Ming Kai Tan

Razy Japir
email: razy.japir@sabah.gov.my

Arthur Y C Chung
email: arthur.chung@sabah.gov.my

RODZAY BIN HAJI Abdul Wahab

Tony Robillard
email: tony.robillard@mnhn.fr

Taxonomy and bioacoustics of scaly crickets (Orthoptera, Mogoplistidae, Mogoplistinae) from Borneo and Singapore

Keywords: calling song, diagnosis, new locality, new species, Ornebius, Southeast Asia

A species of scaly cricket is described here: Ornebius lupus sp. nov. from the mangrove forests in Singapore. Ornebius pullus Ingrisch, 2006 is recorded in eastern Sabah for the first time. The calling song of Cycloptiloides bimaculatus Tan et al., 2021 and Ornebius pullus from Sabah are described. We also revise the diagnosis of Ectatoderus nigrofasciatus Tan et al., 2021 from Brunei Darussalam.

Introduction

Chopard pioneered the study of the taxonomy of Mogoplistinae -or commonly called the scaly crickets-in Southeast Asia (e.g., [START_REF] Chopard | Spolia Mentawiensia: Gryllidae[END_REF][START_REF] Chopard | The Gryllidae of Sarawak[END_REF][START_REF] Chopard | Family Gryllidae: Subfamilies Mogoplistinae, Myrecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Family Oecanthidae, Gryllotalpidae[END_REF]. Since then, the taxonomy of scaly crickets from Southeast Asia has advanced considerably and many species are consequently discovered and described [START_REF] Ingrisch | Neue Grillen von Borneo und aus Thailand (Insecta: Saltatoria: Grylloidea)[END_REF][START_REF] Ingrisch | New Mogoplistinae from the Kinabalu region in Sabah, North Borneo (Insecta: Ensifera: Grylloidea: Mogoplistidae)[END_REF][START_REF] Ingrisch | New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae)[END_REF][START_REF] Tan | New taxa and notes of some described species of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Singapore[END_REF][START_REF] Tan | New species of scaly crickets (Orthoptera: Mogoplistinae) from Bukit Fraser, Malay Peninsula[END_REF][START_REF] Tan | New species of tiny scaly crickets of genus Micrornebius (Orthoptera: Mogoplistidae) from Singapore[END_REF][START_REF] Tan | Two new cricket species (Orthoptera: Gryllidae and Mogoplistidae) from the mangrove areas of Singapore[END_REF][START_REF] Tan | Contribution to the taxonomy of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from Southeast Asia[END_REF][START_REF] Tan | New species of Ornebius (Orthoptera; Mogoplistidae; Mogoplistinae) from Siargao Island of the Philippines[END_REF]Tan et al., , 2021;;[START_REF] He | Taxonomy of scaly crickets (Orthoptera: Mogoplistidae: Mogoplistinae) from China: five new species groups and three new species of the genus Ornebius Guérin-Méneville, 1844[END_REF]. Only very recently, the bioacoustics of scaly crickets from this region was also investigated concertedly for the first time (Tan et al., 2022a under review).

Despite previous efforts to document the orthopteran diversity from the mangroves of Singapore [START_REF] Tan | Two new cricket species (Orthoptera: Gryllidae and Mogoplistidae) from the mangrove areas of Singapore[END_REF][START_REF] Tan | Orthoptera in the mangroves of Singapore[END_REF], we found another species of Ornebius from such a habitat which is new to science: Ornebius lupus sp. nov. We also examined new specimens of three species of scaly crickets from eastern Sabah: Cycloptiloides bimaculatus Tan et al., 2021, Ectatoderus nigrofasciatus Tan et al., 2021 and Ornebius pullus [START_REF] Ingrisch | New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae)[END_REF]. The calling song of Cycloptiloides bimaculatus and Ornebius pullus from Sabah are described for the first time. New locality records for Ornebius pullus and Cycloptiloides bimaculatus in eastern Sabah are also reported. Lastly, we also revise the diagnosis of Ectatoderus nigrofasciatus from Brunei Darussalam.

