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FIGURE 12. Heremusina udeoecetes n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 12. Heremusina udeoecetes n. sp. Waveform plots, based on field recordings at 11 km N.W. along Tanami Desert Road, 20 km north of Alice Springs, in low grassland. Figs. A, D show the characteristic repeated echemes, these having durations ranging between 272–332 ms, each commencing with 2–9 isolated pulses, followed by a sequence of closely spaced pulses; an isolated pulse is emitted at end of each echeme. Figs. B, D are time expanded waveform plots showing more detail of the pulses. The initial pulses (Fig. B) are isolated pulses, these changing to doublet pulses within the continuous echeme segment. Pulses (including doublet pulses) have repetition rates varying between 160–193 Hz. (C), shows the final segment of an echeme in which the pulses alternate between single and double pulses, followed by the single isolated final pulse. Filtered to 3.5 kHz.
FIGURE 15 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 15. Comparison of Heremusina udeoecetes n. sp. song in terms of, (A), waveform plot (temporal structures of song), (B), spectrogram showing timbal emission frequencies, these concentrated in range ~11–17 kHz, but exhibiting no indication of systematic changes of frequencies during song emission, although minor frequency disturbances may occur at the start of each echeme; and (C), amplitude spectrum showing the dominant frequency envelope between 11.5–16.8 kHz, with the estimated dominant frequency at 13.9 kHz. Filtered to 3.5 kHz (A), and 7.5 kHz (C).
FIGURE 11 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 11. Pulse repetition rates (Hz) versus durations (ms) between the doublet primary pulses in the songs of Mugadina marshalli and M. emma. These data are based on multiple recordings, from multiple locations, at multiple dates, including field and container recordings (recordings by AE and LWP). The PRR of the M. marshalli songs are consistently higher than the M.emma songs, with minimal overlap of data. The inset shows only PRR data from recordings made 2 km west of Macalister, southeastern Queensland, where both species were found to occur together. These recordings are container recordings made under identical conditions. They exhibit a marked separation of PRR between the two species at this overlap location.
FIGURE 10 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 10. Waveform (A, B, D) and complementary envelope curves (C, E) of the ticking song of Mugadina emma, all recordings made near the Fairbairn Dam Rd, ~7 km S.W. of Emerald, eastern central Queensland, recorded 'in situ' in an open net in grassland. Fig. 10A presents an overview segment showing the multiple repeated 'ticks', the PRR varying between 2.5 to 4.6 Hz. As in the M. marshalli song, each 'tick' comprises a set of two pulses, with each initial 'primary' pulse followed by a weaker secondary pulse, the intervals between the two pulses being 1.4–3.4 ms, slightly variable, although similar to the M. marshalli data. The detailed pulse structures within the primary and secondary pulses are also similar to those of M. marshalli, and the two sets of pulses are similarly interpreted as time offsets in the firing of the two timbals. Unfiltered.
FIGURE 13. Heremusina udeoecetes n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 13. Heremusina udeoecetes n. sp. Time expanded details of selected pulses shown in Fig. 12, shown as waveform and companion envelope plots. In 13A, B, the initial single pulse is markedly of longer duration compared to each of the following double pulses, suggesting that it results from a partial overlapping of asynchronous pulses from the two timbals; the double clicks are therefore interpreted as temporally more clearly separated pulses emitted by the two timbals. Figs 13C, D are strongly developed doublet pulses, emitted asynchronously by the two timbals. The following smaller peaks which are almost coalesced with the main peak are believed to represent the emission from timbal ribs 2–5. The peaks labeled 'S' are interpreted to be secondary ('OUT') clicks of the timbals, not seen following first peak most likely as they overlap the emission of the second primary peak of each doublet. Fig. 13E, F, time expanded single pulses emitted at start of echeme, these showing merged weaker peaks (probably emissions of ribs 2–5). Secondary peaks, marked 'S', occurring 1.7 ms after the primary peaks, are inferred to represent be 'OUT' clicks of the timbals. Filtered to 3.5 kHz.
FIGURE 29 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 29. Comparison of Xeropsalta festiva n. sp. song in terms of: (A), waveform plot, illustrating the main temporal structures; (B), spectrogram showing the dominant frequency band between 5 and 10 kHz, with no clear or systematic change during the song production; (C), amplitude spectra showing a well defined dominant frequency envelope between 5.1 and 8 kHz, with inferred dominant frequency of 6.6 kHz.
