Yoyetta subalpina, 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 : 313-320

Yoyetta subalpina sp. nov.

http://zoobank. org/NomenclaturalActs/ 1C0EBF61-D05D-4EB6-A792-FFCD6012A489

Figs 2E , 3E , 26– 31 , 52E, 53G

Holotype ♂, 18 km E Cooma , NSW, 36°20'47"S 148°13'57"E, 10.xii.2010, L. Popple, 514-0013 ( AM K.559471) . Paratypes NEW SOUTH WALES: 1♂ 1♀, Piccadilly Circus, Brindabella Ra. , ACT, 35°21'S 148°48'E, 26.xii.1978, D. C. F. Rentz, ANIC Database no. 20 010822 ; 1♀, Snowy River, Mt Kosciusko , 4000FT, 12.xii.1931, L. F. Graham, ANIC Database No. 20 010821 ; 1♂, Old Adaminaby NSW, 24.i.1967, T. G. Campbell, ANIC Database no. 20 010820 ; 2♀♀, NSW Snowy Plains , 19.ii.1969, M. G. Stanger, on Eucalypt, [1♀] ANIC Database no. 20 010823 ; 1♀, Cooma NSW, 10.i.[19]06, Sellar, W. W. Froggatt collection, ANIC Database No. 20 010815 ; 1♂, Tinderry Mountains, Round Flat , netted, 1142 m, 25.xii.2014, 35°42'52.08"S 149°17'40.48"E, D. J. & R. L. Ferguson, ANIC Database No. 20 010718 ; 1♂, Tinderry Mountains, Tinderry Rd verge, netted, 1160 m, 35°43'49.17"S 149°18'07.92"E, 21.xii.2014, D.J. Ferguson, ANIC Database No. 20 010816 ( ANIC) ; 2♂♂ 4♀♀, Mt Kosciusko , NSW, 36°27'S 148°16'E, 18.i.1931 ( HEM 2554–58) ; 1♀, Jindabyne NSW, 36°25'S 148°37'E, 17.i.1953, ( HEM2559 ) ( MMV) ; 4♂♂, 3♀♀, 1 km S. of Brown Mountain, via Nimmitabel , 36°36'16"S 149°22'31"E, 29.xii.2009, L. W. Popple, 514-0003 to 514-0009 ; 1♂, Tom Groggin Road , 18 km ESE of Cooma, 36°20'48"S 148°13'57"E, 10.xii.2010, L. W. Popple, 514-0014 ( LWP) ; 1♀, 7 miles [SW of?] Countegany on Kybean Rd , 16.xii.[19]62, L. R. G[reenup]?, 105 ; 1♂ 8♀♀, Mt David , 30 km SW of Oberon, 26.xii.1985, S. &. B. Underwood ; 1♂ 2♀♀, 12 mi. from Cooma on Nimitabelle [sic.] Rd , 27.xii.1973, Euc.pauciflora , L. R. G[reenup] ; 3♂♂ 2♀♀, Yarrangobilly Caves road, Kosciuszko NP, 35°33.258'S 148°30.842'E, 17.i.2011, 1305 m [elev.], K. Hill, D. Marshall, AU.NS.YAX, C. Simon lab voucher, Yoyetta “double-zit” ; 1♂, same data; legs in ETOH, body pinned, 11.AU.NS.YAX.01 ( MSM) ; 1♂ 6♀♀, Piccadilly Circus, Brindabella