Xenogryllus eneopteroides Bolívar, 1890

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 : 306-310

Xenogryllus eneopteroides Bolívar, 1890

( Figs 1 A–C, 3A–B, 4A, 5A, 6, 7A–C, 8A–B, 9A, 10A, 11A–B)

Xenogryllus eneopteroides Bolívar, 1890: 232 — Karsch 1893: 204; Kirby 1906: 106; Giglio-Tos 1907: 26; Bolívar 1910: 544; Chopard 1954: 80; Chopard 1968: 349; Chiffaud & Gillon 1984 (life history); Paris 1994[1993]: 69 (lectotype designated); Robillard & Desutter-Grandcolas 2004a: 578; 2004b: 273 (morphological phylogeny); 2006: 644; 2008: 67; 2011: 637; Robillard 2006: 673; Robillard et al. 2007: 1267 (song frequency); Nattier et al. 2011: 2201 (molecular phylogeny); Chintauan-Marquier et al. 2016: 62 (molecular phylogeny); Vicente et al. 2017: 2203 (historical biogeography); Cigliano et al. 2018 ( Orthoptera Species File Online).

Xenogryllus eucopteroides— Kevan & Knipper 1961: 368 (misspelling); Chopard 1968: 349.

Type material. Lectotype, ♂, Angola: Duque de Bragança , Bayão (Paris 1994) [ MNCN, examined on photograph].

Additional material examined. Benin: Bas-Dahomey , plateaux de Zagnamado et de Ketou, forêt d’Aquaqueré (saison sèche), 1910 P. Ducorps, 1♀ (MNHN). Cameroun: [L.] Conradt , 1♂, identified Xenogryllus eneopteroides Bol. ( MNHN) . Reg [ion] de Kribi, 1925, littoral, saison humide, Dr. Gromier, 1♂ ( MNHN) . Kamerun [Cameroun], Pipindi [Bipindi], Eitel ded. 1♀, identified Xenogryllus eneopteroides Bol. ( MNHN) . 10 km N. of Yaoundé, E. of Agronomy Station, Nkolbisson , 10.vii.1975, N. D. Jago, 1♂, identified Xenogryllus eneopteroides Bolivar by B. C. Townsend, 1976 ( NHMUK 010926525 ) . Central African Republic: République Centrafricaine, 75 km WNW Mbaiki, N’Gotto [Ngoto], W Lobaye, savane arbustive, 7.iv.1995, nuit [night], L. Desutter-Grandcolas, 1 juvenile, n°14, molecular sample Xen-CA (MNHN-E0-ENSIF3159); 1♂, #1, sur plante [on plant] ( MNHN) . Democratic Republic of the Congo: Congo, 20 km from L. Tumba (towards Coquilhatville ) [Mbandaka], 5.iii.1964, from grassy clearing in forest, 1♂, 1♀, Tyson Roberts, Brit. Mus. 1982-71 ( NHMUK

