Ming Kai Tan

Sigfrid Ingrisch
email: s.ingrisch@leibniz-zfmk.de

Tony Robillard
email: tony.robillard@mnhn.fr

Jessica B Baroga-Barbecho

Sheryl A Yap
email: sayap3@up.edu.ph

New taxa and notes on spine-headed katydids (Orthoptera: Conocephalinae: Agraeciini) from the Philippines

Keywords: new species, resurrected combinations, Southeast Asia, taxonomy, Tettigonioidea

The diversity of the Agraeciini spine-headed katydids (Orthoptera: Tettigonioidea: Conocephalinae) in Southeast Asia re-mains poorly known, with species still awaiting discovery. Recent field collections in the Philippines resulted in the dis-covery of three novel species and redescription of two known species of Agraeciini, namely, (1) Anthracites furvuseques sp. nov. from Siargao in Mindanao, a species close to A. major Hebard, 1922; (2) Axylus mabinii sp. nov. from Luzon: (3) a new variant in Axylus c.f. philippinus (Hebard, 1922); (4) Salomona lumadae sp. nov. from Siargao in Mindanao; and (5) Salomona brevicollis Stål, 1877 comb. resurr. is a valid species distinct from Salomona maculifrons Stål, 1877.

Introduction

The Agraeciini, or spine-headed katydids, is a tribe of cone-headed katydids (Orthoptera: Tettigonioidea: Conocephalinae). It is a very speciose group, with more than 100 genera distributed worldwide [START_REF] Cigliano | Orthoptera species file online[END_REF]. Despite considerable work on the taxonomy of Agraeciini, many species are still unknown and waiting for discovery. This holds true for the species from Southeast Asia, partly because of fragmentary sampling [START_REF] Ingrisch | Monograph of the Oriental Agraeciini (Insecta: Ensifera: Tettigoniidae): Taxonomic revision, phylogeny, stridulation, and development[END_REF][START_REF] Tan | Trends in new species discovery of Orthoptera (Insecta) from Southeast Asia[END_REF], and owing to these katydids being strictly nocturnal or canopy-dwelling and are often difficult to find [START_REF] Ingrisch | Monograph of the Oriental Agraeciini (Insecta: Ensifera: Tettigoniidae): Taxonomic revision, phylogeny, stridulation, and development[END_REF][START_REF] Ingrisch | New taxa of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) from Singapore and Malaysia with a review of the genus Jambiliara[END_REF][START_REF] Tan | New taxa and notes of some described species of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) from Malay Peninsula[END_REF].

The first major monographic works on the Agraeciini in Southeast Asia was done by [START_REF] Willemse | Notes on the genus Salomona Blanchard (Orthoptera, Tettigonioidea, subfam. Agraecinae)[END_REF] and [START_REF] Ingrisch | Monograph of the Oriental Agraeciini (Insecta: Ensifera: Tettigoniidae): Taxonomic revision, phylogeny, stridulation, and development[END_REF], in which the former focused on the genus Salomona Blanchard, 1853, and the latter revised the taxonomy of 40 genera and 160 species. As many as 70 species new to science were also described [START_REF] Ingrisch | Monograph of the Oriental Agraeciini (Insecta: Ensifera: Tettigoniidae): Taxonomic revision, phylogeny, stridulation, and development[END_REF]. Subsequently, different authors contributed to the discovery of additional new species from all over Southeast Asia (e.g., [START_REF] Gorochov | A new genus for a new species of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae)[END_REF][START_REF] Gorochov | Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 2[END_REF]2015;[START_REF] Gorochov | Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 11[END_REF][START_REF] Naskrecki | Studies in the orthopteran fauna of Melanesia: New katydids of the tribe Agraeciini from Papua New Guinea (Orthoptera: Tettigoniidae: Conocephalinae)[END_REF][START_REF] Ingrisch | New taxa of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) from Singapore and Malaysia with a review of the genus Jambiliara[END_REF][START_REF] Tan | New taxa and notes of some described species of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) from Malay Peninsula[END_REF][START_REF] Tan | A new species of spine-headed katydid Mesagraecia Ingrisch, 1998 (Conocephalinae: Agraeciini), with key to species[END_REF]. Furthermore, Ingrisch's work on Agraeciini in Southeast Asia continued with three more volumes [START_REF] Ingrisch | Revision of the genera Paramacroxiphus C.Willemse 1961 and Pseudomacroxiphus C.Willemse 1961 (Orthoptera: Tettigoniidae: Conocephalinae: Agraeciini). Revision of the Indo-Australian Conocephalinae, part 1[END_REF][START_REF] Ingrisch | Revision of the genus Pseudonicsara Karny, 1912 (Orthoptera: Tettigoniidae: Conocephalinae: Agraeciini). Revision of the Indo-Australian Conocephalinae, part 2[END_REF]2015).

Currently, there are 16 genera known from the Philippines, and at least 29 species described from the Philippines [START_REF] Cigliano | Orthoptera species file online[END_REF]. Nonetheless, recent collections in the Philippines by the authors revealed more new species. More materials also allowed us to better understand species boundaries of previously little-known species. In this paper, we describe two new species from Siargao in Mindanao, namely Anthracites furvuseques sp. nov. and Salomona lumadae sp. nov. Another new species is also described from Laguna in Luzon, namely Axylus mabinii sp. nov. Based on specimens from museum collections, we find that Salomona brevicollis Stål, 1877 comb. resurr. is a valid species distinct from Salomona maculifrons Stål, 1877.

Materials and methods

Night surveys involving opportunistic collection were conducted by M.K. Tan, Huiqing Yeo, J.B. Baroga-Barbecho, and S.A. Yap in the coastal forests (Fig. 1) of in Mindanao between 7 and 9 April 2018 and lowland hill dipterocarp of Mount Makiling (150-250 m.a.s.l.) between 10 and 11 April 2018. Specimens collected from the previous survey conducted by T. Robillard, at Laguna in Luzon Island were also examined. The specimens were preserved in absolute analytical-grade ethanol and later pinned and dry-preserved. A single hind leg from each specimen was also preserved in absolute analytic-grade ethanol for future molecular work. Male genitalia were dissected from softened specimens, and genitalia were removed by squeezing the cerci against one another laterally. For dry-pinned specimens, the specimen was first softened using a mixture of absolute ethanol and diluted KOH solution. The male genitalia were then cleaned using aqueous KOH and subsequently preserved in glycerine.

Habitus images were made with a Canon EOS 500D digital SLR camera with a compact-macro lens EF 100mm f/2.8 Macro USM. Close-up images of morphological features (including male genitalia) were done using a Canon EOS 50D digital SLR camera with a macro photo lens MP-E 65mm f/2.8 USM (1-5×). Canon Macro Twin Lite MT-24EX and Canon Macro Ring Lite MR-14EX were used for lighting and flash. Image editing was accomplished using Adobe Photoshop CC 2014. Scales were given with the images.

