Pauropsalta annulata v. lamp, Popple & Popple v. lamp, Popple & McKinnon v. lamp v. lamp, Popple & McKinnon v. lamp, Popple & Hando <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> POPPLE, L. W. Zootaxa 2013 2013-10-25 3730 1 1 102 75V88 POPPLE, 2013 POPPLE 2013 [151,392,897,923] Insecta Cicadidae Pauropsalta Animalia Hemiptera 71 72 Arthropoda species inversa sp. nov.  ( Figs 2N, 4O, 31F, 39C, 43, 46A–B, 47–48)    Pauropsalta annulatasong typeCN: Popple et al.(2008); Popple & Walter (2010).  Types:   Holotype: ♂‘ AUSTRALIA/ Queensland’, ‘ Ceratodus/ 10kmN. of Eidsvold’, ‘  25.i.2003/m.v. lamp’, ‘ L. Popple& W. Popple’, ‘431-0002’, ‘ QM Reg. No. T156331’ (  QM);   Paratypes: QUEENSLAND: 1♀ Ceratodus, 10kmN. of Eidsvold,  25.i.2003, m. v. lamp, Popple & Popple, 431-0011 (  QM);  1♂ 1♀ Eidsvold, 25°22'06"S 151°07'25"E,  24–25.xi.2006, m. v. lamp, Popple & McKinnon, 431-0014 & 431-0058 (  AE); 2 MM Pistol Gap S.of Byfield Qld,  10.i.1970, EB, GH & SM, at light (  ANIC);  6♂ 1♀ Burnett River, 10kmN. of Eidsvold Qld,  28.i.1988, Moulds& Moulds(  MSM);  1♀ Eidsvold, 25°22'06"S 151°07'25"E,  24–25.xi.2006, Popple& McKinnon, m. v. lamp, 431-0057 (  MSM);  4♂ 3♀ GoolgowiN.S.W.,  9.ii.1984, Moulds& Moulds;  1♂ 1♀ Springsure Qld,  15.xii.1979, Valentine(  MSM);  1♂AU.QLD.MOZ, CreekS. of Rollestonon SH55 Qld, 24°28.633'S 148°36.768'E,  11.i.2002, Cooley, Cowan, Hill, Marshall& Moulds(  MSM);  54♂ 12♀ 1♂ 1♀ Eidsvold, 25°22'06"S 151°07'25"E,  24–25.xi.2006, m. v. lamp, Popple & McKinnon, 431-0012 to 431-0056 & 431-0060 to 431-0081 (  LWP);  2♂ Billabong Motor Inn, Mundubbera, 25°35'03"S 151°15'18"E,  8.i.2007, Popple& Hereward, 431-0082 & 431-0083 (  LWP);  1♂ Clermont, 22°49'04"S 147°38'37"E,  18.xi.2005, m. v. lamp, Popple & Hando, 442-0001 (  LWP);  1♂ AUSTRALIA  New South Wales, Mount Hope, 32°50'25"S 145°52'53"E,  21.xi.2010, S12588, Popple& Emery, 431-0084 (  LWP);  NEW SOUTH WALES: 1♂ 1kmS. of Mount Hope, 32°50'38"S 145°52'54"E,  21.xi.2010, S12588, Popple& Emery, 431-0085 (  LWP).   Description. Male( Figs 4O, 31F, 39C, 46A). Head. Dorsal surface black with an inconspicuous dark 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 irregularly distributed over remainder. Genae and vertex black, with long silver pubescence. Mandibular plate black; long silver pubescence. Antennae dark brown to black. Ocelli dull red. Eyes dark red in live insects, fading to dark reddish-brown in some preserved specimens; distinct furrow between eyes and pronotum. Postclypeus shiny black with narrow yellow-brown margins; short yellow pubescence along posterior ventral surface. Anteclypeus black, with long silver pubescence. Rostrum dark brown grading to black apically, with silver pubescence; extending to mid coxae.  Thorax. Pronotum black with silver-yellow pubescence; pronotal collar brown, grading to yellow-brown laterally. Mesonotum often entirely black, sometimes with inconspicuous dark brown dorsolateral fasciae between submedian and lateral sigilla; cruciform elevation reddish-brown to yellow-brown, black anteriorly; wing grooves yellow-brown; silver-yellow pubescence, most evident around cruciform elevation.   FIGURE 46.(A–B)  Pauropsalta inversa  sp. nov., Eidsvold (25°22'S 151°07'E), (A) male, (B) female; (C–D)  P. rubristrigata(Goding and Froggatt), Mt Lofty (34°59'S 138°43'E), (C) male, (D) female; (E–H)  P. ayrensisEwart, (E–F) 1km N. of Maroon Dam (28°11'S 152°39'E), (E) male, (F) female, (G–H) Mount Surprise (18°09'S 144°19'E), (G) male, (H) female. Approximately 1.6x natural size, except (C–D) at 1.0x natural size.  