Xenogryllus marmoratus ( Haan, 1844 )

Jaiswara, Ranjana, Dong, Jiajia, Ma, Libin, Yin, Haisheng & Robillard, Tony, 2019, Taxonomic revision of the genus Xenogryllus Bolívar, 1890 (Orthoptera, Gryllidae, Eneopterinae, Xenogryllini), Zootaxa 4545 (3), pp. 301-338 : 324-328

Xenogryllus marmoratus ( Haan, 1844)

( Figs 1 D–G, 3I–J, 4E, 5E, 7H–I, 8G, 9B, 11CD, 12B, 14)

Gryllus (Phalangopsis) marmoratus Haan, 1844: 235 .

Calyptotrypus marmoratus— Saussure 1878: 714 (the figures do not correspond to the species X. marmoratus , which is consistent with the surprising distribution “Java, Iles de la Sonde” proposed by Saussure, where the species is not distributed); Bolívar 1900 [1899]: 805.

Madasumma marmorata —Chopard 1924: 56: Kirby 1906: 93.

Xenogryllus marmoratus — Chopard 1968: 350; Yin & Liu 1995: 96; Oshiro 1995: 43; Walker 2010: 27; Robillard & Desutter- Grandcolas 2004a: 578 (calling song); 2004b: 273 (morphological phylogeny); 2006: 644 (molecular phylogeny); 2008: 67; 2011: 637; Anso et al. 2016: 9 (molecular phylogeny); Chintauan-Marquier et al. 2016: 62 (molecular phylogeny); Vicente et al. 2017: 2203 (historical biogeography); Schneider et al. 2017 (tympanum morphology) Jing et al. 2018: 274.

Xenogryllus marmoratus marmoratus Gorochov, 1992 , in Gorochov & Belokobylskij 1992: 11 (nominal subspecies); Ichikawa et al. 2000: 276; 2006: 180; Storozhenko & Paik 2007; Storozhenko et al. 2015: 145; Cigliano et al. 2018 ( Orthoptera Species File Online); Olivero & Robillard 2017: 1 (mating behavior).

Xenogryllus marmoratus unipartitus ( Karny, 1915) — Gorochov & Belokobylskij 1992: 10 (subspecies).

Synonym name:

Heterotrypus unipartitus Karny, 1915 — Chopard, 1968: 350.

Common names: Pine cricket (English), matsumushi (Japanese = pine insect, waiting insect); bao ta ling (Chinese = pagoda bell).

Type material. Neotype, ♂ [new designation], Japan: Honshu, collection Finot, identified Calyptotrypus species nova by A. Finot (MNHN-EO-ENSIF1592).

Additional material examined. Japan: Honshu , Mie, ix.1957, F. Ohmachi, 1♂, 1♀, identified Xenogryllus marmoratus de Haan, Xma Robillard morpho (MNHN). Japon, 1♀, Exposition Universelle 1869, #1276-69,