Measurements of dry-pinned specimens were made using a 0.05 mm vernier caliper. Detailed measurements of the stridulatory apparatus were done by taking close-up images using a macro photo lens MP-E 65mm f/2.8 USM (1-5×) and measuring in Adobe Photoshop. The following terms are used in the measurements: BL = body length; BWL = body with wing length; PL = pronotum length; PW = pronotum width; TL = tegmen length; HFL = hind femur length; HTL = hind tibia length; OL = ovipositor length. For BL, it refers to the length from the apex of the fastigium to the apex of the epiproct (female) or subgenital plate (male) [START_REF] Naskrecki | Studies in the orthopteran fauna of Melanesia: New katydids of the tribe Agraeciini from Papua New Guinea (Orthoptera: Tettigoniidae: Conocephalinae)[END_REF]. For BWL, it refers to the length from the apex of the fastigium to the apex of the tegmina (for macropterous species only Brief diagnosis. Pronotum lateral lobes strongly descending posteriorly; pronotum ventral margin concave. Mesosternal lobes with medium long apical cone. Hind knee lobes bispinose. Male cercus blunt, with two internal processes near apex. Male titillators separate with lateral extensions in prolongation of tip of titillators or separate with widened base from apical area of titillators. Female subgenital plate often partly or fully divided along midline and lateral surfaces elevated and modified.

Ovipositor falcate [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF].

Remarks. The genus is currently known from the Philippines and Sulawesi. Ten species were described from the Philippines [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF] Hebard, 1922. The taxonomy of the genus has been revised by [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF]. Diagnosis. The new species differs from all known congeners by the male cercus dorso-ventrally compressed but slanting with distal part conical and with two inner processes pointing inwards in distal half, both processes of male cercus stout and triangular with apex slightly curved and sclerotised, and by the shape of the titillators that are stiffened along rim and provided with two pairs of lateral sclerites that are connected by a simple, narrow bridge. A. furvuseques sp. n. strongly resembles Anthracites major Hebard, 1922 from Surigao, Anthracites flagellatus Ingrisch, 2015 from Misamis Orientalis, and Anthracites nitidus Redtenbacher, 1891 from Mindanao by its black body. However, it differs from A. major by: anterior and middle legs black around knees (unicolourous in A. major); posterior femora with black dorsal stripes (absent in A. major); two inner processes of male cercus similar in size and robustness (instead of the ventral one being decidedly heavier); sternites and subgenital plate brown (instead of black). From A. flagellatus it differs by: male tegmen more tint of black and mirror more angular (instead of rounded in A. flagellatus), shape of male cercus and titillators, and subgenital plate with median carina but without dorsad-pointing lateral spine-like processes. From A. nitidus it differs by two inner processes of male cercus similar in size and robustness (instead of the dorsal one being decidedly heavier), and titillators stiffened along rim. Description. Habitus of male as shown in Fig. 2. Fastigium verticis compressed laterally (Fig. 3A), in dorsal view conical with a fine longitudinal groove, lateral ocelli before middle, apex truncated, shorter than scapus (Fig. 3B). Frons shining with few shallowly impressed dots (Fig. 3A). Pronotum smooth to slightly wrinkled, disc broadly rounded into paranota, apical area rounded and faintly shouldered; anterior transverse sulcus short, posterior transverse sulcus longer and angularly rounded; anterior margin truncated but faintly concave in middle; posterior margin subtruncate (Fig. 3B). Paranota with ventral margin concave, posterior margin swollen (Fig. 3C). Acoustic spiracle large and elongated oval, completely covered by paranota (Fig. 3C). Tegmen micropterous, mirror area almost fully covered by pronotal disc (Fig. 3B), reaching only middle of 2 nd abdominal tergite, with rounded apex; hind wing present, but covered by tegmen (Fig. 3C). Prosternum with two long acute spines pointing nearly ventrad. Mesosternal and metasternal lobes conical with acute apices. Tympanum on anterior tibia slightly swollen with a narrow slit. Anterior coxa with a long and curved spine (Fig. 3C). Femora with the following number of spines on ventral margins: anterior femur 3 external, 6 internal; middle femur 5 external, 3 internal; posterior femur 7 external, 2 internal. Knee lobes of anterior and middle femora obtuse externally and spinose internally; knee of posterior femur bispinose on both sides, longer than those of anterior and middle femora.

Anthracites furvuseques

Male. Mirror on left tegmen small, angularly rounded, circa 1.3 times wider than long (Fig. 3D). Stridulatory file on underside of left tegmen straight; about 1.5 mm long; with about 80 teeth fairly regularly spaced throughout (Fig. 3E). Tenth abdominal tergite at posterior margin truncated, emarginated in middle; posterior end in middle depressed; depression and posterior margin with setae (Fig. 3F). Epiproct tongue-shaped. Cercus short and stout, pointing slightly inwards, dorso-laterally compressed; distal part conical with obtuse apex; with two inner processes pointing inwards in distal half; both processes stout and triangular, apex slightly curved and sclerotised; dorsal process slightly longer than ventral (Figs. 3F-H). Subgenital plate longer than wide, with anterior margin concave, gently converging lateral margins, apical margin deeply excised, excision truncated, with a medial carina running from bottom of excision to centre of plate (Fig. 3I). Styli stout and long with apex rounded (Fig. 3I). Titillators (Figs. 3J-M) separate, bases narrow band-shaped and curved (Fig. 3K); stiffened along rim and provided with two pairs of lateral sclerites (Fig. 3J). Female (Fig. 4A). Tenth abdominal tergite short and transverse, deeply and widely emarginated in middle; forming two broadly rounded lobes (Fig. 4B). Epiproct broadly triangular with basal depression and broadly rounded apex (Fig. 4B). Cerci with apex pointing (Fig. 4B). Subgenital plate large, wider than long, roundly emarginated in middle, with two lateral angular lobes (Fig. 4C). Ovipositor long sabre-shaped; margins smooth (Fig. 4D).

Colouration. Body black with pale legs. Face black, median ocellus yellow; scapus black ventrally and on inner side, pale dorsally and on outer side; antennae black; maxillary palpi with basal three segments pale, apical and subapical segments black; mandible black, clypeus pink-orange, labrum yellow-orange (Figs. 3A-C). Pronotum completely black (Figs. 3B,3C). Tegmen with lateral field bright yellow with irregular black spots, spots smaller closer to costal (anterior) margin; dorsal field mostly black, posterior of mirror also bright yellow with irregular black spots (Figs. 3D,4E). Spines on anterior coxa black (Fig. 3C). Anterior and middle femora black at base and at knee, otherwise pale; anterior and middle tibiae pale with base and apex black; fore and middle tarsi black. Posterior femur pale laterally, with a black longitudinal dorsal stripe that extends from the base to nearly the knee; knees, including lobes black. Posterior tibia black near knee, otherwise pale with black dorsal spines. Posterior tarsus black. Abdominal tergite black, with anterior margin pink when alive, with two faint orange spots in middle of posterior margin. Male tenth abdominal tergite and epiproct also black; brown setae on tenth abdominal tergite (Fig. 3F). Cercus black with brown setae, apices of internal processes brown (Fig. 3H). Sternites and subgenital plate brown. Ovipositor with dorsal valve mostly black, red brown apically, ventral valve mostly red brown with tint of black at base.

