Yoyetta regalis, Emery & Emery & Popple, 2019

Emery, David L., Emery, Nathan J. & Popple, Lindsay W., 2019, A Revision of the Yoyetta abdominalis (Distant) Species Group of Cicadas (Hemiptera: Cicadidae: Cicadettinae), Introducing Eight New Species, Records of the Australian Museum 71 (7), pp. 277-347 : 304-308

Yoyetta regalis sp. nov.

http: //zoobank.org/NomenclaturalActs/ 2428C6E9-758A-4D71-8B4A-AD0233415B46

Figs 2C , 3C , 18–21 , 52C, 53L

Cicadaetta sp. nr abdominalis (“Red ringer”).— Emery et al., 2005: 99, Fig. 6 .

Holotype ♂, Wises Track , Royal National Park, NSW, 34°06'59"S 151°03'30"E, 25.i.2004, S. & D. Emery ( AM K.536051) . Paratypes NEW SOUTH WALES: 1♂, Hartley Vale , 30.xii.1971, S. G.Watkins,donated 2001 ; 1♂, same data, 7.i.1972 ; 1♂, NSW, Blue Mtns, G. Burns, 1905 ( ANIC) ; 1♂, Ingleside , NSW, 33°40'17"S 151°15'14"E, 20.xii.1994, D.Emery ; 1♂, Cataract Dam , NSW, 34°14'39"S 150°49'18"E, 8.xi.1995, D. Emery ; 1♂ 1♀, Kinka Reserve, Terrey Hills , NSW, 33°40'39"S 151°12'00"E, 16.i.1997, D. Emery ; 7♂♂, Booralie Rd, Terrey Hills , NSW, 33°40'34"S 151°12'52"E, 8.i.2004, S. & D. Emery ; 1♂, Wises Track, Royal National Park , NSW, 34°06'59"S 151°03'30"E, 13.i.2006, N.& D. Emery ; 3♂♂, same location, 1–20.i.1998, D. Emery ; 1♀, same location, 8.i.2002, S., N.& D. Emery (DE) ; 1♂, Wentworth Falls , NSW, 21.xii.1923, Harrison ; 1♂, Jenolan , NSW, 7.xii.1929, CRW ; 1♂ 1♀, Monaro , NSW (no name or date) ; 1♂, Cooma, NSW (no name or date); 2♂♂, Buffalo, Vic. , i.1937, K. S. W. Sutton ( McM) ; 1♂, Bendalong , NSW, 8.i.1969, G. Daniels ; 1♂, Sydney ,NSW, 22.i.1953, H.Hughes ; 1♂, Goondera Range, Royal National Park , NSW, 13.xii.1969, G. Daniels ( AM) ; 2♂♂ 1♀, Kuringai Chase National Park , 8.i.2004, S., N.& D. Emery ; 1♂, Wises Track, Audley , 30.xi–1.xii.2003, L. Popple, D.Emery, 512-0001 ; 1♀, same location as previous, 5.ii.2004, S., N. & D. Emery ; 1♀, same location as previous, 24.i.2004, S. & D. Emery ; 1♂, Hat Hill Rd, Blackheath , 22.xii.2002, S., N. & D. Emery ; 1♂, Wambool Nature Reserve , 33°29'43"S 149°45'41"E, open woodland, 8.i.2010, Popple & Emery, 5120-0002 ; 1♂, Blackheath , i.2010, D. Emery,512 ; 1♂, Pulpit Rock via Blackheath , 33.619265°S 150.327337°E, xii.2009, D. Emery, 512 ( LWP) ; 1♂, Curra Moors T [rac]k, R[oyal] Nat. Pk , 13.i.2004, J. R. W .; 4♂♂, Menai 2.ii.[19]79, R. Eastwood ; 1♂, Forestville, Sydney , 12.ii. 1975, C. Holmes ; 1♂, West Head , 20.i.1970, A. B. Rose ; 1♂, Nellie’s Glen, Blue Mountains , 30.xii.1978, L. R. Ring ; 1♂, nr Native Dog Hill , 38 km E of Rylestone, 30.xii.1977, G. Daniels ; 1♂, Mt Keira , 27.xi.1994, R.de Keyzer & A. Sundholm ; 1♂, Gonarra Ridge, Royal National Park , 29.i.1979, G. Daniels ; 1♂, AVALON, 5.XII.[19]57; 1♀, Hat Hill, Blackheath , 30.xi.1975, G. Daniels ; 1♂, 75/203 Fairburn Rd, Wedderburn, S. of Sydney , 34°09.009'S 150°49.789'E, 135 m [elev.], 21.xi.2006, K. Hill, D. Marshall, D. & T. Britton, C. Simon lab Voucher, legs in ETOH, body pinned, 06.AU. NS.BRI.01, “ Yoyetta red ringer”, specimen recorded ( MSM) ; 1♀, Moe, Vic , 38°11'S 146°16'E, 26.ii.1891 ( HEM2327 ) ( MMV) .

