the of meracanthus. Distal part only partly covering presumed synapomorphic tcgmen venation. tymbal cavity in ventral view, though covering Owra shares the 5 apical areas in the wing with most of folded membrane. Lateral margin mak- Chiorocysta. Only one male and one female have ing an almost right angle with basal part of oper- been examined. culum and initially almost straihgt, but angu- Glaucopsaltria Coding and Froggatt larly bending mesiad where operculum curves towards body. Distal margin straight, making a Glaucopsaltria Coding and Froggatt, 1904: 657, sharp angle with straight medial margin. Disto- 659. — Moulds, 1 990: 1 89 — De Boer, 1 992b: 18,19. lateral corner broadly rounded. Operculum 20. — Dc Boer, 1 993a: 1 6- 1 7. — De Boer, 1 993b: 1 42. strongly curved-up along proximal part of its lat- — De Boer, 1995a: 8. — De Boer, 1995b: 6. Brisbane specimen), Glaucopsaltria in synonymy of Chlorocvsta. — Dis- eral margin (especially in tant. 1906: 159. — Metcalf, 1963: 255-256, 473. forming a small lobe (fig. 44 arrow). Abdomen. Strongly inflated, uniformly green Type species. Glaucopsaltria viridis Coding and or discoloured yellow, with traces of green. Sec- Froggatt (monotypic). ond tergite medially about 3 x 1st tergite and auditory capsules. Glaucopsaltria viridis Coding and Froggatt laterally gradually tapering to Seventh tergite wedge-shaped in lateral view; 38-44 Figures very long and convex dorsally (lateral view),

(ilaucopsa/tria viridis Goding and Froggatt, 1904: short ventrally. 566,658. — Kirkaldv. 1907: 308(6). — Moulds, 1990: Genitalia. Pygofer in lateral view as in fig. 39. 190, pi. 22 tig 3. — De Boer, 1995a: 16. Dorsal margin weakly concave, continuous with Chlorocvsta viridis. — Distant. 1906: 159. — Frog- erect caudodorsal beak. Distal margin weakly gatt. 1907:352. — Burns. 1957,643. — Metcalf. 1963: convex between beak and lateral protuberance. Van Laan, 1985: 249. 256. — Duffels and der Ventral margin straight, forming an almost rec- Material examined. Australia. Qld. 24.ii.l946, rf, tangular corner just under lateral protuberance, SEM. Brisbane. 1973. J.B. Vogel. 8, ZMA. Qld. J. A. concavely bent near base of pygofer opening. Grant. BM- CS1RO Expedition, BM 1973-346. rf. Ventral margins converge and form a sharp 4836. BMNH. Rockhampton. Museum Godefroy No angle at base of pygofer opening (fig. 38). Cau- rf det. C viridis. Z1M. dodorsal beak in dorsal view (fig. 41) broad and rectangular at apex. Lateral Description. Body olive green or yellow-green triangular, almost inwards towards distal (for photograph see Moulds, 1990: pi. 22 fig 3). lobe of pygofer bending an angularly rounded and lat- Tegmina slightly shorter than body. Abdomen margin, forming protuberance. Clasper as in 1.9-2.2 x head and thorax. erally flattened and more slender Tegmina and wings. Hyaline. Tegmen ven- Chiorocysta. but with longer Apical part of clasper ation variable and differing even between left apical part (fig. 40). strongly bent down towards smoothly rounded and right tegmen of individuals. Tegmina with with a sharply edged hollow at 13-15 apical areas and a continuous band, at apex and directed side. Dorsodistal corner of some places concisting of a double row, of 7-10 inwardly bending mesiad around aedeagus, sup- subapical areas. Costal area very distinct, broad- clasper aedeagus in upright position. Claspers ening towards tegmen apex. Tegmen with dis- porting almost parallel, only slightly diverging towards tinct hyaline border along hind margin, though apices and fused at base to a fairly low and con- narrower than in wing. Wing with 6 apical tinuous collar around base of anal valves. areas. Medial part of collar curving inwards, slightly Legs. Fore femur with row of 3 erect spines dome-shaped proximally of aedeagus, between (fig. 43). 108 A. J. DE BOER

Figures 38-46. 38-44, Glaucopsallria viridis Goding and Froggatt, 1 904: 38, pygofer in oblique view, Australia; 39, pygofer in lateral view, Brisbane; 40, claspers, Australia; 41, caudodorsal beak in dorsal view, Australia; 42, aedeagus in lateral view, Australia; 43, male fore femur in lateral view, Australia; 44, operculum, Brisbane, arrow indicating upcurving lateral lobe. 45-46 Venustha superba Goding and Froggatt, 1904: 45, male postclypeus in lateral view, Kuranda; 46, male head in dorsal view, Kuranda. PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 109 dorsal crests of claspers. Dorsal margin of Recently Moulds ( 1 990) transferred the genus to clasper forming a sharply edged crest, continu- Chlorocystini since several characters (e.g., fore- ing on lateral surface of clasper base. Aedeagus wing vein Sc separated from costa; apical cells (fig. 42) weakly S-curved, with very short and mostly longer than ulnar cells; and an inflated narrow lateral crests along proximal half. Aed- male abdomen) are "associated with that eagal pore oval. tribe". Measurements, (mean): Body length: 29.8- Venustria has a weakly S-curved aedeagus, 33.8 mm (32.1 mm); tegmen length: 29.5-31.3 with very narrow lateral crests and should there- mm (30.0 mm); head length: 2.1-2.4 mm (2.3 fore be placed in Chlorocystini. The width of the mm); pronotum length: 2.7-2.9 mm (2.8 mm); head and shape of the pronotum of Venustria are mesonotum length: 5. 1-6.0 mm (5.5 mm); head very similar to those of Chlorocysta and its width: 6.4-6.5 mm; width of pronotal collar: metasternum, though more protruding than in 6.8-7.0 mm (6.9 mm). Chlorocysta, does not differ greatly from that of other genera of the tribe. Males of Venustria, Distribution. Along the eastern coast of Qld and however, have a very short and rather solid look- in north-eastern NSW. Moulds (1990) recorded ing abdomen, often shorter than head and tho- the species from Daintree River punt crossing rax together. The eyes are large and angular with and Cairns in northern Qld, from Mackay, Car- inwardly directed proxomedial corners and the lisle Island, Middle Percy Island and Eungella in tegmina are relatively long so that in general central Qld and from the Kolan River north of aspect Venustria looks fairly different from all Gin Gin to Dorrigo. other species of the tribe. Remarks. Glaucopsaltria viridis is larger than the Chlorocysta species with a strongly inflated Venustria superba Goding and Froggatt abdomen. Glaucopsaltria is easily separated Figures 45-55 from Chlorocysta and Owra, since it has six (instead of five) apical areas in the wings. Males Venustria superba Goding and Froggatt, 1904: 565, of G. viridis can be recognized by the shape of the 597, 603, pi. xix figs 7, 7a. — Distant. 1906: 628. — 7th tergite which is very long middorsally but Burns. 1957: 634. — Metcalf, 1963: 203-204. — Duf- 1985: 233. Moulds, 1990: short ventrally so that the pygofer is curved to a fels and Van der Laan, — 180-181, pi. 16, fig. 11. DeBoer, 1995a: 3, 4, 9, 13, ventral position. Females have not been exam- — 15, 74, 77. ined. The local name of this species is Bottle Cicada (Moulds, 1990). Material. Australia. Australien, 1888, v Miiller, 9, SMN. Etty Bay nr Innisfail. 23.ii.1982, M.S. and B.J. Moulds, rf Venustria superba det. M.S. Moulds. ZMA. Venustria Goding and Froggat Cairns, J.F. Illingworth. 1920. 5cf, 9 det. Venustria

