Pauropsalta annulata Goding and Froggatt, 1904

POPPLE, L. W., 2013, <p class = " HeadingRunIn " align = " left "> <strong> A revision of the <em> Pauropsalta annulata </ em> Goding & amp; Froggatt species group (Hemiptera: Cicadidae) based on morphology, calling songs and ecology, with investigations into calling song structure, molecular phylogenetic relationships and a case of hybridisation between two subspecies </ strong> </ p>, Zootaxa 3730 (1), pp. 1-102 : 12-22

Pauropsalta annulata Goding and Froggatt, 1904

( Figs 2A , 3 4A–B , 5A–B , 6A–B , 7–9 )

Pauropsalta annulata Goding and Froggatt, 1904: 620–621 ; Metcalf, 1963: 405; Ewart, 1986: 51; Ewart, 1988: 181, 196–197; Ewart, 1989: 349–355; Moulds, 1990: 139–140.

Pauropsalta annulata song type A: Popple et al. (2008); Popple & Walter (2010).

Material: Lectotype ♂: ‘ Southport / Queens [land]’, ‘ 16 Oct. 1901 ’, ‘ W. W. Froggatt’ (ex. Macleay Museum , Sydney 'on permanent loan' to Australian National Insect Collection , Canberra ; designated by Ewart (1989)); other material: NEW SOUTH WALES: 1♀ Kyogle N.S.W., 7.xi.1983, at black light, Scambler ( MSM); 1♀ Wiangaree State Forest N.S.W., 10.xi.1983, at black light, Scambler ( MSM); QUEENSLAND: 2♂ 2♀ The Amphitheatre, NW. Robinson Gorge N.P. [Qld], 25°12.07’S 148°59.43’E, Open forest (camp), 17.xii.1997, [Ewart] ( AE); 1♂ 1♀ same as previous data, genitalia prep. 433-01 [ LWP] ( AE); 1♂ 1♀ Anstead Qld , 16.xii.1999, Coombs ( MC); 1♂ Armstrong Creek , Dayboro SE. Qld , 3.ix.1983, North ( MSM); 1♀ Atkinsons Dam SE. Qld , 27°28'00"S 152°27'00"E, 18.i.2003, Popple & Popple, m. v. lamp , 432-0035 ( LWP); 1♂ Bee C [ree]k, 25km SW. of Nebo Qld , 6.ii.1991, Moulds & Moulds ( MSM); 1♂ Beerwah Field Study Centre SE. Qld , 26°51'24"S 152°58'38"E, 29.x.2005, Popple , 432-0080; 1♀ [Redvale] 4km W. of Binjour SE. Qld , 25°31'00"S 151°25'00"E, 24–25.i.2003, Popple & Popple, m. v. lamp , 432-0031 ( LWP); 1♀ Birkdale , Brisbane [Qld] ( QM); 1♂ Base of Blackbutt Range W. of Moore SE. Qld , 26°53'13"S 152°12'49"E, 9.i.2002, Popple , Moulds , Moss , Cooley , Marshall & Hill, m. v. lamp , 431-0001 ( LWP); 9♂ 9♀ [Charlevue Creek causeway], below escarpment, Blackdown Tableland Qld , [16– 17]. xii.1985 ( AE); Moss & Ewart; 1♀ same data as previous, genitalia prep. 432-03 [ LWP] ( AE); 1♀ [Stony Creek car park], Blackdown Tableland , 13–20.x.1985, Moss & Ewart ( AE); 1♂ 1♀ Boomer Ra. N.P. NW. of Rockhampton C. Qld, camp, 23°12.88'S 149°44.37'E, 30.ix.1999, Ewart , Burwell , Monteith & Evans ( AE); 1♂ Brisbane Qld , 25.iv.1980, Yeates ( MSM); 1♂ Brisbane Qld , 27.ix.1979, North ( MSM); 1♂ Brisbane Qld , 21.ix.;1980, Semple ( MSM); 1♀ Brisbane Qld , 8.i.1983, South ( MSM); 1♂ Brisbane Qld , 4.x.1980, Campbell ( MSM); 1♂ Brisbane [Qld], 22.x.1912, Pauropsalta encaustica Germar (QM) ; 1♀ Brisbane [Qld] 16.x.1912, Hacker ( QM); 1♀ Brisbane [Qld] 12.i.1914, Hacker ( QM); 16.ii.1925, Hacker ( QM); 1♂ Brisbane [Qld], 8.ii.1922, Hacker ( QM); 1♂ Brisbane , 16.x.1923, Hacker , Pauropsalta annulata det. A. Ewart, 1997 ( QM); 1♀ Brisbane Q., Wild , iii.1890, HAN, Pauropsalta annulata G. & F. (abdomen glued upside down) ( QM); 1♀ Brookfield , Brisbane Qld , 10.xi.1980, Cribb ( AE); 1♂ Brookfield SE. Q., 19.xi.1985, Cribb ( AE); 2♂ 3♀ Brookfield Qld , 9.xii.1995, Coombs ( MC); 1♂ 1♀ same data as previous, 24.xi.1994 ( MC); 1♂ 1♀ same data as previous, 5.i.1994 ( MC); 1♀ same data as previous, 15.xii.1993 ( MC); 1♀ same data as previous, 7.xi.1994 ( MC); 1♂ Brookfield , Brisbane Qld , 19.i.1990, Cribb ( MSM); 1♀ 24.5°S 152.2°E, Bundaberg Qld , xii.1970, Frauca , on tea tree ( ANIC); 1♂ Burnett C [ree]k ( Mt Maroon ) [Qld], 16.xi.1975, half way up Casuarina [tree], on ridge, [Ewart] ( AE); 1♂ Burswood , Bardon [Qld], x.1995, Emery ( DE); 1♂ 3♀ Camp Hill SE. Qld , [27°30'S 153°04'E], 27.x.1998, Popple , at light, 432-0002 & 432-0007 to 432-0009 ( LWP); 1♂ Camp Hill SE. Qld , [27°30'S 153°04'E], 6.xi.1998, Popple , m. v. lamp , 432-0001 ( LWP); 2♂ Carina Heights SE. Qld , [27°29'S 153°05'E], 11.xi.2004, Popple , 432-0043 & 432-0044 ( LWP); 1♀ Carina Heights SE. Qld , [27°29'S 153°05'E], 22.x.2002, Popple , 432-0029 ( LWP); 1♂ Carina Heights SE. Qld , [27°29'S 153°05'E], 22.ix.1998, Popple , 432-0012 ( LWP); 1♂ Carina Heights SE. Qld , [27°29'S 153°05'E], 3.x.2002, Popple , 432-0028 ( LWP); 1♀ Carina Heights SE. Qld , [27°29'S 153°05'E], 4.x.1998, Popple , 432-0003 ( LWP); 1♂ Carina Heights SE. Qld , [27°29'S 153°05'E], xii.2000, Popple & Popple, 432-0011 ( LWP); 1♂ Carina Heights SE. Qld , [27°29'S 153°05'E], x.2006, Popple , 432-0070 ( LWP); 1♂ Carindale SE. Qld , [27°30'S 153°06'E] x.2007, Popple , 432-0059 ( LWP); 2♀ Carnarvon Gorge , 8– 12.xii.1985, Moss & Ewart ( AE); 1♂ Chelmer [Qld], 28.x.1980, [Ewart] ( AE); 1♂ Chelmer [Qld], 2.xi.1980, [Ewart] ( AE); 1♂ Childers Qld , 25°13.8'S 152°20.7'E, 15.xii.2000, Maywald ( MC); 2♀ Coast Ra , 14km S. by E. of Biggenden Qld , 17.xi.1978, Frauca ; 1♀ Comet Qld , 19.iii.1958, Common ( ANIC); 1♂ 1♀ Cooyar park SE. Qld , 26°58'53"S 151°49'50"E, 7.xii. 2006, Popple & McKinnon, 432-0057 & 432-0074 ( LWP); 1♂ Darra SE. Qld , [27°34'00"S 152°57'00"E], x.2006, Lindsay , 432-0073 ( LWP); 2♂ SE. Q.: 27°12’S 152°49’E, 0.5km S. of Dayboro , 21.i.2001, Evans ( QM); 1♀ Doolandella [Qld], 28.xii.1981, [Ewart], genitalia prep. 432-02 [ LWP] ( AE); 1♂ Doolandella [Qld], Brisbane annulata, 9.ii.1983, [Ewart]; 1♂ Doolandella [Qld], 12.xii.1981, to light, [Ewart] ( AE); 2♂ Doolandella [Qld], 12.xii.1981, to light, [Ewart] ( AE); 1♂ 1♀ Doolandella [Qld], 9.ii.1982, [Ewart] ( AE); 1♂ Doolandella [Qld], 29.i.1982, [Ewart] ( AE); 1♂ 1♀ Doolandella [Qld], 9.ii.1982, [Ewart]; 1♀ Doolandella [Qld], 17.ii.1982, [Ewart] ( AE); 1♂ Doolandella [Qld], 8.ii.1983, [Ewart]; 1♂ Doolandella [Qld], 9.ii.1983, [Ewart] ( AE); 1♂ Doolandella [Qld], ii.1983, [Ewart]; 1♂ Doolandella [Qld], 14.x.1983, [Ewart] ( AE); 1♂ annulata, Doolandella [Qld], 5.ii.1983, [Ewart]; 1♂ 1♀ Doolandella [Qld], 9.ii.1982, [Ewart] ( AE); 1♂ Doolandella [Qld], 29.i.1982, [Ewart], genitalia prep. 432-02 [ LWP] ( AE); 1♀ Doolandella SE. Qld , 27.ii.1991, [Ewart] ( AE); 1♂ Doolandella S. Qld, 10–20.i.1996, [Ewart] ( AE); 1♂ 1♀ Doolandella SE. Qld , 28.i.1982, Ewart ( AE); 1♂ Doolandella , Brisbane Qld , 11.i.1985, Moulds & Moulds ( MSM); 1♀ Durack ( Brisbane ) SE. Qld , 27°36'S 152°58'E, xi.2007, Manners , 432-0063 ( LWP); 5♂ 8♀ AUSTRALIA Qld , Eidsvold Caravan Park , 25°22'06"S 151°07'25"E, 24–25.xi.2006, Popple & McKinnon, m. v. lamp , 432-0050 to 432-0056 & 432-0075 to 432-0079 ( LWP); 3♂ 6♀ Eidsvold SE. Qld , 25°22'06"S 151°07'25"E, 27–29.xii.2004, Popple , m. v. lamp , 432- 0045 to 432-0049 & 432-0064 to 432–0067 ( LWP); 1♂ Emerald Qld , 6.ii.1983, North ( MSM); 4♀ same data as previous, 8.xi.1983 ( MSM); 1♂ 1♀ QLD.EEM, approx. 2km E. of Emerald Qld , 23°32.2'S 148°10.3'E, 200m, 3.i.2004, Cooley , Hill , Marshall , Moulds ( MSM); 1♂ Fairbairn Dam via Emerald , 23°39'33"S 148°04'41"E, 9– 10.i.2007, Popple & Hereward, m. v. lamp , 432-0058 ( LWP); 1♂ Ferny Grove Qld , 15.xi.1993, Coombs ( MC); 1♀ Gayndah Qld , 8.i.1983, North ( MSM); 1♀ Qld : 28°03’S 153°07’E, ‘ Glen Witheren’ general, 26–29.xii.2001, Mv light, Monteith , 10352 ( QM); 2♂ Gold Coast [Qld], xii.1995, Emery ( DE); 1♀ Grave Gully , approx. 40km N. of Marlborough Qld , 3.ii.1973, Walford-Huggins & Walford-Huggins ( MSM); 1♂ 1♀ Gunalda Range , [26°00'S 152°34'00"E], 1.i.1994, Moss & Ewart, 432-0005 ( LWP); 1♂ 1♀ Gympie SE. Qld , 26°12'S, 152°40'E, 1.xi.2002, Popple , 432-0024 & 432-0025 ( LWP); 1♂ The Hummock , Bundaberg Qld , 15.xii.2000, Coombs ( MC); 2♂ 1♀ Brisbane ( Indooroopilly ) [Qld], 13.xii.1996, Emery ( DE); 1♂ Indooroopilly [Qld], 30.x.1996, Emery ( DE); 1♂ Indooroopilly 1.xii.1994, Coombs ( MC); 1♂ same data as previous 23.xi.1993 ( MC); 1♂ Indooroopilly , Brisbane Qld , 26.x.1971, North ( MSM); 1♂ Jamboree Heights , Brisbane Qld , 25.x.1990, Lambkin ( ANIC); 1♂ Jamboree Heights , Brisbane Qld , 30.ix.1984, Daniels ( MSM); 1♀ Fairmore St , Kenmore [Qld], 7.ii.1978, [Ewart] ( AE); 1♀ Kinbombi Falls near Goomeri, SE. Qld , 19.xii.1976, Moulds & Moulds ( MSM); 1♀ 38.4km SW. by W. of Kingaroy [SE. Qld], 21.xi.1968, Britton & Misko, m. v. lamp , 432-0017 ( LWP); 1♂ 2♀ Laidley SE. Qld , 27°38'S 152°24'E, 25.xii.2003, Beard , at light, 432-0040 to 432-0042 ( LWP); 3♂ Lake Dyer SE. Qld , 27°37'56"S 152°22'38"E, 22.ii.2003, Popple & MacSloy, 432-0032 to 432-0034 ( LWP); 1♂ 1♀ Lake Dyer SE. Qld , 27°37'56"S 152°22'38"E, 4.x.2003, Popple & MacSloy, 432-0036 & 432-0037 ( LWP); 2♀ Lamington N.P. [Qld], 28°17'S 153°03'E, 18.ix.1999, Maywald ( MC); 1♀ Lawnton , Brisbane Qld , 4.x.1981, North ( MSM); 1♀ Logan Village Qld , 26.i.1912, W.W. Froggatt collection ( ANIC); 1♂ 1♀ 0.5km along Marlong Plains Rd , Mt Moffatt N.P., 2[4]°57.70'S 147°59.86'E, 14.xii.2006, at light, Ewart ( AE); 1♂ Maroon Dam SE. Qld , 28°10'44"S 152°39'21"E, 2.x.2002, Popple , 432-0030 ( LWP); 1♀ McFarlanes Bridge , Brisbane Valley [Qld], 12.x.1975, Casuarina tree, [Ewart] ( AE); 1♂ Meadowlands Park , Carindale , 27°29'32"S 153°07'05"E, 24.x.1998, Popple & Popple, 432-0013 ( LWP); 1♂ 1♀ Minto Crags [South face] S. of Boonah SE. Qld , 28°07'S 152°36'E, 3.xi.2001, Popple , Moss & Stoodley, 432-0015 & 432-0016 ( LWP); 1♀ Moggill , Brisbane Qld , 19.x.1980, Ewart , P. annulata det. A. Ewart ( MSM); 1♂ red eyed annulata, Mon Repos, Bundaberg [Qld], 30.i.1983, [Ewart] ( AE); 2♂ Moogerah Dam [Qld], 23.x.1977, [Ewart] ( AE); 1♂ P. annulata (red eyed), Base Mt Coolum [Qld], 17.x.1981, recorded, [Ewart] ( AE); 1♂ (white eyed), Base Mt Coolum [Qld], 17.x.1981, recorded, [Ewart] ( AE); 1♂ Mt Coottha [Qld], 31.x.1996, Emery ; 1♀ Nerang [Qld], 13.xii.1996, Emery ( DE); 2♂ 1♀ Qld : 27°29’S 152°57’E, Mt Coottha 260m, 10.i.2002, Mv light, Monteith , open forest, 10350 ( QM); 3♂ 1♀ Mt Flinders Rd N. of Peak Crossing SE. Qld , [27°44'S 152°44'E], 12.xii.2002, Popple & Emery, 432-0018 to 432-0021 ( LWP); 1♂ 2♀ Mt Hay , 40km SW. of Rockhampton Qld , 27.xi.1986, Sattler ( MSM); 1♂ 2♀ Jct. of Triple Tracks , Mt Moffatt N.P., 15.xii.2005, at light, Ewart ( AE); 2♂ Mt Moffatt , 0.5km along Marlong Plains Rd , 25°57'42"S 147°59'52"E, 14.xii.2006, Ewart , at light, 432-0061 & 432-0062 ( LWP); 1♂ Mt Moffatt , Junction of Triple Tracks , 24°58'32"S 147°55'53"E, 15.xii.2006, Ewart , at light, 432-0060 ( LWP); 10♂ 7♀ Mt Moffatt N.P. C. Qld, Top Shelter shed, 100m, 10– 12.xii.1987, Monteith, Thompson & Yeates ( MSM); 1♂ Mt Moon Qld (near Boonah), 4.x.1980, Sneath ( MSM); 2♀ Mt Walsh base SE. Qld, 25°33'S 152°03'E, 1–2.xi.2002, Popple & Billington, m. v. lamp , 432-0022 & 432-0023 ( LWP); 3♂ Mundubbera (north), 25°35'03"S 151°18'14"E, 8– 9.i.2007, Popple & Hereward, m. v. lamp , 432-0081 to 432-0083 ( LWP); 1♂ Narangba, 22.x.1983, North ( MSM); 1♀ National Park Q., 24.x.1923, Hacker (damaged) ( QM); 1♀ National Park Q., 23.x.1923, Hacker ( QM); 2♂ 1♀ SE. Q.: 25°42’S 151°26’E, Nipping Gully site 6, 18– 19.xii. 1998, 200m, Monteith, Gough & Maywald, 7532 ( QM); 5♂ 2♀ Qld: 25°41'S 151°26'E, Nipping Gully site 1, 16.i. 2003, 300m, m. v. lamp , open forest, Monteith, 12399 ( QM); 1♀ Ormiston [Qld], 17.xi.1924, Hacker ( QM); 1♂ Oxley SE. Qld, [27°33'S 152°59'E], 2.xii.2003, Popple, 432-0039 ( LWP); 1♂ Redland Bay , SE. of Brisbane QLD, 10.i.1990, Knight ( ANIC); 1♂ St Lucia [Qld], 4.xii.1975, [Ewart] ( AE); 4♂ St Lucia, Brisbane Qld, 17.xi.1990, Knight; 1♂ Indooroopilly, Brisbane Qld, 13.xi.1990, Lambkin ( ANIC); 1♀ St Lucia, Brisbane SE. Qld, 20.xi.1984, de Keyzer ( MSM); 1♂ St Lucia, Brisbane SE. Qld, x.1983, de Keyzer ( MSM); 1♂ same data as previous, 1.xi.1984 ( MSM); 1♂ 1♀ same data as previous, 2.xi.1984 ( MSM); 1♂ St Lucia, Brisbane Qld, 12.ix.1981, North ( MSM); 1♀ St Lucia, Bris. Qld, 11.x.1990, Firth ( MSM); 1♂ St Lucia, Brisbane Qld, 25.xi.1983, Graham ( MSM); 1♂ St Lucia, Brisbane Qld, 9.ix.1983, North ( MSM); 3♂ St Lucia, Brisbane Qld, 3.xi.1990, Burwell ( QM); 3♂ 1♀ Samford [Qld], 2.xii.1978, Ewart ( AE); 1♀ Samford, W. of Brisbane Qld, 5.x.1981, North ( MSM); 1♂ 2♀ Southport Qld, 16.x.1901, Froggatt, 13, W.W.F., 18.i.1907 ( ANIC); Shute H [arbour] Rd, G.B.R. Qld, 27.viii.1980, de Beer ( MSM); 1♂ Somerset Dam [Qld], 18.xi.1979, red eyes when fresh, [Ewart] ( AE); 1♂ Tarragindi Qld, x.1977, [Reeves] ( AE); 1♀ Brisbane, Tarragindi [Qld], 23.xi.1977, [Reeves] ( AE); 1♂ Tarragindi [Qld], 3.ii.1978, [Reeves] ( AE); 1♂ Tarragindi [Qld], 27.xi.1978, [Reeves] ( AE); 2♂ Tarragindi Qld, 4.xii.1978, at fluoro light, Reeves ( AE); 1♂ 1♀ Tarragindi SE. Q., 2–3.xi.1980, Reeves ( AE); 1♂ 1♀ Tarragindi, Brisbane Q., 22.xii.1979, Reeves ( AE); 1♂ Tarragindi, Brisbane Q., 17.xii.1979, Reeves ( AE); 1♂ Tarragindi, Brisbane Q., 13.xii.1979, Reeves ( AE); 1♂ Tarragindi, Brisbane Q., 8.xii.1979, Reeves ( AE); 1♂ Tarragindi, Brisbane Q., xii.1979, Reeves ( AE); 1♀ Tarragindi SE. Q., 16.xi.1980, Reeves ( AE); 1♂ Tarragindi SE. Q., 9.xi.1980, Reeves ( AE); 1♂ Tarragindi SE. Q., 27.xi.1980, Reeves ( AE); 2♀ Tarragindi SE. Q., 23.xi.1980, Reeves ( AE); 1♂ Tarragindi SE. Q., 12.xii.1978, Reeves ( AE); 1♀ Tarragindi [Qld], 15.xi.1979, Reeves ( AE); 2♀ Tarragindi [Qld], 12.xii.1981, [Reeves] ( AE); 1♂ 1♀ Tarragindi [Qld], 28.xi.1978, [Reeves] ( AE); 1♂ Tarragindi Q., 17.i.1978, at light, Reeves ( AE); 1♂ Tarragindi Qld, 13.xi.1979, Reeves ( AE); 1♂ Tarragindi Qld, 1.xi.1978, [Reeves] ( AE); 1♂ 1♀ Tarragindi Qld, 9.xi.1977, [Reeves] ( AE); 1♂ Tarragindi, Brisbane Qld, 23.xi.1977, Ewart, genitalia prep. PAU76, P. annulata det. A. Ewart ( MSM); 1♂ The Gap, Brisbane Qld, 27.x.1962, Fanning ( MSM); 1♀ Third Lagoon Reserve , Sandgate SE. Qld, 27°18'00"S 153°04'00"E, 18.x.1998, Popple & Moss, 432-0006 ( LWP); 1♂ Toogoom, Hervey Bay Qld, 9.i.1979, Peters ( MSM); 1♂ Toowoomba SE. Qld, Picnic Point, 27°35'S 151°59'E, 19.x.2003, Popple & Strange, 432-0038 ( LWP); 1♂ 1♀ Toowoomba SE. Qld, scrub below escarpment, 18.xii.1976, Moulds & Moulds ( MSM); 1♂ Willawong, 27°36'00"S 153°01'00"E, 23.ix.2001, Popple, 432-0014 ( LWP); 1♂ Wishart [Qld], 4.x.1981, [Ewart] ( AE); 1♂ 1♀ Heidkes Rd Woodgate SE. Qld, 25°11'S 152°30'E, 18– 21.ix.2002, Popple, 432-0026 & 432-0027 ( LWP); 7♂ 1♀ Woody Point, Fraser Island [Qld], 15.xii.1977, [Ewart] ( AE).

