— Holotype male, China, Yunnan, Honghe, Pingbian, Daweishan Forest Reserve, 22°56.55'N, 103°41.43'E, 1700–2100 m a.s.l., 15–17 viii 2013, leg. Liu Chunxiang, #CH7660, in ethanol. Depository IZCAS.

Paratypes 11 males (#CH7659, 61-62), 6 females (#CH7663-5), dried/in ethanol, same data as holotype, all in IZCAS except #CH7659, 61, 64 (in CH). In addition sound recordings of two males, not collected.

— Differs from all species of the genus in proportions of anal (1/3) and basal (2/3) part of the stridulatory file and number (about 100; Table 1) and distribution of teeth on the stridulatory file (Figs 2, 3). From E. apicalis, which has a similar stridulatory file, it differs by larger size and the hind wings being clearly longer than the tegmina while slightly shorter in E. apicalis. In general habitus it is most similar to E. fulva, but has a much lower number of teeth on stridulatory file (see Fig. 3) and narrower tegmina (Fig. 4).

— Male. General habitus see Fig. 5.

Fastigium verticis narrower than first segment of antennae, sulcate dorsally, not contacting fastigium frontis. Complex eyes approximately round. Pronotum with disc nearly flat, only last quarter elevated; anterior margin slightly concave, posterior margin truncate with small notch in middle; medial carina inconspicuous; transverse sulcus V-shaped; lateral carina weakly developed; lateral lobe of pronotum much longer than high, anterior margin straight, posterior margin obtuse-rounded, ventral margin oblique downward posteriorly, humeral sinus present but inconspicuous.

Prothoracic spiracle large, but mostly covered by a posterior extension of paranotum.

Tegmen shorter than hind wing, posteriorly with long parallel part (Figs 4A, 5A); venation see Fig. 4. Mirror in stridulatory area of right tegmen with a more or less circular deepening, similar to that in E. fulva (Fig. 6). The vein demarcating the distal end of the stridulatory area curved smoothly, not with an edge as in E. fulva (Fig. 6). Stridulatory file comparatively straight but slightly thinner in middle, with circa 90 densely spaced fine teeth in basal area and about 10 widely spaced large teeth in distal (=apical or anal) area (Fig. 3). Fore coxae without spine, fore tibiae dorsally sulcate and spinuliferous. All femora ventrally spinulose; mid and hind femur with 5-6 spines on outer ventral margins; hind tibiae with many spines on both dorsal margins. Tenth abdominal tergite prolonged backwards, hind margin emarginate; cerci stoutest at base, basal quarter sharply acuminated, then first slightly widening, later gradually narrowed till apex, dorso-ventrally flattened; subgenital plate split for almost apical two thirds, up-curved (Figs 7, 8). Internal genitalia membranous, no sclerotized titillator.

— Body and tegmina green, parts of legs, dorsal side of pronotum and dorsal field of tegmina brown, tegmina with some small brown dots (Figs 4, 5).

— Mostly similar to male except abdominal apex and tegmina (stridulatory organs).

Pronotom with disc nearly flat; ventral and dorsal edge of paranotum parallel.

Right tegmen at posterior edge in basal part with about 9 short cross veins carrying stout spines used to produce the female acoustic response (similar as shown in Heller et al. 1997); more distally small cross veins and even longitudinal veins with many slender spines, probably used to prevent accidental wing opening. In left tegmen, posterior edge in basal part scraper-like in-curved. Supra-anal plate triangular; cerci rather short, conical. Ovipositor fairly broad and short, upcurved, both margins at tip denticulate (Fig. 9). Subgenital plate triangular, apex obtusely rounded.

— Green except a white mid-line on pronotum and dorsal edges of tegmina (Fig. 5B).

>— Thin, flattened, ovoid, typical phaneropterine shape.