Ranges , NSW, 35°21'34"S 148°47'59"E, 10–14.i.2002, D.Emery ; 2♀♀, Kanangra Boyd National Park , NSW, 33°59'18"S 150°02'25"E, 8.i.2010, D. Emery & L. Popple (DE) . VICTORIA: 1♂ Mt Buffalo Nat. Pk , 16.i.1966, T.Weir ( UQIC) ; 1♀, 36°29'S 147°54'E, 8 km SE by E of Nariel Vic. , 16.i.1984, K. H.L. Key ; 1♀, 9 km E of Mt Pendergast, E of Benambra, Vic. , 1325m, 36°53'S 148°05'E, 29.xii.1983, K. H. L. Key, ANIC Database No. 20 010819 ( ANIC) ; 3♂♂ 3♀♀, midway between Harrietville & St Bernard Hospice, 19 Dec 1933, A. Musgrave ( AM K.307236–41) ; 1♂, Sommerville Ave, Lorne, Vic , 15.xii.2002, D. J. Hilton ; 2♀♀, 1 km SE Forrest, Otway Ranges, Vic , 38°54'39"S 143°74'40"E, 5.ii.15, S. Emery & T. Corbin ; 1♂, Kinglake, Vic , 37°28'11"S 145°20'12"E, 17.xii.2016, S. Emery & T. Corbin ; 2♂♂, Country Fire Authority Station, Toolangi , 37°32'32"S 145°28'29"E, 11.i.2017, S. Emery & T. Corbin (to light) ; 2♂♂, same location, 23.i.2017, N., C. & D. Emery (DE) ; 2♂♂ 9♀♀, Mt Hotham Alpine Reserve , 1,830 m [elev.], 2.i.1976, M. S. & B. J. Moulds (1♂ 1♀ in cop) ; 4♂♂, same data, 1.i.1976 , 1♂ in cop; Mt Hotham Village , 36°59'S 147°11'E, 1–13.i.1990, G. R. Brown & M. A. Terras ( MSM) ; 1♂, Kinglake, Vic , 37°32'S 145°20'E, 4.xii.1927, ( HEM2611 ) ; 5♀♀, Mt Buffalo, Vic , 36°43'S 146°46'E, 29.xii.1951 ( HEM 2612–16) ; 1♂, Country Fire Authority Station, Toolangi , 37°32'32"S 145°28'29"E, 27.xii.2018, S. Emery & T. Corbin (to light; T 22389) ; 5♂♂ 3♀♀, Allambee, Victoria, 14.i.2019, J. Creen ( HEM5765–7 [♀], HEM5768–72 [♂]) ( MMV) . AUSTRALIAN CAPITAL TERRITORY. 1♀, 35°21S 148°48E, Piccadilly Circus, Brindabella Ra. A.C.T., 26.xii.1978, D. C. F. Rentz ( ANIC) ; 1♂, Mt Franklin Rd, Brindabella Ranges , 35°32'S 148°46'E, 5.xii.2004, B.Kenway,514-0001 ; 1♂, Piccadilly Circus, Brindabella Range , 35°21'37"S 148°47'59"E, 28.xii.2009, L. W. Popple, 514-0002, LT2009371 ; 1♂ 1♀, 1.5 km N. of Bull’s Head , 35°22'39"S 148°48'16"E, 15.i.2011, L. W. Popple, 514-0015, 514-0017, LT2009371 ; 1♀, Mt Franklin area , 35°29'23"S 148°45'48"E, 15.i.2011, L. W. Popple, 514-0018, LT2009371 ( LWP) .