Description. Male. ( Figs 3A, 3C , 4A–B , 5A–B , 6A–B ). Head. Dorsal surface black with a pale brown to medium brown triangular depression between the lateral ocelli, pointing anteriorly, with flat side against posterior margin of head; long silver pubescence behind eyes, with shorter yellow pubescence over remainder. Genae and vertex black, with long silver pubescence. Mandibular plate black with an outer narrow pale marginal ridge, extending to suture between gena; long silver pubescence. Antennae dark brown to black. Ocelli pinkish-red. Eyes red in live specimens, darkening to reddish-brown or brown in dried specimens (with some live specimens from coastal districts having white eyes); distinct furrow between eyes and pronotum. Postclypeus black anteriorly, with narrow reddish-brown margin and a small yellow-brown dorsal spot on midline (only in paler specimens) extending onto dorsal surface; short and long silver pubescence along ventral surface except down midline; short yellow pubescence along posterior ventral surface. Anteclypeus black, with long silver pubescence. Rostrum pale brown grading to black apically, with silver pubescence; extending to mid coxae.

Thorax. Pronotum black or shiny black (in specimens from Queensland's central highlands only); central fascia brown to yellow-brown (when present), forming a smear that narrows anteriorly, not extending to posterior or anterior margins; silver-yellow pubescence, especially along oblique fissures; pronotal collar brown, often reddish and pale brown to black laterally. Mesonotum, including submedian and lateral sigilla, entirely black, or sometimes with narrow, dark brown (or occasionally medium brown), dorsolateral triangular fasciae present in area between submedian and lateral sigilla and occasionally additional fasciae also present along lateral margins in some specimens; cruciform elevation brown to pink-brown, black anteriorly; ridges between wing grooves pallid; long silver-yellow pubescence, most evident around cruciform elevation.