Measurements (^ indicates measurement for male holotype). BL: male 26.0-29.3^, female 30.7; PL: male 9.4^-9.8, female 9.2-9.4; PW: male 6.2^-6.5, female 6.9-7.0; TL: male 5.4-6.0^, female 5.4-5.8; HFL: male 21.8-22.1^, female 22.3-23.2; HTL: male 22.0^-22.3, female 21.7-23.2; OL: female 16.3-16.5 mm.

Etymology. The species name refers to the Dark Knight (in Latin; dark = furvus, knight = eques). This species is named after the fictional superhero character Batman in the Dark Knight Trilogy directed by Christopher Nolan. The species' black colouration resembles the dark suit of the vigilante while the titillators also shows certain resemblance to the iconic Batman mask (Fig. 3J) and logo (Fig. 3M) in the film.

Natural history. The new species is a common Agraeciini species found in the coastal vegetation on the island (Fig. 1) at night. The species was observed to feed on small snails (Fig. 5A) and possibly seed pods of ground orchids (Spathoglottis sp.) (Fig. 5B).

Genus Axylus Stål, 1877

Type species: Axylus castaneus Stål, 1877 Brief diagnosis. Mesosternal lobes with medium long to long apical cone. Hind knee lobes bispinose. Male cercus conical or laterally compressed with a proximal and a distal internal process. Male titillators separate with lateral extensions branching from apical area; with a pair of lateral sclerites. Ovipositor elongate and little curved or falcate and highest just behind mid length [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF].

Remarks. The genus is currently known from east of the Wallace Line (sensu Huxley) which includes the Philippines. Seven species were described from the Philippines [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF] (Hebard, 1922); Axylus taylori (Hebard, 1922). The taxonomy of the genus has been revised by [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF].

Axylus mabinii Tan & Robillard new species (Figs. 6,7) Material examined. Holotype (male, TR347): Philippines, Luzon, Laguna, UPLB University Land Grant, secondary forest at night, N14°23'56.9" E121°32'47.2", 376 m.a.s.l., 4 July 2011, coll. T. Robillard (UPLBMNH). Allotype: 1 female (TR348), same locality as holotype, mating with holotype on palm tree, coll. T. Robillard (MNHN-EO-ENSIF4390).

Paratypes: 1 female (TR398) and 1 male (TR399), Philippines, Luzon, Laguna, UPLB University Land Grant, herbaceous area near station, N14° 24' 02.5" E121° 32' 30.3", 327 m.a.s.l., mating, 5 July 2011, coll. T. Robillard (MNHN-EO-ENSIF4391-43920).

Diagnosis. The new species differs from all known congeners by black patterns on the pronotum: two black spots on the anterior angle of disc and two larger black lateral bands on the posterior area of disc, and by the male cercus that is very stout and conical, laterally compressed with obtuse apex and provided with a broad baso-internal process and a slender ventral process. The new species is most similar to Axylus taylori (Hebard, 1922) from Luzon and Axylus nigrifrons (Brunner v. Wattenwyl, 1898) from Lombok in the shape of the female ovipositor and in the colour patterns on frons and pronotum, but differs by shape and processes of the male cercus and the titillators. Also the colour pattern on pronotal disc differs from that of A. taylori by the anterior and posterior pairs of spots being not continuous and from that of the A. nigrifrons by additional posterior pairs of black spots. The shape of the female ovipositor is also similar to that of Axylus inferior (Brunner v. Wattenwyl, 1898) from Sulawesi but A. mabinii sp. n. differs distinctly by the male cercus and by colour pattern. Description. Habitus of male as shown in Fig. 6A. Fastigium verticis compressed laterally (Fig. 7A), in dorsal view conical, shorter than scapus, apex obtuse, with ocelli at base (Fig. 7B); ventral area compressed and ventral margin separated by a weak sinuosity from fastigium frontis (Fig. 7A). Vertex with a pair of irregular sulci (Fig. 7B). Frons shining with few shallowly impressed dots and wrinkles especially towards clypeus. Pronotum subsmooth, disc rounded into paranota, apical area subflat and shouldered; two transverse sulci not interrupted in middle; anterior margin faintly emarginated in middle; posterior margin rounded (Fig. 7B). Paranota strongly descending posteriorly (typical of genus), with anterior angle rounded; ventral margin concave and considerable descending posteriorly; posterior angle angularly rounded; auditory swelling long-ovoid; humeral sinus weak (Fig. 7C). Macropterous, tegmen surpassing abdominal apex and slightly longer than hind wing, apex rounded. Prosternum with two long acute spines. Mesosternal lobes long-conical, apex subacute; metasternal lobes short conical, apex obtuse; medial plate with an obtuse tubercle. Anterior coxa with a long and curved spine (Fig. 7C). Internal and external tympana on anterior tibia conchate with narrow openings. Femora with the following number of spines on ventral margins: anterior femur 7 external, 7 internal; middle femur 6 external, 3 small internal at the base; posterior femur 10-13 external, 7 internal (2 distal spinose and 5 basal minute nodular). Knee lobes of anterior and middle femora obtuse externally and short spinose internally; of posterior femur bispinose on both sides. Tibiae with the following number of spines on ventral margins: anterior tibia 7 external, 7-8 internal; middle tibia 9 external, 8 internal. Male. Stridulatory file on underside of left tegmen slightly sinuous; about 1.55 mm long; with teeth increasingly wider and more widely spaced towards middle compared to basal end; with about 64 teeth (Fig. 7D). Tenth abdominal tergite at posterior margin truncated, faintly and broadly emarginated in middle, forming faint lateral lobes (Figs. 7E,7F). Epiproct broadly tongue-shaped, concealed by tenth abdominal tergite (Fig. 7F). Cerci very stout and conical (Fig. 7G), laterally compressed; apex obtuse; with a broad baso-internal process pointing anteriorly and internally; ventral of this process with a slender process pointing internally; both processes with apex sclerotised and acute (Figs. 7E,7F,7H). Subgenital plate barely surpassing cerci, with rounded lateral carinae and a narrow but distinct median carina; apex widely excised; styli short with apex obtuse. Titillators large, elongate, separate; basally with rounded apex; converging in middle and nearly parallel to one another before diverging apically; slender posteriorly and produced into an acute apex, curved dorsad just before apex (Fig. 7I-K). Dorsal sclerites slender and curved, with posterior margin undulated (Fig. 7I). Lateral sclerites semi-membranous, with a rounded nodular sclerite (Fig. 7K).

Female (Fig. 6B). Tenth abdominal tergite excised in midline, apical margin on each side truncate. Epiproct broadly rounded. Cerci with apex pointing. Subgenital plate small, broad, triangular, with apex rounded. Ovipositor compressed, elongofalcate but basal area substraight; dorsal margin substraight in middle, only slightly curved apically; ventral margin curved after basal area; widest slightly after middle, margins smooth (Fig. 7L).

