Bioacoustics of Cardiodactylus muria

High speed video observations at 1250 frames per second show that each calling song corresponds to only one FW closure (i.e. one syllable), the gaps within the syllables are made by the jerking closing movement of the FWs. At 20°C, these mono-syllabic calling songs are emitted in irregular groups of 7–12 (m = 10.6 ± 2.2, n = 5) that lasts for 77 ± 13 s with a period of 208 +/- 50 s. Each syllable shows an indented temporal profile due to the jerking FW closures, with the following characteristics: syllable duration = 48 ± 2 ms; syllable period (within the echemes / syllable groups) = 8.0 ± 2.3 s; syllable duty cycle = 0.65 %. The dominant frequency is 13.28 ± 0.22 kHz and corresponds to the third frequency peak of the song, with a clear harmonic series.

Description. Average size for the species group. General coloration brown to dark brown with contrasted vivid yellow areas ( Fig. 42A–D ). Head dorsum with a large brownish area comprising short dark brown lines posterior to eyes and 4 more or less distinct dark brown bands, faint posteriorly, the lateral bands complete, the median ones punctuated ( Fig. 6C ). Fastigium dark brown to black. Scapes orange brown with a transverse whitish ring. Antennae brown. Face most often brown with a thin yellow transverse band ventral to eyes. Mouthparts mostly yellow brown. Lateral sides of head gray brown. Maxillary palpi yellow brown. Pronotum: Dorsal disk anterior and lateral areas yellow with variable dark brown symmetrical patterns and black spots; posterior area dark brown to black. Lateral lobes dark orange brown, ventral margins yellow brown. Legs I–II yellow brown to orange brown. FIII yellow brown to orange brown; knees dark brown; tibiae faintly ringed with dark brown at spur level; tarsomeres III-1 yellow brown with both ends dark brown. Hind wing tail dark brown to black, 1.5 longer than pronotum. Cerci brown with faint dark brown rings. Abdomen mostly dark brown. Male: FW coloration mostly dark brown to black with yellow areas including bases of anal veins (4A completely yellow), angle of 1A, first third of CuA, harp veins, chords, veins within a crescent-shaped area posterior to mirror. Translucent areas include harp anterior region, chord area and area posterior to mirror. M/R area dark brown, R/Sc area orange brown. Sc orange brown, its posterior part yellow. Projections of Sc and more ventral veins of lateral field yellow, cells between them dark brown. FW venation ( Fig. 10C ): 1A slightly bisinuated; CuP missing but claval fold present in harp. Stridulatory file with about 200 teeth on the transverse and curved parts of 1A without discontinuity, and 15 more teeth on top of a bump near 1A base ( Table 11 , n = 4). Harp with 2 w-shaped veins. Mirror area: mirror (d1) longer than wide, oval, crossed near mid-length by a transverse vein, the posterior part variably subdivided by an accessory veins; d2 as wide as d1, variably subdivided by accessory veins; e1 crossed by several accessory veins. Apical field with 4–5 cell alignments posterior to mirror (n = 4). Lateral field with 7–9 (m = 8, n = 5) projections of Sc and 4–5 (m = 4.2, n = 5) ventral veins. Subgenital plate yellow brown. Male genitalia ( Fig. 42E–G ): Region of pseudepiphallus posterior to mid-length narrowing almost rectangular. Dorsal ridges divergent posteriorly, their anterior part with dorsal asymmetrical expansions folded laterally, the left one larger and more anterior, the right one smaller and more posterior; pseudepiphallus slightly bisinuated laterally. Pseudepiphallic sclerite with flat and wide latero-anterior expansions, membrane between them with thick dense setae. Posterior pseudepiphallic apex rounded, with a small rounded translucent area. Rami parallel, with large preapical plates, apical stems slightly convergent. Pseudepiphallic parameres median lobe rectangular. Ectophallic arc complete, narrow and curved, without posterior expansion. Bases of ectophallic apodemes with a posterior membranous lobe but with no clear anterior sclerotization. Ectophallic fold with weak lateral sclerites; apex trilobate, membranous, median lobe oval. Endophallic sclerite well sclerotized, with long posterior arms and a triangular posterior expansion. Endophallic apodeme with wide lateral lamellas and a strong medio-dorsal crest. Membrane of endophallic cavity smooth. Female: FW coloration orange brown to dark brown, including veins and cells ( Fig. 11H ). Base of FWs whitish, including bases of anal veins. Whitish lateral band on FW anterior part, including anterior half of CuA and M and associated transverse veins, their posterior half orange brown. R/Sc area orange brown to dark brown, veins orange brown. Projections of Sc and more ventral veins yellow brown, the cells between them dark brown. FW venation ( Fig. 11H ): 10–11 (m = 10.2, n = 5) strong longitudinal veins on dorsal field; lateral field with 7–9 (m = 8.2, n = 5) longitudinal veins including 4–5 projections of Sc and 3–4 ventral veins. Ovipositor long, apex with both dorsal and ventral edges denticulate ( Fig. 13M ). Female genitalia ( Fig. 15C ): Copulatory papilla trapezoidal with baso-lateral sclerotizations; apex membranous, folded ventrally and related to a medio-ventral crest; apex with a posterior sclerotized area near base. Juvenile: Light brown mottled with dark brown. Measurements. See Table 11 . PronL PronW FWL FWW HWT FIIIL FIIIW TIIIL Male holotype 3 4.8 15.3 4.5 2.1 16.1 4.1 14.1 Males (n = 5) 3–3.4 4.8–5.3 15.2–17.9 4.3–5.1 2.1–5.2 16–19 4.1–5 14.1–17 (Male mean) (3.2) (5) (16.3) (4.8) (4.3) (17) (4.4) (15.7) Female allotype 3.5 5 17.4 4.6 5.5 17 4.6 16.3 Females (n = 5) 3.5–3.7 5–5.6 16–18 4.1–4.9 4.1–5.5 17–18.7 4.5–4.9 15.1–16.7 (Female mean) (3.6) (5.4) (17) (4.5) (4.5) (17.7) (4.7) (16) TIIIs TaIIIs ST (n = 4) OL Ias Ibs Oas Obs Tt Lt Bt Male holotype 7 4 18 5 3 ? ? ? - Males (n = 5) 7–10 4–6 12–18 5–8 2–4 172–189 12–22 9–24 - (Male mean) (8) (6) (14) (8) (3) (180) (19) (15) - Female allotype 9 6 12 10 5 - - - 17.2 Females (n = 5) 7–9 5–6 9–13 8–10 3–5 - - - 17.2–19.1 (Female mean) (8) (6) (14) (9) (4) - - - (18.4) Habitat and life history traits. Cardiodactylus muria is a nocturnal species living in or near secondary forested areas. Males produce calling songs at night, from tree branches and leaves ( Fig. 43 ). Juveniles are found day and night on low vegetation. Behavior. Calling song ( Fig. 44 ): High speed video observations at 1250 frames per second show that each calling song corresponds to only one FW closure (i.e. one syllable), the gaps within the syllables are made by the jerking closing movement of the FWs. At 20°C, these mono-syllabic calling songs are emitted in irregular groups of 7–12 (m = 10.6 ± 2.2, n = 5) that lasts for 77 ± 13 s with a period of 208 +/- 50 s. Each syllable shows an indented temporal profile due to the jerking FW closures, with the following characteristics: syllable duration = 48 ± 2 ms; syllable period (within the echemes / syllable groups) = 8.0 ± 2.3 s ; syllable duty cycle = 0.65 %. The dominant frequency is 13.28 ± 0.22 kHz and corresponds to the third frequency peak of the song, with a clear harmonic series.