Kuranda 1 ml. E, 1 l.iii.1964, I.F.B. Venustria Goding and Froggatt, 1904: 565, 596. — superba. BPBM. and M.S. Upton, d, CSIRO. Distant. 1906: 128, 129. — Metcalf, 1963: 203. — Common Moulds, Duffels and Van der Laan, 1985: 233. — Description. Body light brown (for photograph 1991: 1 990: 32. 1 80. — De Boer, 1 990: 64. — De Boer. see Moulds, 1 990: pi. 16, fig. 1 1 ). Females about 2-3. — De Boer. 1992a: 164. — De Boer, 1992b: 19. as long as males, but with more robust head and — DeBoer, 1993a: 16-17. — De Boer, 1993b: 142.— thorax and longer tegmina. Abdomen of males De Boer, 1994a: 3. — De Boer, 1994b: 130. — De relatively short, 0.8-1.1 X head and thorax Boer. 1995a: 1, 3. 4, 8, II. — De Boer, 1995b: 5. 0.9-1.0 X. Tegmina fairly Vcnustra [sic]. — Kato. 1932: 181. together, of females long in males and females, being 1.3-1.5 X body Venustria superba Goding and Type species. length. Froggatt, 1 904 (monotypic). Head (fig. 46): Ochraceous brown, darkened Remarks. Distant (1906) included Venustria in towards anterior margins of vertex lobes, with his division Taphuriaria (now the tribe Taphu- row of 3 dark spots on vertex lobe between lat- slightly broader than rini, which has as principal characters: "Eyes eral ocellus and eye. Head projecting beyond anterior margin of pronotum; anterior part of pronotum, 2.4-3. 1 X as broad pronotum subquadrate, not distinctly narrowed as long and 2.1-2.4 X as broad as distance anteriorly; abdomen about as long as space between eyes. Head distinctly shorter than dis- between apex of head and base of cruciform tance between eyes. Postclypeus 2.6-3.2 X as elevation; a more or less distinct posterior wide as long and smoothly rounded anteriorly, metasternal process visible in males between or its anterior margin convex and almost continu- anterior margins of vertex lobes. at the base of the opercula" (Distant, 1906). ous with A. .1. DE BOER

Figures 47-55. Vcnmtria superba Goding and Froggatt. 1904: 47, female genital segment. Cairns; 48, female operculum. Cairns: 49, female caudodorsal beak in dorsal view. Cairns; 50, pygoferin lateral view, Kuranda; 51, pygofer in oblique view, Kuranda; 52. aedeagus in lateral view, Etty Bay; 53, male caudodorsal beak in dorsal view, Kuranda; 54. male fore femur in lateral view, Kuranda; 55. male operculum, Etty Bay. PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 111

Postclypeus (fig. 45) weakly swollen ventrally. tomedial and medial margins straight, disto- anterior margin (lateral view) weakly convex. medial corner broadly rounded, medial corner Lateral sides of" postclypeus with about 12 fur- more narrowly rounded, almost rectangular. rows and 5-6 very short and irregular rows of Meracanthus very short, hardly reaching beyond short parallel ridges on weakly inflated crest basal part of operculum. Opercula widely separ- along the lorum. Vertex smooth with weak ated medially. Female operculum (fig. 48) very medial furrow and without diverging furrows. small. Basal part as in males, medially distinctly Vertex in males 1.8-2.2 x, in female 2.3-2.4 X longer than laterally. Distal part shorter than as wide as long and. in both sexes, 1.4-1.8 X as basal part, its straight lateral margin forming a wide as width of eye. Eyes large and angular; broadly rounded angle with weakly convex medial margins of eyes strongly converging: distal margin. shortest distance between eyes at their proxo- Abdomen. Light brown, silvery pilose and medial corners. Ocelli small and wide apart. with dark brown midventral band. Males with Distance between lateral ocelli 0.8-1.0 X dis- darkened 8th and sometimes 7th, tergite and tance between lateral ocellus and eye and 2.8- ventrolateral row of slightly darkened spots on 3.4 X width of frontal ocellus. segments 3-7. Male abdomen more solid in Thorax. Pronotum ochraceous, with dark aspect than in other species of Chlorocystini, streaks in and along oblique fissures and a nar- hardly swollen and almost without fold between row light ochraceous or greenish coloured tergites and sternites; nearly circular in cross sec- medial band bordered by vaguely darkened tion. First tergite very short and partly hidden streaks. Pronotum without medial fissure, 2.3- under metanotum. Anterior margin of 2nd ter- 2.7 X as wide as long and, across the collar, 1.1- gite concave medially. Lateral parts of 2nd