Legs. Coxae black with longitudinal pale brown fasciae along fore coxae; fore femora pink with broad, dark brown longitudinal fasciae; mid and hind femora dark brown to medium brown, becoming pale brown apically; fore and mid tibiae dark brown, pale brown dorsally; hind tibiae pale brown; tarsi and claws medium to pale brown, paler on hind legs; spines dark brown.

Wings. Fore wing venation pale reddish-brown to brown, becoming darker towards apical cells and ambient veins; fore wing costal veins pale brown; slight angulation of fore wings at node; pterostigma pale reddish-brown; hind wing infuscation at the juncture of the anal lobe and wing margin, surrounding the distal termination of vein 2A, distinct.

Timbals. Long ribs 1–2 (and occasionally 3) fused ventrally; long ribs 1–4 fused dorsally to basal spur. Long rib 5 does not extend ventrally beyond adjacent intercalary rib. Four intercalary ribs developed equally and clearly visible.

Opercula. Roughly sickle-shaped, obliquely elongated; central region domed, shiny black; medial margin (and sometimes also lateral margin) pallid; parallel to body axis.

Abdomen. Tergites black anteriorly, grading sharply to dark brown or medium brown dorsolaterally, with contrasting yellow to yellow-brown posterior margins between tergites 3 to 7; diffuse, medium brown colouration extending broadly around posterior half of tergite 8; silver-yellow short pubescence. Sternites pale-brown with prominent black medial fascia or entirely black with anterior sternites showing some medium to pale brown colouration at margins (in melanistic specimens, which are also mostly black dorsally, without thoracic fascia).