Colouration. Yellow brown. Face black, fastigium verticis yellow brown, median ocellus yellow, genae yellow brown; scapus and antennae yellow brown; maxillary palpi yellow brown, clypeus and labrum orange, mandible black (Fig. 7A). Pronotum yellow brown, with two black spots at anterior angles of disc and two larger black lateral bands in posterior area of disc; all black spots slightly continue onto paranota (Figs. 7B,7C). Tegmen yellow brown (Fig. 6). Legs mostly yellow brown (Fig. 6). Abdominal tergites and sternites yellow brown. Ovipositor ventro-basally yellow, otherwise red brown (Fig. 7L).

Measurements (1 male holotype, 1 female paratype). BL: male 34.5, female 34.6; BWL: male 54.9, female 56.3; PL: male 11.7, female 11.0; PW: male 7.5, female 8.8; TL: male 42.8, female 44.8; HFL: male 28.0, female 28.8; HTL: male 27.1, female 28.2; OL: female 21.1 mm.

Etymology. This species is named after Apolinario Mabini y Maranan (1864-1903), a Filipino revolutionary leader and hero who confronted both the Spanish and American colonial rules.

Axylus c.f. philippinus (Hebard, 1922) (Figs. 8, 9) Material examined. 1 male (TR349) and 1 female (TR350): Philippines, Luzon, Laguna, UPLB University Land Grant, 4.vii.2011, secondary forest at night, N14°23'56.9" E121°32'47.2", 376 m.a.s.l., coll. T. Robillard (MNHN). Remarks. Our specimens resemble Axylus philippinus in habitus (Figs. 8, 9A, 9B), shape of the male cercus (Figs. 9E,9F) and the female ovipositor (Fig. 9K) but differ as follows: frons red brown, of same colour as genae (Figs. 9C, 9D), instead of black distinctly different from genae; male subgenital plate angularly and deeply excised (Fig. 9G) instead of roundly and less deeply excised; female subgenital plate not regularly curved from one side to the other (Fig. 9J). Since the titillators of congeners of Axylus can differ drastically in shape, the similarities of the titillators between that of our specimens (Figs. 9H,9I) and those examined in [START_REF] Ingrisch | A revision of the Axylus group of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) and of some other species formerly included in Nicsara or Anthracites Revision of the Indo-Australian Conocephalinae, part 3[END_REF] suggest that despite the differences in other morphology, our specimens from Laguna, UPLB University Land Grant are probably a variant or subspecies of Axylus phillipinus. To test the species boundaries, we reckon that incorporation of molecular data could be useful.

Genus Salomona Blanchard, 1853

Type species: Salomona marmorata (Blanchard, 1853) Brief diagnosis. Large, robust; large, flat, and broad frons, often rugose or punctate; short fastigium of vertex; male subgenital plate with long styli; falcate or subfalcate ovipositor [START_REF] Naskrecki | Studies in the orthopteran fauna of Melanesia: New katydids of the tribe Agraeciini from Papua New Guinea (Orthoptera: Tettigoniidae: Conocephalinae)[END_REF].

Remarks. The genus is relatively widespread and can be found in Southeast Asia, Australia, and western Pacific islands. Four species were described from the Philippines: Salomona conspersa Stål, 1877; Salomona maculifrons Stål, 1877; Salomona tetra Walker, 1869; Salomona vittifrons Walker, 1869. The taxonomy of the genus has been revised by [START_REF] Willemse | Notes on the genus Salomona Blanchard (Orthoptera, Tettigonioidea, subfam. Agraecinae)[END_REF]. Diagnosis. This new species differs from all known congeners by colour patterns of the tegmen and by the shape of the male cercus that is provided with a stout, bulbous, baso-internal process that bears a smaller anterior bulbous process. The presence of a baso-internal process on the male cerci similar to that of the new species occurs in only a few other species from Papua New Guinea: Salomona godeffroyi (Pictet, 1888) from which the new species differs by the frons that is rugose instead of subsmooth and the male cerci are not curved internally near apex; from Salomona karnya [START_REF] Willemse | Notes on the genus Salomona Blanchard (Orthoptera, Tettigonioidea, subfam. Agraecinae)[END_REF] it differs by the baso-internal process of the cercus being not elongated; from Salomona limbata Karny, 1907 it differs by the tenth abdominal tergite that does not have a long apical process in the middle; from Salomona megacephala (Haan, 1843), which is also similar by the presence of small lateral depressions on the tenth abdominal tergite, it differs by the absence of a distal internal process at the cerci; from Salomona solida (Walker, 1869) it differs by the frons that is not red and by the baso-internal process of the cerci that are more rounded instead of cubical and have only one anterior bulbous process instead of two; and from Salomona ustulata Redtenbacher, 1891 it differs by the rugose and yellow instead of smooth and red frons and the cerci are stouter and conical instead of elongate. Description. Male habitus as shown in Fig. 10. Fastigium verticis in dorsal view elongated conical, apex truncate, about as long as scapus, with ocellus in middle (Fig. 11A). Vertex subrugose (Fig. 11A). Frons with large impressed dots, genae rugose (Fig. 11B). Pronotum rugose with impressed dots, disc broadly rounded into paranota, apical area subflat and shouldered (Fig. 11C); anterior margin straight but faintly concave in middle; posterior margin broadly rounded (Fig. 11A). Acoustic spiracle elongated, taller than long, completely covered by paranota. Tegmen macropterous, barely reaching abdominal apex and just as long as hind wing, apex rounded. Prosternum with two stout spines with subobtuse apices. Mesosternal and metasternal lobes conical with blunt apices. Femora with the following number of robust spines on ventral margins: anterior femur 8 external, 9 internal of increasing size distally; middle femur 7 external of increasing size distally, 3 internal at base; posterior femur 12 external of increasing size distally, 0 internal. Knee lobes of anterior and middle femora obtuse externally and long spinose internally; of posterior femur spinose on both sides. Tibiae with the following number of spines on ventral margins: anterior tibia 6 external, 6 internal; middle tibia 10 external, 8 internal.

Male. Stridulatory file on underside of left tegmen straight; about 1.89 mm long; with teeth increasingly wider and more widely spaced towards middle (compared to basal end); with circa 70 teeth (Fig. 11D). Tenth abdominal tergite depressed in middle, with one oval depression near middle on each side of depression; posterior margin prolonged into two lateral lobes with deep rounded excision in middle; posterior lobes triangular with obtuse apex (Fig. 11E). Epiproct broadly tongue-shaped, depressed in middle (Fig. 11F). Cerci conical, slightly compressed towards lateral area, internal margin straight with surface depressed, external margin curved, apex subacute; with a stout baso-internal process which is bulbous and bearing a smaller anterior bulbous process (Figs. 11E-H). Subgenital plate wider than long, apical margin deeply, narrowly and roundly excised; styli conical and very short (Fig. 11I).