1 .2 X as wide as head. Amplified lateral corners tergite swollen anteriorly and almost adjacent to of pronotal collar larger than in Chlorocysta. tymbal, leaving only a narrow gap between tym- Mesonotum ochraceous brown, with reticulate bal and 2nd tergite. Auditory capsules globularly pattern of ochraceous stripes and brown specks, swollen and distinctly elevated relative to con- forming a pair of paramedian semicircular spots necting bar between abdomen and tymbal. Ter- at pronotal margin and a pair of converging lat- gite part between auditory capsule and sternitc 2 eral bands from pronotal margin to corners of almost straight, with distinct crest along tymbal cruciform elevation. 2 dark spots in front of cavity. First sternite globularly swollen between cruciform elevation. opercula. Female abdomen about as large as that Legs. Fore femur (fig. 54) with row of 3 erect of male, but more strongly tapering towards spines, diminishing in length towards tibia. apex. Female pygofer (fig. 47) long and slender Most proximal spine very long, longer than with distinct thorn-shaped protuberance at ven- distance to middle spine. tral margin. Ovipositor sheaths reaching to apex Tegmina and wings. Hyaline, but slightly of caudodorsal beak. Female caudodorsal beak bronzed in apical areas. Venation reddish in dorsal view (fig. 49) triangular and sharply brown. Tegmen with 9 apical areas and distinct pointed at apex. costal area. Veins CuA and M adjacent, but not Male genitalia. Pygofer in lateral view as in fig. fused near basal area. Wing with 6 apical 50. Dorsal margin almost straight, but concave areas. to base and weakly convexly bent into stout and Tymbals. 4 transverse sclerotized ridges span- slightly posteriorly curved caudodorsal beak. ning the tymbal from dorsal to ventral margin, a Distal margin angularly convex, concavely bent 5th ridge almost reaching ventral margin and a into caudodorsal beak and ending in almost 6th, most proximal, ridge spanning about three- right angle on broad and angular lateral pro- quarters of tymbal width. 5 short intercalary trusion. Ventral margin strongly concave. Lat- ridges seem to form a lateral band across eral lobe of pygofer with very small angularly tymbal. swollen lateral protuberance, distinctly reaching Pygofer open- Opercula. Male operculum (fig. 55) covering beyond distal margin of pygofer. broad (fig. 5 ); ventral margins ending wide greater part of tymbal cavity in ventral view and ing 1 extending medially of mcracanthus. Basal part apart at straight basal margin. Caudodorsal beak of operculum medially distinctly longer than lat- in dorsal view (fig. 53) triangular and sharply Clasper (cf. fig. very short erally, abruptlv broadening at about third of its pointed at apex. 51) width and strongly vaulted. Distal part angularly and stout, with rounded lobate and downwardly sharply edged oblong. Lateral margin long and straight, angu- directed apical part and broad, directed, clasper hollow. Dorsal larly bent into weakly convex distal margin. Dis- inwardly 112 A. J. DE BOER

in Cys- margin of" elasper sharply curving upwards rax, in females 0.9-1.1 X. Male tegmen proximally, merging with broad ring-shaped tosoma 0.8-1.1 X, in Cystopsaltria 1.4 X body

.2- . (fig. nar- claspcr base. Basal part of elasper very broad in length, in females 1 1 .4 X Head 69) lateral view, distinctly swollen around base of rower than anterior part of pronotum, with long anal valves, especially mid-between claspers, and angularly protruding postclypeus. Flead possibly representing the remnants of a medial 1.9-2.3 X as broad as long. Postclypeus dis- uncus lobe. Acdcagus (fig. 52) weakly S-curvcd, tinctly protruding, its sharply edged anterior strongly swollen in proximal half and curving margin forming a right angle at apex. Postcly- down towards apex. Acdeagus strongly widening peus 1.6-2.1 X as broad as long, distinctly near base, forming 2, somewhat lobate, basola- swollen ventrally (fig. 70), its anterior margin teral protuberances Hanking a large hollow at the (lateral view) convex or forming an obtuse angle fold between acdeagus and its basal plate. These at half length. Lateral sides of postclypeus some- protuberances continue into short lateral crests times slightly bulbous, with about 9-12 weak along proximal part of acdeagus. Acdeagus with furrows and a weakly inflated crest along lorum. very slender, almost membranous, laminiform Vertex wrinkled, with distinct medial fissure, ventrolateral crests, just medially of these lateral but without diverging fissures between ocelli. crests, which curve into the ventral hollow Central part of vertex only slightly elevated, between the basolatcral protuberances. Aed- often even somewhat pressed down and con- eagal pore almost round, weakly incised cave. Vertex in Cystopsaltria narrower than in ventrally. Cystosoma. Vertex 1.6-2.1 X as broad as long. &• Measurements (mean t sd): Body length Distance between eyes 0.9-1.1 X length of head 20.6-25.2 mm (x23.4 mm ± 1.3), 9: 23.4 and and 1.1-1.4 X postclypeus width. Ocelli wide

26.2 mm; legmen length d": 3 1 .0-34.6 mm (x33.5 apart in Cystosoma, more closely together in mm i 1.1), 9: 35.1 and 35.3 mm; head length d; Cystopsaltria. Distance between lateral ocelli

2.4-3. 1 mm (x2.6 mm), 9; 2.5 and 2.6 mm; pron- 0.8-1.1 X distance between lateral ocellus and otum length d: 2.9-3.6 mm (x3.3 mm), 9: 3.4 and eye, and, in Cystosoma 2.4-3.7 X, in Cystopsal- 3.7 mm; mesonotum length d: 6.0-7.8 mm (x7.0 tria 1.9-2.2 X width of frontal ocellus. Prono- mm), 9: 6.8 and 8.0 mm; head width d; 6.7-7.7 tum with grooved and pitted surface and distinct mm (x7.5 mm), 9: 8.0 and 8.1 mm; width of medial furrow. Amplified lateral corners of pronotal collar d; 7.6-8.8 mm (8.5 mm), 9: 8.8 pronotal collar forming an inflated crest along and 9.2 mm. anterior margin, which is continuous with the sharply edged anterolateral margin of prono- Distribution. Endemic to the south-eastern cor- tum. Mesonotum with a more smooth surface ner of the Cape York Peninsula, north-eastern and with very narrow cruciform elevation, nar- Old. Moulds (1990) recorded the species from rower than long across its centre. Fore femur Mt Hartley approximately 30 km south of (fig. 60) with row of 3 spines, diminishing in Cooktown to the Kirrama Range north-west of length towards tibia, most proximal spine Cardwell. strongly bent and adjacent to femur, reaching to Remarks. The species can be recognized by about half-way the distance to middle spine (see slightly bronzed tegmina with 9 apical areas. arrow). Tegmina green, opaque rusty brown or Males are characterised by short lobate claspers greenish tinged in museum material, fairly slen- and females are easily recognized by the sharp, der and pointed at apex. Venation reticulate in thorn-shaped, protuberance at the ventral mar- distal half of tegmen. This reticulation includes gins of the pygofcr (fig. 47). The local name of ulnar areas in Cystopsaltria, but Cystosoma has this species is Frog Cicada (Moulds. 1990). normally developed ulnar areas. Costal area and border along hind margin of tegmen very nar- Cystosoma and Cystopsaltria row. Wings hyaline, in Cystosoma with 7-10, in Description. Body reddish brown, without any Cystopsaltria with 12-14 apical areas, often distinct colour markings. Surface of head and somewhat reticulate near wing apex and with pronolum roughly wrinkled and pitted. Head fairly broad hyaline border along hind margin. and pronolum eonically shaped; anterior mar- Tymbal with 6-10 transverse sclerotizcd ridges gins of postclypeus and vertex lobes forming an from dorsal to ventral margin. Male operculum almost continuous straight line with lateral mar- fairly small. Basal part of operculum hardly gin of pronotum, though interrupted by eyes. vaulted, but in males forming a large, often glo- Females smaller than males. Abdomen of males bularly swollen, protuberance at distolateral cor- strongly inflated, being 1.3-2.0 X head and tho- ner. Distal part of male operculum angularly PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