Other material. NEW SOUTH WALES: 2♂♂, Wises Track, Royal National Park , NSW, 34°06'59"S 151°03'30"E, 22.xii.2002, S., N. & D. Emery ; 2♂♂, same location, 17.ii.2005, S. & D. Emery ; 4♂♂ 1♀, same location, 1.i.2006, S. & D.Emery ; 1♀, same location 22.i.2007, N. & D. Emery ; 5♂♂ 1♀, same location, 29.xi.2003, N.& D. Emery & L. Popple ; 2♂♂ 3♀♀, same location, 5.i–5.ii.2004, S., N.& D. Emery ; 13♂♂, 3♀♀, same location, 5–25.i.2004, S., N.& D. Emery ; 1♀, Hat Hill Rd, Blackheath ,NSW, 33°36'59"S 150°18'29"E, 14.xii.2002, D. Emery ; 6♂♂ 1♀, same location, 15.xi.09, N. & D. Emery ; 2♀♀, same location, 6.xii.2009, D. Emery ; 1♂, Waterfall , NSW, 21.xi.2006, R.Chin ; 1♀, Capertee , NSW, 33°08'25"S 149°58'47"E, 9.xii.2009, D.Emery ; 1♂, 4 km N Wambool Nature Reserve , NSW, 8.i.2010, D. Emery & L. Popple ; 5♂♂ 1♀, Anvil Rock , NSW, 33°35'44"S 150°50'19'E 12–16.xii.2012, N.& D. Emery ; 7♂♂, Lockley’s Pylon track, Leura , NSW, 33°36'05"S 150°20'00"E, 16.xii.2012, N. & D. Emery ; 1♂, Boodi National Park , NSW, 33°30'41"S 151°24'03"E, 100 m, 24.xi.[20]13, N. Emery ; 1♂, same location, 5.1.14, N. Emery ; 4♂♂, Anvil Rock, Blackheath , NSW, 33°35'44"S 150°50'19"E, 971 m, 4.xii.14, D. & C. Emery ; 1♂ 1♀, Wentworth Falls , NSW, 820 m, 4.xii.[20]14, D. & C. Emery ; 1♀, Evans Lookout, Blackheath , NSW, 33°06'60"S 150°18'29"E, 5.xii.[20]14, C. & D. Emery ; 2♂♂, 15 km SW Nowra , NSW, 35°00'35"S 150°28'37"E, 280m, 1.i.[20]15, S., N., C. & D. Emery ; 1♂, Towlers track, Kuringai-Chase National Park ., NSW, 33°37'33"S 151°16'39"E, 8.i.[20]15, D. Emery (DE) .

Distribution, habitat and seasonality. Occurring within central New South Wales from Brisbane Waters and Royal National Parks west to Wambool Nature Reserve near Bathurst and south to around Nowra, with isolated old specimens collected from eastern Victoria ( Fig. 18 ). Males occur mainly on the trunks and upper branches of smoothbarked eucalypts. Populations may be sympatric but are generally not syntopic with Y. spectabilis sp. nov., which prefers the tops of shrubs in open woodland. However, in open woodland, females of Y. regalis sp. nov. may also be located close to the ground on woody shrubs. They can be encountered from late November to January.