— Body: male 14–20, female 17–22; pronotum: male 3.7–4.8, female 3.9–4.2; tegmen: male 26.7–31, female 22–25.3; tegmen width (greatest): male 5.1–6.5, female 4.9–5.1; hind wing: male 33–38; female 26–29.4; post femur: male 20.9–24, female 20.5–23.4; length of ovipositor: 6.5–8.1; width of ovipositor 1.9–2.1, length of egg 4.7–4.8; width of egg 1.6–1.8 (n=12 males, 6 females, partly dried, partly in ethanol).

— Time-amplitude-pattern. In Ectadia diuturna sp. n. one song unit typically lasted about 30 to 60 s (all data for 25°C except otherwise mentioned; up to 80 s in the field at 18°C). It was followed by the next unit after a silent interval as short as some seconds, but also as long as some minutes. The shortest intervals were observed when males and females were in acoustical contact. Each unit contained four phrases each of which consisted of a different combination of micro- and macrosyllables. A simple and clear example is shown in Fig. 10A. However, often the different phrases were connected by transitional phrases. A song unit started with a series of tick-like microsyllables, separated by intervals of about 150 ms (phrase A; see Fig. 11 for details). Sometimes the intervals between the ticks were larger and more irregular. Between these loud syllables (occasionally?) short and soft sounds were observed. At the end of a phrase typically two tick sounds followed each other quite closely (10 ms interval) a few times. Such phrases were also heard isolated outside complete song units, possibly representing unsuccessful attempts to start a song unit. After this phrase a series of macrosyllables followed (phrase B), in which always two types of impulse groups alternated, a long series (ca. 70 impulses; duration 120 ms), sometimes split into two or more by one or several unusually large intervals, and a short series (ca. 55 densely packed impulses; duration 30-40 ms). Both groups differed distinctly in carrier frequency (see below). In amplitude, this phrase was always lower than the loud phrase A. Its duration was the most variable of all phrases, from nearly missing up to ten seconds. Phrase C consisted of a long series of stereotypic elements (period about 500 ms), each containing two pairs of impulse groups as seen in phrase B. These pairs were separated from the next pair by an interval of about 200 ms. During this interval one (or two) loud microsyllables were produced and additionally a series of softer isolated impulses. While in Fig. 10 an abrupt change from phrase B to C can be seen, in many other recordings there is a long transition with phrase C elements appearing more and more frequently (see e.g. Fig. 12). Phrase D finally consisted of several repetitions of the impulse series from phrase C (period about 400 ms) without macrosyllables.

The females responded near the end of a song unit (Fig. 12). They answered mostly not only once, but several to many times (1–25 responses per song unit). The responses occurred nearly always during phrase D, and often also additionally at the end of phrase C. Very rarely they were heard during phrase A, in this case the females possibly assuming a late phrase D from the previous song unit. In many phaneropterine species, the females respond with a species-specific delay after the male song or some parts of it (e.g. Heller and Helversen 1986). This is probably also the case in E. diuturna sp. n., but the response delay is difficult to measure exactly, because often several possible trigger events are closely together, and the response consists of several loud impulses distributed over more than 100 ms. The delay is obvious in the range of 150–250 ms. In any case, often very soft impulses (more than 20 dB softer than the main response) were observed that occurred much faster (70–80 ms) than the loud impulses.

— During one song unit, parts with quite different spectral composition were observed. The short, tick-like elements of which phase A and D consisted and which occurred also in phase C, had always a quite similar spectrum. Its peak was situated at about 20 kHz, in the lower half with strong components starting abruptly at about 5 kHz, in the higher half continuously decreasing in power until 40 kHz (Fig. 13). Also the softer and shorter impulse groups found in phase B and C were similar to this pattern presenting a relatively narrow peak at 20 kHz and a side-peak at 40 kHz. However, the longer and often louder impulse groups from phase B and C differed distinctly. Their spectrum contained two clearly separated peaks. Besides a high and narrow peak at 10 kHz it showed a broad maximum at 60 kHz (Fig. 13). These impulse groups alternated with the shorter and softer impulse groups (see fig. 5 in Heller et al. 2015). The female response had its maximum between 10 and 20 kHz (recorded in audible range only).

