Four New Species of Cicadas in the Yoyetta abdominalis (Distant) Species Group (Hemiptera: Cicadidae: Cicadettinae) from Southeastern Australia Author Popple, Lindsay W. Biodiversity & Geosciences, Queensland Museum, South Brisbane QLD 4101, Australia Author Emery, David L. Sydney School of Veterinary Science (SSVS), University of Sydney NSW 2006, Australia text Records of the Australian Museum 2020 2020-08-12 72 4 123 147 journal article 10.3853/j.2201-4349.72.2020.1765 2201-4349 4654453 B4BECFF5-5972-4F1B-819A-380798AE9082 Yoyetta loftyensis sp. nov. urn:lsid:zoobank.org:act: DFC5114D-A217-4281-9BD1-76A43C10D063 Figs 1 , 4C , 5C , 6C , 12–15 Holotype ♂ , Australia , SA , AU. SA .VHN, 9.8 km NW. of Victor Harbor Rd on Myponga Rd , 229 m , 35°25.733'S 138°34.818'E , 30.xi.2006 , Marshall & Hill , SAMA Database No. 20-017507 ( SAM ) . Paratypes — SOUTH AUSTRALIA : 1♂ , Australia , SA , AU. SA . BIR , 2.3 km S. Birdwood on road to Torrens , 391 m , 34°50.459'S 138°57.687'E , 29.xi.2006 , D. Marshall , K. Hill (C. Simon lab voucher specimen 06.AU. SA . BIR .01) ; 1♀ , same location and date as previous ( MSM ) ; 1♀ , same data as holotype ; 1♂ , S. of Balhannah on Jones Rd. , 1.2 km S Junction Rd. , 351 m , 35°00.244'S 138°49.500'E , 8.i.2011 , Marshall & Hill (C. Simon lab voucher specimen 11.AU. SA . ADE .01) (DE) ; 1♂ , Collected by G. Rayson , location Stack Jarra Stakes f[ro]m W.A. to Belair RR, 23-11-77, National Parks & Wildlife Service , SAMA Database No. 20-014465 ; 1♂ , Mt. Barker S.A. , 4. Dec. [19]64, N. McFarland , SAMA Database No. 20-014439 ; 1♂ , S. Aust , Mt. Lofty , Jny. [18]85, A. White , SAMA Database No. 20-014440 ; 1♂ , Sth Aus , Reed Beds , Male , 9-Dec-70, W White , SAMA Database No. 20-017497 ; 1♂ 1♀ , Morphett Vale , Dec. 1889 , J. W. Mellor ; 1♂ , 22-11-1958 , Belair , S.A., S. Dawson SAMA Database No. 20-017501, 20-017499 ; 2♀♀ , Wilmington , Rev. Burgess , 16.12.86, SAMA Database No. 20-014442, 20-014443 ; 1♂ 1♀ , S. Australia , Rev. A. P. Burgess , SAMA Database No. 20-014444, 20-014445 ; 2♀♀ , Mt Lofty Rge , S. H. Curnow ; 1♀ , S. Aust , Mt Lofty , Dec-1891 , SAMA Database No. 20-017500, 20-017502 ( SAM ) . Figure 7 . Yoyetta douglasi sp. nov. : illustration of male genitalia; (a) pygofer, viewed laterally from the left; (b) pygofer, viewed ventrally; (c) aedeagus, viewed laterally from the left; (d) apex of theca, viewed laterally from the left and (e) apex of theca, viewed dorsally. 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 Halls Gap (37°14'S 142°32'E). Figure 8 . Yoyetta enigmatica sp. nov. : (A) male, Lamington National Park (28°13'S 153°13'E), dorsal habitus; (B) male, Lamington National Park, ventral habitus; (C) typical habitat, Lamington National Park; (D) typical habitat at Mt Kaputar (30°17'S 150°10'E). Scale bars = 10 mm. Etymology . Named after the Mt Lofty Ranges south-east of Adelaide, South Australia , where this species is most prolific. Distribution, habitat and seasonality . This species is restricted to the Greater Adelaide region from near Victor Harbour north through the Mt Lofty Ranges to near Williamstown, with a potentially isolated population in the southern Flinders Ranges at Wilmington ( Fig. 1 ). It has also been heard calling in Adelaide Botanic Garden. Populations are found in open eucalypt forest and occasionally in parks with tall eucalypts ( Fig. 12E, F ). The adults occupy the upper and outer branches where they tend to remain fairly sedentary. Males move singing stations irregularly. Adults have been found during November and December. Figure 9. Male calling song structure of Yoyetta enigmatica sp. nov. illustrated in waveform plots, including: (A) typical repeated echemes (in flight); (B) expanded section from (A) showing the detailed structure of a single echeme. The final subfigure (C) is a spectrogram displaying song frequency. This specimen was