Yoyetta denisoni ( Distant, 1893 )

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 : 281-288

Yoyetta denisoni ( Distant, 1893)

Figs 1 , 2B , 3B , 4–6 , 10A , 52B, 53A

Melampsalta denisoni Distant, 1893: 78 , plate VII, figs 14, 14a, 14b (female); Goding & Froggatt, 1904: 636, plate xviii, figs 7, 7a; Distant, 1906: 174; Ashton, 1912: 24; Ashton, 1914: 354; Greenup, 1964: 23.

Cicadetta denisoni .— Moulds, 1990: 156–157, plate 17, figs 8, 8a, plate 24, fig. 14; Naumann, 1993: 22, 80, 149; Moss & Popple, 2000: 57; Moulds & Cowan, 2002: 26; Williams, 2002: 15; Arensberger et al., 2004: 558, 566; Emery et al., 2005: 109.

Not Cicadetta denisoni .— Ewart, 1986: 51–52, table 1.

Yoyetta denisoni ( Distant, 1893) .— Moulds, 2012: 24, 237; Sanborn, 2014: 554.

The original description of Y. denisoni ( Distant, 1893) refers to a male specimen and is based on material “contained in the Brussels Museum”. Therefore, this specimen must be treated as the holotype (below). There are also specimens of both sexes from the same locality in the NHM (listed below) with red “type” labels. However, as noted by Moulds (1990), the type locality (Port Denison near Bowen in Queensland, Australia) must be regarded as erroneous.

Material examined. Holotype male, [handwritten] “ Melampsalta denisoni Dist. ” / [typeset] “ Coll. R. I. Sc. N. B., Australie”, “ Coll. Camille van Voixem ”, [handwritten] “ Port Denison,W.68” / [typeset] “Type” / “ Holotype ” ( RBINS). 1♂ 1♀, same location and collection data as holotype [locality erroneous] ( NHM). NEW SOUTH WALES: 1♂, Batemans Bay,1.ii.20,ANIC database no. 20 010775; 1♂, New South Wales, Kioloa Creek crossing, 2.5 km W.on Dangerboard Road , temperate rainforest, 35°32'51"S 150°20'54"E, 27.i.2014, D. J. & R. L. Ferguson; ( ANIC); 1♂ 1♀, Tumbarumba , NSW, 35°51'54"S 148°06'54"E, 750m, 31.xii.13, A. Kwok, S. Travers; 2♂♂, Mt. Gibraltar , Bowral , NSW, 34°27'44"S 150°25'40"E, 800 m, 23.xii.13, N. & D. Emery; 1♀, same location, 10.xii.2017, L. Nolan (DE); 2♂♂, NSW Dampier SF Diamond Ck Survey Tk. 0.5 km N of Junctn with Little Sugarloaf Rd ( Deua National Park ), 17.xii.1991, F.Douglas ( AM K.264315, AM K.307123); 1♀, Wagga, 2.xii.1965, S. Hamlyn ( UQIC); 4♂♂, Lemann Compound, ca 10 km SE of Bowral, NSW, 34°32.50'S 150°29.20' E, 24.xii.1996, C. Lemann & S. McEvey, 12892, 12895–97; 1♀, Jenolan SF, 29.xii.1968, C. E. Chiswick; 1♂ 1♀, 17 miles from Cooma, on Bungarby Road, Euc.dives , 9.xii.1961, L. R. G[reenup] ( MSM). AUSTRALIAN CAPITAL TERRITORY: 1♂, 1♀, Tidbinbilla Nature Reserve , ACT, 9.iii.1978, P. Omay ( AM); 1♂ 1♀, Black Mtn, Canberra ACT, 15.xii.1964, L. J. Chinnielz, ANIC database nos 20 010698, 20 010701; 1♂, 35°17'S 149°06'E, Black Mt,ACT, 16.i.1987, D.C.F. Rentz,ANIC Database No. 20 010697; 1♂, Bl[ac]k Mt, 26.xii.71, E. Maldor, H.-M., K.-O., ANIC database no. 20 10694 ( ANIC); 1♂, Frith St, Acton, 35°15'58"S 149°06'34"E, 22.xi.2010, L.W. Popple, D.Emery, 510-0005; 1♂, same data as previous, 24.xi.2010, L. W. Popple 1♂, same data as previous, 6.xii.2010, Emerging, 510-0006; 3♂♂, same data as previous, 2.xii.2016, netted, 510-0012, 510-0014, 510-0015; 1♀, Frith St, Acton, 35°15'58"S 149°06'34"E, 24.xi.2010, 509-0010; 1♂ 1♀, same data, 6.xii.2010 ,510-0006, 509-0011 ( LWP); 2♂♂, same data as previous, 510-0013, 510-0016 ( QM); 1♂, Frith Rd,O’Connor,ACT, 35°16'07"S 149°06'32"E, 24.xi.2009, D.Emery & L. Popple (DE); 1♂, Aranda, 8.xii.2002, Allan Robertson; 1♀, Mt Majura , 27.xi.1977, G. Daniels ( MSM).