Genitalia. Pygofer black anteriorly, grading to brown posteriorly; upper lobes prominent, erect, terminally rounded; lower lobes distinct, bulbous, with an inner tooth weakly to strongly developed on each lobe; inner lobes enlarged, acute, posteriorly tapering; dorsal beak prominent, sharply pointed, slightly curved in lateral view; claspers with a pair of hooked processes; aedeagus with dorsal pseudoparameres that bifurcate and join theca near gonocoxite IX; pseudoparameres apically curved, tapering; theca sclerotized, gradually curved posteriorly, apex projecting further dorsally than ventrally, slightly greater in vertical width than shaft, with a variable number of small spines on lateral sides, 8–14 medium spines on posterior lateral sides and 2–4 large spines on posterior ventral edge.

Female ( Fig. 3B, 3D ). Similar colouration and patterning to male, often paler, but not in all specimens. Head colouration similar to that of male, except often with yellow-brown vertex lobes. Pronotum black; central fascia (when present) yellow to dark brown, often broader than that of male and reaching anterior margin, extending further posteriorly and dorsolaterally (in pale specimens) to form a pair of outward-pointing, wedge-shaped patterns. Mesonotum with submedian and lateral sigilla black; dark brown to yellow-brown colouration present over much of the remainder; typically with an overall greater extent of pale brown colouration than in male. Legs similar in colour to those of male, with tarsi sometimes paler. Abdomen with tergites similar in colouration to those of male; abdominal segment 9 sandy-brown with a pair of longitudinal near-dorsal black fasciae that extend to the anterior edge and ventrolaterally to some extent; sternites sandy-brown with a broad and more or less regular median black fascia from sternites II to VII; ovipositor sheath extending approximately 0.5 mm beyond termination of abdominal segment 9.

Measurements. N= 30 ♂ 30 ♀. Ranges and means (in parentheses), mm; BL: ♂ 11.7–14.7 (13.5); ♀ 12.2– 14.6 (13.9); FWL: ♂ 13.0–16.8 (15.4); ♀ 13.8–18 (17); FWB: ♂ 4.7–6.1 (5.6); ♀ 5–6.4 (5.8); HW: ♂ 3.7–4.5 (4.2); ♀ 3.8–4.8 (4.4); PW: ♂ 2.8–3.7 (3.3); ♀ 3.0–3.9 (3.5); AW: ♂ 3.3–4.3 (3.9); ♀ 3.2–4.0 (3.7); OL: ♀ 4.3–5.1 (4.8).

Distinguishing features. Both sexes of P. annulata can be distinguished from P. rubristrigata by their smaller size (fore wing length less than 20 mm). Males can be distinguished from P. corymbiae , P. decora and P. kobongoides , by the pale, rather than mostly black or dark brown medial margin of the opercula and from P. subtropica and P. torrensis by the rounded, rather than slightly hooked, posterior ventral termination of the upper pygofer lobes. Males differ from those of P. granitica by the sharp transition in colouration of the lateral abdominal tergites from black to pale brown, without a conspicuous three-step grade in tone from black to medium brown to pale brown. They differ from P. inversa by timbal long rib 5 not extending ventrally beyond the adjacent intercalary rib and by having both dorsal and ventral projections (rather than just the former) associated with the apex of the theca, and from P. ayrensis and P. simplex by having pale, rather than dark, upper pygofer lobes. Pauropsalta annulata males share most features in common with those of P. tremula and P. blackdownensis ; however, individuals never share the combination of distinct mesonotal fasciae (in the area outside of the submedian and lateral sigilla) and sternites that are close to being entirely black, as seen in the latter two species. Females are less easily distinguished, except where there are differences in ovipositor length (refer to descriptions of remaining taxa below). Differences between P. annulata and P. notialis (including subspecies P. n. notialis and P. n. incitata, and the hybrids between these two subspecies) are more subtle and details are given in the descriptions that follow. In particular, some male specimens of P. annulata may be impossible to distinguish by morphology from P. n. notialis , and both sexes do not always differ in appearance from P. n. notialis x incitata hybrid individuals (see below).

Notes on Geographical Variation. Specimens of P. annulata from the Central Highlands and Coalfields district of Queensland, between Emerald, Charlevue Creek and Clermont (see grey shaded area in Fig. 7 ), consistently have a glossy textured thorax and head; whereas those from the rest of the distribution are dull or matte textured over the entire body ( Figs 3C–D ; cf. Figs 3A–B ). These key out in a different couplet in the morphological key (above). The Mt Moffatt and Expedition Range populations (to the south and south-west of the grey shaded area in Fig. 7 ) are comprised entirely of dark or melanistic coloured males and females. The thorax and abdominal sternites of these insects are entirely black, except for the cruciform elevation on the thorax and the extreme lateral edges of the sternites. Such melanism is also present, but uncommon in subcoastal districts, and especially so in females. A white-eyed male phenotype (live specimens only) is moderately common, but largely coastal in distribution, between Bundaberg and Coolangatta in south-eastern Queensland.