Description

Male ( Figs 2C , 3C , 19A,B , 52C). Head almost as wide as mesonotum, mainly black, with an ochraceous central triangular marking posterior to ocelli, widest at posterior margin, reducing anteriorly, and a second small ochraceous spot between ocelli and eye in some specimens; ocelli pink to pale red; dorsal postclypeus black centrally, dark brown laterally, ventral surface black with black transverse grooves, lateral and posterior borders ochraceous; anteclypeus black, rostrum black at base, dark brown centrally, black at apex, reaching anterior margins of hind coxae; lora black with ochraceous lateral margin, gena black; eyes dull black; antennae black, supra-antennal plates black, reddish area at junction with pedicels.

Thorax mainly dull black with variable brown patterning. Pronotum black, lateral half or anterior margin brown, with a thin central brown line tapering and not reaching anterior or posterior margins, brown with patchy black markings over raised lateral areas of pronotum, fissures mainly black; pronotal collar black, posterior margin brown, margins of lateral angles brown, paranota black with anterior angles brown. Mesonotum black with dark brown coloration between sigella in some specimens, parapsidal suture dark brown, scutal depressions and surrounds black; cruciform elevation arms black, lateral depressions brown. Metanotum black.

Legs. Coxae mainly black, fore coxae black anteriorly, reddish-brown posteriorly with a black lateral stripe, mid and hind coxae black, coxal membranes red, basisterna black; meracantha small, narrow, orange, black at base, pointed, overlapping one-third of opercula; trochanters orangered with black medial stripe; fore femora with lateral and posterior areas red, black medially and around base of femoral spines; femoral spines erect, reddish to black at base tending dark brown at tips, mid and hind femora black anteriorly, remainder orange-brown, with variable black stripe laterally, distal femoral joints orange-red; tibiae black proximally, tending brown towards tarsi; tarsi reddish-brown becoming black towards claws; claws dark brown, black at tips.

Wings with fore wing costal veins orange-brown, darker central rib; pterostigma mottled red; basal cell yellow-brown to translucent with black anterior border; basal membranes bright orange; other veins dark brown to black, with eight apical cells; hind wing plagas white along margins of anal cell 3 and vein 2A, central area of jugum variably clear, clear posteriorly, some reddish stippling at base, with six apical cells.

Opercula ( Fig. 3C ) medium, spatulate, following body axis ventrolaterally, depressed centrally, variably black over basal (a) viewed laterally from the left; (b) viewed ventrally; (c) aedeagus; and (d) apex of

theca. Characters as depicted in Fig. 4 . Specimen from Blackheath (33°35'S 150°50'E).

half, orange across remainder, clearly separated.

Timbals ( Fig. 2C ) with five distinct long ribs; long ribs 1–3 extending across surrounding membrane and fused dorsally along basal spur; long rib 4 unfused to basal spur; long rib 5 independent of basal spur, comparatively shorter, extending ventrally across half of membrane; prominent intercalary rib between long ribs 3 and 4; large ridged dome on posterior timbal plate extending across two-thirds of timbal; apodeme pit oval-shaped and conspicuous.

Abdomen with tergite 1 black with brown-ochraceous border around timbal cavity; tergite 2 black; tergites 3–7 black with orange posterior margins, extending laterally to epipleurites, increasing anteriorly on tergites 6–7 on either side of midline; tergite 8 shiny black, with variable brown lateral markings. Epipleurites with medial black strip, orange over reminder. Sternite II mainly black, central ochraceous spot; sternite III black medially, orange-red over central third posterior to timbal cavity, black laterally; sternites IV–VI orange, posterior halves becoming progressively reddish, black midline markings over anterior two-thirds of each sternite, decreasing posteriorly; sternite VIII orange brown, faint black longitudinal markings either side of midline, with brownish pubescence ( Fig. 52C).

Genitalia ( Fig. 20 ). Pygofer black, posterior border below dorsal beak and around upper lobe variably brownochraceous; dorsal beak black, anal styles pale brown; upper lobe mainly black, variably pale brown on posterior borders; basal lobe black. Uncus brownish; in lateral view beak-like; lobes in ventral view bulbous, with tapering lateral termination; claspers clearly divided, short, with apices gradually tapering laterally.Aedeagus with pseudoparameres extending around two-thirds the length of theca; theca recurved ventrally at 120° towards apex, with a transparent flange along the outer margin of recurvature, this broadly smooth, similar width to theca, terminating before apex; apex short, knob-like, with distinct cornuti around base.