Legs. Coxae black, yellow-brown apically; fore femora black with narrow, dark brown longitudinal fasciae, pale brown apically; mid and hind femora black, pale brown apically; fore and mid tibiae medium to dark brown; hind tibiae pale to medium brown; fore and mid tarsi medium brown; hind tarsi pale brown; claws and spines dark brown.  Wings. Fore wing venation medium to dark brown, becoming darker towards apical cells and ambient veins; fore wing costal veins pale to medium brown; slight angulation of fore wings at node; pterostigma 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 sometimes 3) fused ventrally; long ribs 1–4 fused dorsally to basal spur. Long rib 5 typically extending further, ventrally, than adjacent intercalary rib.  Opercula. Roughly sickle-shaped, obliquely elongated; central region domed, black; medial margin pallid; parallel to body axis.  Abdomen. Tergites mainly black with narrow, contrasting yellow-orange posterior margins, wider on tergite 8; short silver pubescence conspicuous laterally. Sternites yellow to yellow-brown with a black medial fascia.  Genitalia. Pygofer black anteriorly, grading to yellow or yellow-brown apically; upper lobes prominent, erect, narrow, terminally rounded; lower lobes distinct, bulbous, without an inner tooth on each lobe; inner lobes enlarged, acute, posteriorly tapering; claspers with a pair of hooked processes; aedeagus with dorsal pseudoparameres that bifurcate and join theca near gonocoxite IX; pseudoparameres apically curved, tapering; theca gradually curved posteriorly, sclerotized, widening apically; apex abrupt and sheer, ornamented with numerous small spines and fewer medium-large spines (sometimes absent), with dorsal edge projecting conspicuously further posteriorly (> 0.1mm) relative to ventral edge.  Female( Fig. 46B). Similar colouration and patterning to male, but darker brown overall. Head similar in colour to that of male with eyes varying from dark red to brown. Pronotum black; central fascia inconspicuous, dark brown, forming a smear that tapers proximally and distally, not reaching anterior or posterior margins. Mesonotum similar in colour to that of male, with barely visible (or absent) dark brown dorsolateral fasciae in area between submedian and lateral sigilla, and additional fasciae along lateral margins; when present, dorsolateral fasciae triangular, sometimes fused medially; lateral fasciae narrow, barely visible (or absent). Legs similar in colour to those of male. Abdomen with tergites similar in colour to those of male, with posterior tergite margins typically yellow-brown to medium brown; abdominal segment 9 medium brown to yellow-brown with a pair of longitudinal, near-dorsal, black fasciae that extend to the anterior edge and ventrolaterally to some extent; sternites yellow-brown to medium brown with a broad, black fascia distributed medially along sternites II to VII, and which tapers posteriorly; ovipositor sheath extending < 0.5 mmbeyond termination of abdominal segment 9.  Distinguishing features. The extensive black colouration of the males, along with the lack of conspicuous mesonotal fasciae, distinguishes  P. inversafrom all others in the  P. annulatagroup, apart from some specimens of  P. annulata, P. n. notialis, P. n. incitata,  P. subtropicaand  P. torrensis. Males can be distinguished from these remaining taxa by timbal long rib 5, which extends further ventrally than the adjacent intercalary rib, and by the apex of the theca, which has a prominent dorsal, but no ventral, projection. Females, being similar in appearance to the males, can also be distinguished by their extensive black colouration, with the exception of some dark specimens of  P. annulata. They differ from melanistic individuals of  P. annulataby having pale brown sternites with a black medial fascia, in contrast to sternites that are almost entirely black.  