oblong, curved to close against the body and buted to Chlorocystini (sensu stricto). Cystos- covering most of tymbal cavity in ventral view. oma and Cystopsaltria have a weakly S- curved Opercula widely separated medially, by broad aedeagus with winged lateral crests, the synapo- and rounded first sternite. Meracanthus short, morphy of that tribe. The shapes of pygofer and not reaching distal margin of operculum. claspers also agree with such an allocation. Fur- Female operculum shorter than that of male, thermore, for the discriminating characters used with a more elongate lateral crest of basal part. by Moulds, the inflated male abdomen of Cys- Male abdomen very delicate and strongly tosoma and Cystopsaltria is larger but otherwise inflated, without distinct folds between tergites very similar to that of most species of Chloro- and sternites; almost circular in cross section. cystini and different from that of Hemidictya, First tergite in male fairly long, not hidden under and the head of Cystosoma and Cystopsaltria is metanotum. Proximal margin of second tergite not as notably different as that of several other convex medially and almost straight between genera of Chlorocystini. auditory capsules and sternite 2 in Cystosoma Hemidictya frondosa Burmeister, 1835 and (fig. 68), though weakly curved in Cystopsaltria Hovana distanli (Brancsik, 1893), the two (fig. 82) and forming a fairly distinct ridge along remaining species of the Hemidictyini, are tymbal cavity. Lateral parts of 2nd tergite undoubtedly closely related and probably sister weakly swollen at anterior margins and adjacent species, interesting from a biogeographical point

to tymbals. Sternites 1 and 2 adjacent. Auditory of view, since the former comes from Brazil and capsules in males weakly developed, hardly pro- the latter from Madagascar. These two are truding, but distinctly elevated relative to con- characterised by (1) very broad, opaque yellow- necting bar between tymbal and abdomen. green and apically pointed tegmina with reticu- Female abdomen more robust than that of male, late venation in the distal halves, (2) a costa but much smaller with short and broad genital strongly widened and flattened in its proximal segment. Ovipositor sheaths not reaching to half, (3) a distally elongate and sharply pointed apex of bluntly rounded caudodorsal beak (fig. mesonotum reaching to the 2nd abdominal seg- 63). Male pygoferglobularly rounded, with short ment, and (4) a strongly streamlined head and caudodorsal beak. Beak not curved over basal pronotum, with anterior margins of postclypeus part of claspers or anal valves. Lateral lobes of and vertex lobes forming a nearly straight and pygofer curved inwards and forming bluntly almost continuous line with margins of the eyes rounded lateral protuberances. Claspers broad and lateral edges of pronotum. The last charac- at base and hook-shaped; sharply curving down ter is also found in the African genus Lacetas at half-length. Apical part of clasper with large, indicating that Lacetas should possibly be sharply edged, clasper hollow. Claspers weakly included in Hemidictyini. diverging towards rounded apices. Basal parts of Cystosoma and Cystopsaltria share opaque claspers forming a continuous but very low ring- apically pointed and reticulate tegmina and a shaped collar around base of anal valves. Aed- sharp lateral edge of the pronotum with Hemid- eagus weakly S-curved. Aedeagal pore round. ictya and Hovana. Whether reticulate tegmen venation can be regarded synapomorphic is dif- Tribal placement. Distant (1905) included Cys- ficult to decide since its extent varies between general aspect the iosoma in his division Hemidictyaria (presently the four genera. However, in are very dif- tribe Hemidictyini) which has as its principal tegmina of Hemidictya and Hovana character a narrow head. This division also con- ferent from those of Cystosoma and Cystopsal- broader, tained Hemidictya Burmeister, 1835, Havana tria. Tegmina of the former two are have costa. while Distant, 1905 and seven genera that were later more squarish and a widened fuse well before reaching brought into the tribe Prasiini (e.g., Arfaka the veins CuA and M Distant, 1905; lruana Distant, 1905; Jacatra the basal area. sharp lateral edge of pronotum is defi- Distant. 1905; Lacetas Karsch, 1890; Lembeja The Sapantanga nitely different, sharper, in Hemidictya and Distant, 1 905: Prasia Stal, 1 863 and Head and pronotum of these two Distant, 1905). Recently Moulds (1990) trans- Hovana. species are more strongly streamlined than in ferred Cystopsaltria from Chlorocystini to Hem- Cystosoma and Cystopsaltria; even the eyes are idictyini based on the narrow head, inflated flattened and contribute to this streamline. A male abdomen and reticulate tegmina, charac- lateral pronotum edge similar as in Cystosoma ters very similar to Cystosoma. Cystopsaltria was found in several other A study of male genitalia, however, shows that and Chlorocystini (e.g., Mirabilopsaltria Cystosoma and Cystopsaltria should be attri- species of 14 A.J. DE BOER

viridicata (Distant, 1897), M. humilis (Blote, — DeJong. 1982: 182. — Duffels and Van der Laan 1985: 315. — Moulds, 1990: 192. — De Boer, 1990: 1 960), Gymnotympana monlana Dc Boer, 1 995.

1 1 1 9. 64. — De Boer, 1 992a: 1 64. — De Boer. 992b: 8, G. olivacea Distant, 1905 and G. verlaani Dc — — De Boer, 1 993a: 1 6- 1 7. — De Boer, 1 993b: 142 Boer, 1 995; see De Boer, 1 995a; De Boer, 1 995b, De Boer, 1995a: 8, 12, 16. — Dc Boer. 1995b: 3. 1996). saundersii Westwood. Study of male genitalia of Hemidictya fron- Type species. Cvstosoma ;/avtf (unpublished) does not suggest relationship 1842. to Chlorocystini since there are considerable dif- Diagnosis. Head with flattened vertex and large, ferences in shapes of pygofer, claspcrand aedea- angularly protruding, postclypeus. Pronotum gus between species of that tribe. with distinct medial fissure and rough, pitted, Mottophyly. Cystosoma and Cystopsaltria are surface. Tegmina opaque green, apically easily recognized by the following synapomor- pointed. Tegmen venation reticulate, with many phies: apically pointed tegmina and venation cross veins in apical areas, ulnar areas of normal reticulate in the distal halves of tegmina ( 1 and 2 shape. Veins CuA and M fusing at corner of in fig. 56). The hyaline wings too, tend to be basal area. Male abdomen strongly inflated. somewhat reticulate towards their apices and Remarks. Cystosoma is presumed to be mono- have more than 6 apical areas. The species phyletic, the fusion of the veins CuA and M at further share: 1, a very stout and angularly pro- the corner of the basal area is the supposed apo- truding postclypeus, in lateral view not unlike morphy for the genus. that of rhaumastopsaltria, but more strongly protruding and often with convexly swollen Cystosoma saundersii Westwood sides; 2, a sharp lateral edge of the pronotum in Figures 57-68 dorsal view, though interrupted by the eyes, almost continuous with the anterior margins of ( "tcada saundersii Westwood, 1 842: 1 1 8. postclypeus and vertex lobes; and 3, a very slen- Cystosoma saundersii. — Westwood, 1842: 118. — der cruciform elevation on the mesonotum, nar- Coding and Froggatt, 1 904: 566. 662, 663. — Distant, rower than long across the centre. These charac- 1906: 185. — Burns. 1957: 670. — Metealf, 1963: 435. — Duffels and Van der Faan, 1985: 315. Moulds, ters also occur in Prasiini, the presumed sister —