Female ( Fig. 19 C–F). Head colour on dark specimens similar to male ( Fig. 19C ), orange-brown on lighter specimens with black marking around ocelli and supra-antennal plate, black spot at medial border of eye.

Thorax with a much greater range of brown coloration than males, mainly black on some specimens, with pronotal collar black; on brown specimens, black markings restricted to lateral and posterior margins around a broad brown midline strip, variably along fissures, pronotal collar brown, with lateral angles black; mesonotum black on dark specimens, cruciform elevation black, lateral depressions with variable brown mottling, metanotum black; mesonotum brown on pale specimens, patchy black markings over submedial and lateral sigilla, cruciform elevation yellow-ochraceous, lateral depressions ochraceous, anterior and posterior depressions dark brown to black, scutal depressions black, metanotum black.

Abdomen. Colour variation similar to thorax. On dark specimens, similar to male; sternite VII ochraceous with variable black mark on anterior margin; abdominal segment 9 black dorsally, becoming orange laterally with lateral black spot. On pale specimens, tergite 1 black; tergites 2–8 orange brown, anterior margins black, reducing posteriorly, black midline markings reducing posteriorly, becoming blacker towards lateral margins, black transverse stippling along central segment of tergites; epipleurites almost entirely pale brown; sternite II black; sternites III–VI pale brown with midline black marking; sternite VII pale brown with anterior black line directed posterolaterally; abdominal segment 9 brown with three longitudinal black stripes on midline and either side of midline, darker brown spot in lateral depression. On all specimens, dorsal beak black; ovipositor light brown, becoming black at tip, extending> 1 mm beyond apex of abdominal segment 9. Anal styles orange-red; ovipositor sheath dark brown to black.

Measurements (in mm; range with mean in parentheses: 12 males, 12 females). Body length: male 23.5–25.2 (24.8); female 25.6–29.0 (27.5). Fore wing length: male 26.8–29.8 (28.5); female 29.7–33.5 (32.1). Head width: male 7.1–7.7 (7.4); female 7.9–8.2 (8.0). Pronotum width: male 6.7–7.6 (7.1); female 7.4–8.6 (8.0). Abdomen width: male 6.7–7.3 (6.9); female 6.9–8.0 (7.6). Ovipositor length 8.8–10.6 (9.7).

Etymology. From the Latin “ regalis ”, meaning royal or kingly, reflecting the stately coloration and song of this species as well as its abundance in the Royal National Park, south of Sydney.

Distinguishing features. Within the Y. abdominalis species group, Y. regalis sp. nov. is superficially similar in appearance to Y. abdominalis , Y. electrica sp. nov, Y. grandis sp. nov., Y. spectabilis sp. nov., Y. serrata sp. nov., Y. subalpina sp. nov. and Y. verrens sp. nov. Nevertheless, Y. regalis sp. nov. is readily distinguished from Y. abdominalis , Y. electrica sp. nov, Y. spectabilis sp. nov and Y. serrata sp. nov. by the following combination of characters: (1) large size with body length> 23 mm, (2) lack of symmetrical, orange triangular patterns on tergites 4–7, (3) ovipositors of females extend> 1 mm beyond the posterior abdomen. The forewing basal membrane of Y. regalis sp. nov. is orange, in contrast to the grey or pale orange coloration in Y. grandis sp. nov. Geographically, populations of Y. regalis sp. nov. do not overlap with those of Y. grandis sp. nov. and only marginally with Y. subalpina sp. nov. Variations in the phenotype of male Y. regalis sp. nov., Y. subalpina sp. nov. Y. verrens sp. nov. and, Y. grandis sp. nov. make differentiation difficult. However, the black midline markings on the sternites are far more prominent in Y. regalis sp. nov. than in Y. subalpina sp. nov. or Y. grandis sp. nov. In Y. regalis sp. nov., the white colouration on the hind wing plaga is bold and extends to cover the entire jugum. Contrastingly, in Y. verrens sp. nov. the colouration is dull white and restricted along the margins of anal cell 3 and vein 2A. In female specimens, the ovipositors extend beyond the abdomen around 1.0 mm for Y. subalpina sp. nov., 1–1.5 mm for Y. regalis sp. nov. and> 1.5 mm for Y. grandis sp. nov. The length of the ovipositor of Y. regalis sp. nov. is> 8.8 mm, <8.6 mm for Y. subalpina sp. nov. and> 10.0 mm for Y. grandis sp. nov. and Y. verrens sp. nov.