Additional records from song recordings NEW SOUTH WALES: Rocky Pic Road, Talleganda National Park , NSW, 35°37'12"S 149°29'54"E, 5.xii.2009, L. W. Popple . VICTORIA: Ridge Rd, Mt Dandenong , 600 m, 37°49'28"S 145°21'34"E, 15.xii.2017, T. Corbin .

Distribution, habitat and seasonality. Cool temperate alpine areas from Kanangra-Boyd National Park in central New South Wales south to the Upper Yarra region in central Victoria, with an isolated population in the Otway Ranges ( Fig. 26 ). Adults occur mainly on smooth-barked eucalypts on the main trunks and upper branches. They can be encountered from December to February.

Description

Male ( Figs 2E , 3E , 27 A– B, 27F, 51G, 52E). Head as wide as mesonotum, black, with an ochraceous diamond-shaped marking on midline, posterior to ocelli; ocelli pink to pale; dorsal postclypeus black with central ochraceous triangular marking, apex directed anteriorly; ventral postclypeus black with anterior brown marking on midline, and ochraceous, lateral and posterior margins; anteclypeus black, rostrum brown at base, becoming black at apex, reaching the middle of hind coxae; lora black with ochraceous anterior margin, gena black; eyes dull black; antennae black, supra-antennal plates black, ochraceous around base of pedicel.

Thorax predominantly black. Pronotum black, some specimens with brown in fissures; pronotal collar black, posterior margin ochraceous in some specimens, margins of lateral angles light brown, paranota black, anterior angles ochraceous in some specimens. Mesonotum black; cruciform elevation arms black, lateral depressions tinged brown. Metanotum black.

Legs. Coxae mainly black, fore coxae black with proximal ochraceous triangular stripe on posterior surface; mid and hind coxae black; coxal joint and membranes red, basisterna black; trochanters black with central segment orange-red; meracantha small, narrow, orange, black at base, pointed, minimally overlapping opercula; fore femora striped longitudinally orange-red and black, black along base of femoral spines femoral spines erect, black at base tending dark brown at tips; mid and hind femora black, orange-brown longitudinal

(a) viewed laterally from the left; (b) viewed ventrally; (c) aedeagus; and (d) apex of theca.

Characters as depicted in Fig. 4 . Specimen from Piccadilly Circus (35°22'S 148°48'E).

stripe on medial aspect, distal femoral joint orange-red; fore tibiae black, brown at distal tip; mid tibiae black, with medial ochraceous stripe; hind tibiae black proximally ochraceous over distal half; fore tarsi black with brown distal margins; mid and hind tarsi blackish-brown becoming black towards claws; claws dark brown, black at tips.

Wings with fore wing costal veins black, browner central rib, becoming ochraceous distally; pterostigma mottled red; basal cell translucent with black anterior border; basal membranes bright orange; other venation dark brown to black; with eight apical cells; hind wing plagas white over entire anal cell 3 and medial third of ac2(v), thin along vein 2A, an irregular grey-black marking at proximal end of jugum, reddish stippling at base of ac2(v), veins 2A and 3A pale, remainder dark brown to black, with six apical cells.

Opercula ( Fig. 3E ) medium, spatulate, following body axis ventrolaterally, depressed centrally, variably black over basal half, orange-brown across remainder, clearly separated.

Timbals ( Fig. 2E ) with five distinct long ribs; long ribs 1–4 extending across surrounding membrane and fused dorsally along basal spur; long rib 5 independent of basal spur, comparatively shorter, extending ventrally across half of membrane; large ridged dome on posterior timbal plate extending across three-quarters of timbal; apodeme pit ovalshaped and conspicuous.

Abdomen. Tergite 1 black, with orange-red membranes around timbal cavity; tergite 2 black; tergites 3–7 black with orange posterior margins, extending laterally to epipleurites, increasing on tergites 6–7 on either side of midline; tergite 8 shiny black, with ventral margins orange. Epipleurites with medial black strip, orange over reminder. Sternite II mainly black, with central orange spot; sternite III black medially, becoming orange laterally along posterior margin; sternites IV–VI orange, posterior halves becoming progressively reddish, diffuse black midline markings over anterior twothirds of each sternite, decreasing posteriorly, sternite VIII dull orange-red, with black pubescence.

Genitalia ( Fig. 28 ). Pygofer black, upper lobe mainly black, apex brown, basal lobe black; dorsal beak black, dark brown posterior margin, anal styles orange-red. Uncus reddishbrown; in lateral view beak-like and elongated; lobes in ventral view narrow, with rounded lateral termination; claspers clearly divided, short, bulbous, with apices tapering laterally.Aedeagus with pseudoparameres almost extending as far as theca; theca recurved ventrally at around 180° towards apex, with prominent transparent flange along distal margin of recurvature, smooth and broadening to> 3× width of theca; remainder of theca is short, with <20% of theca extending apically beyond margin of recurvature; apex blunt, with 2–3 cornuti on ventral surface and several small cornuti on dorsal side at tip.