Female. Unknown. Colouration. Pale brown when alive (Fig. 10), yellow brown when preserved in ethanol. Face pale brown, fastigium verticis with a black spot ventrally, antennal socket ventrally with two black spots; scapus pale brown, pedicel pale brown with a ventral black spot, antennae yellow brown; maxillary palpus, clypeus and labrum orange, mandible black (Fig. 11B). Pronotum pale brown (Fig. 11C). Tegmen generally with cells black, most obvious in costal area (anterior area) (Fig. 11J); veins in costal area turquoise, otherwise red brown (Fig. 10). Femora and tibiae pale brown and brown respectively, with black ventral spines; tibiae with two dorsal black spots at base and before first ventral spine; stout spine of femoral knee lobe also black. Hind femur basally green, medially yellow green, distally green, knee dark red brown. Abdominal tergites pale brown; sternites very pale brown. Abdominal apex pale brown (Figs. 11E-I). Measurements (holotype). BL 37.6;BWL 41.0;PL 10.3;PW 6.5;TL 28.8;HFL 17.9;HTL 17.5 mm. Etymology. This species is named after the indigenous people in the southern Philippines, including Siargao Island; lumad means indigenous in Cebuano, an Austronesian language spoken in the Philippines. This species is dedicated to the Lumads.

Natural history. The new species can be found among coastal vegetation.

Salomona brevicollis Stål, 1877 comb. resurr.

Material examined. Images of type specimen (female, NRM-ORTH0001233) in Cigliano et al., 2018 (Orthoptera Species File Online).

Salomona maculifrons Stål, 1877 (Fig. 12) Remarks. Salomona brevicollis was described based on a female but was later synonymised under Salomona maculifrons by [START_REF] Willemse | Notes on the genus Salomona Blanchard (Orthoptera, Tettigonioidea, subfam. Agraecinae)[END_REF]. After examining both male (Fig. 12A) and female specimens (Fig. 12B) of S. maculifrons from UPLBMNH, we found that the female holotype of S. brevicollis differs from that of S. maculifrons by the black pattern on the frons that is more elaborated instead of resticted to a single oval spot in the latter [Fig. 12C]), the fastigium verticis is shorter and stouter instead of long, conical and with acute apex [Fig. 12C]), the female ovipositor appears wider (higher) in subapical than in subbasal area (Fig. 12D). The colour pattern of the frons on S. maculifrons does not vary drastically between specimens, indicating that the different colour pattern in the female holotype of S. brevicollis is a genuine species differences, rather than a conspecific variation.

Material

FIGURE 1 .

 1 FIGURE 1. Environment of the coastal forest in Siargao Island where the Agraeciini were collected.

FIGURE 2 .

 2 FIGURE 2. Anthracites furvuseques sp. nov. live male habitus in dorsal view. The male was lying on Pandanus foliage.

FIGURE 3 .

 3 FIGURE 3. Anthracites furvuseques sp. nov. male: face in anterior view (A); head and pronotum in dorsal (B) and lateral (C) views; left tegmen in ventral view (D); stridulatory file on left tegmen in ventral view (E); abdominal apex in dorsal (F) and posterior (G) views; cercus in dorsal view (H); subgenital plate in ventral view (I); titillators (J-M). Scale bars: 5 mm (B, C), 2 mm (F, G, I), 1 mm (A, D, H).

FIGURE 4 .

 4 FIGURE 4. Anthracites furvuseques sp. nov. female: habitus in lateral view (A); abdominal apex in dorso-posterior (B), ventral (C), and lateral (D) views; tegmina in dorsal view (E). Scale bars: 10 mm (A), 5 mm (D), 2 mm (B, C, E).

FIGURE 5 .

 5 FIGURE 5. Anthracites furvuseques sp. nov. juveniles feeding on (A) small snail and (B) seed pod of Spathoglottis ground orchid.

FIGURE 6 .

 6 FIGURE 6. Axylus mabinii sp. nov. male (A) and female (B) habitus in lateral view. Scale bars: 10 mm.

FIGURE 7 .

 7 FIGURE 7. Axylus mabinii sp. nov.: face in anterior view (A); head and pronotum in dorsal (B) and lateral (C) views; stridulatory file of left tegmen in ventral view (D); male abdominal apex in dorsal (E), dorso-posterior (F), lateral (G), and posterior (H) views; titillators in posterior (I), ventral (J) and dorsal (K) views. Scale bars: 5 mm (B, C, L); 2 mm (A, E-H); 1 mm (D, I-K).

FIGURE 8 .

 8 FIGURE 8. Axylus philippinus male (A) and female (B) habitus in lateral view. Scale bars: 10 mm.

FIGURE 10 .

 10 FIGURE 10. Salomona lumadae sp. nov. live male habitus on a branch.

FIGURE 11 .

 11 FIGURE 11. Salomona lumadae sp. nov. male: head and pronotum in dorsal (A) and lateral (C) views; face in anterior view (B); stridulatory file of left tegmen in ventral view (D); abdominal apex in dorsal (E), dorso-posterior (F, G), and ventral (H, I) views; tegmen in lateral view (J). Scale bars: 5 mm (A, C, J), 2 mm (B, E-I), 1 mm (D).

FIGURE 12 .

 12 FIGURE 12. Salomona maculifrons male (A) and female (B) habitus in lateral view; face of female in anterior view (C); female abdominal apex in lateral (D) view; female subgenital plate in ventral view (E). Scale bars: 5 mm (A, B), 2 mm (C, D), 1 mm (E).

  ).

	Depositories
	UPLBMNH	Museum of Natural History, University of the Philippines Los Baños, Philippines
	ZRC	Zoological Reference Collection, Lee Kong Chian Natural History Museum (former Raffles Museum of
		Biodiversity Research), National University of Singapore
	MNHN	Muséum national d'Histoire naturelle, Paris, France.
	Systematics part Orthoptera:
	Tettigonioidea Subfamily
	Conocephalinae Tribe Agraeciini
	Genus Anthracites Redtenbacher, 1891

Type species: Anthracites nitidus Redtenbacher, 1891

:

  Axylus brachypterus Ingrisch, 2015; Axylus castaneus Stål, 1877; Axylus furcatus Ingrisch, 2015; Axylus montanus Ingrisch, 2015; Axylus negros Ingrisch, 2015; Axylus philippinus

  examined. 1 female, Philippines, Luzon, Los Baños, Mount Makiling, October 1974, coll. A.A. Barroso; 1 male, Philippines, Luzon, Los Baños, Puting Lupa, 2 June 1983, coll. B.M.A. Laigao; 1 female, Philippines, Luzon, Los Baños, Soledad, February 1992, coll. R. Rubles (all UPLBMNH).

Acknowledgements

The authors are thankful to Huiqing Yeo for field assistance and for the help in the sorting and examination of specimens. We are also grateful to the UPLB Museum of Natural History for allowing us access their orthopteroid collections, especially for the assistance of Mr. Orlando L. Eusebio and Mr. Cristian C. Lucañas during the visit of MKT at UPLB. The authors would also like to thank Siargao Islands Wildlife Conservation Foundation, Inc., Mr. Jose Macavinta, Ms. Deny Comon, MODECERA project, Mayor Alfredo Coro, Jr., and the municipality of Del Carmen for the accommodation and allowing us to conduct research on their area. The work in the Philippines was also granted by the Orthoptera Species File Grant 2018. The work of MKT was also supported by the Lady Yuen Peng McNeice Graduate Fellowship of the National University of Singapore.