Calling song ( Fig. 21 , 53L ). Males of Y. regalis sp. nov. produce a distinct, rapidly emitted, calling song with correspondingly rapid and regular amplitude modulations ( Fig. 21 ). Each separate phrase appears to comprise a single echeme, with the most conspicuous amplitude inflections occurring at the beginning and end of each echeme ( Fig. 21A ). Smaller amplitude modulations occur in the intervening duration between these inflections. This gives the song a “swirling” quality. Closer inspection of the echeme reveals that it is composed of well-defined syllables ( Figs 20B , 21C ), some of which are organized into a series of closely–spaced macrosyllables immediately prior to each minor amplitude modulation ( Fig. 21B ). The amplitude inflections at the beginning and end of each echeme have a denser (less defined) syllable structure ( Fig. 21D ). Recordings of individuals around Sydney Basin (n = 10) illustrate significant variation in the length of the single echeme phrases. Phrases at the start of a bout of calling song range from c. 5–12 s duration. Phrases for the remainder of the song are slower (between c. 13.5 s duration). When males are actively calling, a gap of 0.08– 1.8 s duration typically separates each series of phrases. In some instances, these single echeme phrases are preceded (or succeeded) by intervening short chirps (short echemes or long macrosyllables, 0.06– 0.08 s duration, separated by gaps of c. 1–5 s), which may serve to elicit a response from a nearby female (not illustrated). These short chirps are produced with increasing frequency immediately before taking flight. It has been noted that males in Royal National Park accelerate the emission rate of their song immediately before taking flight, and tick in flight.

The highest frequency plateau ranges from 8.2–15.4 kHz, with a dominant frequency of 10.3–13.3 kHz ( Fig. 21E ).

The rapid amplitude modulations render the song of Y. regalis sp. nov. distinct among its relatives. One other species, Y. spectabilis sp. nov. produces similar modulations; however modulations occur at a much slower rate than in Y. regalis sp. nov.

Figures

Figure 2. Photos of male left timbals, with dorsal edge at left and posterior margin at bottom: (A) Yoyetta timothyi sp. nov., Gordon (33°76'S 152°15'E); (B) Y. denisoni (Distant), O’Connor (35°16'S 145°06'E); (C) Y. regalis sp. nov., Blackheath (33°35'S 150°50'E); (D) Y. spectabilis sp. nov., Barren Grounds (34°41'S 150°44'E); (E) Y. subalpina sp. nov., Cooma (36°20'S 148°14'E); (F) Y. verrens sp. nov., Torrington (25°15'S 153°11'E); (G) Y. abdominalis (Distant), Hobart (42°52'S 147°19'E); (H) Y. kershawi (Goding & Froggatt) stat. rev., comb. nov., Toolangi, (37°32'S 145°28'E); (I) Y. serrata sp. nov., Nimmitabel (36°31'S 149°14'E); (J) Y. aaede (Walker), Mt Lofty (34°59'S 138°43'E); (K) Y. electrica sp. nov., Arrawarra (30°02'S 153° 11'E); (L) Y. grandis sp. nov., Forrest (38°31'S 143°43'E). Scale bars = 200 µm; J= 500 µm.

Figure 3. Photos of male left opercula, with dorsal edge at left and posterior margin at bottom: (A) Yoyetta timothyi sp. nov., Gordon (33°76'S 152°15'E); (B) Y. denisoni (Distant), O’Connor (35°16'S 145°06'E); (C) Y. regalis sp. nov., Blackheath (33°35'S 150°50'E); (D) Y. spectabilis sp. nov., Barren Ground (34°41'S 150°44'E); (E) Y. subalpina sp. nov., Cooma (36°20'S 148°14'E); (F) Y. verrens sp. nov., Torrington (29°15'S 151°42'E); (G) Y. abdominalis (Distant), Hobart (42°52'S 147°19'E); (H) Y. serrata sp. nov., Nimmitabel (36°31'S 149°14'E); (I) Y.electrica sp. nov.,Arrawarra (30°2'S 153°11'E); (J) Y. grandis sp. nov., Forrest (38°31'S 143°43'E); (K) Y. aaede (Walker), Mt Lofty (34°59'S 138°43'E); (L) Y. kershawi (Goding & Froggatt) stat. rev., comb. nov., Toolangi, (37°32'S 145°28'E). Scale bars; A–J =200 µm; K=500 µm.