I 990: 1 93-- 1 pi. 23 figs I , I a-b. — De Boer, 1 995b: group of Chlorocystini and might therefore be 96, 5. plesiomorphic. Cvtosoma [sic] saundersii. — Musgrave, 1953: 13, 1

C. saundersii fig. (yslosomu laundersit [sic]. — Schrenimer, 1957: C. schultzi 19. Fig. 11, 46. C. immaculata ~eE Material examined. Australia. Australia, Saunders, cf, Figure 56. Cladogram of the species of Cystosoma and cf ( yslosomu saundersii det. Edm. Schmidt, IZW. Bris- Cystopsaltria. Numbers refer lo apomorphies dis- bane, 1973, J. B. Vogel, 3tf, ZMA; Brisbane. Kenmorc. cussed in the text. xii.1972, J.B. Vogel, 3rf, 9, ZMA. Cairns, 9 km W., ii-

iv. 1 985, 9. ZMA. Cunninghams Cap, 700-750 m. 20- Phytogeny. Although Cystosoma schultzi and 30.xi.1963, J.L. Cressitt,

1 Metealf. I 963: 433-434. 85. — — Boulard. 1 979; 46. hardly elevated, ocelli somewhat pressed PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 115

Figures 57-68. Cystosoma saundersii Westwood, 1 842: 57, pygofer in oblique view, Tully; 58, pygofer in lateral view, Tully; 59. male caudodorsal beak in dorsal view, Tully; 60, male fore femur in lateral view, Tully, arrow indicating strongly bent proximal spine; 61, aedeagus in dorsal view, Brisbane; 62, aedeagus in lateral view, Brisbane; 63. female genital segment in lateral view, Cunningham's Gap; 64, female caudodorsal beak in dorsal view. Cunningham's Gap; 65, female caudodorsal beak in dorsal view, Brisbane; 66, clasper. Tully; 67, female operculum. Brisbane; 68, male operculum, Tully. 116 A. J. DE BOER inward. Surfaces of vertex and dorsal part of almost rectangular distomedial corner and postclypeus very rough, covered with small pits, broadly rounded distolateral corner. and anterior parts of vertex lobes wrinkled. Vertex Abdomen. Ochraceous or green and postclypeus with distinct medial fissure. unmarked. Male abdomen strongly, almost glo- Vertex with semicircular furrow around frontal bularly inflated, distinctly broader than thorax ocellus, but without diverging fissures between and strongly convex dorsally; higher than thorax ocelli. Postclypeus distinctly swollen ventrally, in lateral view. First tergite not extending under tergite anterior margin (lateral view) angularly convex metanot um, more than half as long as 2nd at about half-length. Sides of postclypeus with middorsally. Auditory capsules weakly inflated elevated relative to connecting bar 1 1 weak furrows and a smooth and narrow, but distinctly weakly inflated, crest along lorum. between tymbal and abdomen. Tymbal cavity Thorax, Pronotum pitted, smooth in medial very narrow; auditory capsules fairly close furrow and with long wrinkles across collar. together and tergite part between auditory' cap- Mesonotum smooth and unmarked. sule and 2nd sternite very short. Second tergite Tegmina and wings: Tegmina opaque green- straight between auditory capsule and 2nd brown and pointed at apex. Basal half of tegmen, sternite and forming a fairly distinct crest along including ulnarareas, with regular venation, dis- tymbal cavity. Female abdomen long and slen- tal half of tegmen reticulate, with many cross der, with conically protruding auditory capsules. veins. Veins M and CuA fused at, or very close Female pygofer (fig. 63) short and weakly to, corner of basal area (cf. fig. 81). Wings swollen, in lateral view with weakly convex dor- hyaline, with irregular venation, 7 or 8 apical sal margin. Ovipositor sheaths not reaching to areas and 2 or 3 subapical areas. Apex of wing apex of caudodorsal beak. Female caudodorsal tending to be reticulate, with several cross beak in dorsal view swollen triangular and veins in 1st, 2nd and sometimes 3rd apical pointed at apex (fig. 65) but rounded in speci- area. men from Cunningham's Gap, southern Qld Legs. Fore femur (fig. 60) with 3 spines, most (fig. 64). proximal spine strongly bent, adjacent to Male genitalia. Pygofer in lateral view as in fig. femur. 58. Dorsal margin weakly convex, almost Tymbals, Large, in lateral view covering more straight, continuous with straight caudodorsal than half of body width. 5 transverse sclerotized beak. Distal margin weakly convex, concavely ridges spanning the tymbal from dorsal to ven- bent into straight margin of beak. Lateral lobe of tral margin and a 6th most proximal ridge span- pygofer slightly curving inwards and forming a ning about three-quarters of tymbal width. 6 weakly developed, slightly swollen and rounded distinct intercalary ridges form a lateral band lateral protuberance. Ventral margin forming a across tymbal. broad and rounded corner just below this pro- Opercula, Male operculum (fig. 68). Basal part tuberance and angularly convex, but concave of operculum with a knobby lateral protuber- towards pygofer base. Pygofer globularly ance at distolateral corner and weakly vaulted, swollen; pygofer opening very broad, broadest lateral vaulting almost absent. Distal part of between lateral protuberances of pygofer lobes, operculum rather large relative to basal part, ventral margins of pygofer converging to a con- almost as long as wide and almost completely vex basal margin at base of pygofer opening; covering tymbal cavity in ventral view. Disto- ventral part of pygofer opening broadly U- medial corner of operculum extending medially shaped (fig. 57). Caudodorsal beak in dorsal beyond meracanthus. Lateral margin long, run- view (fig. 59) very stout and broadly rounded at ning straight to base of protuberance at distola- apex. Gasper (fig. 66) broad, square-shaped at teral corner of basal part and convexiy bent into base, with stout triangular, downwardly directed straight and about equally long, distal margin. apical part, curving inwards distally of aedeagus Distomedial corner blunt, medial margin and forming an angular dorsodistal corner sup- straight and slightly directed mesiad. Meracan- porting aedeagus in upright position. Claspers thus reaching to about three-quarters of opercu- strongly diverging towards rounded apices. Api- lum length. Female operculum (fig. 67) narrower cal part of claspcr with distinct, sharply edged and much shorter than in male. Basal part of clasper hollow. Aedeagus (fig. 62) very stout and female operculum about as long as in male, but almost straight, slightly swollen and weakly more distinctly vaulted and with long and dis- upcurved at three-quarters of its length, but tinct crest along rectangular distolateral corner. recurving towards apex. Aedeagus with very Distal part very short and erect, oblong, with slender lateral crests and a pair of broad and 1