recorded in the field at Dawson Springs, Mt Kaputar (30°17'S 150°10'E) by LWP. Calling song . The calling song of Y. loftyensis sp. nov. is moderately high pitched, with a frequency plateau between approximately 10 and 15 kHz and dominant frequency of approximately 12 kHz. It contains repeated patterns of syllables (each 0.008 –0.015 s duration) that are grouped in phrases of two subtly different structures: a “normal phrase” and a “cueing phrase”. In both cases, each phrase commences with two (rarely one or three) preceding syllables, separated by a gap of 0.04– 0.09 s , followed by a shorter gap (0.01– 0.07 s ), then an echeme (which could equivocally be interpreted as a dense syllable sequence; 0.26– 0.71 s duration) and then two (rarely three) subsequent syllables, separated by successively longer gaps (0.03– 0.08 s , increasing to 0.07– 0.13 s ). The cueing phrase is characterized by the longer gap after the final syllable (0.09– 0.22 s ). In contrast, the normal phrases lack this longer gap, which makes the beginning and ending of each phrase less conspicuous ( Fig. 13 ). Occasionally, a one-off extended echeme (or dense syllable sequence; 3–19 s duration) is produced within the structure of a normal phrase or cueing phrase ( Fig. 13A ). The two different phrase types may be produced interchangeably and there is little difference in overall duration between normal and cueing phrases (each lasting approximately 0.5– 1.1 s ). A comparison of cueing phrase structures recorded from across the geographical distribution of this species is provided in Fig. 14 . Figure 10 . Waveform plots comparing different recordings of the calling song of Yoyetta enigmatica sp. nov. from: (A) Mt Kaputar (30°17'S 150°10'E); (B) Mt Kaputar; (C) Lamington National Park (28°13'S 153°13'E); (D) Springbrook National Park (28°13'S 153°16'E). All recordings were made in the field by LWP. Singing occurs almost exclusively when males are stationary. However, a phrase with an extended echeme may also be produced during short flights between singing stations. It is inferred that the female wing-flick response would be timed during these long gaps at the end of each cueing phrase. Morphology . Male ( Figs 4C , 5C , 6C , 12 A–B, 15). Head almost as wide as mesonotum, black; a central triangular yellow-brown fascia on midline posterior to ocelli, base on posterior margin, reducing anteriorly, ocelli pink to pale; dorsal postclypeus black, with a narrow yellow-brown fascia on midline, apex directly anteriorly, ventral surface black, ochraceous spot on anterior midline, with black transverse grooves, lateral margins brown-ochraceous; anteclypeus black, rostrum pale brown at base, black around apical third, brown centrally becoming dark brown to black at apex, reaching anterior of hind coxae; lorum black, gena black; eyes dull black; antennae black, supra-antennal plates black. Thorax mainly dull black. Pronotum black, with a central yellow-brown fascia extending from behind anterior margin almost to margin of pronotal collar; patchy dark brown markings over raised lateral areas of pronotum; paramedial fissures black, lateral fissures variably dark brown; pronotal collar dark brown to black, central third variably brown, margins of lateral angles brown, lateral areas black. Mesonotum black, parapsidal sutures black, scutal depressions and surrounds black; cruciform elevation arms black, lateral depressions light brown. Metanotum black, midline black, pale brown over central third, becoming black laterally. Figure 11. Yoyetta enigmatica sp. nov. : illustration of male genitalia; (a) pygofer, viewed laterally from the left; (b) pygofer, viewed ventrally; (c) aedeagus viewed laterally from the left; (d) apical portion of theca, viewed laterally from the left; (e) terminal apex of theca; (f) apical portion of theca, viewed ventrally. Characters depicted as