Notes on Geographical Variation. Specimens of  P. inversafrom western New South Walesconsistently have a glossy textured thorax and head; whereas those from Queenslandare dull or matte textured over the entire body.  Measurements. N= 30 ♂ 18 ♀. Ranges and means (in parentheses), mm; BL: ♂11.3–15.0 (12.2); ♀11.1–14.2 (12.8); FWL: ♂13.4–16.3 (15.0); ♀14.9–17.7 (16.0); FWB: ♂4.8–5.6 (5.1); ♀4.9–6.1 (5.5); HW: ♂3.7–4.3 (3.9); ♀3.8–4.6 (4.2); PW: ♂3.0–3.5 (3.3); ♀3.2–3.7 (3.4); AW: ♂3.2–4.2 (3.7); ♀3.3–4.0 (3.7); OL: ♀3.7–4.5 (4.0).   Etymology. From Latin, meaning inverted. For most species, there are short and long silent gaps in the lilting component and the long gaps typically occur after the short echeme. However, in  P. inversa, the long gaps always occur after the long echeme. As a consequence, in an oscillogram the call of this species appears to be temporally inverted relative to the calls of other species in the  P. annulataspecies group.   Distribution and Ecology( Figure 43). Central and southern inland Queenslandfrom Clermont south across the Great Dividing Range to Augathella, near Tambo, Mt Moffatt (Carnarvon National Park) and St George, and east to the Fitzroy, Dawson, upper Burnett and upper Mary River catchments of south-east Queensland, including Byfield, Theodore, Dawson Range, Kalpowar, Auburn River near Mundubbera and near Gympie. A disjunct population also occurs in central inland New South Walesfrom Mount Hope south to Goolgowi. In Queensland, males sing from the upper branches and main trunks of tall eucalypts, such as  E. tereticornis,  C. tessellarisand, on occasion,  E. moluccana, whereas in New South Wales, adults appear to be associated with  Callitris columellaris. Adults are active from late September to at least the end of January with peak numbers usually between mid– December and New Year.  Geological and Pedological Associations. Adults are associated with tree species that are generally found in loamy alluvial soils of variable depth and on undeveloped soils in sloping terrain.  Calling Song( Figs 2N, 47–48).  Pauropsalta inversahas prominent buzzing and lilting components in its song. It is distinguished from other species in the  P. annulatagroup by the long silent gap that follows the long echeme (in all other species of the  P. annulataspecies group the longer gap follows the short echeme or the echemes or the gaps that follow them are of equal duration). The lilting component comprises a long echeme (27–126 syllables, 0.270 – 1.403s) followed by a long silence of variable duration ( 0.130 – 1.912s), a short echeme (2 syllables, 0.018 – 0.035s), and a short silence ( 0.054 – 0.136s) (all statistics, n=74 individuals). As found in most other species, the buzzing component is a long drawn-out single echeme that appears equivalent to the long echeme in the lilting component. Observations on the attraction of males to simulated female wing-flicks suggest that the female responds during the silence that follows the short echeme (D. Marshall pers. comm., 1.vii.2013), which corresponds with other species in the  P. annulataspecies group. The calling song has a phrase repetition rate of 0.472 – 4.070s and a syllable repetition rate of 90–110 Hz. In Queensland, the dominant frequency plateau extends from ~9.5–15.0 kHz, with a highest amplitude dominant frequency between 11.3–12.1 kHz. In the disjunct New South Walespopulation, the dominant frequency plateau extends from ~8.8–14.0 kHz, with a highest amplitude dominant frequency between 9.7–11.4 kHz.   FIGURE 47.Male calling song structure of  Pauropsalta inversa  sp. nov.illustrated in expanded waveform plots (explained in Fig. 8), showing both buzzing and lilting components. 