0 10926531, 010926572). Kalembe, 15.ix.1947, Miss. Tanganika, 1♀ (RBINS). Lukafa [Lukafu], Congo Belge , 6.xii.1938, H.J. Brédo, 1♀, R.Mus.Nat. Belg. I.G.12.204 ( RBINS) . Belgian Congo, Aru , iii.1936, 5 ♂, 2♀, H.J. Bredo, dry Acacia bush ( NHMUK 0 10926536, 0 10926527, 0 10926578, 0 10926580, 0 10926539, 0 10926566, 010926569). Niangara, v-vi-1915, 28°0 E 3°40’ N, 1♂ ( AMNH) . Haut-Ituri: Faradje, 1♂, Blommaert , Musée du Congo, identified Xenogryllus eneopteroides by L. Chopard, R. Det. N 2814 ( MRAC) . S.E. Katanga: Ngaye : xixii.1931, 1♂, R.P. Claquin, Musée du Congo, identified Xenogryllus eneopteroides by L. Chopard, R. Det. N 2814 ( MRAC) ; 1932, 1♂, 1♀, R.P. Claquin, Musée du Congo, identified Xenogryllus eneopteroides by L. Chopard, R. Det. DD3509 ( MRAC) . Haut-Uele , vi.1925, 1 ♀, Musée du Congo, identified Xenogryllus eneopteroides by L. Chopard, R. Det. N 2814 ( MRAC) . Kibali-Ituri: Geti , ii–iv.1937, 1♂, Ch. Scops, Musée du Congo, identified Xenogryllus eneopteroides by L. Chopard, R. Det. B 5318 ( MRAC) . Gabon: Plateaux Batéké, piste Ekalla , 30 km W Leconi [Lekoni], milieu herbeux, 19.vi.1994, L. Desutter-Grandcolas: 1 juvenile, n°61, jour [day], litière [leaf litter], molecular sample Xen-GA (MNHN-EO-ENSIF3442); 1♂, n°3, nuit [night], sur plante—base tronc [on plant, low on tree trunk] ( MNHN) . Bas Ogooué , 1♂, identified Xenogryllus eneopteroides Bol. by L. Chopard ( MNHN) . Ntoum , 19.vii. [19]85, A. Pauly, 2♂, 2♀, 2 juveniles, lumière [at light] ( RBINS) . Ghana: Eastern region, Hansowodze , 2 mls from Kade A.R.S. [Agricultural Research Station], 19.vii.1963, I. K. Bacheampong, 4♂ ( NHMUK 0 10926524, 0 10926567, 0 10926528, 010926571). Ghana, Togoland plateau, grassy, summit above [Nkonya] Wurupong, 23.xii.1959, N.D. Jago, 1♂, Brit. Mus. 1965-474 ( NHMUK 010926535 ) . Eastern region, Kade Agr. Res. Stn [Agricultural Research Station], 11.vii.1963, I. K. Bacheampong, 2♂, Brit. Mus. 1965-474 ( NHMUK 0 10926526, 010926538). Transvolta, Togoland, Amedzofe: 3.v.1959, montane grass, 1♀, N.D. Jago, Brit. Mus. 1965-474 ( NHMUK 010926529 ) ; 25.v. [19]59, N.D. Jago, Brit. Mus. 1965-474 ( NHMUK 010926579 ) . Western Reg. [ion] mls E of Elmina, 1°20W 5°05.5N, xii.1960, 1 ♂, 1♀, N.D. Jago, identified Xenogryllus eneopteroides by P.C. Tinning, 1966, Brit. Mus. 1965-474 ( NHMUK 0 10926509, 010926511). T.v.T [Trans-Volta Togo], Ho-Hohoe rd. 10 miles from Volta bridge, iii.1960, N.D. Jago, Brit. Mus. 1965-474 ( NHMUK 010926553 ) . Western Reg. Asankragua-Enchi rd., Tano R. Ferry , 24.ix.1962, 1♂, N.D. Jago, Brit. Mus. 1965-474 ( NHMUK 010926510 ) . Guinea: Kaoulenta [Kéoulenta], [Mount] Nimba ( Guinée ), ii–vi.[19]42, M. Lamotte, 1♂, 1♀, 2 juveniles ( MNHN) . Nimba (Guinée), [Mount] Pierre Richaud , 900 m, M. Lamotte ii–iv.