FIGURE 9 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 9. Mugadina marshalli song. Waveform (A, B, D) and complimentary envelope (C, E) plots, of the ticking song of M. marshalli, recorded in open net, in grassland near Kaimkillenbun, southeast Queensland (A to C) and Yelarbon, S. Queensland, in situ with parabola in grassland (D, E). Fig. 9A shows an overview plot of the repeated pulses ('ticks'), the pulse repetition rates (PRR) varying between 4.9–5.4 Hz. Figs. 9B, C, are time expanded illustrating details of an individual pulse, showing that when viewed in detail, each actually comprises two dominant (primary) pulses, each followed by a weaker, but clearly separate pulse, the time gaps being 1.9 and 2.8 ms. These are interpreted as secondary pulses, produced by timbal 'OUT' (relaxation) movement. The two initial strong pulses exhibit weaker and almost coalesced pulses, which is suggested to result from the 'firing' of long ribs 2 to 5 (implying that the dominant pulses are produced by the 'IN' movements of timbal 1). The presence of two complete sets of pulses that comprise each 'tick' are inferred to result from a very short but significant delay in the contraction (firing) between the two timbals. Figs. 9D, E, represent time expanded plots from another M. marshalli from a separate location, showing the similar arrangement and structures of the pulses, although the gaps between the primary and secondary pulses are significantly longer. Filtered to 1 kHz.
FIGURE 8 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 8. Map of Central and Northeastern Australia, showing the currently known occurrences of Heremusina udeoecetes n. sp., H. pipatio n. sp., Xeropsalta thomsoni n. sp., X. aridula n. sp., and X. rattrayi n. sp. The heavy dot-dash line shows the approximate northern and eastern boundaries of the ecoregion which defines the dominance of deserts and xeric shrublands within inland Australia (based on National Reserve System map, published by the Australian Government). The fine dotted lines are major roads, the names of some prominent towns listed. Lake Eyre and associated ephemoral lakes are shown by the darker shading. The approximate outlines of the modern dunefields comprising much of the Simpson and Strzelecki deserts are shown by the lighter shading, these based on 'Google Earth' images.
FIGURE 7 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 7. Pygofer and male genitalia, ventral views, respectively, of the five new species described in this paper; (A), (B), H. udeoecetes n. sp. and H. pipatio n. sp.; (C) to (E), X. thomsoni n. sp. (see also Pl. 4), X. aridula n. sp., X. rattrayi n. sp. Sizes as in Fig. 6.
FIGURE 28. Xeropsalta festiva n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 28. Xeropsalta festiva n. sp., field recording (L.W.Popple), Hilston, N.S.W. Fig. 28A, overview waveform plot of multiple echemes with the intervening, isolated, high amplitude double pulses. (B), time expanded view of an echeme showing the abrupt initial start, followed by a short gentle decrease, and then progressively increasing amplitude extending to the abrupt termination. The strongly developed pulse doublets emitted before and after the echeme are clearly shown. (C), detailed time expanded plot of selected segment of echeme (see (B) for location) showing three macrosyllables, each, with 6 pulses. Macrosyllable RR range between 132–139 Hz. (D), detailed time expanded plot of a double pulse showing the pulse structures. Each of the two dominant pulses are themselves doublets with a separation of 1 ms, and each is evidently followed by a much weaker secondary pulses (labeled as 'S' in plot D, timbal relaxation 'OUT' pulses), emitted between 2.1–2.4 ms after the primary pulses. The two sets of doublet pulses are interpreted as the results of small, but asymmetries of the firing of two timbals. Filtered 2 kHz.
