Phaloria chopardi ( Willemse, 1925 ) Desutter-Grandcolas, 2009

Desutter-Grandcolas, Laure, 2009, New and little known crickets from Espiritu Santo Island, Vanuatu (Insecta, Orthoptera, Grylloidea, Pseudotrigonidium Chopard, 1915, Phaloriinae and Nemobiinae p. p.), Zoosystema 31 (3), pp. 619-659 : 630-637

Phaloria chopardi ( Willemse, 1925) n. comb.

( Figs 3 ; 4 ; 5 A-D; 6)

Podoscirtus chopardi Willemse, 1925: 521 .

Munda chopardi – Chopard 1968: 412.

This species was known up to now by one female only ( Willemse 1925). Several males have been found during SANTO 2006, which allow to propose an amended diagnosis and description.

HOLOTYPE. — Female, Jan 8 1923, N Hebrides [ Vanuatu], c. 500 ft, E. c’st Esp[iritu] Santo I., open forest near Hog Harb[our], dd J.R. Baker and & Percy Sladen Mem. FD. 1923 / Santo 8 Jan 1923 [manuscript label] / TYPE female Willemse Podoscirtus chopardi . Tr. Ent. Soc. L. 1925, p. 521. [white label with red margin] / Podoscirtus chopardi nov. sp. female Det. C. Willemse / Type [red

New crickets ( Insecta, Orthoptera, Grylloidea ) from Espiritu Santo, Vanuatu label] / Jan 8 1923, N Hebrides [ Vanuatu], c. 500 ft, E. c’st Esp[iritu] Santo I., open forest near Hog Harb[our], dd J.R. Baker and & Percy Sladen Mem. FD. 1923 [same as above] / TYPEORTH: 908 Podoscirtus chopardi Willemse’s HOPE DEPT.OXFORD. ( UMO). Type in bad condition (legs III missing, right FW broken at base, right HW damaged, ovipositor slightly damaged, antennae missing except left antenna base, colouration unclear).

OTHER MATERIAL EXAMINED. — 4 ♂♂, 5 ♀♀. Vanuatu. [Sanma province], Espiritu Santo [Island],

Peavot, 14°59’37”S, 166°47’4”E, 38 m alt., rive S de la rivière principale, terrasse, forêt secondaire (ancien jardin), nuit, sur plante, 2 m H, sous feuille d’arbre, 20.X.2006, 1 ♂ (fn TR254), 1 ♀ (fn TR253), observed in copula (MNHN-ENSIF2102, 2103). — Idem, nuit, sur plante, 21.X.2006, L. Desutter-Grandcolas, 1 ♂ (fn 65) (MNHN-ENSIF2104).

Butmas, 600 m alt., forêt du plateau de Tankara, 15°21’56”S, 166°59’E, jour, sur plante, 13.X.2006, T. Robillard, 1 ♀ (fn TR21) (MNHN-ENSIF2109). — Nuit, lisière proche du camp, herbes hautes en bord de piste, sur feuille, 14.X.2006, T. Robillard, 1 ♂ (fn TR75) (MNHN-ENSIF2106). — Jour, sur plante de sousbois, 1 m H, 15.X.2006, T. Robillard, 1 ♀ (fn TR116) (MNHN-ENSIF2110). — Nuit, végétation secondaire, bord de piste, 17.X.2006, L. Desutter-Grandcolas, 2 ♀♀ (fn 5-6) (MNHN-ENSIF2108, 2107).

Big Bay, Matantas, Vatthé Conservation Area, 15°20’S, 166°95’E, nuit, sur plante, 27.X.2006, T. Robillard, 1 ♂ (fn TR398) (MNHN-ENSIF2105).

DISTRIBUTION. — Vanuatu, Espiritu Santo island: species known from western (Peavot, Vatthé Conservation Area) and eastern (Hog Harbour, type locality) coastal areas around Big Bay and in the inland (Butmas).