Materials and Methods

Specimens from Sabah were collected during recent fieldwork (see Tan et al., 2022b). The specimens were preserved in absolute analytical-grade ethanol and later pinned and dry-preserved. Close-up images of morphological features (including male genitalia) were done using a macro photo lens MP-E 65 mm 1:2.8 USM (1-5×). Canon Macro Twin Lite MT24EX was used for lighting and flash. Image editing and line drawings were accomplished using Adobe Photoshop CC 2014. Measurements were done using ImageJ 1.51j8 (Wayne Rasband, Research Services Branch, National Institute of Mental Health, Bethesda, MD, USA). Terminology used to describe the male genitalia follows [START_REF] Ingrisch | New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae)[END_REF].

A single leg was preserved in absolute analytic-grade ethanol for future molecular work. The specimens will eventually be deposited in: 2). The calling song generally consists of a sequence of echemes which can be highly variable in duration (8-26 s). The echeme sequence typically begins with the echemes more erratic in terms of number of syllables, syllable durations and intervals. Each echeme is made up of 4-10 syllables (= pulses), and has an average duration of 140. 9±65.3 ms (78.0-249.1 ms). Consecutive echemes have an average interval of 371. 6±86.3 ms (285.6-474.3 ms). The average syllable duration is 14.1±1.5 ms (11.9-15.6 ms) and the average interval between consecutive syllables is 7.2±1.6 ms (5.4-9.7 ms). The dominant frequency is 8.34 kHz.

Compared to the continuous trill of Cycloptiloides timah [START_REF] Ingrisch | New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae)[END_REF] from Singapore (see Tan et al., 2022a), the calling song of C. bimaculatus also differs by a distinctly shorter syllable duration (14.1 ms in C. bimaculatus vs. 25.7 ms in C. timah) and by the intervals between consecutive syllables (7.2 ms vs. 104.5 ms). The dominant frequency is also distinctly lower (8.3 kHz vs. 9.1 kHz). It should be noted that the average temperature of the recordings for the two species are different (26.5°C in C. bimaculatus vs. 30.5°C in C. timah), which may have contributed partly to the differences in the call parameters between the two species. Remarks. When Ectatoderus nigrofasciatus Tan et al., 2021 was first described, the basal part of the male genitalia of the holotype was not examined (Tan et al., 2021: Fig. 7). The apical part of the male genitalia, previously thought to be the complete genitalia, was considered to resemble that of Ectatoderus argentatus Ingrisch, 2006 from Thailand. However, after examining more specimens collected from Brunei Darussalam, it was found that the basal half of the male genitalia of E. nigrofasciatus has in fact the medial valve curved to a spiral. This character is more similar to that in Ectatoderus angusticollis [START_REF] Chopard | Family Gryllidae: Subfamilies Mogoplistinae, Myrecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Family Oecanthidae, Gryllotalpidae[END_REF] from Singapore (Fig. 3). Below, the diagnosis is revised accordingly.

Ectatoderus nigrofasciatus

Revised diagnosis. The species is characterised by the scapus being dorsally black, but ventrally and laterally brown, and the antennae being entirely black basally; the head has a distinct black band behind eyes; and the male phallic complex is very large and forming a spiral. The new species is most similar and closest in distribution to E. angusticollis. In both species, the pronotum is more strongly widening toward apex, and the TIs and TIIs are provided with black dorsal stripes. The male phallic complex is very similar to that of E. angusticollis, including the spiral-shaped medial valve and the shape of the ventro-external sclerite. The species differs from E. angusticollis by the male phallic complex being exceptionally larger despite of having equal body size (Fig. 3); the scapus and antennae being black (instead of testaceous); and the head being provided with a distinct black band behind eyes. The new species is also similar to E. argentatus having the pronotum strongly widening apically and the frons dark; it differs by the scapus and basal antennal segments being darkened instead of yellowish brown, and by the male phallic complex forming a spiral at base instead of being merely curved.

Distribution Remarks. The specimen collected from Sabah exhibits some differences compared to O. pullus collected from Belait (Brunei Darussalam) and Singapore: legs with more whitish marmoration, supra-anal plate whitish when alive. However, we could not identify clear-cut differences in the genitalia, including both the sclerotized median valve and the membranous lateral valves between the specimen from Sabah and those from Brunei Darussalam (Fig. 5). As Tan et al. (2021) have also suggested that there might be a polymorphic species and that more evidence was needed (e.g., DNA or acoustics) to better understand the species boundaries. At the moment, the external morphology and genitalia morphology are not congruent between specimens from Brunei Darussalam, Singapore and Sabah; we also do not have acoustic data for the Brunei population; and no molecular data are available.