FIGURE 23. Xeropsalta aridula n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 23. Xeropsalta aridula n. sp. Time expanded waveforms and complimentary envelope plots of two selected segments of echemes. Figs. 23 A, B, recording near start of an echeme, showing the details of four macrosyllables, these each characterised by high amplitude pulses emitted at the end of each macrosyllable. Although each macrosyllable has six higher amplitude main pulses, the pulses show a marked tendency to occur in sets of 4 or 3 pulses. The initial smaller pulse of each macrosyllable is followed closely by a gap (0.8–0.9 ms) and then a set of 4 pulses. These pulse patterns are inferred to result from a slight offset (asynchrony) of pulse emissions between the two timbals, resulting in partial pulse overlaps and the grouping of pulses into 4 or 3 sets, the start of each macrosyllable is marked by 3 partially merged pulses. These 3-pulse sets are inferred to result from the complete overlap of a second peak with the following first peak, resulting in the higher amplitude central peaks. Figs. 23C, D are cage recordings, in which at the end of echemes, the normal 2-timbal song is modified by the cessation of one timbal, causing a drop of the RR of the pulse doublets by half. In these 1 timbal segments, the doublet pulses show the initial high amplitude pulse and following low amplitude peak (0.8–0.9 ms offset). One interpretation is therefore that the initial strong peak results from the timbal rib 1, the weaker peak from the flexing of ribs 2 to 5. The alternate explanation is that the second weaker peaks are the secondary relaxation 'OUT' pulses. Filtered A, B, to 1 kHz. C, D unfiltered.
FIGURE 6 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 6. Pygofer and male genitalia, lateral views, respectively, of the five new species described in this paper; (A), (B), Heremusina udeoecetes n. sp. and H. pipatio n. sp.; (C) to (E), Xeropsalta thomsoni n. sp. (see also Pl. 4), X. aridula n. sp., X. rattrayi n. sp. Pygofer lengths are, respectively, 1.60, 1.70, 1.86, 1.91, and 1.9 mm.
FIGURE 4 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 4. Left timbal, posterior margin to right, dorsal margin at top, of the five new species described in this paper; (A), (B), Heremusina udeoecetes n. sp. and H. pipatio n. sp.; (C) to (E), Xeropsalta thomsoni n. sp., X. aridula n. sp., X. rattrayi n. sp., respectively. For clarity, only selected long ribs and one short (intercalary) rib are labelled, as are the timbal plate and apodeme. 5 long ribs are present, only numbers 1 and 5 labelled.
FIGURE 3 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 3. Male opercula, all left opercula, of the five new species described in this paper; (A), (B), Heremusina udeoecetes n. sp. and H. pipatio n. sp.; (C) to (E), Xeropsalta thomsoni n. sp., X. aridula n. sp., X. rattrayi n. sp., respectively. Opercula nomenclature shown in (E).
FIGURE 2 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 2. Lateral abdomen views of the five new species described in this paper; (A), (B), Heremusina udeoecetes n. sp. and H. pipatio n. sp.; (C) to (E), Xeropsalta thomsoni n. sp., X. aridula n. sp., X. rattrayi n. sp., respectively.
FIGURE 26. Xeropsalta rattrayi n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 26. Xeropsalta rattrayi n. sp. Waveform and companion envelope plots (A, B) of a time expanded segment from the extended echeme shown in Fig. 25B (which shows the precise location within the echeme), and Figs. 25C, D, the time expanded termination of a short echeme (shown in detail in Fig. 25D). Figs, 26A, B illustrate three macrosyllables, each with 6 sets of pulse doublets. The strongest amplitude pulses mark the start of each macrosyllable, these, however, grouped into alternating sets of 3 to 4 pulses (see especially Fig. 26B, the numbers above the pulses indicating the number of pulses in each set). The sets of pulses between the high amplitude sets at the start and end of each macrosyllable also show a tendency for alternation between 4 and 3 pulses, although the pulse structures are somewhat blurred and merged. Fig. 26C, D, waveform and envelope plots of the 'inflated' termination of the short echeme as shown in Fig 25 D, with higher amplitude sets of pulses tending to occur at start of each macrosyllable. The pulses between the start and end of each macrosyllables also exhibit an apparent alternation between 4 and 3 pulses. More detailed examination, however, reveals that the '3-pulse' sets do possess weak doublets, the middle pulses containing these doublets clearly of slightly longer duration (0.6–0.7 ms) than the other pulses (0.3–0.5 ms), indicating merging of pulses. The apparent complexity of the pulse structure is, as also seen in X. aridula songs, attributed to the partial and/or complete overlaps between the double pulses. This is seen in the relative lengths of the pulse sets, those with 4 pulse sets of slightly longer durations than the 'apparent' 3-pulse sets. Mean macrosyllable RR's are 117–132 Hz. Unfiltered.