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 Eidsvold (25°22'S 151°07'E).   FIGURE 48.Waveform plots illustrating the lilting component of the male calling song of  Pauropsalta inversa  sp. nov.from six different localities, including: (i) Eidsvold (25°22'S 151°07'E), (ii) Mundubbera (25°35'S 151°18'E), (iii) Mount Hope (32°50'S 145°53'E), (iv) Goolgowi (33°59'S 135°43'E), (v) Clermont (22°50'S 147°38'E), and (vi) Mount Moffatt (24°52'S 148°01'E). Mean phrase repetition rates (PRR) for each recording are provided to the right of each plot for reference. Recordings (iii) and (iv) were made with RS5, while the remainder were made with RS1 (see methods).  The duration of silence that follows the long echeme in each phrase of the lilting component varies considerably, to some extent within and among individuals, but even more conspicuously between populations. For example, in Queensland, recordings from Augathella, near Tambo, Springsure, Clermont, Rolleston, Wyseby, Mt Moffattand St George show a conspicuously long silence ( 0.505 –1.912s), whereas those from Middlemount, Blackwater and the Burnett and Mary Rivercatchments further east have a much shorter silence ( 0.130 –0.379s). In western New South Wales, the silence after the long echeme also varies with an intermediate duration at Goolgowi ( 0.530 –0.855s) and a shorter duration again at Mount Hope( 0.145 –0.214s), localities that are only 155 kmapart. In Queensland, the variation from east to west may be clinal. Alternatively, it may be related to the abundance of males in any particular area at the time when recordings were taken. For the instance, males were observed to be quite common in the Burnett catchment and also at Mount Hopewhere some of the most rapidly emitted call sequences have been recorded. In contrast, at Clermont and at Goolgowi, where some of the most drawn-out sequences were recorded, males were generally observed to be quite widely scattered through their habitat. Further investigations would be required to unravel the cause(s) and potential implications of this variation. 2003-01-25 QM L. Popple & W. Popple' & QM Reg. No. T Australia Eidsvold' Queensland' Ceratodus 71 72 1 1 holotype 2003-01-25 QM Australia Eidsvold Ceratodus 71 72 1 1 Queensland paratype 2006-11-24 2006-11-25 2006-11-24 AE Australia -25.368334 Eidsvold 21 151.12361 71 72 2 1 1 Queensland paratype 1970-01-10 ANIC MM Pistol Gap S. Australia Byfield Qld 71 72 1 Queensland paratype 1988-01-28 MSM Moulds Australia Eidsvold Qld Burnett River 71 72 7 1 6 Queensland paratype 2006-11-24 2006-11-25 2006-11-24 MSM Popple & McKinnon Australia -25.368334 Eidsvold 21 151.12361 71 72 1 1 Queensland paratype [460,1133,1286,1311] 1984-02-09 Moulds Australia Goolgowi 71 72 7 3 4 Queensland paratype 1979-12-15 MSM Valentine Australia Springsure Qld 71 72 2 1 1 Queensland paratype 2002-01-11 MSM Cooley & Cowan & Hill & Marshall & Moulds Australia Creek -24.477217 Qld 1 148.6128 Rolleston 71 72 1 1 Queensland paratype 2006-11-24 2006-11-25 2006-11-24 LWP Australia -25.368334 Eidsvold 21 151.12361 71 72 68 13 55 Queensland paratype 2007-01-08 LWP Popple & Hereward Australia -25.584167 Mundubbera 21 151.255 Billabong Motor Inn 71 72 2 2 Queensland paratype 2005-11-18 LWP Australia -22.81778 Clermont 21 147.64362 71 72 1 1 Queensland paratype [246,447,1502,1527] Australia AUSTRALIA 71 72 1 1 paratype S12588 2010-11-21 LWP Popple & Emery Australia -32.840275 Mount Hope 20 145.8814 71 72 1 New South Wales paratype S12588 2010-11-21 LWP Popple & Emery Australia -32.843887 Mount Hope 20 145.88167 71 72 1 1 New South Wales paratype Mt Moffatt 0.145 Mount Hope 78 -0.214 Mary River 75 76 1 Queensland