Furthermore, the specimen from Sabah also shares some similarities (but also with clear differences) in the external morphology of Ornebius minusculus [START_REF] Chopard | Spolia Mentawiensia: Gryllidae[END_REF] from Sipura Island (Sumatra) (http://coldb.mnhn.fr/catalognumber/mnhn/eo/ensif4531), although the genitalia for this species has not been examined. As such, without more specimens from Sabah to examine variations, we refrained from describing it as a new species, tentatively identified it as O. pullus and preferred to revise its taxonomy when the chance arises.

Distribution. Brunei Darussalam (Belait and Brunei-Muara Districts), East Malaysia (Sabah State) [new locality record], Singapore.

Type locality. BRUNEI DARUSSALAM, Brunei-Muara District, near bridge over river Sungai Lubang Barus on road coming from Tutong.

Calling song (1 ♂, in captivity, 31.5°C) (Fig. 4). The calling song generally consists of a sequence of syllables (= pulses), which can be highly variable in duration (2-14 s or 4-25 syllables). The average syllable duration is 37.5±5.0 ms (31.0-46.0 ms) and the average interval between consecutive syllables is 501.2±20.4 ms (469.3-538.4 ms). The dominant frequency is 7.97±0.13 kHz .

Compared to the calling song of Ornebius pullus from Singapore (see Tan et al., 2022a), the calling song of this specimen from Sabah has a similar call structure (i.e., the calls of both species are made up of a sequence of syllables). The biggest difference is in the interval between consecutive syllables being distinctly longer (501.2 ms vs. 400.0 ms) than that from Singapore. The syllable is only marginally longer in duration on average (37.5 ms vs. 32.4 ms) and the dominant frequency is not different (8.0 kHz vs. 8.1 kHz). Diagnosis. The new species has genitalia which appear most similar to those in the species group Ornebius (rufonigrus) [START_REF] Ingrisch | Neue Grillen von Borneo und aus Thailand (Insecta: Saltatoria: Grylloidea)[END_REF], but differs by the dark-grey colouration; the maxillary palps being not so elongated (more similar to Ornebius alvarezi Tan, Ingrisch, Baroga-Barbecho & Yap, 2019 from the Philippines; elongated oblong in Ornebius dumoga [START_REF] Ingrisch | New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae)[END_REF] from Sulawesi and in Ornebius consternus Ingrisch, 2006 from Sulawesi), the FW colouration darker (orange in O. dumoga and in Ornebius rufonigrus [START_REF] Ingrisch | Neue Grillen von Borneo und aus Thailand (Insecta: Saltatoria: Grylloidea)[END_REF] from Indochina and Malay Peninsula) and less covered by the pronotum (compared to O. consternus). The male genitalia of this new species differ from congeners from the species group by the internal sclerites at base being smaller instead of expanded.

Ornebius lupus

We also compared the new species with other species currently not placed in any species group (i.e., with unknown male genitalia). Our species resembles species with wholly dark/ black or smoky FWs, such as Ornebius obscuripennis [START_REF] Chopard | The Gryllidae of Sarawak[END_REF] from Sarawak and Ornebius fuscipennis [START_REF] Chopard | Spolia Mentawiensia: Gryllidae[END_REF] from Sumatra (including Padang, Pulau Sipura, Pulau Telo and Batu Islands). The new species differs from O. obscuripennis by the pronotal lateral lobe and lateral parts of the head being dark-grey (i.e., unicolorous with dorsal parts) instead of forming a yellow lateral band. It also differs from O. fuscipennis by much smaller body size (BL 8.4 vs. 10.5 mm; PL 2.8 vs. 4.5 mm; TL 2.5 vs. 2 mm; FIIIL 4.4 vs. 7 mm); by the FW more uniformly coloured instead of smoky with posterior margin blackish; by the apical segment of the maxillary palps longer than subapical and third; by the pronotal disc not covered with yellow-brown scales; and by the legs that are not unicolourous yellowish.

