Yoyetta verrens, 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 : 332-338

Yoyetta verrens sp. nov.

http://zoobank.org/NomenclaturalActs/ 025D81F6-5094-41D4-B0F9-35BCADE515DC

Figs 2F , 3F , 41–47 , 53F

Holotype ♂, Blatherarm Camp, Torrington, 29.2523°S 151.7072°E, 4–7.i.2016, L. Popple, at light, 515-0001 ( AM K.549270) . Paratypes NEW SOUTH WALES: 1♀, same data as holotype, 515-0005 ( AM K.549271) ; 1♂, same data as holotype, 515-0003 (DE) ; 2♂♂, same data as holotype, 515-0002 [genitalia prep.], 515-0004 ; 1♂, Butler Mine track, Torrington , 29.2367°S 151.6466°E, 5.i.2016, L.W.Popple, 515-0006, SL100704 ( LWP) .

Additional locations with audio recordings. QUEENSLAND: Daves Creek, Lamington National Park, 28°13'21"S 153°12'35"E, 12.xii.2017; Canyon Parade, Springbrook National Park , 28°13'12"S 153°16'22"E . NEW SOUTH WALES: L.W. Popple; Basket Swamp National Park , NSW, 28°54'34"S 152°09'12"E, 21.xi.2001, L. W. Popple .

Distribution, habitat and seasonality. Restricted to moderately high altitude areas (above c. 600 m) the Queensland / New South Wales border region from the Macpherson Range southwest to Torrington State Conservation Area ( Fig. 38 ). Adults are active in the middle to upper canopy of eucalypt

forest and are rarely observed at close quarters. They are present from November to January.

Description

Male ( Figs 2F , 3F , 42 A–B, 43–47). Head almost as wide as mesonotum, black, with a central dark brown fascia posterior to ocelli; ocelli pink; dorsal side of postclypeus black anteriorly, brown over remainder; ventral side of postclypeus black with black transverse grooves, lateral and posterior borders brown; anteclypeus black, rostrum black at base, dark brown centrally, black at apex, reaching anterior edge of hind coxae; lora black; gena black; eyes dull black; antennae black, supra-antennal plates black, brown at base of pedicel.

Thorax mainly black with variable brown patterning. Pronotum black, a thin central brown line on anterior half, dark brown on lateral angles; fissures mainly black; pronotal collar black. Mesonotum black, posterior margin dark brown centrally, becoming black laterally, cruciform elevation arms black, lateral depressions brown. Metanotum black.

Emery et al.: Yoyetta denisoni-abdominalis cluster 333 Legs. Coxae mainly black; coxal membranes red; basisterna black, katepisterna brown posteriorly; meracantha small, narrow, cream, black at base, pointed, minimally overlapping opercula; fore femora orange anteriorly, black medially, brown strip laterally, black posteriorly and around base of femoral spines; femoral spines erect, brown at base, becoming black at tips; mid and hind femora black, orangebrown medially, distal femoral joint orange; fore- and midtibiae black laterally, orange-brown over remainder; hindtibiae orange brown; tarsi orange-brown becoming black towards claws; claws dark brown, black at tips.

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

Opercula ( Fig. 3F ) medium, spatulate, following body axis ventrolaterally, depressed centrally, black over basal half, stippled brown-cream across central areas, posterior margins orange yellow, clearly separated.

Timbals ( Fig. 2F ) 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; 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 1–2 black; tergites 3–7 black with orange posterior margins, extending anteriorly to cover lateral margins; tergite 8 shiny black, lateral face dark brown posteriorly, lateral margins orange. Epipleurites with medial anterior strip, orange over reminder. Sternite II mainly black; sternite III black, orange posterior margin; sternites IV–VII orange; sternite VIII dark brown anteriorly, becoming dark orange posteriorly, with brownish pubescence.

Genitalia ( Fig. 43 ). Pygofer black; dorsal beak with ochraceous tip, anal styles orange; upper lobe black, ochraceous and rounded posteriorly; basal lobe black. Uncus dark brown; lateral process beak-like, hooked ventrally; lobes in ventral view bulbous, with tapering lateral termination; claspers black, apposed at base, clearly divided anteriorly, short, with apices rounded, gradually tapering laterally. Aedeagus with pseudoparameres extending around two-thirds the length of theca; theca recurved ventrally at 120° towards apex, with spectacular transparent flange along the outer margin of recurvature, this broadly smooth along theca with prominent lateral ornamentations, together>5 times width of theca, terminating at apex; apex short, knoblike, with 4–6 distinct cornuti around base.

Female ( Fig. 42 C–D). Similar to male, but with overall lighter colouration.

Head mainly brown with black markings surrounding ocelli and extending to supra-antennal plate; a black spot on anterior side of each compound eye; postclypeus ochraceous on dorsal side, black on ventral side with central ochraceous spot anteriorly, also ochraceous on extreme margins; anteclypeus and rostrum black; gena black; mandibular plates mainly ochraceous, black posteriorly.