identified Dionymus marmoratus by L. Chopard (MNHN-EO-ENSIF1593). Japon [Japan, no precise locality and date], 1♂, 1♀, identified Dionymus marmoratus by L. Chopard, Xma Robillard morpho; 1♂, 2♀ (MNHN); 1♀ (NHMW); 1♂, 20.679, identified X. marmoratus unipartitus by Karny (NHMW); 4♀, identified Calyptotrypus species nova, collection Finot (MNHN). Kobe, H. Fruhstorfer, 1♀, 24.125 (NHMW). Kyoto, I. Yamashiro, 1♂, Xma Robillard morpho MEB, MEB Ziegler; 1♀; 1♀, molecular sample T. Robillard 2004 (MNHN). Kioto [Kyoto], Y. Hirase, 1♂, identified Madasumma marmorata (Haan) and Calyptotrypus marmoratus (Haan) by Hebard, 1924 (MNHN). Kanagawa Prefecture, Kanate, Ooi-Cho Ashigara-kami-gun [35° 23' 00"N, 139° 08' 00"E], H. Sakai, 14.viii.2010, 1♂, 2♀, 1 juvenile, identified Xenogryllus marmoratus by A. Ichikawa (MNHN); 1♂, enregistrement appel TR-male2 [call recording—MNHN-SO-2016-14364], identified Xenogryllus marmoratus by A. Ichikawa (MNHN-EO-ENSIF1704); 1♂, enregistrement appel TR-male1 [call recording, MNHN-SO-2016-14365], identified Xenogryllus marmoratus by A. Ichikawa (MNHN-EO-ENSIF1598). Japon, environs de Tokyo, J. Harmand, 1906 (MNHN). Kinki Distr.[ict], Wakayama Pref.[ecture], Hashimoto city, 19.x.1986, A. Ichikawa, 1♂, identified X. marmoratus by A. Ichikawa (RBINS). Tsushima, H. Fruhstorfer, septoct [ix–x], 24.123 (NHMW). China: Guanxi, Jin Xiu, 10.x.1981, E110°11’ N24°07’, 1♀ (14062635) (SIPPE). Longzhou, 1995-viii18 /23, Xianwei Liu, Weinian Zhang, Xinbao Jin, 1♀ (14062635) (SIPPE). Chongqing, Beibei, E106°23’ N29°48’, 1.x.2000, Zhou, 1♂ (14080765) (SIPPE). Anhui, Huangshan, E118°19’ N29°43’ [Shanghai market], viii.2012, T. Robillard, 1♂ (TR40), enregistrement appel [call recording MNHN-SO- 2018-36] (MNHN-EO-ENSIF1702); 1♂ (TR2), enregistrement appel [call recording MNHN-SO- 2018-52], molecular sample X12 (MNHN-EO-ENSIF1594). Jiangsu, Zhenjiang, 32°12'0.00"N 119°27'0.00"E [Shanghai market], viii.2012, T. Robillard, 1♂ (TR41), enregistrement appel [call recording, MNHN-SO- 2018-38] (MNHN-EO- ENSIF1707); 1♂ (TR3), enregistrement appel [call recording MNHN-SO-2018-], molecular sample X13 (MNHN- EO-ENSIF3562). Shanghai, Padang Shanghai, E121°32' N31°13' [Shanghai market], viii.2012, T. Robillard, 1♂, molecular sample X11, enregistrement appel [call recording, MNHN-SO- 2018-37] (MNHN-EO-ENSIF1599); 1♂ (MNHN). Shanghai, Prov. Klange[?], Musée Meude, O. Piel, 30.viii. [19]30, 1♀, #1623, 1♂, 6.ix. [19]30 (MNHN). Guangdong, Shenzhen Distr., Xichong, 27.ix.2014, coll. Zhang Tao, 4♂, 2♀ (SNNU). Henan, Xinxian Distr., 4– 8.ix.2014, coll. Ma Libin, 2♀ (SNNU). Hainan, Wuzhishan Mt., 12.viii.2010, coll. Jiang Chaozhong, 1♂ (NWAFU). Zhejiang, Tianlongshan Mt., 8.vii.2009, coll. He Zhuqing, 1♂ (ECNU). Shaanxi, Ankang, 10.viii.2017, coll. Ma Libin, 1♂ (SNNU). Taiwan, Takao, Formosa, Sauter xi.[19]07, type of Heterotrypus unipartitus , det. Karny, identified Dionymus formosanus n. sp. Type by T. Shiraki, (DEI, Eberswalde), identified Xenogryllus marmoratus unipartitus (Karny) by A. V. Gorochov (DEI) [examined on photo]. Sri Lanka: Ceylon, Kandy, H. Rolle Berlin SW.11, 1♂ (MNHN). South Korea: data from website “South Korea: Orthopteroids of Korea” (http://www.jasa.pe.kr/pulmuchi/korthoptera/Xenogryllus-marmoratus.htm).

Distribution. Japan, China (including Taiwan and Hainan), South Korea (http://www.jasa.pe.kr/pulmuchi/ korthoptera/Xenogryllus-marmoratus.htm), India (specimens observed by Chopard (1969)), Sri Lanka (1 male specimen).

Emmended diagnosis. Species characterised by its small size ( Fig. 1 E–G), head dorsal coloration with a faint median dark band on vertex, and male genitalia with very long and thin pseudepiphallic lophi ( Fig. 8G ); in male FWs ( Fig. 5E ), mirror wider than long, less rounded than in X. maichauensis , X. ululiu and X. transversus .

Redescription. Species of small to average size, the smallest for the genus ( Fig. 1 E–G). Coloration almost homogeneously light ochre. Median dark band on head dorsum faint on vertex ( Fig. 4E ), prolonged anteriorly by black fastigium, with three narrow lateral bands on each side. Fastigium straight, not widened anteriorly. Antennae light brown, first segment darker. Face relatively flat in lateral view ( Fig. 3J ). Pronotum dorsal disc light brown, with a median brown band variably marked prolonging head coloration; lateral margins not carinated, with a narrow yellow line; lateral lobes ochre, progressively lighter ventrally.

Male. FWs light brown, translucent ( Fig. 5E ); dark coloration anterior to 1A sometimes extended on angle of 1A. Transverse part of file almost straight, with 435 stridulatory teeth (n=1) on transverse part of 1A. Harp as wide as long. Fused part of cells c1 and b1 as long as individualised part of c1. Cell d2 wider than in other species. Mirror wider than long, little rounded, its inner limit forming a wide angle. Apical field short, including five cell alignments.