Female ( Fig. 27 C–E). Head, thorax and abdomen of dark specimens similar to male. Paler, brown-coloured females apparently occur at a ratio of approximately 1:2.

Head in dark specimens, similar to male; in paler specimens mainly brown with black markings around ocelli extending to the supra-antennal plate, brown at anterior margin, black spot at medial border of eye; dorsal postclypeus ochraceous centrally, dark brown laterally; ventral postclypeus black with central ochraceous spot anteriorly; anteclypeus and rostrum similar to male; gena variably ochraceous and black, lateral margins ochraceous posteriorly, black anteriorly.

Thorax in dark specimens, similar to male. In paler specimens; pronotum with black markings restricted to lateral and posterior margins of a broad ochraceous midline strip over central third, and variably along fissures and lateral margins; pronotal collar brown, with lateral angles black; mesonotum brown with black submedian and lateral sigilla, and a diffuse black central triangular marking with base around scutal depressions, apex directed anteriorly on midline to posterior third of submedian sigilla; cruciform elevation black on central, medial portion and lateral margins, yellow-ochraceous on posterior margins, 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; abdominal segment 9 black dorsally with ochraceous triangular markings either side of midline, tapering posteriorly, ending midway to dorsal beak, becoming orange laterally, reddish on ventral margin. On brown specimens; tergite 1 black, tergite 2 black on midline, orange merging to black laterally; tergites 3–8 black with orange posterior margins, increasing posteriorly, extending laterally to epipleurites; epipleurites with medial black strip, orange over remainder; sternite II black; sternites III–VI pale brown with midline black marking; sternite VII pale brown with anterior black spot either side of midline; abdominal segment 9 orange-brown with three longitudinal black stripes faint on midline and prominent either side of midline. On all specimens, dorsal beak orange to black at base, black at tip; ovipositor dark 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: 10 males, 8 females). Body length: male 25.0–25.1 (25.1); female 24.0–27.0 (26.0). Fore wing length: male 27.7– 28.5 (28.1); female 29.4–33.2 (31.9). Head width: male 7.0–7.6 (7.3); female 6.8–7.6 (7.4). Pronotum width: male 6.7–7.4 (7.1); female 6.5–8.4 (7.5). Abdomen width: male 7.4–8.0 (7.7); female 7.5–8.2 (7.8). Ovipositor length 7.6–8.4 (8.0).

Etymology. Since most specimens have been collected in cool temperate areas, the species name reflects the climatic situation where this species is typically found.

Distinguishing features. Yoyetta subalpina sp. nov. displays concentric orange bands on the posterior margins of tergites 3–7; these are absent on Y. denisoni , Y. kershawi stat. rev., comb. nov., Y. timothyi sp. nov. and Y. grandis sp. nov. It 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 membranes of Y. regalis sp. nov. and Y. subalpina sp. nov. are bright orange contrasting with the grey basal membranes of Y. grandis sp. nov. in southern Victoria and the dull orange basal membranes of specimens of Y. grandis sp. nov. in the Grampians Ranges. Morphologically, male specimens of Y. subalpina sp. nov., Y. verrens sp. nov. and Y. regalis sp. nov. are difficult to differentiate, but the follow features can be used to achieve the correct identification. In Y. subalpina sp. nov., the white colouration on the hind wing plaga is bold and extends to cover the entire jugum (cf. in Y. verrens sp. nov. the colouration is dull white and restricted along the margins of anal cell 3 and vein 2A). In Y. subalpina sp. nov., the sternites are predominantly orange, sometimes with diffuse black spots medially (cf. in Y. regalis sp. nov. where the sternites have bold black medial markings that occupy more than half of each sternite). In female specimens, there is substantial colour variation morphologically (orangebrown to black) within species, but the ovipositors extend beyond the abdomen around 1.0 mm for Y. subalpina sp. nov., 1–1.5 mm for Y. regalis sp. nov.,> 1.5 mm for Y. grandis sp. nov., and> 2 mm for Y. verrens 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.