Thorax. Pronotum mainly brown with an ochraceous midline; black markings along fissures and surrounding midline; pronotal collar brown, with extreme edges of lateral angles dark brown; mesonotum brown with black submedian and lateral sigilla, and a diffuse black central marking extending anteriorly from cruciform elevation; cruciform elevation black on central, medial portion and lateral margins, yellow-ochraceous on posterior margins and

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

theca. Characters as depicted in Fig. 4 . Specimen from Torrington (25°15'S 151°44'E).

lateral depressions; scutal depressions black; wing grooves ochraceous with central black spot; metanotum lurid brown.

Wings match description given for male, with venation mainly a paler brown.

Legs. Coxae mainly brown; coxal membranes orange; basisterna dark brown to black, katepisterna mainly brown; meracantha as in male; fore femora black anteriorly with an orange stripe, mainly brown posteriorly and around base of femoral spines; femoral spines erect, brown at base, becoming black at tips; mid and hind femora dark brown to black anteriorly, brown posteriorly; distal femoral joint orange; fore- and mid-tibiae dark brown to black anteriorly, orange-brown over remainder; hind-tibiae pale orangebrown; tarsi orange-brown becoming black towards claws; claws dark brown, black at tips.

Abdomen. Tergite 1 black, tergite 2 mainly black with reddishbrown posterior margins on dorsolateral sides; tergites 3–7 black with reddish-brown posterior margins, grading to orange-brown and broadening ventrally; tergite 8 mainly orange-brown, black along anterior margin; epipleurites mainly orange-brown, with diffuse black interiorly; sternite II black with orange black posterior margins ventrolaterally; sternites III–VI orange-brown with narrow black midline; sternite VII orange-brown with anterior black spots on either side of midline; abdominal segment 9 orange-brown with three longitudinal black stripes, one on midline and two broader examples either side of midline; dorsal beak black; ovipositor sheath extending approximately 2 mm beyond apex of abdominal segment 9; anal styles orange.

Measurements (in mm; range with mean in parentheses: 5 males, 1 female).

Body length: male 22.1–24.7 (23.3); female 26.1. Fore wing length: male 27.8–29.6 (28.6); female 31.8. Head width: male verrens sp nov. These include (A) two phrases from Torrington (25°15'S 151°44'E);

(B) one phrase expanded (from A above); (C) two phrases from Blatherarm (29°15'S

151°42'E); (D) one phrase expanded (from C above). Recordings were made in the

field by LWP using RS3 (see Methods and terminology).

6.4–6.6 (6.5); female 7.1. Pronotum width: male 6.4–7.0 (6.6); female 7.2. Abdomen width: male 5.7–7.1 (6.5); female 6.4; ovipositor length (female): 9.7.

Etymology. The Latin name “ verrens ” means “sweeping”, which represents the nature of the calling song of this species in flight.

Distinguishing features. Males of Y. verrens sp. nov. have contrasting orange bands on the posterior margins of tergites 3–7, a feature that readily distinguishes them from Y. denisoni , Y. kershawi stat. rev., comb. nov. and Y. timothyi sp. nov. They can be readily distinguished from Y. abdominalis , Y. aaede , Y. spectabilis sp. nov., Y. electrica sp. nov. and Y. serrata sp. nov. by (1) their larger size with body length> 22 mm, and (2) lack of symmetrical, orange triangular patterns on dorsolateral sides of tergites 4–7. Males of Y. verrens sp. nov. differ from those of Y. grandis sp. nov. and Y. subalpina by the colour of the hind wing plaga, which is dull white and restricted along the margins of anal cell 3 and vein 2A in Y. verrens sp. nov., whereas in Y. grandis sp. nov. and Y. subalpina this colouration is brighter and extends over the entire jugum. In addition, the orange-red coloration on the opercula of Y. verrens sp. nov. contrasts with the grey coloration on Y. grandis sp. nov. and the bright orange colour of the fore wing basal membranes on Y. verrens sp. nov. are dull orange to grey in Y. grandis sp. nov. Females can be distinguished from all of the aforementioned species by their characteristically long ovipositor sheath, which extends> 2 mm beyond the apex of abdominal segment 9.

Calling song ( Figs 44–47 , 53F ). Males of Yoyetta verrens sp. nov. call whilst stationary and also in flight. When stationary there are two different modes of song production, one which we term the “ratchet” mode ( Figs 44 , 47 ) and another which is described as the “cascade mode” ( Fig. 44–46 ). In flight, a simple series of echemes is produced, each ending with an acute amplitude modulation as illustrated in Fig. 46 , which gives the song a “sweeping” quality.

Several recordings have been obtained from the type locality and nearby vicinity in Torrington State Conservation area ( Figs 44–47 ). Based upon these recordings (n=11), the ratchet mode comprises monotonously repeated phrases, composed of a syllable sequence (7–55 single syllables, doublets, or rarely triplets, each 0.01– 0.03 s duration, separated by gaps of 0.073 –0.142 s) and an echeme (0.1– 0.428 s duration), followed by a gap (0.09– 0.18 s duration) ( Figs 44 , 47 ). The echemes at the end of each phrase show an acute amplitude modulation, which is most obvious in longer duration examples (> 0.3 s). These longer examples are often produced in the transition from the ratchet mode to the cascade mode .