— The analyzed species is characterized by a male chromosome number 2n=29 and sex determination system X0. All chromosomes are acrocentric and the X chromosome is the largest element in the set. C-banding of mitotic metaphase revealed constitutive heterochromatin blocks (thin C-bands) in the paracentromeric region in most chromosomal elements, with the exception of one small pair of autosomes (thick C-bands). Constitutive heterochromatin polymorphism involving telomeric C-bands was located in the first-sized pair (Fig. 14A). The observed rDNA-FISH signal on this largest pair, similar to C-bands, varied in size, suggesting the occurrence of polymorphism in copy number of rDNA sequences (Fig. 14B).

— Common in low bushes around buildings (Fig. 5C).

— The name of the new species refers to its long lasting song: diuturna. Latin adjective, meaning long lasting

— Known only from Yunnan, China.

Figure 2. 

Inter-tooth spacing in male stridulatory files of Ectadia species.

Figure 3. 

Stridulatory file of E. diuturna sp. n. and E. fulva (articulation of tegmen to the right). A. File of E. diuturna sp. n. on the underside of left male tegmen. Scale 1 mm. B–C. Replica of the file of B. E. diuturna sp. n. in comparison to that of C. E. fulva.

Figure 4. 

Transmitted light scan of left tegmina. A. Ectadia diuturna sp. n., B. E. fulva (the fold in the basal part is an artifact resulting from the strong curvature of the tegmen). Scale 10 mm.

Figure 5. 

Habitus and habitat of E. diuturna sp. n. A. male (photo by Guoqing Ma), B. female, C. habitat at type locality. The photo of the male is shown under E. fulva in Kang et al. (2014).

Figure 6. 

Stridulatory area with mirror at base of right tegmen. A. E. diuturna sp. n., B. E. fulva. Scale 1 mm.

Figure 9. 

Lateral view of female ovipositor. A. E. diuturna sp. n., B. E. fulva. Scale 1 mm.

Figure 7. 

Lateral view of male subgenital plate. A. E. diuturna sp. n., B. E. fulva. Scale 1 mm.

Figure 8. 

Ventral view of male subgenital plate. A. E. diuturna sp. n., B. E. fulva. Scale 1 mm.

Figure 10. 

Oscillograms of male calling songs, comparative overview. A.E diuturna sp. n., B.E fulva, In A, the different phases of one song unit are indicated.

Figure 11. 

Oscillograms of male calling song of E. diuturna sp. n. A. Overview with indication of the origin of the details (B–E). In B, D and E. the origin of the spectra shown in Figure 12 is indicated.

Figure 12. 

Oscillograms of male calling song and female response of E diuturna sp. n., recorded simultaneously. A, B. Overviews with indication of the origin of the details (C–D).

Figure 13. 

Power spectra of different parts of the calling song of E. diuturna sp. n.

Figure 14. 

Male chromosomes of Ectadia diuturna sp. n. A. C-banded metaphase; B. FISH with 18S rDNA (green) probe in metaphase/anaphase (A – the left) and prometaphase (B – the right). Arrows indicate telomeric located C-bands (A) and cluster of 18S rDNA (B) in the heteromorphic large-sized chromosomes (marked with “1”); X, sex chromosome.

— For the behavioral studies four males and three females (out of series of nine each) were used, all offspring from animals from Thailand, Chiang Mai, Doi Suthep-Pui, 18°48'N, 98°55'E, 1100–1150 m a.s.l., 13.04.1995, ex ovo, bred in lab., leg. S. Ingrisch. Stridulatory files in six males, China, Yunnan Province, Jinping County, Fenshuiling National Nature Reserve, 22.88178768°E, 103.23377388°N, August 2012.