Male genitalia ( Fig. 7 H–I): Pseudepiphallic sclerite longer than rami; pseudepiphallic lophi very long ( Fig. 8G ), narrow and sclerotized, slightly convergent, ended by an apical lamella curved dorsally and forming a small notch. Pseudepiphallic sclerite with wide lateral membranous lobes. Basal reinforcement of pseudepiphallic sclerite strong. Rami short with convergent hook-like apex. Pseudepiphallic parameres with a ventral transverse crest absent in other species; inner apex of of pseudepiphallic parameres sharp, slightly sclerotized apically. Ectophallic apodemes strong, not lamellate. Ectophallic lateral arms strong, fused to lateral arms of ectophallic fold, and with lateral arms of endophallic sclerite, forming together a wide gutter. Ectophallic fold entirely sclerotized ventrally. Endophallic sclerite small, apodeme made of a thin anterior dorsal crest and narrow lateral lamellas.

Female. FWs dorsal field with 8–9 strong longitudinal veins (m = 8, n = 10) and faint transverse ones. Subgenital plate not indented apically ( Fig. 9B ). Ovipositor longer than FIII, its apex rounded and smooth.

Female genitalia: Copulatory papilla ( Fig. 11 C–D) thin and conical, widened basally; sclerotization strong basally and apically, its membrane plicate.

Life history traits. X. marmoratus lives on low vegetation in bushes and grass in secondary habitats and along roads. The species has a nocturnal activity; males call at night from the vegetation. See Olivero & Robillard (2017) for description of peculiar mating behaviours.

Calling song ( Figs 12B , 14 ). At 26.5°C, the calling song of X. marmoratus is made of relatively long syllables, each corresponding to one FW closure. Syllables form a two-part echeme: 1–3 singleton syllables (m = 2, n = 10), followed by a chirp made of three syllables, sometimes duplicated. The amplitude profile of the call shows that the singletons are usually less loud than the chirp. Total echeme duration is 711 ± 33 ms, for an echeme period of 6776 ± 368 ms (echeme duty cycle = 10.5%). The syllables have the following characteristics: 1 st syllable duration = 51.8 ± 1.5 ms (period = 318.3 ± 35.6 ms); 2 nd syllable duration = 50.3 ± 1 ms; chirp duration = 183.8 ± 1 ms; chirp syllable duration = 49.5 ± 1.9 ms (period = 68 ± 0.8 ms). The frequency spectrum shows a pure tone fundamental peak corresponding to the dominant frequency at ca. 5.9 ± 0.2 kHz, with a clear harmonic series, the third harmonic (ca. 18 kHz) being almost as powerful as the fundamental peak.

Measurements. See Table 6.

Taxonomic discussion. The types of De Haan, supposed to be in Leiden, were not found in RMNH in 2006 (TR, pers. obs.), with no record mentioning them as loaned (Rob De Vries, curator of the Orthopteran collection in RMNH, pers. com. 2006; confirmation by Luc Willemse, current curator of the Orthopteran collection in RMNH, pers. com. 2018). Specimens possibly belonging to the original type series were also searched for in other European museums, but could not be found. We thus designate a neotype from Japan that will serve as a reference for future systematic works.

Gorochov (1992) distinguished two subspecies, X. m. marmoratus ( Haan, 1844) and X. m. unipartitus ( Karny, 1915) . In his study, he considered Gryllus (Phalangopsis) marmoratus Haan, 1844 (type from Japan), as the nominal subspecies Xenogryllus marmoratus marmoratus , while he established the subspecies X. m. unipartitus based on the species Heterotrypus unipartitus Karny, 1915 , which was described from one female from Taiwan (Takao, examined on photo). The type of H. unipartitus clearly corresponds to a female of Xenogryllus , however, given the continuous variation observed across the wide distribution of X. marmoratus , and the limited information available based on female morphology, there is no reason to maintain these two subspecies. We thus synonymise the subspecies under X. marmoratus , and H. unipartitus becomes a junior synonym of X. marmoratus , as previously proposed by Chopard (1968).

Figures

FIGURE 1. (A–C) Xenogryllus eneopteroides: male in dorsal (A) and lateral (B) views, female in dorsal view (C); (D–G) Xenogryllus marmoratus: male in dorsal (D) and lateral (E) views, female in dorsal (F) and lateral (G) views; (H–I) Xenogryllus lamottei n. sp.: male holotype in dorsal (H) and lateral (I) views; (J–K) Xenogryllus maniema n. sp.: male paratype in dorsal (J) and lateral (K) views; (L–O) Xenogryllus maichauensis: male in dorsal (L) and lateral (M) views, female in dorsal (N) and lateral (O) views. Scale bar: 1 cm.