Additional published records. NEW SOUTH WALES: Raspberry Lookout, Gibraltar Range , 29°33'S 152°15'E ( Moss and Popple, 2000) .

Additional locations with audio recordings. NEW SOUTH WALES: Forest Track, Royal National Park, NSW, 34°09'04"S 151°01'24"E, 25.xi.2012, B. Smith; Girrakool Picnic Area , Brisbane Waters National Park , NSW, 33°25'54"S 150°16'35"E, 16.xii.2012, B. Smith; Old Grafton Rd, Glen Elgin ,

NSW, 29°39'20"S 152°02'38'E, 7.i.2016, L. W. Popple; Jindabyne district , NSW, 36°31'58"S 148°41'36"E, 30.xii.2009, L. W. Popple .

Distribution, habitat and seasonality. Yoyetta denisoni is found along the ranges and subcoastal forested areas of New South Wales from Gibraltar Range to the south coast

and inland to the Southern Tablelands ( Fig. 5 ). Populations also occur prominently in the Australian Capital Territory and adjacent ranges ( Fig. 5 ). Adults are active in the upper canopy of eucalypt forest and are difficult to attract or capture. They are present from November to January. There is also a single record from as late as March, although this is considered anomalous.

Description

Male ( Figs 1 A–D, 2B, 3B, 52B). Head approximately as wide as mesonotum; predominantly dull black, with a brown to orange-brown central marking on posterior dorsal area extending between lateral ocelli and through dorsal postclypeus; ocelli pink to red; postclypeus predominantly black, with circular dull reddish-brown markings above either side of central orange line, black transverse grooves below with orange-brown lateral and posterior margins, rounded laterally; lora black; gena black; anteclypeus black; rostrum black anteriorly, variably ochraceous centrally, black at apex, extending to posterior margin of middle coxae; eyes brown to dull black; antennae black, supra-antennal plates black.

Thorax almost entirely black; pronotum predominantly black, paramedian and lateral fissures black with intermediate ridges variably dull orange-brown laterally; pronotal collar black, sometimes with central lateral section ochraceous posteriorly. Mesonotum black, cruciform elevation arms black lateral depression orange-brown; scutal depressions black; parapsidal sutures black; metanotum black.

Legs with coxae black; coxal joints bright orange-red; meracantha small creamy, pointed, black at base, partly overlapping opercula; trochanters orange-red; meron reddish; fore femora striped black and red; mid and hind femora with lateral and anterior surfaces black, inner side ochraceous to dark brown; femoral joints orange to pale yellow; tibiae black to dark brown laterally becoming ochraceous medially and towards base, with fore tibial spines black, mid and hind tibial spines variably pale brown and black at tips; tarsal joints orange; tarsi dark brown, becoming black towards claws; claws brown, black at tips.

Wings with fore wing costal veins black; pterostigma with reddish-brown mottling; basal membranes red to orange; basal cell translucent brown; veins mainly black to dark brown, with eight apical cells; hind wing plaga white, extending over medial third of anal lobe, remainder of anal lobe diffuse white, veins mainly brown, CuA black proximally, brown after junction, with six apical cells.