Lastly, the new species also differs from a syntopic species, Ornebius tampines Tan & Robillard, 2012 by the overall body and FW colouration, the body shape is more typical of Ornebius, instead of slenderly built, the shape of the male genitalia and by the calling songs (see Tan et al., 2022a). It is also somewhat similar to a sympatric species, O. pullus by the wholly black FWs, but differs by the lack of white margins along the lateral lobe and the posterior margin of the dorsal disc of the pronotum, by the shape of male genitalia and by the calling songs (see Tan et al., 2022a) Etymology. The species name refers to the dark-grey colouration, resembling the grey-wolf, Canis lupus. Description. Small species; habitus of male as shown in Fig 1C ,1D. Dorsum of head flattened, covered with dark-grey scales (Figs. 6A). Frontal rostrum 2.0 times wider than scapus (Fig. 6A). Scapus dark-grey (Fig 6A ); basal antennal segments black, thereafter brown with some segments black (Figs 6B, 6C). Frons and mouthparts of dark colour (Fig. 6B). Maxillary palps pale grey; with apical (= fifth) segment longer than subapical (= fourth) segment, third segment of subequal length as subapical segment; apical segment pyriform (Fig. 6C). Lateral parts of head, including genae, of dark colour, without bands (Fig. 6C). Pronotal disc brown and wholly covered with dark-grey scales, about 1.3 times longer than wide, with anterior margin narrow and straight; lateral margin widening slightly posteriorly; covering only base of mirror of FW; posterior margin convex (Fig. 6D). Lateral lobe of pronotum also wholly covered with dark-grey scales (Figs 1D,6C). TI with internal tympanum small and oval; without external tympanum. FIII 1.4-1.5 times longer than TIII; TIII 2.6 times longer than MTaIII. Femora generally pale, covered with marmoration of dark-grey scales except in basal parts. Tibiae and tarsi generally grey. Abdominal tergites covered with dark-grey scales; sternites pale. Abdominal apex darkened. Cercus covered with grey scales.

Male. FW mostly infumated black, dorsal field darkest at posterior margin (Fig. 6D); lateral field likewise in some areas with infumated white veins (Fig. 6E). Supra-anal plate with last abdominal tergite and epiproct distinctly separated by a transverse suture. Last abdominal tergite transverse, posterior margin abruptly indented in middle; with oval pale spot in middle (not obvious in dry-pinned specimen) (Fig. 6F vs. 6G); posterior margin on either side of the indentation with two patches of short and strong setae. Supra-anal plate less transverse than last abdominal tergite, black coloured, apex subtruncate (Figs 6F,6G). Subgenital plate black (Fig. 6H). Paraproct process black, elongate, cylindrical and tapering into a subacute apex (Fig. 6H). Genitalia as in Fig. 7. Central lobe of phallus with lateral valves (lv) elongate, apical area curved slightly ventrad in lateral view; tapering into obtuse apex, in apical area with inner margin curved in dorsal view; both sides together almost forming a dorsally and ventrally open tube. External sclerite (es) well developed, with external part more strongly sclerotized than inner part (in dorsal view). Medial valve (mv) membranous with base widened (in dorsal view) and upcurved (in lateral view). Internal sclerites (is) at base small but widened and curled, afterwards straight and elongate. Ecology. This species is found among foliage of mangrove trees, occurring in syntopy with another mangrove-specialist, Ornebius tampines.

Distribution. SINGAPORE Type locality. SINGAPORE, Pasir Ris mangrove Calling song. Described in Tan et al. (2022a). The calling song of this species is similar to that of Ornebius rufonigrus Ingrisch, 1987 from the same species group with regard to echeme duration and dominant frequency, although the new species produces echeme-sequences composed of triple-chirps instead of double-chrips as in O. rufonigrus.

FRCForest

  Abbreviations for general morphology: I, II, III = front, median, hind respectively (femora, legs, tibiae) F = femur FW = forewing (tegmen) T = tibia Ta = tarsus MTa = metatarsus Abbreviations for measurements: BL = body length PronL = pronotum length PronW = pronotum width FWL = forewing length FWW = forewing width FIIIL = length of hind femur TIIIL = length of hind tibia MTaIIIL = length of hind metatarsus The basic cricket song terminology follows Baker & Chesmore (2020) and Tan et al. (2022a). Analyses of the parameters of the temporal and frequency domains follow Tan et al. (2022a, b) using Raven Lite 2.0.0 and R software version 4.1.3.