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 117

rounded dorsal ridges (fig. 6 ). 1 Aedeagal pore smooth in medial furrow and with long wrinkles broad and rounded. on collar. Mesonotum smooth and unmarked. Measurements (mean ± sd): Bodv length d; Tegmina and wings: Tegmina opaque green- 42.0-51.0 mm (47.2 mm ± 2.6), 9: 30.0-33.8 brown and pointed at apex. Basal half of tegmen, mm (31.3 mm ± 1.4); tegmen length d; 40.3- including ulnar areas, with regular venation, api- 47.8 mm (43.9 mm ± 2.4), 9: 37.7-42.2 mm cal half reticulate, with many cross veins. Veins (40.2 mm 1.7); head length d; 3.2-3.9 ± mm (3.5 M and CuA fused at corner of basal area (fig. 8 1 ). mm), 9: 3.4-3.9 mm (3.6 mm); pronotum length Wings hyaline, with almost regular venation and d: 4.2-4.8 mm (4.6 mm), 9: 4.2-4.5 mm (4.4 7 apical areas. A fairly narrow hyaline border mm); mesonotum length d; 7.0-8.3 mm (7.8 along hind margin of wing, though distinctly mm), 9: 6.6-8.1 mm (7.4 mm); head width d; broader than opaque border of tegmen. 7.1-7.8 mm (7.5 mm), 9: 6.8-7.8 mm (7.4 mm); Tymbals. Large, in lateral view covering more width of pronotal collar d: 10.8-11.9 mm (1 1.4 than half of body width. 6 transverse sclerotized mm), 9: 9.7-1 1.5 mm (10.7 mm). ridges spanning the tymbal from dorsal to ven- tral margin and a 7th most proximal ridge span- Distribution. Eastern Qld and north-eastern ning about three-quarters of tymbal width. 7 NSW. Moulds (1990) recorded the species from distinct intercalary ridges form a lateral band the Atherlon Tableland in northern Qld, the across tymbal. Clarke Range, Eungella plateau and Mackay in Opercula. Male operculum (fig. 71) with Central Qld, inland at Carnarvon Range and weakly vaulted basal part as in C. saundersii. from Kroombit Tops in southern Qld to Syd- Distal part of operculum very different from ney. that of C. saundersii, oblong and rather large Remarks. Cystosoma saundersii is the largest relative to basal part, completely covering tym- species of Chlorocystini, easily recognized by its bal cavity in ventral view and distinctly extend- enormously inflated abdomen, hence the local ing medially beyond meracanthus. Lateral mar- name "Bladder Cicada" (Moulds, 1990). gin long, gradually and concavely bending into crest around distolateral corner of basal part and Cystosoma schmeltzi Distant convexly bending into long and weakly convex distal margin. Distomedial corner broadly Figures 69-71, 73-75, 81 rounded, medial margin straight. Meracanthus Cystosoma schmeltzi Distant, 1882: 32. pi. vii figs reaching to about two-thirds of operculum 11,1 la-b. — Coding and Froggatt, 1904: 566: 662. length. Female operculum as in C. saundersii 664. — Distant, 1906: 185. — Burns, 1957; 670. — with erect and oblong distal part (cf. fig. 67). Metcalf, Duffels and Van der Laan. 1963: 436. — Abdomen. Male abdomen ochraceous to 1985: 316. — Moulds, 1990: 192-193, pi. 23 figs 2. green and unmarked, distinctly inflated, but not 2a. as globular as in C. saundersii; with dorsal mar- Material. Australia. Dama, Cape York, 9, ZMB; Gayn- gin (lateral view) not distinctly elevated relative ZIM. dah, Mus Godefroy No 17630, 9 paratype, to dorsal margin of thorax. First tergite not hid- Wacol, 28.ii.1970, H. Sas, rf, RMNH. den under metanotum and medially only slightly Description. Body yellow-green but green when shorter than 2nd tergite. Auditory capsules alive and unmarked (for photographs see weakly developed, hardly protruding and not distinctly elevated Moulds, 1990: pi. 23 figs 2, 2a). Male abdomen visible in dorsal view, but 0.9- relative to connecting bar between tymbal and 1 .4 X head and thorax together, in females cavity very narrow; auditory 1.0 x. Tegmina of male 1.0 X body length, of abdomen. Tymbal fairly together and tergite part females 1.4 x. capsules close auditory capsule and 2nd sternite very Head (fig. 69): Ochraceous with traces of olive between auditory green, vertex lobe with blackish spot between short. Second tergite straight between 2nd sternite and forming a fairly eye and lateral ocellus. Head heavily wrinkled capsule and crest along tymbal cavity. Female geni- between ocelli and on anterior parts of vertex distinct short and weakly swollen as in C. lobes, vertex with distinct medial fissure. Post- tal segment Ovipositor sheaths not reaching to clypeus distinctly swollen ventrally, anterior saundersii. of caudodorsal beak. Female caudodorsal margin (lateral view) angularly convex at about apex in dorsal view swollen triangular and half-length. Lateral parts of postclypeus with 1 beak weak furrows and a smooth and narrow, weakly pointed at apex. Male genitalia. Pygofer in lateral view as in fig. inflated, crest along lorum (fig. 70). margin concave near base, but Thorax. Pronotum pitted as in C. saundersii, 73. Dorsal Figures 69-77. ( 'ysiosoma schmeltzi Distant, 1 882: 69, male head in dorsal view; 70, male postclypeus in lateral view; 71, male operculum; 72, base of right tegmen, male (Cystopsaltha immaculata); 73,pygofer in lateral view; 74. male caudodorsal beak in dorsal view; 75, pygofer in oblique view; 76, clasper; 77, aedeagus in lateral view. PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 1 19 strongly convex to apex of stout and short cau- Cystopsaltria immaculata dodorsal beak. Distal margin straight, forming Goding and Froggatt an obtuse angle with margin of beak, angularly Figures 72, 78-80, 82-87 bending outwards at distal end. into weakly developed, slightly swollen and rounded lateral Cystopsaltria immaculata Goding and Froggatt. protuberance. Ventral margin weakly convex, 1904: 566, 661. pi. xvii figs I. la. — Distant. 1906: but forming a broad and rounded inwardly 1 60. — Burns, 1 957: 644-645. — Metcalf. 1 963: 260. curved corner just below this protuberance. — Duffels and Van der Laan, 1985. 250. — Moulds. 1990: 197-198, pi. 23 figs Pygofer globularly swollen; pygofer opening 3, 3a. very broad, broadest between lateral protuber- Material. Australia. Cairns. 9 km W.. ii-iv.1985. 39. ances of pygofer lobes. Ventral margins converg- ZMA. Kamcrunga near Cairns, N. Qld., lO.i. 1977. ing to a very short and convex basal margin at M.S. and B.J. Moulds, ef, ZMA. Mt Windsor Table- base of pygofer opening; ventral part of pygofer land. NW of Mossman, 30.xii.l980. M.S. and B.J. Moulds, 9 Cystopsaltria immaculata G. and F. det. opening U- shaped (fig. 75). Caudodorsal beak M.S. Moulds. ZMA. in dorsal view (fig. 74) very stout and broadly rounded at apex. Claspers (fig. 76) parallel to Description. Bodv light brown (for photographs rounded apices, broad, square-shaped at base, see Moulds, 1990: pi. 23 figs 3, 3a). Male abdo- with long and slender, downwardly directed api- men strongly inflated 1.3 X head and thorax cal part, curving inwards at angular dorsodistal together, in females 1.0-1.1 X.Tegminaofmale corner and supporting aedeagus in upright pos- 1.4 X body length, in females 1.3-1.4 X. ition. Claspers fused at base to low collar around Head. As in Cystosoma, but with somewhat base of anal valves. Apical part of clasper with narrower vertex; ocelli more closely together. distinct but slender and sharply edged clasper Distance between lateral ocelli 1.9-2.2 X width hollow. Aedeagus (fig. 77) slightly upcurved, but of frontal ocellus. recurving near apex, with very slender lateral Thorax: Pronotum less distinctly pitted than crests and pair of broad rounded dorsal ridges. in Cystosoma, in male smooth, but with a dis- Aedeagal pore broad and oval. tinct medial furrow.