Calling song ( Figs 29–31 , 53G ). The male calling song of Y. subalpina sp. nov. is a simple repetition of phrases, each

comprising an echeme followed by a macrosyllable ( Fig. 29A ). Closer inspection of the echeme ( Fig. 29B ) reveals that it comprises a series of macrosyllables that increase in amplitude and coalesce at the highest point of amplification, typically occurring between 0.16 and 0.43 s duration (n = 13). A short gap of 0.01– 0.04 s duration separates the echeme from the proceeding macrosyllable. The macrosyllable is typically louder than the echeme. It lasts between 0.05 and 0.09 s ( Fig. 29C ). Gaps of approximately 0.5 s duration separate each phrase. After warming up a little, males sometimes produce soft ticking (syllable trains) in the gaps between each phrase, with longer syllable trains between bouts of phrases ( Fig. 30 ).

Males from most locations across the geographical distribution of this species had a frequency plateau of 8.4–11.1 kHz and a dominant frequency ranging between 9.4 and 10.1 kHz ( Fig. 31 ). The exceptions were males from Talleganda that produced a slightly lower frequency plateau, ranging from 7.1–9.7 kHz with a dominant frequency of 8.4–9.0 kHz (n = 4; Fig. 30 ).

Within the Y. abdominalis species group, the calling song of Y. subalpina sp. nov. is most similar to Y. verrens sp. nov. The main difference in the call of Y. subalpina sp. nov. is the addition of a macrosyllable following the echeme, which gives it a distinct two-note signature (cf. the single echeme sweeping notes of Y. verrens 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 26. Map of eastern Australia showing the geographical distribution of Yoyetta subalpina sp. nov. (open triangles), Y. serrata sp. nov. (closed squares) and Y. grandis sp. nov. (open circles).

Figure 27. Yoyetta subalpina sp. nov., (A) male, Cooma (36°20'S 148°13'E), dorsal habitus; (B) male, Cooma, ventral habitus; (C) female, Piccadilly Circus (35°21'S 148°48'E), dorsal habitus; (D), female, Piccadilly Circus, ventral habitus; (E), female, brown form, Piccadilly Circus, dorsal habitus; (F), live male specimen, Apollo Bay (38°45'S 143°33'E), photo by Adrian Tritschler. Scale bar (for preserved specimens) = 10 mm.

Figure 28. Yoyetta subalpina sp. nov., illustration of male pygofer and internal genitalia;

Figure 29. Typical male calling song structure of Yoyetta subalpina sp. nov., illustrated in waveform plots, including (A) two phrases; (B) expanded section from A showing a single phrase with an echeme followed by a long macrosyllable; (C) expanded section from B showing more detail from the echeme and the macrosyllable. The final subfigure (D) is a spectrogram displaying song frequency. The specimen was recorded in the field at Piccadilly Circus (35°22'S 148°48'E) by LWP using RS2 (see Methods and terminology).

Figure 30. Comparative male calling song structure of Yoyetta subalpina sp. nov., illustrated in waveform plots, including (A) a 20 s section containing echemes, interspersed with a “soft ticking” segment; (B) expanded “soft ticking” section (fromA)showing 10 syllables.The final subfigure (C) is a spectrogram displaying song frequency.The specimen was recorded in the field at Talleganda (35°37'S 149°30'E) by LWP using RS2 (see Methods and terminology).

Figure 31. Comparative structures of the callings songs for Yoyetta subalpina sp. nov., from a range of locations, including: (i) Brindabella Range (35°22'S 148°48'E); (ii) Kanangra Boyd National Park (33°53'S 150°03'E); (iii) Square Rock (35°31'S 148°55'E); (iv) Talleganda National Park (35°37'S 149°30'E); (v) Brown Mtn (36°36'S 149°23'E). Specimens were recorded in the field by LWP using RS2.

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.