Ranjana Jaiswara
email: ranjana.jaiswara@gmail.com

Jiajia Dong
email: jiajia.dong@mnhn.fr

Tony Robillard
email: tony.robillard@mnhn.fr

Revision of the genus Pseudolebinthus (Orthoptera, Gryllidae, Eneopterinae) with the description of a new species from Southeast Africa

Keywords: Xenogryllini, taxonomy, endemic distribution, Xenogryllus, Afrotropical region

 P. whellani Robillard, 2006. The genus was initially diagnosed based exclusively on male morpho-logical features. In this study, we add a new species to the genus, P. gorochovi Robillard sp. nov. and revise the diagnostic features using both male and female characters. We also update identification keys for the species of the genus.

Introduction

Subfamily Eneopterinae is known for its disjuct distribution [START_REF] Vicente | In and out of the Neotropics: historical biogeography of Eneopterinae crickets[END_REF]; its members are either endemic to a geographical region or islands [START_REF] Nattier | Secondary Sympatry Caused by Range Expansion Informs on the Dynamics of Microendemism in a Biodiversity Hotspot[END_REF][START_REF] Dong | Biogeographic patterns and diversification dynamics of the genus Cardiodactylus Saussure (Orthoptera, Grylloidea, Eneopterinae) in Southeast Asia[END_REF] or widely distributed across continents [START_REF] Robillard | Redescription of two Cardiodactylus species (Orthoptera, Grylloidea, Eneopterinae): the supposedly well-known C. novaeguineae (Haan, 1842), and the semi-forgotten C. guttulus (Matsumura, 1913) from Japan[END_REF]Jaiswara et al. in press). Genus Pseudolebinthus [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF] is one of the endemic taxa, found only from few places in the southeast of Africa.

Pseudolebinthus is an intriguing genus known from very few specimens and characterized by unique morphological features such as asymmetrical male forewings and harp veins shaped as elongated balloons. Phylogenetic studies based on morphological [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF]) and molecular markers [START_REF] Vicente | In and out of the Neotropics: historical biogeography of Eneopterinae crickets[END_REF]) revealed that Pseudolebinthus is sister to the widely distributed genus Xenogryllus. Until now, the genus included two species, P. africanus [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF]P. whellani Robillard, 2006, known only by males.

In this study, we carry out a taxonomic revision of Pseudolebinthus and enrich the species list by describing a new species, Pseudolebinthus gorochovi Robillard sp. nov. from southeast Africa. In addition to male characters, female sex is described, including external and genitalic characters to append the diagnostic features of the genus.

Material and methods

Material. The material examined comes from the following natural history collections: Museum national d'Histoire naturelle, Paris, France (MNHN); Natural History Museum, London, United Kingdom (NHMUK); Naturhistorisches Museum, Vienna, Austria (NHMW); Zoological Institute, Russian Academy of Sciences, S. Petersburg, Russia (ZIN).

In lists of materials, square brackets are used for additional information not mentioned on specimen labels or for translation of key information present on labels in languages other than English.

Taxonomic description. Description of species follows terminologies as proposed by [START_REF] Robillard | Revision of the cricket genus Cardiodactylus (Orthoptera, Eneopterinae, Lebinthini): the species from both sides of the Wallace line, with description of 25 new species[END_REF]. Observations of external morphological characters and dissection of male and female genitalia were performed using Leica stereomicroscopes MZ16. Terminologies for male FW venation follow Ragge (1995) and [START_REF] Robillard | Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae)[END_REF]. Male and female genitalia were dissected either from dry preserved or freshly killed specimens. Male genitalia were dissected by making a small slit between paraproct and subgenital plate. Female copulatory papilla was dissected by cutting the membrane between ovipositor and subgenital plate. Dissected genitalia were cleared in 10% cold KOH solution and preserved in glass vials containing glycerine. Terminologies for genitalia follow [START_REF] Org> Desutter | Structure et évolution du complexe phallique des Gryllidea (Orthoptera) et classification des genres néotropicaux de Grylloidea. 1re partie[END_REF], modified in [START_REF] Desutter-Grandcolas | Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera)[END_REF] and [START_REF] Robillard | Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae)[END_REF]. Imaging of male and female genitalia were made using an AmScope MU1000 digital camera (www.Amscope.com) or with a Canon EOS 40D Digital SLR camera. To highlight the structural components of male and female genitalia, water solution containing a drop of JBL Punktol was used. To fix orientations and stabilization of genitalia for photography, a clear and viscous Power Plast Hand Sanitizer was used following [START_REF] Su | A simple and quick method of displaying liquid-preserved morphological structures for microphotography[END_REF].

Abbreviations used in taxonomic descriptions. General morphology: FI, FII, FIII, fore, median, hind femur; FW, forewing; TI, TII, TIII, fore, median, hind tibia; Tarsomere I/II/III-1: basal segment of fore, median and hind leg tarsomere.

Tegminal venation: 1A-4A, first to fourth anal veins; CuA, anterior cubitus; CuA1, CuA2, ... first, second, ... branches of CuA; CuP, posterior cubitus; M, median vein; R, radial vein; Sc, subcostal vein and its branches; c1-3, first to third cells of C alignment; d1 cell (mirror), first cell(s) of D alignment; d2, second cell of D alignment; e1, first cell of E alignment.

Species abbreviations used in figures: Paf, Pseudolebinthus africanus; Pwh, Pseudolebinthus whellani; Pgo, Pseudolebinthus gorochovi sp. nov.

Systematic part

Orthoptera

Gryllidae

Eneopterinae Saussure, 1874 Xenogryllini Robillard, 2004 Genus Pseudolebinthus [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF] Type species: Pseudolebinthus africanus [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF]Pseudolebinthus Robillard, 2006: 672-Robillard & Desutter-Grandcolas 2008: 67;[START_REF] Vicente | In and out of the Neotropics: historical biogeography of Eneopterinae crickets[END_REF]Vicente et al. : 2202 (historical biogeography) (historical biogeography); Cigliano et al. 2018 (Orthoptera Species File Online).

Emended diagnosis. Genus with species of small size, differing from Xenogryllus by shorter FWs, long and slender abdomen, male FWs asymmetrical, venation with unique swollen oblique veins. Eyes larger than in Xenogryllus, located laterally. Fastigium forming a wide rectangular rostrum, little prominent on face. Face flat when viewed laterally. Pronotum dorsal disc almost rectangular, its lateral margins not carinated, its posterior margin straight. TII with one inner apical spur. FIII wide basally, shorter and more muscular than in Xenogryllus. Male. Metanotal glandular structures well developed, as in Xenogryllus. FWs short, not reaching mid-length of abdomen and not widened basally as in Xenogryllus. FW cells smooth, without longitudinal wrinkles on its surface. Left and right FWs asymmetrical, left darker and more sclerotized than right, thin and transparent. File vein with a rounded angle. Harp with characteristic swollen oblique veins (shaped as elongated balloons) and without incomplete oblique vein between diagonal and first complete oblique (incomplete veins present in Xenogryllus); harp distal angle flat. CuA vein at the bottom of a deep longitudinal furrow. Mirror small, wider than long, not rounded. Cell d2 not differentiated. Apical field very short, made only of few cells of E alignment. Hind wings present but shorter than FWs. Male genitalia. Pseudepiphallic sclerite shorter than rami; lateral membranous lobes present but narrow; basal margin straight and thickened. Pseudepipallic lophi parallel, well sclerotized dorsally, membranous ventrally; lophi shrunk at mid-length, their basis twice as wide as apical part, their apex usually bilobate, the dorsal lobe ended by a small dorsal expansion. Pseudepiphallic parameres with wide and short conical ornamentation, trilobate, the posterior lobes forming a hook oriented posteriorly. Ectophallic apodemes long, forming wide triangular lamellas. Ectophallic arc faint, slightly sclerotized medially. Ectophallic fold short and triangular, reaching basis of pseudepiphallic parameres; with two lateral sclerites; not fused to endophallic sclerite as in Xenogryllus. Endophallic sclerite wide and flat, trifid posteriorly, its median part triangular and short, lateral parts long, reaching basis of ectophallic apodemes. Endophallic apodeme comprising mostly two wide lateral lamellas.