Distribution and Ecology ( Figure 7 ). Pauropsalta annulata has a subtropical coastal and subcoastal distribution from Shute Harbour (near Proserpine), Eungella and Clermont south to Carnarvon National Park in central Queensland and further to near Kyogle in northern New South Wales. It is a conspicuous species along the coast from Gladstone south to the border region of Queensland and New South Wales, including Bundaberg, Wide Bay, Gympie, Greater Brisbane, Redland City and the Gold Coast. It is also quite prolific subcoastally around Monto, Eidsvold, Gayndah, Goomeri, Kingaroy, Cooyar, Toowoomba, Esk, Gatton, Aratula, Beaudesert and patchily to the north and south-east of Boonah. Populations occur primarily in association with Eucalyptus tereticornis , E. crebra and Allocasuarina littoralis ( Popple & Walter, 2010) . The localised population on Blackdown Tableland is associated with Eucalyptus sphaerocarpa . At Mt Moffatt in the Carnarvon National Park, P. annulata is primarily associated with E. conica , with some individuals also on E. microcarpa and E. populnea ( Popple & Walter, 2010) . Throughout its distribution, particularly in towns and cities, this species is sometimes associated with other plant species (mostly Myrtaceae ), especially in suburban gardens. Adults sit in the outer foliage of trees and are wary, with males taking frequent flights between singing stations.

Geological and Pedological Associations. Found in association with silt loams, silty clay loams and sandy loams derived from Tertiary basalt and in the transitional area between soils derived from basalt and those derived from quartz, feldspar-rich sandstones, rhyolite and granite. Upland localities where P. annulata occurs also include plateaus with soils that display little profile development. It is occasionally found in some sandy soils that form part of coastal dune complexes, but only on relatively dry sites where silt and clay make up at least 10% of the soil content. There appears to be an association with soils of reasonable depth (i.e.> 0.5 m to 2 m).

Calling Song ( Figs 2A , 8–9 ). The calling song of P. annulata comprises two components, lilting and buzzing, which alternate during song production. Singing may commence with either song component and song production often persists when the insect flies between calling sites. The lilting component is made of repeated phrases, each of which includes a long echeme (2–4 syllables, 0.020 – 0.047 s) followed by a brief silence (0.012 – 0.037 s), a short echeme (1 syllable, 0.008 – 0.015 s) and a longer silence (0.045 – 0.125 s) (all statistics here and below, n=131 individuals). phrase repetition rates vary between 0.105 – 0.196 s, and typically fall within the range of 0.110 – 0.145 s. The buzzing component comprises a sustained repetition of the short echeme used in the lilting component (1 syllable, 0.009 – 0.015 s) with brief silences (0.012 – 0.026 s) that punctuate each syllable. The song has a frequency plateau between 6.5–10.5 kHz, with a highest amplitude dominant frequency between 7.3–9.1 kHz and a syllable repetition rate of about 90–110 Hz. A single recording of the male–female duet has been made for this species ( Fig. 9v ). Females respond with a soft wing-flick during the silence at the end of each lilting component phrase ( Fig. 1 ), 0.028 – 0.058 s after production of the short echeme. No response is given by the female during production of the buzzing component. The majority of the original recordings illustrated by Ewart (1989) never included the buzzing component, probably because these cicadas make the call only in nature and not in the plastic boxes used in those recordings. The only recording that did include the buzzing component was from the northern base of the Blackdown Tableland (Charlevue Creek), and this was a field recording ( Ewart, 1989). At the top of the Blackdown Tableland and at Mount Moffatt, males produce the buzzing component only on rare occasions.

Figures

FIGURE 1. Oscillograms showing examples of calling songs from cicada species in the Pauropsalta annulata species group, to illustrate the acoustic terminology used in this paper. Most P. annulata cicadas produce a biphasic song. A 'buzzing' component alternates with a 'lilting' component and each of these is illustrated with examples from three species, all labelled with terms relevant to each. The buzzing component is on the left and the lilting component (not necessarily conspecific) on the right.

FIGURE 2. Oscillograms providing a comparison of the calling song structure of each taxon (and hybrid population) in the Pauropsalta annulata species group: (A) Pauropsalta annulata Goding and Froggatt, Eidsvold (25°22'S 151°07'E), (B) P. notialis notialis subsp. nov., Concord (33°51'S 151°06'E); (C) P. n. incitata subsp. nov., Eidsvold (25°22'S 151°07'E); (D) P.n. notialisxincitata hybrid, Upper Yarraman (26°54'S 151°54'E); (E) P. tremula sp. nov., Lake Broadwater (27°21'S 151°06'E); (F) P. blackdownensis sp. nov., Blackdown Tableland (23°44'S 149°07'E); (G) P. simplex sp. nov., Atherton (17°16'S 145°29'E); (H) P. granitica sp. nov., 7km SE. of Mount Carbine (16°34'S 145°10'E); (I) P. subtropica sp. nov., Auburn River National Park (25°43'S 151°03'E); (J) P. torrensis sp. nov., Torrens Creek (20°47'S 145°01'E); (K) P. decora sp. nov., 82km N. of St George (27°23'S 148°52'E); (L) P. kobongoides sp. nov., Weengallon (28°22'S 149°03'E); (M) P. corymbiae sp. nov., Bull's Gully via Adavale (25°58'S 144°59'E); (N) P. inversa sp. nov., Eidsvold (25°22'S 151°07'E); (O) P. rubristrigata (Goding and Froggatt), Nimmitabel (36°31'S 149°14'E); (P) P. ayrensis Ewart, Eidsvold (25°22'S 151°07'E). All recordings were made with RS1, except (M) and (O), which were obtained by A. Ewart with RS3 and the author using RS5, respectively (see methods).