in Fig. 7. Specimen from Dawson Springs, Mt Kaputar (30°17'S 150°10'E). Figure 12. Yoyetta loftyensis sp. nov. : (A) holotype male, 9.8 km NW. of Victor Harbour Road (35°26'S 138°35'E), dorsal habitus; (B) holotype male, ventral habitus; (C) female, 2.3 km S. of Birdwood (34°50'S 138°58'E) dorsal habitus; (D) female, 2.3 km S. of Birdwood, ventral habitus; (E) typical habitat, Adelaide; (F) typical habitat, Adelaide. Scale bars = 10 mm. Legs. Coxae black, joints orange-red; basisterna black, bordered orange-red; trochanter black with diffuse brown areas; meracantha small, narrow, pale cream, black at base, pointed, marginally overlapping opercula; fore femora mainly orange-red, with black longitudinal markings on inner ventral sides and around base of femoral spines; femoral spines erect, black; mid femora black mainly orange red, with diffuse black areas on anterior sides, hind femora orange-brown with black markings on anterior apices, joints orange-red; fore tibia mainly black, dark reddish-brown on inner sides; mid tibia black, tending dark reddish-brown towards anterior apices; hind tibia dark reddish-brown with diffuse black areas; fore and mid tarsi dark reddish brown, tending black on outer sides; hind tarsi pale orange-brown, becoming dark brown towards claws; spines orange-brown; claws dark brown, black at tips. Wings with fore wing costal veins black anteriorly, orangebrown central rib, becoming brown distally, pterostigma mottled red, basal cell pale yellow, translucent, black anterior border, arculus black, basal membranes orange, other veins dark brown to black, proximal segment of vein CuA pale, becoming brown distally with eight apical cells; hind wing plaga white to whitish-cream over jugum and vein 3A, thin along vein 2A, subcostal vein pale, other veins brown, with six apical cells. Opercula ( Fig. 4C ) covering abdominal cavity, spatulate, following body axis ventrolaterally, depressed centrally, variably black at base, pale cream across remainder, tinged orange in some specimens, clearly separated. Timbals ( Fig. 4D ) 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 two-thirds of timbal; apodeme pit oval shaped, inconspicuous. Abdomen with tergite 1 black, membrane anterior to timbals orange-red, tergite 2 black, tergites 3–7 black with orange posterior margins, extending laterally to epipleurites, progressively increasing on either side of midline to reach anterior borders of tergites 4–7, posterior margins yellow; tergite 8 black, with red markings on lateral margins ( Fig. 6C ). Epipleurites 1–3 black anteriorly, orange red posteriorly, epipleurites 4–6 mainly orange. Sternite I black, sternite II black medially, orange-red over central third posterior to timbal cavity, black laterally, sternite III orange-red with central black spots on anterior and posterior margins, sternites IV–VI orange, sternite VII mainly orange, red posteriorly, sternite VIII fiery red, with pale brown pubescence. Figure 13. Male calling song structure of Yoyetta loftyensis sp. nov. illustrated in waveform plots, including: (A) a single cueing phrase commencing with an extended dense syllable sequence/long echeme, followed by three normal phrases; (B) a series of four normal phrases, followed by a cueing phrase, then two sets of alternating normal and cueing phrases; (C) expanded section from (B) showing two normal phrases (demarcated by solid black bar above) and one cueing phrase (demarcated by dashed line above). The final subfigure (D) is a spectrogram displaying song frequency. This specimen was recorded in the field at 9.8 km NW. of Victor Harbour Road (35°26'S 138°35'E) by D. Marshall and K. Hill. Figure 14. Male calling song structure of Yoyetta loftyensis sp. nov. A comparison of phrase structures recorded from multiple sites, including: (A) Mt Lofty (34°59'S 138°43'E); (B) Hale Conservation Park (34°41'S 