Measurements. Body length d; 27.6 mm, 9: Tegmina and wings. Tegmina opaque gree-

21.0 and 23.5 mm; tegmen length d: 28.2 mm, 9: nish brown and reticulate as in the 2 foregoing 29.0 and 32.3 mm; head length d: 2.7 mm, 9: 2.7 species, but with very different pattern of veins. 1 Distinctly cross veins and larger part of and 2.8 mm; pronotum length d ; 3.6 mm, 9: 2.8 more and 3.0 mm; mesonotum length d; 6.4 mm, 9: 6.2 tegmen reticulate; no regular ulnar areas. Veins and 6.3 mm; head width: 5.4 mm, 9: 5.3 and 5.5 M and CuA reach basal area separately. Radial mm; width of pronotal collar d: 8.0 mm, 9: 7.5 area divided by extra vein, parallel to costa and and 7.9 mm. fusing with vein M near corner of basal area (fig. Distribution. Eastern Qld and northern NSW. 72). Tegmen with distinct, almost hyaline, cor- Moulds (1990) recorded the species from Moss- dial fold. Wings hyaline, with irregular venation man Gorge and Forty Mile Scrub, to inland as in C saundersii, but with more (12-14) apical northern NSW south to Gunnedah. areas. Apex of wing tending to be reticulate, with several cross veins, especially near apex of first 4 is distinctly smaller than Remarks. C. schmeltzi or 5 apical areas. C. saundersii and has a less strongly inflated Tymbals. Large, covering more than half of is "Lesser Bladder abdomen. The local name body width in lateral view. 8 transverse sclero- Cicada" (Moulds, 1990). tized ridges spanning the tymbal from dorsal to Cystopsaltria Goding and Froggatt ventral margin, a 9th ridge spanning about three-quarters of tymbal width and a 10th most 1904: 559. 566. Cystopsahria Goding and Froggatt, proximal ridge reaching to about half of tymbal 661. — 1906: 154, 160. — Metcalf, 1963: Distant, width. 9 weak intercalary ridges form a lateral 260. — Duffels and Van der Laan, 1985: 250. — band across tymbal. Moulds. 1990: 196-197. — De Boer. 1992b: 18-19. — 142.— De Opercula. Male operculum (fig. 82) resem- DcBoer, 1993a: 1 6- 1 7. — De Boer, 1993b: bling that of €. schmeltzi. Distal part of opercu- Boer, 1995a: 8. — De Boer, 1995b: 3. lum oblong and rather large relative to basal immaculata Goding Type species. Cystopsaltria part, covering greater part of tymbal cavity in and Froggatt, 1904. ventral view and distinctly extended medially mcracanthus. Lateral margin long, Remarks. Cystopsaltria is a monotypic genus, beyond obtuse angle at about third its length. closely related to Cystosoma. forming an A. J. DE BOER

Figures 78-87. CystopsaltriaimmaculataGodingand Froggatt, 1904: 78, pygoferin oblique view; 79, pygoferin lateral view; 80, male caudodorsal beak in dorsal view; 81, base of right tegmen, male (Cystosoma schmeltziy, 82, male operculum; 83, female caudodorsal beak in dorsal view. Cairns; 84, clasper; 85, aedeagus in lateral view; 86, aedeagus in oblique view; 87, female operculum, Cairns. PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 121 angularly bending into crest around distolateral tegmen length Cicadidae. tinct but slender and sharply edged clasper hol- Proceedings of the Roval Society of Victoria (n.s.) low. Aedeagus (figs 85, 86) very stout, swollen 24: 221-229, pis xlix-li. Ashton, H., 1914. Catalogue of the Cicadidae in the and slightly upcurved in proximal half, but more South Australian Museum, with descriptions of slender in distal half and recurving near apex. several new species. Transactions of the Royal Aedeagus without distinct lateral crests, but with Society ql South Australia 38: 345-358. pi. xvii. a dorsal ridges, ending in pair of broad rounded Boer. A.J. dc. 1990. Aedeastria, a new cicada genus weakly outcurving subapical appendages. Aed- from New Guinea, its phylogeny and biogcogra- eagal apex incised. phy (Homoptera, Tibicinidae), preceded by a dis- Measurements (mean ± sd): Body length <$: cussion on the taxonomy of New Guincan Tibi- Bcaufortia 40 (3): 63-72. 35.0 mm, 9: 31.5-35.1 mm (33.7 mm ± 1.4); cinidae. 122 A.J. DF BOER