AMENDED DIAGNOSIS. — Species close to Phaloria solomonica Gorochov, 1996 from the Solomon Islands, from which it differs by its smaller size, the smaller ovipositor of the females (between 11.3 and 12.7 mm, compared to 14.4 mm in P. solomonica ), female tegminal venation (more numerous transverse veins on dorsal field, Fig. 4F, G; R less bifurcated), and male genitalia ( Fig. 5 A-D): pseudepiphallic lophi well developed, long and membranous (lacking in P. solomonica , see Gorochov 1996: fig. 365); ectophallic fold more deeply indented than in P. solomonica (compare to Gorochov’s figure 367); ectophallic parameres thin and relatively short, not going beyond pseudepiphallic sclerite, but longer than in P.solomonica . Due to the discoveries of additional species, the diagnostic characters mentioned by Willemse (1925) revealed uninformative for species recognition (proportions of maxillary palpi joints, position of median ocellus on fastigium, female FW venation, colouration).

REDESCRIPTION

Size medium. Overall colouration light ochre, the head and pronotum darker dorsally, the legs lighter with indistinct darker rings and spots, highly pilose.

Head ( Fig.3 ). Fastigium flattened dorsally, slightly narrower than the scape. Ocelli small, the distance between the median and one lateral ocellus hardly greater than the distance between lateral ocelli; median ocellus subapical in position, protruding. Eyes small, but protruding laterally and anteriorly. Head dorsum rounded. Maxillary palpi: joint 5 longer than joint 3, which is longer than joint 4; joint 5 widened from mid length to the tip, truncated in oblique. Pronotum. Anterior and posterior margins of dorsal disc slightly concave and convex respectively.

Legs. TI with 2 very small apical spurs, both ventral in position; tympana both open, small and oval, the outer less than half the surface of the inner; tympana prolonged by a small more distal depression.TII with 3 small apical spurs, the upper outer spur missing; upper inner spur longer than ventral spurs. TIII with 4 pairs of subapical spurs; within each pair, inner spur lower on the tibia; inner spurs longer, and more apart from the main axis of the tibia, than the outer spurs. TIII serrulated over their whole length; inner spines number: 0 between apex and spur 1, and between spurs 1 and 2; 1 or 2 between spurs 2 and 3; 2 or 3 between spurs 3 and 4; 8-13 above subapical spur 4; outer spines bigger and more numerous: 0 or 1 between apex and spur 1; 2-4 between spurs 1 and 2; 3-5 between spurs 2 and 3; 4-7 between spurs 3 and 4, 10-13 above subapical spur 4. TIII with 3 apical spurs on each side ( Fig. 4A, B ); outer spurs very small, the median the longest; inner spurs much longer, the median twice as long as the ventral one, the upper the longest, as long as basitarsus. Basitarsi III asymetrical, their dorsal outer side crested with 4 or 5 spines in addition to apical one; spines bigger toward basitarsus apex. FWs and HWs well developed in both males and females; HWs somewhat shorter or longer than FWs, both about body size; FWs covered with numerous, very short golden setae.

Colouration

In holotype, no obvious colouration pattern on head and legs; dorsal disc of pronotum perhaps darker on posterior third. In other specimens, face, clypeus, labre and cheeks whitish; a small oblique, brown line converging from the lower angle of each eye toward the epistemal suture, but not going further than mid face. Ocelli whitish. Head dorsum dark ochre. Antennae light ochre. Palpi light ochre; tip of joints 4 and 5 somewhat darker. Mouthparts

ps

yellowish. Dorsal disc of pronotum dark ochre, with yellowish inscriptions; lateral lobe with a large yellow spot. Legs I and II light ochre; one darker subapical ring on femora; 3 darker dorsal spots on tibiae, in addition to one little spot near the knees. Legs III ochre but more spotted with dark;TIII with 5 dark spots/rings, one at the level of each pair of subapical spurs and one below the knee; subapical spurs brown, yellow near the apex, black at apex; apical spurs yellowish, their apex black; FIII darker, without a very clear pattern of colouration: more or less, one dark subapical ring, surrounded by

G

yellowish areas, and 2 dark patches on inner and dorsal sides, these patches more or less prolonged by dark stripes on outer face. Basitarsi III whitish, brown basally and apically.