138°55'E); (C) 9.8 km NW. of Victor Harbour Road (35°26'S 138°35'E); and (D) 2.3 km S. of Birdwood (34°50'S 138°58'E). All examples are cueing phrases, apart from the one in (B) with a black scale bar above, which is a normal phrase. Recordings (A) and (B) were made in the field by LWP. Recordings (C) and (D) were made by D. Marshall and K. Hill. Genitalia ( Fig. 15 ). Pygofer shiny black upper lobe mainly black with red posterior margin; basal lobe ochraceous; dorsal beak black, dark brown posterior margin, anal styles orange, red at tip. Uncus orange-brown; in lateral view beak-like and stumpy; lobes in ventral view bulbous, with rounded lateral termination; claspers clearly divided, triangular with apices gradually tapering laterally. Aedeagus with pseudoparameres extending around one third the length of theca; theca recurved ventrally at 180° towards apex, with transparent flange along outer margin of recurvature, broadly smooth, around 2× width of theca, terminating adjacent to apex of theca; apex short, tapered, sclerotized, angled 30° ventrally, with two lateral rows of prominent cornuti at base, numerous cornuti over apex. Female ( Fig. 12 C–D). Head and thorax similar to male, with darker brown markings along parapsidal sutures. Abdomen . Tergites similar to male. Sternite I–II black, posterior margins ochraceous, sternites III–VI orangered with midline black marking decreasing to sternite IV, sternite VII pale brown with anterior black spot either side of midline. Abdominal segment 9 with central black triangular marking, base at anterior margin, reducing over middle third, expanding over dorsal beak, flanked laterally by orange stripe and lateral black area, extending laterally over anterior third, remainder reddish-orange laterally with posterior black spot. Dorsal beak black, ovipositor dark brown, becoming black at tip, not extending beyond apex of abdominal segment 9. Anal styles orange-red; ovipositor sheath black, becoming paler brown ventrally, gonocoxites black laterally, ochraceous along midline. Figure 15. Yoyetta loftyensis sp. nov. : illustration of male genitalia; (a) pygofer, viewed laterally from the left; (b) pygofer, viewed ventrally; (c) aedeagus, viewed laterally from the left; (d) apex of theca. Characters depicted as in Fig. 7. Specimen from 9.8 km NW. of Victor Harbour Road (35°26'S 138°35'E). Measurements (in mm; 10 males , 8 females ). Body length: male 20.9–24.7 (22.7); female 22.3–24.0 (28.5). Fore wing length: male 25.5–28.7 (26.9); female 28.3–31.2 (29.4). Fore wing width: male 9.2–10.6 (9.8); female 9.7–11.3 (10.5). Head width: male 6.0–7.0 (6.6); female 6.6–7.1 (6.9). Pronotum width: male 5.9–7.7 (6.9); female 6.4–8.0 (7.5). Abdomen width: male 6.1–7.2 (6.5); female 6.4–7.5 (7.0). Ovipositor length: female 6.0–8.4 (7.7). Distinguishing features . Within the Y. abdominalis species group, the calling song of Y. loftyensis is most similar to the call of Y. serrata Emery, Emery & Popple from south-eastern Australia . The longer minimum gaps between syllables (> 0.16 s ; Emery et al. 2019 ) distinguishes Y. serrata from Y. loftyensis . The former also lacks the characteristic structure of the cueing phrases produced by Y. loftyensis (see Figs 13 , 14 ). Morphologically, Y. loftyensis sp. nov. is closest to Y. abdominalis (Distant) , which also occurs in south-eastern South Australia , and Y. serrata . It can be distinguished easily from Y. serrata by the colour of the fore wing basal membranes, which are orange (cf. pale whitish-grey). Males can be distinguished from Y. abdominalis by examining the apex of the theca, which is tapered and angled 30° ventrally (cf. short and club-like). Both sexes can be distinguished from fresh specimens of by Y. abdominalis by the lack of conspicuous golden hairs on the head and thorax. Otherwise, females cannot be distinguished reliably from Y. abdominalis .