FIGURE 3. Pauropsalta annulata Goding and Froggatt (A) male, Lake Dyer (27°38'S 152°22'E); (B) female, Lake Dyer; (C) male, Charlevue Creek; (23°41'S 145°08'E); (D) female, Charlevue Creek. Approximately 1.6x natural size.

FIGURE 4. Illustrations of male right timbals, with dorsal edge at left and posterior margin at bottom: (A–B) Pauropsalta annulata Goding and Froggatt, Lake Dyer (27°38'S 152°22'E) and Charlevue Creek (23°41'S 149°08'E) respectively; (C) P. notialis notialis subsp. nov., Concord (33°51'S 151°06'E); (D) P. n. incitata subsp. nov., Barakula State Forest via Chinchilla (26°15'S 150°49'E); (E) P. n. notialisxincitata hybrid, Atkinsons Dam (27°26'S 152°27'E); (F) P. tremula sp. nov., Lake Broadwater (27°21'S 151°06'E); (G) P. blackdownensis sp. nov., Blackdown Tableland (23°47'S 149°00'E); (H) P. simplex sp. nov., Atherton (17°16'S 145°29'E); (I) P. granitica sp. nov., 7km SE. of Mount Carbine (16°34'S 145°10'E); (J) P. subtropica sp. nov., 1km N. of Auburn River National Park (25°43'S 151°03'E); (K) P. torrensis sp. nov., Torrens Creek (20°47'S 145°01'E); (L) P. decora sp. nov., 82km N. of St George (27°23'S 148°52'E); (M) P. kobongoides sp. nov., 65km NNE. of St George (27°32'S 148°50'E); (N) P. corymbiae sp. nov., Bull's Gully via Adavale (25°58'S 144°59'E); (O) P. inversa sp. nov., Eidsvold (25°22'S 151°07'E); (P) P. rubristrigata (Goding and Froggatt), Mt Lofty (34°59'S 138°43'E); (Q–R) P. ayrensis Ewart, (Q) 1km N. of Maroon Dam (28°11'S 152°39'E), (R) Mount Surprise (18°09'S 144°19'E). Scaling is the same for all illustrations except (P), which is scaled differently to the rest.

FIGURE 5. Illustrations of male right opercula, with medial margin at bottom. (A–B) Pauropsalta annulata Goding and Froggatt, (A) Lake Dyer (27°38'S 152°22'E), (B) Charlevue Creek (23°41'S 149°08'E); (C) P. notialis notialis subsp. nov., Concord (33°51'S 151°06'E), (D) P. n. incitata subsp. nov., Barakula State Forest via Chinchilla (26°15'S 150°49'E); (E) P.n. notialisxincitata hybrid, Atkinsons Dam (27°26'S 152°27'E); (F) P. tremula sp. nov., Lake Broadwater (27°21'S 151°06'E); (G) P. blackdownensis sp. nov., Blackdown Tableland (23°47'S 149°00'E); (H) P. simplex sp. nov., Atherton (17°16'S 145°29'E); (I) P. granitica sp. nov., 7km SE. of Mount Carbine (16°34'S 145°10'E).

FIGURE 6. Illustrations of male pygofers and internal genitalia, viewed ventrally (left) and laterally from left (right): (A–B) Pauropsalta annulata Goding and Froggatt, (A) Carina Heights (27°30'S 153°06'E), (B) Charlevue Creek (23°41'S 149°08'E); (C) P. notialis notialis subsp. nov., Concord (33°51'S 151°06'E), (D) P. n. incitata subsp. nov., Dogwood Creek via Miles (26°39'S 150°11'E); (E) P. n. notialisxincitata hybrid, Atkinsons Dam (27°26'S 152°27'E); (F) P. tremula sp. nov., Lake Broadwater (27°21'S 151°06'E).

FIGURE 7. Map of eastern Australia showing the geographical distribution of Pauropsalta annulata Goding and Froggatt. Large triangles represent specimen records (see material examined), and small triangles are aural records (some recorded). The grey shaded area shows the distribution of the central highlands population (Clermont–Emerald) of P. annulata in Queensland (see description and morphological key).

FIGURE 8. Male calling song structure of Pauropsalta annulata Goding and Froggatt illustrated in waveform plot form, showing both buzzing and lilting components. Each lower plot is expanded from the one above, to illustrate the finer temporal structure of each song comparison. The spectrogram at the bottom of the figure displays song frequency, which exhibits no modulation between the song components in this species. This specimen was recorded in the field at Carina Heights (27°30'S 153°06'E).

FIGURE 9. Waveform plots illustrating the lilting component of the male calling song of Pauropsalta annulata Goding and Froggatt from six different localities, including: (i) Carina Heights (27°30'S 153°06'E), (ii) Eidsvold (25°22'S 151°07'E), (iii) Woodgate (25°08'S 152°29'E), (iv) Emerald (24°52'S 151°51'E), (v) Blackdown Tableland (23°48'S 149°05'E), and (vi) Mount Moffatt (25°01'S 147°57'E). Mean phrase repetition rates (PRR) for each recording are provided to the right of each plot for reference. The Blackdown Tableland recording (v) is a duet that includes female response, with each female wing-flick indicated by an arrow.