The cascade mode commences with a series of three or four long echemes, increasing successively in duration (each 0.47 s to 1.94 s duration), and each separated by gaps of 0.146 – 0.302 s. In a similar structure to the call produced in flight, each of the long echemes ends with an acute amplitude modulation, although in this case there are differences in the duration of the different echemes, with the last two echemes being longer than the preceding one(s). This is then often followed by a soft, short echeme (0.12– 0.18 s duration), then a gap of 0.05– 0.34 s. A syllable sequence (as described for the ratchet mode, but often longer and with gaps between echemes being 0.1– 0.3 s duration) then completes the phrase. A breakdown of this song mode is provided in Figs 45 , 46 .

Few recordings of this species have been obtained outside of the type locality. A single recording of the cascade mode from Basket Swamp (approximately 55 km north-east of the type locality) were found to fall broadly within the range of calling song variation present at the type locality in terms of temporal structure ( Fig. 46 ). The only localities from which recordings have been taken is at Daves Creek in Lamington National Park and nearby at Canyon Parade, Springbrook National Park. These recordings (two of a male in flight and another of the cascade song mode) were found to be broadly similar in temporal structure to the type locality; however they are too faint for the purposes of illustration.

There were found to be no apparent modulations in song frequency between the two modes of the calling song. In the vicinity of the type locality, the male calling song has a highest amplitude frequency plateau that typically falls approximately between 8.5 and 13.0 kHz, with dominant frequencies of 10.7–11.1 kHz ( Fig. 44 ). Recordings from north-east of the type locality were found to have a higher overall frequency. To the north, at Basket Swamp, the dominant frequency of the only available recording is 11.7 kHz, which is only slightly higher. Further afield at Daves Creek, the frequencies ranged even higher (plateau of 9.4–14 kHz, with dominant frequency between 11 and 12.4 kHz.

No recordings of male-female duets have been obtained for this species. However, observation in the field based on successful simulation of female wing-flick responses suggest that the females respond during the gaps that follow each long echeme that ends with an acute amplitude modulation. These echemes are produced: (1) as the ratchet mode transitions toward the cascade mode, (2) during the cascade mode itself, and (3) in flight. The males appeared to be responsive to simulated wing-flicks during each of these calling song modes/situations.

The calling song of Yoyetta verrens sp. nov. is unlike others in the Y. abdominalis species group. It shares some features in common with Y. subalpina sp. nov.; however, it can be distinguished easily by its more simplistic sweeping echemes (cf. two-note phrases in Y. subalpina 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 38. Yoyetta aaede (Walker) (A) male, Mt Lofty (34°59'S 138°43'E), dorsal habitus; (B) male, Mt Lofty, ventral habitus; (C), holotype female, Adelaide, dorsal habitus; (D), holotype female, ventral habitus (holotype photos by Ken Merrifield); (E), male, live specimen Adelaide (34°56'S 138°39'E), photo by NJE; (F), female, live specimen, Adelaide, photo by NJE). Scale bars = 10 mm.

Figure 41. Map of southeastern Australia showing the geographical distribution of Yoyetta aaede (Walker) (open circles), Y. verrens sp. nov. (open triangles) and Y. electrica sp. nov. (closed squares).

Figure 42. Yoyetta verrens sp. nov., (A) male, Torrington (29°15'S 151°42'E), dorsal habitus; (B) male, Torrington, ventral habitus; (C) female, Torrington, dorsal habitus; (D), female, Torrington, ventral habitus. Scale bar = 10 mm.

Figure 43. Yoyetta verrens sp. nov., illustration of male pygofer and internal genitalia;

Figure 44. Typical male calling song structure of Yoyetta verrens sp. nov., illustrated in waveform plots and a spectrogram from recordings in the field at Torrington (29°15'S151°42'E) by LWP using RS3 (see Methods and Terminology).This includes (A) a broad plot commencing with the “cascade” mode and progressing into the “ratchet” mode; (B) a segment (from A above) showing the beginning of the ratchet mode; (C) another expanded segment (from A) illustrating the structure and marked amplitude modulation of a single echeme from the cascade mode; (D) a spectrogram displaying song frequency.

Figure 45. Comparisons ofthe “cascade” mode of thecalling song ofYoyetta verrens sp. nov., illustrated in wave plots from three localities, including (i) Torrington, (29°15'S 151°42'E); (ii) Blatheram (29°15'S 151°42'E); (iii) Basket Swamp (28°55'S 152°09'E). Each was recorded by LWP using RS3 (see Methods and terminology).

Figure 46. Wave plots illustrating the “cascade” mode of the calling song of Yoyetta verrens sp nov., produced in flight. The illustration (A) includes a simple series of 3 echemes, each ending with an acute amplitude modulation, giving the song a sweeping quality.The lower plot (B) is expanded from the one above to illustrate the detailed structure of a single echeme. Recorded in the field at Torrington (29°15'S 151°42'E; 515-184 Torrington2) byLWP using RS3 (see Methodsand Terminology).

Figure 47. Examples from the “ratchet” mode from the calling song of Yoyetta

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.