basin evolution in Indonesia. Marine and Pet- Boer. A.J. de. 1 99 1 . Seottotympana, a new cicad genus and 1-21. from New Guinea, with the description of three roleum Geology (MPGD) 8(1): new species, their taxonomy and biogeography Distant. W.L.. 1882. On some undescribed Cicadidae (Homoptera, Tibicinidae). lieaufortia 42 (1): I- from Australia and the Pacific Region Proceed- London 1881: I 1. ings of the Zoological Society of 7-14. Boer. AJ.de, 1 992a. The taxonomy and biogeography 125-134, pi. 7 figs of the viridis group of the genus Baeturia Stal. Distant, W.L.. 1905. Rhynchotai notes. — XXXIV. 1866 (Homoptera. Tibicinidae). Bijdragen tot de Annals and Magazine of Natural History (7) 16:

Dierkunde i>\ (3): 163-183. 203-216.

Boer, A.J. de, 1 992b. The taxonomy and biogeography Distant. W.L., 1 906. A synonymic catalogue 0)'Homop-

of the genus Thaumasiopsaliria Kirkaldy. 1900. tera. Part I. Cicadidae. Trustees British Museum: 7- (Homoptera. Tibicinidae). Hemitonia 43 (3): 1 London. 44. Distant, W.L.. 1 907. Rhynchotai notes.- XLIII. Annals Boer. A.J. de. 1993a. Guineapsaltria. a new genus of and Magazine of Natural History (7) 20: 41 1- the Australian-New Guinean region (Homoptera, 423 Tibicinidae). with notes on its taxonomy and bio- Duffels, J. P., 1 977. A revision of the genus Diceropyga geography. tot dc 63(1): 15- Bijdragen Dierkunde Stal. 1870 (Homoptera, Cicadidae). Monogra- 41. fieen van de nederlandse entomologisehe Verenig- Boer, A.J. de. 1993b. Ten species added to the genus ing 8: 1-227. Aedeasiria De Boer. 1990, with the description of Duffels, J. P., 1986. Biogeography of Indopacific eight new species and notes on the taxonomy and Cicadoidea. a tentative recognition of areas of biogeography (Homoptera, Tibicinidae). Beaufor- endemism. Cladisties 2 (4): 318-336. tia 43 (9): 140-167. Duffels. J. P. and Boer. A.J. dc, 1990. Areas of endem- Boer. A.J. de. 1 994a. The taxonomy and biogeography ism and composite areas in East Malesia. Pp. of the loriae group of the genus Baeturia Stal. 1 866 249-272 in: Baas. P.. Kalkman, C. and Geesink. (Homoptera. Tibicinidae). Tijdsehrifi voor Ento- R. (eds). The plant diversity of East Malesia: Pro- mologie 137: 1-26. ceedings ofthe Flora Malesiana symposium com- Boer. A.J. de. 1 994b. The taxonomy and biogeography memorating Professor Dr. C. G.G.J, van Steenis, of the exhausta group of the genus Baeturia Stal, Leiden. August 1989. Kluwer Academic Publish- 1866 (Homoptera, Tibicinidae). Beaufortia 44 ers: Dordrecht. (5): 127-158. Duffels. J. P. and van der Laan. P. A.. 1985. Catalogue Boer. A.J. dc. 1994c. The taxonomy and biogeography of the Cicadoidea (Homoptera. Auchenorhynga) of the guttulinervis group of the genus Baeturia 1956-1980. Series Entomologiea 34: i-xvi, 1- Stal. 1866 (Homoptera. Tibicinidae). Bijdragen 414. tot de Dierkunde 64 (2): 87-100. Froggatt, W.W.. 1907. Sub-order ii.- Homoptera Aus- Boer. A.J. de. 1995a. The taxonomy phylogcny and tralian insects. William biogeography of the cicada genus Gymnotympana Brooks and Co.:Sydney. xiv + 449 pis 1-37. Stal. 1861. (Homoptera. Tibicinidae). Invert- pp.; ebrate Taxonomy 9(1): 1-81. Coding. F.W. and Froggatt. W.W., 1904. Monograph of the Australian Cicadidae. Proceedings of the Boer. A.J. de. 1 995b. The taxonomy and biogeography 18- of the cicada genus Papuapsaltria gen. n. (Homop- Linnean Society ofNSW 29 (3): 561-670; pis tera, Tibicinidae). Tijdsehrifi voor Entomologie 19. 138: 1-44. Hamilton, W.B., 1979. Tectonics of the Indonesian Boer, A.J. de. 1995c. Islands and cicadas adrift in the region. United States Geological Survey Profe- West-Pacific: biogeographic patterns related to sional Papers 1078: i-ix. 1-345. plate tectonics, lijdschrift voor Entomologie 138: Holloway, J.D.. 1979. A survey of the Lepidoptera, 169-244. biogeography and ecology of New Caledonia. Boer, A.J. de. 1995d. The phylogcny and taxonomic Series Entomologiea 15: i-xii, 1-588. status of the Chlorocystini (Homoptera, Tibicini- Holloway. J.D., 1984. Lepidoptera and the Melane- dae). Bijdragen tot de Dierkunde 65: 201-231. sian Arcs. Pp. 129-169 in: Radovsky, F.J., Raven, Boer. A.J. de, 1996. Mirabilopsaltria a new cicada P.H. and Sohmer. S.H. (eds), Biogeography ofthe genus from New Guinea, its taxonomy and bio- tropical Pacific, proceedings of a symposium. geographv. (Homoptera. Tibicinidae). Tropical Association of Systematics Collection: /.oology 9: 349-379. Lawrence. Boulard. M.. 1979. Revision dc la faune cicadeenne Jong, M.R. de. 1982. The Australian species of the des lies Maurice et Rodrigucs. Bulletin de la genus Lembeja Distant 1892 (Homoptera, Tibi- Sociele Enlomologique de France 84: 27-47. cinidae). Bijdragen tot de Dierkunde 52 (2): 1 75- Burns, A.N., 1957. Check list of Australian Cicadidae. 185. Enlomologi.se/ie. trheilen auddem Museum Georg Jong, M.R. de. 1985. Taxonomy and biogeography of Erey 8 (2): 609-678. Oriental Prasiini 1: The genus Prasia Sal. 1863 Daly. M.C.. Cooper. M.A.. Wilson. I., Smith. D.G. and (Homoptera. Tibicinidae). Tijdsehrifi voor Ento- Hooper. B.G.D.. 1991. Ccnozoic plate tectonics mologie 128: 165-191. PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 123

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