[19]42, 1♀, identified Xenogryllus eneopteroides Bol. by L. Chopard; 1♀ ( MNHN) . Nimba (Guinée), Lamotte , Amiet , Vanderplaetsen , xii.56 – v.57: Ziéla , 17.ii. [19]57, 1 juvenile; Ziéla , 20.iv. [19]57, 1 juvenile; sav. [savannah] entre Ziéla et Gbakbré, 9.xii.1956, 1 juvenile; Zgpo. [Zouguépo], 900 m, androp, 18.iv. [1957], 2♀; sav. [savannah] Ziéla , 3.ii. [19]57, 1 juvenile; sav. [savannah] Ziéla , 20–25.xii. [19]56, 4 juveniles; sav. [savannah] Kéoulenta , 5.xii. [19]56, 2 juveniles; forêt Ziéla , 19.ii. [19]57, 1 juvenile; 1 juvenile, A.V. n°23, identified Xenogryllus eneopteroides Bol. larva by L. Chopard; 1 juvenile, A.V. n°38; 1 juvenile, 87CD; 14.iii. [19]57, 1 juvenile, fge n°47 ( MNHN) . Mt. Nimba, Mt. Leclerc [Mount Jean Charles Leclerc], 1500 m, 23.iii.1991. M. Lamotte, n°319, 1♀ ( MNHN) ; 1♀, molecular sample X22 -XenGU (MNHN-EO-ENSIF1494). Nimba (Guinée), M. Lamotte, xii.56 – v.57, 29. x.[1956], 1♂, herbes A2 Ziela , identified Xenogryllus eneopteroides Bol. by R. Roy, 1959 ( MNHN) . Guinée Française, Diéké [Diecke], P. Chabamaud, 1920, 1♀, identified Xenogryllus eneopteroides Bol. by L. Chopard ( MNHN) . Ivory Coast: Lamto (Toumodi), Coll. E.N.S. Paris, Lamotte et collab.: 1– 10.ii. [19]62, 3♂, 2♀; 1963, 1♂, Xen-Robillardmorpho ; 25–31.xi. [19]62, 1♀; 15–20.viii. [19]62, 1 juvenile; iv.[19]62, 1♀; 7.xi. [19]62, 1♀; 20.xi. [19]62, 1♀; vi.[19]62, 1♀; 4.vii. [19]62, 1♀, mare sav. inondée [inundated savannah] ( MNHN) . Forêt à 15 km de Lamto , 10.ix.1968, C. Girard, 1♀, 1 juvenile ( MNHN) . Forêt de Yape , 1♂ ( MNHN) . Toumodi , xii.1930 – iv.1931, Ch. Alluaud & P. A. Chapuis, 1♂ ( MNHN) . Danané , i.1939, 1 ♀ ( MNHN) . Lamto (Toumodi), C. Girard : 12.ii.1968, savane [savannah], lisière forêt galerie [hedge of gallery forest] 6♂, 3♀, 5 juveniles; 1♂, identified Xenogryllus eneopteroides Bol. by R. Roy , 1974; 12.iii.1968, lisière, forêt, savane, 1♂, molecular sample XenCI (MNHN-EO- ENSIF1481); 8♂, 2♀, 7 juveniles; 1♂, photos MEB [SEM photos] (MNHN-EO-ENSIF1518); 15–30.vii.1968, savane [savannah], lisière forêt galerie [hedge of gallery forest] 1♂, 2♀, 3 juveniles; 27–28. ii.1968, 2 ♀, 1 juvenile, à la lumière; 10–20. iv.1968, 1 ♂, à la lumière ( MNHN) . Lamto (Toumodi), Paul Planquette : 1♀, PNB 18; 1 juvenile, PNB 13B; 15.i. [19]64, 1♀, PNB 64 ( MNHN) . Lamto ( Toumodi ), 30.vi. [19]62, Coll. E.N.S. Paris, Lamotte et al., 1♂, identified Xenogryllus eneopteroides by L. Chopard ( MNHN) . Nimba [Mount]: Lamotte , 1946, Z4, 1♀, identified Xenogryllus eneopteroides by R. Roy, 1967 ( MNHN) . Lamto , 1963, 1♂ ( MNHN) . Assinie , 3295-85, Chaper, 1♂ ( MNHN) . Réserve du Banco [Banco national park], R. Paulian & G. Delamare, 2♂, 2♀, 2 juveniles ( MNHN) . Bouaké (C.I.), 22.v. [19]62, coll. E.N.S. Paris, Lamotte et collab., 1♂ ( MNHN) . Taï , 18.viii.1978, G. Couturier, 1♀ ( MNHN) . Lamto , 1962, P. Le Gall, 2♂, 2♀ ( MNHN) . Kenya: CRS [Catholic Relief Service] Tebere , 0039.5S 3723E, 10.i. [19]72, I.A. D.R. 72/58, 1♀, Brit. Mus. 1982-71 ( NHMUK 010926573 ) . Nigeria: Lagos, Ikoyi , viii. 1951, L. Bala, 1♂, identified Xeneogryllus eneopteroides by L. Chopard ( MNHN) . U.C. [University College] Ibadan , 5.ix.1952, Tephrosia, leaf, Teph-66 coll. G.H. Caswell, 1♀ ( MNHN) . Ibadan , 6.vi. [19]23, 1♂, 1605 ( NHMUK 010926581 ) . Calabar, Nigeria , 10.iv. [18]82, 10.30 A.M., W. edge of [?t] Kwa swamp, E. edge of UNICAL staff quarters; about 50 cm up on base of tangle of turns e.g. Clitonia? rubiginosa, notes 72, 1♀, C.I.E. coll A.156 76 Pres by Comm Inst Ent B.M. 1985-1( NHMUK 010926537 ) . Replublic of Congo: Sibiti , xi-1963, mission A. Descarpentries et A. Villiers 1963–1964, 2♂, 1♀ ( MNHN) . Baie de Lagoa , envoi H. Deyrolle, 1♀ ( MNHN) . Mayumbe, Luki, L. Tiebers [?], 1♀, Musée du Congo ( MNHN) . Odzala , x.1963, 3 ♂, 3♀, 1 juvenile ( MNHN) . Congo , 1963, 1♀ ( MNHN) . Ile [island] M’Bamou , 26.vii.1968, Fidèle, 1♀ ( MNHN) . Boma , Sundi, P. Rolin, 1♂ ( RBINS) . Banana, F. Busschodts , 4♀ ( RBINS) . Sierra Leone: Piste Bandankoro , 29.v.1963, savane de plaine ayant brûlé [burnt lowland savannah], mission ENS-IFAN aux Monts Loma, 1♂, identified Xenogryllus eneopteroides by R. Roy, 1992; 2♂, 3♀ ( MNHN) . Piste Keimadugu , savane [savannah], 520 m, 29.v.1963, mission ENS-IFAN aux Monts Loma, 2♂, 1♀ ( MNHN) . Firawa , savane [savannah], 1.vi.1963, mission ENS-IFAN aux Monts Loma, 1♂, 1♀ ( MNHN) . Njala, E. Hargreaves : 21.vii. [19]32, B.M. 1938-149, identified X. eneopteroides Bol. by L. Chopard, pres Imp. Inst. Ent. Brit. Mus. 1965-474 ( NHMUK 010926552 ) ; x.1936, 1 ♂, pres Imp. Inst. Ent. Brit. Mus. 1965-474, 1♂, identified Xenogryllus eneopteroides by B. Uvarov ( NHMUK 010926534 ) ; 23.v. [19]25, 1♂, identified X. eneopteroides Bol. by L. Chopard, B. M. 1938-149 ( NHMUK 010926530 ) ; xi.1935, 1 ♂, pres Imp. Inst. Ent. Brit. Mus. 1936-252, 1♂, identified Xenogryllus eneopteroides by B. Uvarov ( NHMUK 010926576 ) . Uganda: Entebbe , 20.xi.1970, W.J. Bailey, B.M.1977-246, identified Xenogryllus eneopteroides by B. C. Townsend, 1978 ( NHMUK 010926570 ) . Kisaru , 19.vii.1983, 1♂, H.B. Johnson, at light ( NHMUK 010926551 ) . Zambia: Lusaka, about fifty metres from the Great East Road in Chalimbana, Chongwe Distr., 19.xii.2013, William van Niekerk, 1♂, identified X. eneopteroides by William van Niekerk [online photograph]. Kapiri , 13.xi.1913, 2♀, L. Charliers, Musée du Congo, identified Xenogryllus eneopteroides by L. Chopard, R. Det. N 2814 ( MRAC) .