Figure 4. Yoyetta denisoni (Distant):illustration of male pygofer and internal genitalia;

Figure 6. Male calling song structure of Yoyetta denisoni (Distant) illustrated in waveform plots, including (A) typical rapid chirping song (in flight); (B) expanded section fromA showing three macrosyllables; (C) expanded section from B showing detailed structure of a macrosyllable. The final subfigure (D) is a spectrogram displaying song frequency. This specimen was recorded in the field at Canberra (35°16'S 149°07'E) by LWP using RS2 (see Methods and terminology).

Figure 18. Map of southeastern mainland Australia showing the geographical distribution of Yoyetta regalis sp. nov. (closed squares), Y. abdominalis (Distant) (open circles) and Y. spectabilis sp. nov. (open triangles).

Figure 19. Yoyetta regalis sp. nov., (A) male holotype, Royal National Park (34°06'S 151°03'E), dorsal habitus; (B) male, ventral habitus; (C) female, dark specimen, Royal National Park dorsal habitus; (D) female, dark specimen, ventral habitus; (E), female, lighter specimen, Royal National Park, dorsal habitus; (F) female, lighter specimen, Royal National Park, ventral habitus. Scale bar = 10 mm.

Figure 20. Yoyetta regalis sp. nov., illustration of male pygofer and internal genitalia;

Figure 21. Male calling song structure of Yoyetta regalis sp. nov., illustrated in waveform plots, including (A) a complete single echeme phrase with amplitude inflections indicated by downward pointing arrows (further detail in text); (B) expanded section from the middle of the echeme (inA) showing minor amplitude modulations and intervening macrosyllables; (C) further expanded detail of a minor amplitude modulation showing clean syllable structure; (D) expanded detail of an amplitude inflection showing dense structure. The final subfigure (E) is a spectrogram displaying song frequency. This specimen was recorded at Wises Track, Audley (34°08'S 151°03'E) by LWP using RS1 (see Methods and terminology).

Figure 53. A comparative sample of wave plots illustrating the temporal song structure of cicadas in the Yoyetta abdominalis (Distant) species group (for species where song recordings were available). These include (A) Y. denisoni (Distant), recorded in the field at Canberra (35°16'S 149°07'E) by LWP using RS2; (B) Y.timothyi sp. nov., recorded in the field at Glen Elgin (29°39'S 152°03'E) by LWPusing RS3; (C) Y.electrica sp. nov., recorded in captivity atArrawarra (30°02'S 153°11'E) by LWP using RS2; (D) Y.hunterorum (Moulds), recorded in the field at Garie Beach (34°11'S 151°04'E) by B. Smith using RS5; (E) Y. grandis sp. nov., recorded in the field at Brindabella Range (35°20'S 148°50'E) by LWP using RS2; (F) Y. verrens sp. nov., recorded in captivity at Torrington (25°15'S 151°44'E) by LWP using RS3; (G) Y. subalpina sp. nov., recorded in the field at Talleganda (35°37'S 149°30'E) by LWP using RS2; (H) Y. aaede (Walker), specimen captured in Adelaide (34°56'S 138°39'E) and recorded in captivity by NJE using RS6; (I) Y. abdominalis, recorded in the field at Nimmitabel (36°31'S 149°14'E) by LWP using RS2; (J) Y.serrata sp. nov., recorded in the field at Brown Mountain (36°36'S 149°23'E) by LWP using RS2; (K) Y. spectabilis sp. nov., recorded in the field at Kanangra Boyd National Park (33°53'S 150°03'E) by LWP using RS2; and (L) Y. regalis sp. nov., recorded at Wises Track,Audley (34°08'S 151°03'E) by LWP using RS1.An explanation of recording systems (RS1–6) is provided in Methods and terminology.