Male

Metanotum and abdomen without evident glandular structures. FWs and HWs well developed, HWs almost as long as FWs. FW dorsal field venation standard for the genus ( Fig. 4C ); CuP vein prolonged down to mid harp; CuA bifurcated 6 times after the mirror; apical field with clear cell alignments; lateral field smaller than dorsal field, its venation as on Fig. 4D, R with more than 20 bifurcations. Stridulatory apparatus: harp with 8 veins; mirror crossed by two veins (male MNHN-ENSIF2102); file with 34-37 teeth (mean 36, n = 4), the teeth located only on inner half of 1A ( Fig. 6A ), well separate from each other and slightly oblique (i.e. not transverse to FW surface, Fig. 6B ). Subgenital plate long, not very high ( Fig. 4E ), its apex deeply indented. Colouration: FWs light ochre, the veins darker in apical field; plectrum area whitish. Sternites light ochre, with an indistinct median dark band.

Male genitalia. Male genitalia long and of regular width; rami twice as long as pseudepiphallic sclerite. Pseudepiphallic sclerite very largely and deeply indented, making two lateral “arms” which entirely enclose the long and membranous lophi ( Fig. 5A ); posterior margin of pseudepiphallus, around the lophi, thickened. Pseudepiphallic parameres well developed, well sclerotized in their distal third only, and comprising 4 distinct lobes: a dorsal lobe, thin and transverse; two median, plate-like lobes, which are the most evident part of the parameres, and a latero-ventral, mostly membranous, lobe, comprising a free, membranous apical bud, and a small, spiny plate ( Fig. 5B, D ). Ectophallic invagination sclerotized laterally only (no sclerotized arc); ectophallic apodemes thick. Ectophallic fold long and wide, its lateral margin sclerotized from the ectophallic invagination and prolonged by a pair of free, thin ectophallic parameres ( Fig. 5C ); apex of ectophallic fold deeply indented and well separate from the lateral ectophallic parameres ( Fig. 5A ). Endophallic sclerite well developed, triangular; endophallic apodeme comprising a high, undulated longitudinal crest along the whole length of endophallic sclerite, and a pair of transverse sclerites along the anterior margin of endophallic sclerite ( Fig. 5A ). Endophallic membrane invaginated posteriorly to endophallic sclerite, making a kind of small, half-closed “dorsal cavity”.

Female

FWs longer than body; HWs slightly longer than FWs. Venation ( Fig. 4F, G ): longitudinal veins parallel and strong, separated by numerous, fainter transverse veins; A3 bifurcated twice, near its base, CuP twice and CuA three times. Lateral field: area between R and MA wide; R with 11 bifurcations. Ovipositor apex: dorsal valves with 4 large crests, in addition to the curved apex; ventral valves with 4 transverse crests, in addition to acute apex, the 3 most apical higher than wide, the most basal low and very wide ( Fig. 4H, I ). Subgenital plate large, its distal margin deeply concave ( Fig. 4J ).

Colouration: FWs light ochre; longitudinal veins ochre; CuP and CuA dark brown basally and at mid length respectively; transverse veins darker; in lateral field, MA base dark brown.

Female genitalia. Copulatory papilla always short, thin and hardly sclerotized dorsally only, and shorter in female holotype; lateral margins subparallel, apex rounded more or less bent ventrally; membrane ventral to copulatory papilla forming a kind a plicated pouch ( Fig. 4K, L ).

Measurements

See Table 4.

VARIATION

Eyes often dark brown in their lower half. Dark patches of leg colouration more or less evident. In male FW, mirror crossed by an additional, bifurcated vein ( Fig. 4M ). Veins joining at the anterior angle of the mirror either fused before the mirror ( Fig. 4M ), or separate ( Fig. 4C ). In females, FW transverse veins almost as strong as longitudinal veins.

REMARK Two males originating from Big Bay (Matantas, Vatthé Conservation Area, 15°20’S, 166°95’E, nuit, sur plante, 27.X.2006, T. Robillard, 1 ♂ [fn TR399] [MNHN-ENSIF2111]; nuit, sur plante de sous-bois, L. Desutter-Grandcolas, 1 ♂ [fn 55] [ MNHN- ENSIF2112 ]) differ slightly from the above mentioned males, especially by their more contrasted colouration (FIII distinctly annulated, yellow spot on lateral lobe of pronotum and oblique brown lines on the face more obvious), their slightly lower number of teeth in the stridulatory file (30 and 34), and their male genitalia (ectophallic apodemes shorter and thinner, ectophallic fold somewhat longer, longitudinal crest of endophallic apodeme almost missing). Their identification as P. chopardi n. comb. cannot be ascertained yet. See Table 5 for measurements .