Female. FWs very short, forming lateral triangles not reaching posterior margin of first tergite. Ovipositor apex almost not differentiated, long and lanceolate. Female copulatory papilla triangular, its apex very thin and pointed.

Additional description. In addition to the characters described by [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF], the female sex is described here for the first time.

Female. General shape similar to male (Fig. 1C-D). FWs very short (Fig. 3B), shorter than in males, forming small lateral sclerotized triangles not reaching posterior margin of first tergite; with strong longitudinal veins and faint transverse ones. Abdomen very long and thin. Subgenital plate with a weak apical indentation (Fig. 6B). Ovipositor flattened laterally, apex almost not differentiated, long and lanceolate (Fig. 6A).

Female copulatory papilla (Fig. 6C-E). Triangular, its apex very thin and pointed; sclerotization strong except apex, membranous.

Distribution. Eastern Sub-Saharan Africa Key to known species of Pseudolebinthus Male. Scutellum basal edge sharply raised. FW dorsal field with strong transverse veins. Mirror small, hardly distinct from surrounding cells and crossed by one strong accessory vein; c1 twice as long as c2. Sc with three projections along its length (four in P. whellani, two in P. gorochovi sp. nov.). Male genitalia. Pseudepiphallic lophi (Fig. 5C) longer and thinner than in P. whellani, their apical lobes almost completely fused, as in P. gorochovi sp. nov. Median part of endophallic sclerite well developed.

Pseudolebinthus whellani Robillard, 2006 (Figs 2D-F; 5B)

Pseudolebinthus whellani [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF]: 679-Cigliano et al. (2018) (Orthoptera Species File Online).

Material examined. Holotype. MALAWI: ♂, Zomba Mountain, 28.ii.1964, Min. Agr. Rhod., ex-Nyas collection, Salisbury, S.R., accession no. 11775 (Whellan), identified Lebinthus whellani by L. Chopard (NHMUK).

Type locality. Malawi, Zomba Mountain. Distribution. Southern Malawi.

Diagnosis. Emended from [START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF]. Brown coloration little contrasted; FIII uniformly brown. Dorsal margin of eyes with small ommatidia wide.

Male. Basal edge of scutellum slightly raised. FW dorsal field with faint transverse veins. Mirror larger than in P. africanus, well differentiated and crossed by an incomplete accessory vein. c1 cell as long as c2. Vein Sc with four projections along its length (three in P. africanus, two in P. gorochovi). Male genitalia. Pseudepiphallic lophi (Fig. 5B) shorter and wider than in P. africanus, with two apical lobes well developed, the dorsal one oriented posteriorly, the ventral one oriented inward. Male. FW dorsal field with faint transverse veins; cell c1 wide; c2 square, twice wider than c1, prolonging shape of mirror (c1 and c2 of similar width and smaller in P. whellani). Sc with two projections at one third of its length and near apex (three in P. africanus, four in P. whellani). Male genitalia with very short pseudepiphallic lophi, twisted ventro-apically as in P. africanus; their apex not bilobate (Fig. 5A), as in P. africanus (apex bilobate in P. whellani).

Pseudolebinthus gorochovi

Description. Species of average size for the genus, coloration contrasted, mostly orange brown with dark brown, black and yellow patterns (Fig. 1A-D). Eyes large, lateral, occupying almost half of head height in lateral view (Fig. 2A &C); dorsal margin of eyes with small ommatidia wide. Vertex with four wide longitudinal dark brown bands fused on dorsum (bands visible basally only); area posterior to eyes yellow, without brown bands. Fastigium as long as wide (Fig. 2B), slightly thinner apically than in P. africanus (as in P. whellani), dark brown with a yellow transverse stripe apically. Median ocellus surrounded by a black mark posteriorly. Face almost homogenously brown with faint pale spots; frons with a yellow transverse band below median ocellus, underlined by two back spots (forming a black transverse band in female allotype). Mouthparts yellow, mottled with dark brown; maxillary palpi yellow with dark rings. Lateral part of head dark brown with a median yellow spot; posterior margin of head and area below eye yellow. Scape yellow brown with black patterns on facial part. Antennae with first article black, remainder orange brown. Pronotum disc mostly dark brown, with a darker spot near posterior margin; lateral lobes black with silver setae, with a yellow spot on ventro-anterior corner; posterior corner yellow mottled with black. Fore and median legs mostly yellow brown; apex of tibiae and tarsomeres black. FIII external face bicolor, its dorsal half dark brown, its ventral half yellow.

Male. FWs brown, with dark brown veins (Fig. 3A). 1A angle forming a wide curve. CuP visible in anterior region of harp. Harp very wide, with two characteristic black swollen oblique veins. Mirror as in P. whellani, well differentiated but not rounded, wider than long, crossed by incomplete accessory veins. Cell c1 wide basally, thinner apically; c2 large and square, twice wider than c1, prolonging shape of mirror. Chord cells straight. Apical field with faint transverse veins. M/R area and veins whitish. R/Sc area whitish dorsally (including R), black ventrally (including Sc). Sc with two projections at one third of its length. Lateral field with five longitudinal veins ventral to Sc.

Male genitalia (Fig. 4A-C). Pseudephiphallic sclerite wider than long (not including rami); pseudepiphallic lophi very short (Fig. 5A), divergent basally and slightly convergent apically, their median part twisted ventro-apically; apex with a very small dorsal expansion. Pseudepiphallic parameres typical of the genus, its posterior lobes short. Ectophallic apodemes large and lamellate. Endophallic sclerite with a small median area.

Female. FWs with five strong longitudinal veins (Fig. 3B), the median one whitish. Ovipositor as long as FIII.

Measurements (in mm). See Table 1.

Discussion

Despite extensive studies based on large collections to revise the Xenogryllini genera (Jaiswaira et al. in press), very little information is available concerning Pseudolebinthus. This genus is distributed only in the South Eastern part of Africa, in Malawi, Tanzania and Zimbabwe, and the three documented species show a high endemicity. Although no ecological data is available about any of the species, the few localities from where they are reported correspond to low altitude mountains covered with forests. We can thus hypothesise that the species have a forested habitat similar to that of most brachypterous Lebinthini [START_REF] Robillard | The brachypterous Lebinthini crickets from Papua New Guinea, with description of two new genera and four new species (Orthoptera: Gryllidae: Eneopterinae)[END_REF]. Such habitats could in turn be responsible for more restricted distributions than for species of Xenogryllus, explaining why so few specimens are documented.