— The species has been sufficiently redescribed by Liu et al. 2004. It may be added that the fore wings of the male are vaulted, bulging laterally in proximal half of tegmen length, flat with rounded tip thereafter; those of the female are shortened with acute tip. In both sexes the hind wings surpass the fore wings. The species shows a green-brown color polymorphism in both sexes. Of the specimens collected in the field, two males and two females were green and two males and one female brown. The brown color is of lighter shade in females (more ochreous) than in males (medium brown with dark pattern). In the offspring of a green female bred in lab, 19 males and 12 females became green, four males and nine females brown. Body, legs, fore wings and the projecting part of the hind wings were either all green or all brown, in two females pale yellowish brown. The broad dorsal field of the male tegmen including the stridulatory area was always dark brown in both color morphs while the narrow dorsal field of the female tegmen was yellow in green females or pale brown bordered by a dark brown line in brown females. In resting position, E. fulva sits with the antennae and the fore legs stretched anteriorly in the body axis, the mid and the hind legs are stretched oblique posteriorly, while the abdomen and the wings are pointing dorso-posteriorly. That behavior resembles the situation in Ducetia (e.g. D. japonica) or Elimaea species.

— Time-amplitude-pattern. The male calling song consisted of song units each lasting about 1.5 s repeated after an interval of about 4 s (Fig. 16A). Each was made of a series of microsyllables (about 15), repeated with 20 Hz, followed by a long series of impulses with decreasing intervals. This song unit is produced by opening the tegmina very widely, then closing and opening them several times only in part, resulting in the microsyllables. The microsyllables are not tick-like sounds as in E. diuturna sp. n., but consist of compact series each with a few impulses. Then the male closed the tegmina completely and very slowly. The contact of the scraper with the fine basal part of the file produced a long series of heavily damped impulses (Fig. 16; see also fig. 7 in Heller et al. 2014).

The female responded immediately after the end of the male impulse series (Fig. 16), typically with one or a few impulses, occasionally with one or a small series up to a few hundred ms later in addition. Sometimes responses were observed even before the male series had ended.

— Both parts of the song unit had a quite similar spectral composition with two peaks (Fig. 17). Besides a narrow low-frequent peak around 10 kHz there was a broad maximum at about 50-60 kHz. The female response showed a peak at 20 kHz, in width and placement intermediate between the two male peaks.

Besides the acoustical signals soundless vibratory body movements were observed in both sexes.

— In eight tests, a male (mean body mass 199 ± 16 mg; n=7) and a female (mean body mass 626 ± 71 mg; n = 6) were placed together for mating. Four couples mated with the males transferring only very small spermatophores (1.3 ± 0.6 mg; no data on mating duration available; Fig. 18). Obviously the spermatophores consisted only from a pair of ampullas without spermatophylax.

— Postembryonic development occured over six nymphal instars as in many other Phaneropterinae (Fig. 19). Development from hatching to adult moult took 52–68 days at 20–23°C (Ingrisch 1998). Nymphs can be green or light brown. First instars appear green colored.

— See Fig. 1.

Figure 15. 

Habitus of Ectadia fulva. A. male, B. female.

Figure 16. 

Male calling song A. and female response B. of Ectadia fulva. Oscillograms of stridulatory movement and song [synchronous registration of left tegmen movement and sound (upper line: upward deflection represents opening, downward closing; lower line: sound)]. Male and female were recorded separately.

Figure 17. 

Power spectra of different parts of the male calling song and of the female response of E. fulva.

Figure 18. 

Female Ectadia fulva with spermatophore. A. overview, B. detail.

Figure 19. 

Nymphs of Ectadia fulva. A. First stage, B. second stage, C. third stage, D. 4^th stage, E. 5^th stage, male, F. 5^th stage, female, G. 6^th stage, female, H, I. 6^th stage, male.