CALLING SONG

The male MNHN-ENSIF2102 has been recorded in semi-captivity by night (temperature 27.3°C, microphone at about 20 cm), being raised in a small mosquito net cage selected to avoid echoes during recording.

The calling song is made by the repetition of short buzzing echemes, separated by long intervals ( Fig. 6C ). Echeme duration ranges from 760 ms to 1 s (mean 866 ms, n = 20); that of the intervals ranges from 2.14 s to 8.06 s (mean 4.43 s, n = 20). Each echeme is composed of several tens of subechemes emitted at a very quick rate, and which exhibits clear amplitude modulations ( Fig. 6D ). Most subechemes have a total number of elementary oscillations which exceeds the number of stridulatory teeth (here 38). A subecheme thus cannot be produced by one complete FW closure movement. Unless echoes have nethertheless damaged the amplitude pattern of the eschemes, it must then be hypothesized that each subecheme is produced by a complex FW closure movement involving one or several partial reopening(s) of the FWs, and so that each subecheme comprises two or three syllables emitted extremely rapidly ( Fig. 6F, G ). Subecheme duration ranges from 9.41 ms (2 syllables) to 18.20 ms (4 syllables), with a mean duration of 14.45 ms (n = 20; mean number of syllables = 3). Subechemes are separated by very short intervals lasting from 1.02 to 4.75 ms (mean 2.54 ms, n = 20). Within echemes, the interval between syllables is extremely short, less than 3 ms and most often less than 2 ms. An alternative hypothesis would be that both FW closing and FW opening are used for stridulation, as in ensiferan Tettigoniidae ; the movement of sound production is anyway extremely fast in this species (K.-G. Heller pers. comm.).

The dominant frequency of the call is its fundamental frequency.Up to 8 harmonics are documented on frequency spectrum ( Fig. 6E ); harmonics H2 to H5 are almost equally powerful, and H6 almost lacking. Dominant frequency ranges from 5222 Hz to 5425 Hz (mean 5292, n = 20).

Figures

FIG. 3. — Phaloria chopardi (Willemse, 1925) n. comb., ♂ MNHN-ENSIF2102. Scale bar: 3 mm.

FIG. 4. — Phaloria chopardi (Willemse, 1925) n. comb.: A, B, tibia III, inner subapical and apical spurs (A), outer apical spurs (B); C, D, male forewing venation,dorsal field (C) and lateral field (D); E, male subgenital plate, lateral view; F, G, female forewing venation, holotype (F) and ♀ MNHN-ENSIF2103 (G); H, I, holotype ovipositor, ventral (H), lateral (I) views of right valves, ventral valves marked with grey; J, holotype subgenital plate; K, L, holotype copulatory papilla, dorsal (K), lateral (L) views; M, variation in male forewing dorsal field venation, ♂ ENSIF2104. Scale bars: A-G, J, M, 1 mm; H, I, K, L, 0.2 mm.

FIG. 5. — A-D, Phaloria chopardi (Willemse, 1925) n. comb.; A-C, male genitalia, ♂ MNHN-ENSIF2102, in dorsal (A), ventral (B) and lateral (C) views; D, right pseudepiphallic paramere, ventral view; E, P. nigricollis n. sp., male genitalia, ♂ MNHN-ENSIF2113, right pseudepiphallic paramere, ventral view; F, G, P. nigricollis n. sp. (?), female copulatory papilla in lateral (F) and dorsal (G) views, ♀ MNHN-ENSIF2114. Scale bars: A-C, F, G, 1 mm; D, E, 0.1 mm. Abbreviations: see text.

FIG. 6. — Phaloria chopardi (Willemse, 1925) n. comb., stridulatory file and calling song: A, B, stridulatory file of ♂ MNHN-ENSIF2104 (A), with detailed view of stridulatory teeth (B); C-G, calling song,spectrogram showing two successive echemes (C), oscillograms of a complete echeme (D), of a 105 ms part of the same echeme (F), of one subecheme comprising three syllables (G); E, power spectrum of an echeme. Scale bars: A, 100 μm; B, 10 μm; F, 5 ms; G, 2 ms.