FIGURE 3. Heads of Xenogryllus in facial (left panel) and lateral views (right panel): (A–B) Xenogryllus eneopteroides; (C–D) Xenogryllus lamottei n. sp.; (E–F) Xenogryllus maichauensis; (G–H) Xenogryllus maniema n. sp.; (I–J) Xenogryllus marmoratus. (K–L); Xenogryllus mozambicus n. sp.; (M–N) Xenogryllus ululiu; (O–P) Xenogryllus transversus. Scale bar: 1 mm.

FIGURE 4. Heads of Xenogryllus in dorsal views: (A) Xenogryllus eneopteroides; (B) Xenogryllus lamottei n. sp.; (C) Xenogryllus maichauensis; (D) Xenogryllus maniema n. sp.; (E) Xenogryllus marmoratus; (F) Xenogryllus transversus; (G) Xenogryllus ululiu; (H) Xenogryllus mozambicus n. sp. Scale bar: 1 mm.

FIGURE 5. Male FW venation in dorsal view: (A) Xenogryllus eneopteroides; (B) Xenogryllus lamottei n. sp.; (C) Xenogryllus maichauensis; (D) Xenogryllus maniema n. sp.; (E) Xenogryllus marmoratus; (F) Xenogryllus mozambicus n. sp.; (G) Xenogryllus transversus; (H) Xenogryllus ululiu. Scale bar: 5 mm.

FIGURE 7. Male genitalia: (A–C) Xenogryllus eneopteroides, dorsal (A), ventral (B) and lateral (C) views; (D–E) Xenogryllus lamottei n. sp., ventral (D), lateral (E) views; (F–G) Xenogryllus maniema n. sp., ventral (F), lateral (G) views; (H–I) Xenogryllus marmoratus, ventral (H), lateral (I) views; (J–K) Xenogryllus mozambicus n. sp., ventral (J), lateral (K) views; (L– N) Xenogryllus maichauensis, pseudepiphallic lophi in ventral (L–M) and lateral (N) views, male from South China (L), male from Thailand (M, N); (O–P) Xenogryllus transversus, dorsal (O), ventral (P) views; (Q–R) Xenogryllus ululiu, pseudepiphallic lophi in dorsal (Q) and ventral (R) views. Scale bar: 1 mm.

FIGURE 8. Right pseudepiphallic lophi of male genitalia in dorsal view: (A–B) Xenogryllus eneopteroides, male from Central African Republic (A), male from Ivory Coast (B); (C) Xenogryllus lamottei n. sp.; (D) Xenogryllus maichauensis; (E) Xenogryllus maniema n. sp.; (F) Xenogryllus mozambicus n. sp.; (G) Xenogryllus marmoratus; (H) Xenogryllus ululiu; (I) Xenogryllus transversus. Symbols: Preapical hook-like expansion figured in gray or with a black arrow; dotted parts represent membranous areas; thin lines represent folds and reliefs in sclerites. Scale bar: 1 mm.

FIGURE 9. Apex of female subgenital plate and base of ovipositor in ventral view: (A) Xenogryllus eneopteroides; (B) Xenogryllus marmoratus; (C) Xenogryllus maichauensis; (D) Xenogryllus mozambicus; (E) Xenogryllus transversus; (F) Xenogryllus ululiu. Scale bar: 1 mm.

FIGURE 11. Female copulatory papilla: (A–B) Xenogryllus eneopteroides, in ventral (A) and lateral (B) views; (C–D) Xenogryllus marmoratus, in ventral (C) and lateral (D) views; (E–G) Xenogryllus maichauensis, in ventral (E), dorsal (F) and lateral (G) views; (H–J) Xenogryllus mozambicus n. sp., in ventral (H), dorsal (I) and lateral (J) views; (K–L) Xenogryllus ululiu, in ventral (K) and lateral (L) views; (M–O) Xenogryllus transversus, in ventral (M), dorsal (N) and lateral (O) views. Scale bar: 0.5 mm.

FIGURE 12. Comparison of calling songs, oscillogram of five echemes: (A) Xenogryllus maichauensis; (B) Xenogryllus marmoratus; (C) Xenogryllus mozambicus n. sp.; (D) Xenogryllus transversus; (E) Xenogryllus ululiu. Scale bar: 1 s.

FIGURE 14. Calling song of Xenogryllus marmoratus: (A) oscillogram of 5 echemes; (B) detailed oscillogram (upper panel) and sonogram (lower panel) of 1 echeme; (C) detailed oscillogram of one syllable; (D) frequency spectrum.