Type locality. Angola, Duque de Bragança , Bayão (=Kalandula) .

Distribution. Sub-Saharan Africa: Angola, Benin, Cameroun, Central African Republic, Democratic Republic of the Congo, Gabon, Ivory Coast, Kenya (one large female specimen with unclear identification), Nigeria, Republic of Congo, Sierra Leone, Uganda, Zambia.

Emended diagnosis. Species of average size, close to X. mozambicus n. sp. and X. maniema n. sp., from which it differs by face almost flat in lateral view (more rounded in other species), shorter male FWs (short apical field), and shape of lophi in male genitalia. Differing from X. lamottei n. sp. and Asian species by following characters: well-carinated lateral angles of dorsal disc of pronotum (also carinated in X. mozambicus and X. maniema ), male genitalia and very short ovipositor.

Redescription. Species of average size ( Fig. 1 A–C), coloration gray brown little contrasted. Eyes large, lateral, occupying almost half of head height in lateral view ( Fig. 3 A–B). Face almost flat in lateral view, with typical whitish mask underlined by a black line below eyes and on mandibles. Pronotum dorsal disc strongly carinated laterally ( Fig. 3B ), coloration dark brown or brown, most often with a median dark brown band extended laterally near anterior margin, forming a T shape; lateral lobes almost homogeneously dark brown. First article of antennae dark brown.

Male. FWs very wide, dark coloration anterior to 1A including angle of 1A ( Fig. 5A ). File with 658 stridulatory teeth (n=2) on transverse part of 1A. Harp longer than wide. Cell c1 narrowed posteriorly, its fusion with cell b1 slightly shorter than its individualized part. Cell c2 large, d2 long and narrow. Mirror large, wider than long and little rounded, its inner angle forming an angle. Apical field short, wider than long, including four or five cell alignments.

Male genitalia ( Figs 7 A–C, 8A–B): Pseudepiphallic lophi as long as rest of pseudepiphallic sclerite, forming a long rectangle, sometimes slightly widened preapically; lophi with a narrow membranous inner margin setose basally ( Fig. 8 A–B); their bases fused until mid-length; apex pointed and divergent, ended by a thin lamella without hook-like inner dorsal expansion (in dorsal view, anterior base of lamella suggests that lamella may partly correspond to modified inner dorsal expansion of other species); ventral blade of lophi without longitudinal wrinkles, but with a strong ventral transverse carina, as in X. mozambicus n. sp. Pseudepiphallic parameres with a strong rectangular ventral lobe and a basal membranous lobe. Rami strong, their apex forming wide convergent hooks. Ectophallic apodemes strong, not lamellate. Ectophallic lateral expansions, lateral sclerites of ectophallic fold and endophallic sclerite partly fused, forming a wide ventral sclerotized plate, trifid apically; endophallic apodemes made of wide lateral lamellas and a narrow dorsal crest.

Female: ( Fig. 1C ). Dorsal disc of pronotum almost rectangular, its posterior margin slightly bisinuate. FWs dorsal field with 7–10 longitudinal veins (m=8, n=10). Subgenital plate with a shallow apical indentation ( Fig. 9A ). Ovipositor ( Fig. 10A ) very short, about one third of FIII length.

Female genitalia. Copulatory papilla ( Fig. 11 A–B) conical and narrow, its apex rounded; well-sclerotized except base and apex.

Life history traits. According to Chiffaud & Gillon (1984), X. eneopteroides is found in Ivory Coast at the end of the dry season, in parts of savannah protected from fires. Their study revealed that the species feeds only on leaves and flowers of dicotyledones which are specific to savannah.

Calling song. According to the sonogram showed in Desutter (1983), analyzed by Robillard & Desutter- Grandcolas (2004a, 2011), the calling song of X. eneopteroides is a short echeme made of five long syllables (syllable duration = 65 ms, period = 106.5 ms. Echeme duration = 611 ms, echeme period = 1062 ms. The dominant frequency is low (3.6 kHz) and corresponds to the first peak of the spectrum.

Measurements. See Table 2.

Taxonomic discussion. The material examined reveals numerous slight differences in male genitalia ( Fig. 8 A– B) and size along the wide geographic distribution of the species. In particular, specimens from Congo differ from the material from the western coast of Africa. However, these differences are not sufficiently clear or are based on too few individuals to delimit different species yet. More information and sampling will be necessary to improve the study Xenogryllus in Africa.

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 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 10. Apex of female ovipositor: (A) Xenogryllus eneopteroides; (B) Xenogryllus maichauensis. 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.