Given the morphological and distributional particularities of Pseudolebinthus, it is necessary to obtain additional information about the biology, ecology and bioacoustics of these remarkable species. Such information is mandatory to pursue the exploration of the systems of communication and their evolution, investigated at the scale of the subfamily Eneopterinae (e.g., Robillard & Desutter-Grancolas 2004;[START_REF] Ter Hofstede | Evolution of a communication system by sensory exploitation of startle behavior[END_REF].

FIGURE 1 .

 1 FIGURE 1. (A-D) Pseudolebinthus gorochovi sp. nov.: male in dorsal (A) and lateral (B) views, female in dorsal (C) and lateral (D) views. Scale bar: 5 mm.

FIGURE 2 .

 2 FIGURE 2. Head in dorsal (left), facial (middle) and lateral views (right): (A-C) Pseudolebinthus gorochovi sp. nov. (D-F) Pseudolebinthus whellani; Scale bar: 1 mm.

FIGURE 4 .

 4 FIGURE 4. Male genitalia of Pseudolebinthus gorochovi sp. nov. in dorsal (A), ventral (B) and (C) lateral views.

1 .

 1 Dorsal margin of eye with small ommatidia thin. Male FW venation: mirror small, hardly distinct from surrounding cells. Male genitalia: pseudepiphallic lophi long and thin (Fig.5C) .......................................................................................................P. africanus Robillard, 2006 -Dorsal margin of eye with small ommatidia wider. Male FW venation: mirror larger, wider than long, well differentiated. Male genitalia: pseudepiphallic lophi shorter ............................................................................................................................................................................... 2 2. Coloration little contrasted; FIII homogeneously brown on external face. Male FW venation: c1 and c2 cells sub-equal, thin. Vein Sc with four projections along its length. Male genitalia: apex of pseudepiphallic lophi bilobate (Fig. 5B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. whellani Robillard, 2006 -Coloration more contrasted; FIII external face bicolor, dorsal half dark brown, ventral half yellow. Male FW venation: c1 cell wide, c2 twice wider than c1, square and prolonging shape of miror. Vein Sc with two projections along its length. Male geni-talia: apex of pseudepiphallic lophi twisted ventrally, not bilobate (Fig. 5A) .................................................................................................. P. gorochovi Robillard sp. nov.

	Pseudolebinthus africanus Robillard, 2006
	(Fig 5C)

Pseudolebinthus africanus

[START_REF] Robillard | Phylogenetic systematics of Pseudolebinthus, a new genus of Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae) from south-east Africa[END_REF]: 677-Cigliano et al. 2018

(Orthoptera Species File Online) Material examined. Holotype. ZIMBABWE: ♂, Mt. Melsetter 5000 ft [1500 m], Melsetter Dist. Dept. Agric. S. Rhodesia. xi.1957, accession no. 11776 (Whellan) identified Lebinthus africanus by L. Chopard (NHMUK, Brit Mus 1982-221). Paratype. ZIMBABWE: 1♂, Chirinda Forest, Melsetter Dist. Dept. Agric. S. Rhodesia. x.1948, accession no. 11774 (Whellan) 1# identified Lebinthus africanus by L. Chopard (MNHN-EO-ENSIF10684). Type locality. Zimbabwe, Mount Melsetter 5000 ft. Distribution. Zimbabwe Diagnosis. (Emended from Robillard 2006). Coloration contrasted; FIII uniformly brown. Dorsal margin of eyes with small ommatidia thin.

Table 1 )

 1 Robillard sp. nov. Pseudolebinthus sp.-Vicente et al. 2017: 2203 (historical biogeography). The species is dedicated to Dr. Andrej Gorochov.Diagnosis. Species close to P. whellani, from which it differs by contrasted coloration, FIII external face bicolor, male venation and genitalia.

	(Figs 1A-D; 2A-C; 3A-B; 4A-C; 5A; 6A-E; Material examined. Holotype. MALAWI: ♂, N. Malawi, 100 km N. Mzuzu, Uzumara Mt. 10°52'18"S 34°07'45"E, 1930 m,
	21-22.xii.2010, A. Sochivho leg, molecular sample X17 (ZIN).
	Allotype. TANZANIA: ♀, Tanganyika-Terr., Matengo-Hochland wsw.v. Songea, Ugano 15[00]-1700 m [Mbinga], 20-
	30.xi.[19]35, Zerny, identified Lebinthus zernyi by L. Chopard (NHMW).
	Paratype. MALAWI: 5♂, same locality as holotype, 2-4.x.2018, T. Robillard, K. Salazar, R.J. Murphy (MNHN).
	TANZANIA: 1♀, same information as allotype (MNHN-EO-ENSIF4389).
	Type locality. Malawi, Uzumara Mount, 100 km N. Mzuzu, 10°52'18"S 34°07'45"E, 1930 m.
	Distribution. Northern Malawi, Western Tanzania.
	Life history traits and calling song. Unknown.
	Etymology.

TABLE 1 .

 1 Measurements of Pseudolebinthus gorochovi Robillard sp. nov.

		BL	PronL	PronW	FWL	FWW	FIIIL		FIIIW
	Holotype male	14.7	2.3	3.5	6.3	3.4	11.7		3.4
	Female allotype	13.4	2.2	3	0.6	-	10.7		2.7
	Female paratype	13.4	2	3.4	0.7	-	10.7		3.2
		TIIIL		TIIIs			TaIIIs		OL
			Ias	Ibs	Oas	Obs	Ids	Ods	Ols
	Holotype male	9.9	11	10	14	11	0	4	4	-
	Female allotype	?	?	?	?	?	?	?	?	11.2
	Female paratype	9.7	6	9	12	10	0	4	4	11.6

Legends: BL, body length in dorsal view; FIIIL, length of FIII; FIIIW, width of FIII; TIIIL, length of TIII; FWL, forewing length; FWW, forewing width (at the level of maximal width at about 1/3 of FWL); Ias, inner spines on TIII dorsal side above the spurs; Ibs, inner spines on TIII dorsal side between the spurs; Oas, outer spines on TIII dorsal side above the spurs; Obs, outer spines on TIII dorsal side between the spurs; OL, ovipositor length; PronL, pronotum length; PronW, pronotum width; TaIIIs, spines of third hind tarsomere, not including the apical spines: Ids, inner dorsal spines; Ods, outer dorsal spines; Ols, outer spines on lateral side of TaIII.

Acknowledgements

We thank the Orthopterist's Society for supporting RJ with one-month research grant to visit MNHN in 2017 and work on the project. This work was conducted in the context of the PhD thesis of JD, which was funded by China Scholarship Council (CSC), the Innovation Funds of Graduate Programs of Shaanxi Normal University [2012CXB019], the National Science Foundation of China [Grant No. 31402006]. We thank Andrej Gorochov (ZIN) for providing samples from the collections of Saint Petersburg. We also thank Simon Poulain (MNHN) for his help photographing the specimens, and Karen Salazar (MNHN) for her help with line drawing figures.