Opercula ( Fig. 3B ) medium, spatulate, following body axis ventrolaterally; black at bases, red over remainder, including crests; clearly separated.

Timbals ( Fig. 2B ) with five distinct long ribs, long ribs 1–4 each spanning across timbal membrane and fused dorsally along basal spur; long rib 4 narrowing over third quarter; long rib 5 noticeably shorter than other long ribs; large ridged dome on posterior timbal plate extending across two-thirds of timbal; apodeme pit oval-shaped and conspicuous.

Abdomen with tergites 1–8 black. Epipleurites black. Sternite II black, partly ochraceous around inner surface (a) viewed laterally from the left; (b) viewed ventrally; (c) aedeagus; and (d) apex of

theca. Characters include: aed, aedeagus; bpl, basal lobe of pygofer; cl, clasper; db,

dorsal beak; ps, pseudoparameres; th, theca; upl, upper lobe of pygofer. Specimen from

O’Connor (35°16'S 145°06'E).

of tympanal cavity; sternite II black with small central ochraceous marking; sternites III–VII black; sternite VIII black on anterior margin, red over remainder, with black to pale brown pubescence ( Fig. 52B).

Genitalia ( Figs 4 , 10A ). Pygofer mainly black; dorsal beak black, upper lobe prominent, pointed, red at tip; basal lobe dark brown to black; anal styles red. Uncus orange-red, in lateral view beak-like and stumpy; lobes in ventral view less produced, rounded laterally; claspers divided, closely applied, with apices gradually tapering laterally. Aedeagus with pseudoparameres extending around half the length of theca; theca gradually recurved ventrally at 180° towards apex, with translucent flanges along outer margin of recurvature, these smooth dorsally,> 2× diameter of theca, tapering towards apex of theca; apex transparent, sclerotized, bifurcate, each branch directed laterally and dorsally around 90°, with multiple small cornuti on outer edge of each branch. Female ( Fig. 1 E–G). Head and thorax similar to male (except in brown specimens).

Wings similar to male. Fore wing basal membranes red to pink.

Abdomen with tergites 1–8 black (brown posteriorly in brown specimens). Sternites I–VIII black or mainly black; abdominal segment 9 with symmetrical dorsal triangular markings anteriorly, ochraceous and merging centrally, apices directed posteriorly over anterior third, remaining dorsal surface black dorsal beak black, lateral and ventral surface predominantly fiery red, with lateral black spot; anal styles and ovipositor sheath red; ovipositor dark brown to black, extending up to approximately 0.5 mm beyond dorsal beak.

Measurements (in mm; range with mean in parentheses: 17 males, 4 females). Body length: male 23.7–27.6 (25.7); female 25.6–26.9 (26.5). Fore wing length: male 28.9–32.5 (32.2); female 31.0–34.1 (32.0). Forewing width: male 9.6–11.2 (10.6); female 10.9–12.2 (11.3). Head width: male 6.5–7.9 (7.1); female 7.1–7.8 (7.3). Pronotum width: male 7.0–8.2 (7.7); female 8.1–8.4 (8.2). Abdomen width: male 7.2–8.7 (8.0); female 7.3–8.3 (8.0). Ovipositor length: 8.5–8.8 (8.6).

Colour forms. Females of Y. denisoni are typically black; however a brown form has also been collected in the Canberra region within the Australian Capital Territory. The black form is closely similar to the male, with bright red basal membranes ( Fig. 1E,F ). It is found throughout the distribution of the species. The brown form is characterized by the predominantly brown colour of the head and thorax ( Fig. 1G,H ; similar to the brown form of Y. timothyi sp. nov., illustrated in Fig. 7C,D ). In this form the fore wing basal membranes are red to pink. Only three specimens of the brown form have been collected, two from O’Connor and one from Mt Majura in the Australian Capital Territory. Notably, females of the more typical black form from have been collected from Black Mountain, which is adjacent to O’Connor. A single male specimen that may be a brown form has also been collected from Kioloa, New South Wales (in ANIC). It has some brown to dark brown colouration on the thorax, but is otherwise typical of Y. denisoni , with bright red fore wing basal membranes.