FIGURE 5 in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 5. Right fore and hind wings of the five new species described in this paper; (A), (B), Heremusina. udeoecetes n. sp. and H. pipatio n. sp.; (C) to (E), Xeropsalta thomsoni n. sp., X. aridula n. sp., X. rattrayi n. sp., respectively.
FIGURE 19. Xeropsalta thomsoni n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 19. Xeropsalta thomsoni n. sp. Song details as in Figs. 18A to D. Fig. 19A, time expanded details of the macrosyllable structures within a single echeme, the initial five macrosyllables each comprised two nearly merged syllables. The final two macrosyllables are each comprised of three nearly merged syllables. Fig 19 B, C, waveform and envelope plots, time expanded, of the final macrosyllable shown in Fig. 19A, showing the three syllables, each with 6 clearly defined pulses, the mean duration of the individual pulses 0.37 ms, and PRR ranging between 1069–1085 Hz. Fig.19D, E, waveform and envelope plots of segment of extended macrosyllable (see Fig. 18C for location), emitted prior to the break-up of the song into discrete echemes. This shows a short segment of continuous song, in which four macrosyllables are discernable, each macrosyllable finishing with a high amplitude pulse, each macrosyllable with three clearly defined sets of double pulses, the strongest followed by weaker secondary pulses (timbal relaxation 'OUT' pulses). The arrangement of the pulses, including the secondary pulses, indicates an apparent groupings with 2, 4, and 3 pulses, shown in the plot. Filtered 3 kHz.
FIGURE 22. Xeropsalta aridula n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 22. Xeropsalta aridula n. sp., field recordings (with parabola), Cravens Peak area, eastern Simpson Desert, far western Queensland, low grassland (following heavy rains), mostly disturbed vegetation, locally dominated by introduced button grass (Dactyloctenium radulans). (A) waveform overview of a complete phrase showing the repetitive discrete echemes, these ranging between 236–375 ms in duration, with RR of 2.6 Hz. The initial echemes are longer than the following echemes. (B) Another waveform plot showing echemes which are fragmented into shorter phrases, each comprised varying numbers of echemes, commonly observed in the X. aridula n. sp. songs. (C) time expanded waveform of initial echeme shown in Fig. 22B, illustrating the increasing pulse amplitudes, reaching a maximum at ~400 ms, followed by a slight amplitude decline, the termination marked by a sharp break, followed by a discrete macrosyllable. Fig. 22D, E, further time expanded waveform and envelope curve of the end of an echeme (excluding the final discrete macrosyllable), showing the detailed pulse structures, with the development of well defined macrosyllables, each comprising 6 primary pulses, each immediately followed by (in fact partially merged with) slightly weaker second pulses. These weaker pulses could either represent the slight time offset (asynchrony) of the emission of pulses from the two timbals, this thought to be most likely, or are secondary pulses produced by timbal relaxation ('OUT' clicks) of timbals, this thought to be less applicable due to the relatively strong development of these second pulses (see Fig. 23C, D below).
FIGURE 14. Heremusina udeoecetes n in Two new genera and five new species of Mugadina - like small grass cicadas (Hemiptera: Cicadidae: Cicadettini) from Central and Eastern Australia: comparative morphology, songs, behaviour and distributions
FIGURE 14. Heremusina udeoecetes n. sp., waveform (A, B, D) and envelope (C, E) plots from parabola recordings made 3 km east of Hermannsburg, N.T., ~112 km WSW of Alice Springs; 14A shows an echeme segment with two initial isolated pulses, followed by rapidly repeated doublet pulses, the PRR measured at 192 Hz. Figs. 14B, C, are time expanded plots of the pulse doublets showing both the peaks to be strongly developed, with a weak secondary peak 1.5 ms following the second peak, evidently not so clearly developed after the first peak (the doublet inferred to be emitted asynchronously from both timbals); Fig. 14D, E illustrates single pulses occurring at start of echeme showing only weakly developed secondary peaks (marked 'S'). These plots also shows small but increasingly stronger peaks (arrowed) almost joined to the preceding primary peaks, these thought to represent the initial development of doublet peaks (i.e. increasing asynchrony of timbal emissions). Filtered to 7.5 kHz (A), and 9 kHz (B to E).
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