Distinguishing features. Specimens of Yoyetta denisoni can be distinguished morphologically from other species of Yoyetta , apart from Y. timothyi sp. nov. and Y. kershawi stat. rev., comb. nov., by their size (forewing length> 27 mm) and the abdominal tergites, which are uniformly black without orange or yellow markings or contrastingly posterior margins. They can be distinguished from Y. timothyi sp. nov. by examining the basal membranes of the fore wings, which are bright red to pink (cf. orange to pale orange-white in Y. timothyi ). Males can be distinguished from Y. kershawi stat. rev., comb. nov. by the presentation of the terminal aedeagus ( Fig. 10A ), which is strongly bifurcate and splayed into two lateral arms (cf. simple or weakly bifurcate, without lateral arms in Y. kershawi stat. rev., comb. nov.; Fig. 10C ). Females of Y. denisoni are indistinguishable from Y. kershawi stat. rev., comb. nov., although the latter species is slightly separated geographically, being restricted to Victoria and adjacent areas in far southeastern New South Wales.

Calling song ( Figs 6 , 53A ). Yoyetta denisoni males call when in flight or stationary; however, the call is predominantly made in flight. The call can be described as a rapid chirping song. Each “chirp” is a syllable or short macrosyllable that inflects noticeably in pitch and also to some degree in amplitude within its brief duration. Field recordings are often difficult to obtain as this species mainly calls in flight. Several recordings have been made in Canberra , Royal National Park , Berowra Waters National Park , near Jindabyne and near Glen Innes , all in New South Wales. Based on the recordings (n = 9), the (macro)syllable typically ranges from 0.011 –0.020 s duration followed by a silent gap of 0.127 –0.172 s duration ( Figs 6 A–C). Field recordings have a frequency plateau of 3.8–8.2 kHz and a dominant frequency from 4.5–6.7 kHz ( Fig. 6D ) .

Of the species with known calling songs, the call of Y. denisoni is closest to Y. timothyi sp. nov. and Y. electrica sp. nov., which each also produce calls in flight, though at a slower rate and faster rate, respectively. Further differences are given under the calling song descriptions of those species.

Figures

Figure 1. Yoyetta denisoni (Distant) (A), holotype male, Port Denison [locality considered to be erroneous], dorsal habitus; (B) holotype male, ventral habitus; (C), holotype male, lateral aspect; (D) male, O’Connor (35°16'S 145°06'E), dorsal habitus; (E), female, Bowral, NSW, 34°27'S 150°25'E, dorsal habitus; (F), female, ventral habitus; (G), female, brown form, dorsal habitus, O’Connor; (H), female, brown form, ventral habitus. Scale bars = 10 mm.

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 5. Map of southeastern mainland Australia showing the geographical distribution of Yoyetta denisoni (Distant) (open circles), Y. timothyi sp. nov. (open triangles), Y. kershawi (Goding & Froggatt) (closed squares).

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 7. Yoyetta timothyi sp. nov., brown forms: (A) male holotype, Waitara (33°42'S 151°06'E), dorsal habitus; (B) male holotype, Waitara, ventral habitus; (C) female, Terrey Hills (33°40'S 151°12'E), dorsal habitus; (D), female paratype, Terrey Hills, ventral habitus; (E) male, Crescent Head (31°17'S 152°57'E), live specimen (photo by NJE); (F) female, Medway (34°29'S 150°16'E), live specimen (Photo by NJE) Scale bars = 10 mm.

Figure 10. Photostack comparisons of aedeagus of (A) Yoyetta denisoni (Distant), holotype (photo by J. Constant); (B) Y. timothyi sp. nov., Gordon (33°45'S 151°08'E); (C) Y. kershawi (Goding & Froggatt), Toolangi (37°32'S 145°28'E). Scale bars as indicated.

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.