  Ectatoderus nigrofasciatusTan et al., 2021: 412 Ectatoderus nigrofasciatus -Tan et al. 2022c: 600 Ectatoderus sp. -Tan & Wahab 2018: 132 Specimen examined. Holotype: BRUNEI DARUSSALAM • ♂; Belait District, Jalan Labi near Andulau Forest Reserve, N4.63354, E114.51047, 76.6±5.0 m.a.s.l.; 7 July 2019, 20h42; on rattan foliage; coll. M.K. Tan; BRU.19.70 (UBDM) Additional specimens examined. BRUNEI DARUSSALAM • 1♂; Temburong District, Kuala Belalong Field Studies Centre, along Sungei Mata Ikan, N4.54742, E115.15715, 73.8±27.9 m.a.s.l.; 22 September 2016, 22h33; on a foliage of a tree after rain; coll. M.K. Tan; KB.16.4 (MHNH) • 1♂; Temburong District, KualaBelalong Field Studies Centre, along Sungei Mata Ikan, N4.54733, E115.15720, 113.4±14.2 m.a.s.l.; 23 September 2016, 19h00; on a foliage of a tree after rain; coll. M.K.Tan; KB.16.15 (ZRC)

  Female. Unknown. Measurements (2♂, in mm). ♂ Holotype: BL = 8.4; PronL = 2.8; PronW = 2.2; FWL = 2.5; FWW = 2.3; FIIIL = 4.4; TIIIL = 3.1; MIIIL = 1.2. ♂ paratype: BL = 8.4; PronL = 2.8; PronW = 2.2; FWL = 2.5; FWW = 2.3; FIIIL = 4.4; TIIIL = 2.9; MIIIL = 1.1.

Figure captions FIGURE 1 .

 captions1 Figure captions

FIGURE 2 .

 2 FIGURE 2. Cycloptiloides bimaculatus Tan et al., 2021 ♂ calling song: oscillograms showing two echemesequences (A) and three echemes (B); spectrogram of the same echemes (C); power spectra of a syllable (D) where the grey line represents the power [dB] (axis name and scale in grey) and the black solid peak represents the relative amplitude (axis name in black).

FIGURE 3 .

 3 FIGURE 3. ♂ genitalia of Ectatoderus nigrofasciatus Tan et al., 2021 (A, B) and Ectatoderus angusticollis Chopard, 1969 (C). (A) shows only the apical half of the genitalia of the holotype. Scale bar: 1 mm.

FIGURE 4 .

 4 FIGURE 4. Ornebius pullus Ingrisch, 2006 ♂ calling song: oscillograms showing three echemes (A) and three syllables (B); spectrogram of the same syllables (C); power spectra of a syllable (D) where the grey line represents the power [dB] (axis name and scale in grey) and the black solid peak represents the relative amplitude (axis name in black).

FIGURE 5 .

 5 FIGURE 5. Ornebius pullus[START_REF] Ingrisch | New taxa and notes on some previously described species of scaly crickets from South East Asia (Orthoptera, Grylloidea, Mogoplistidae, Mogoplistinae)[END_REF] male genitalia of specimen from Singapore (A), Belait (Brunei Darussalam) (B, C) and Sabah (D, E). Genitalia were cleaned with KOH (A-D) and with membraneous parts still present after dissection (E). The genitalia of the respective specimens were ordered according to the FWs colouration: the specimen from Singapore has a wholly black/ dark FW, followed by specimens from Belait (dark brown FW with a black posterior band) and specimen from Sabah (brown FW with a black posterior band). Scale bar: 1 mm.

FIGURE 6 .

 6 FIGURE 6. Ornebius lupus sp. nov. ♂ holotype: head and pronotum in dorsal view (A); head in anterior (B) and lateral (C) views; FWs in dorsal (D) and lateral (E) views; abdominal apex in dorsal (F, G) and lateral (H) views. (F) shows the specimen preserved in ethanol and (G) shows the specimen dry-pinned. Scale bar: 1 mm.

FIGURE 7 .

 7 FIGURE 7. Ornebius lupus sp. nov. ♂ holotype genitalia in dorsal (A, B) and lateral (C) views. es = external sclerite, is = internal sclerite, lv = lateral valve, mv = medial valve. Scale bar: 1 mm.

Taxonomic Part Superfamily Grylloidea Laicharting, 1781 Family Mogoplistidae Costa, 1855 Subfamily Mogoplistinae Costa, 1855 Cycloptiloides bimaculatus Tan, Japir & Chung, 2021 (

	Rainforest Discovery
	Centre; N5.87580, E117.94299, 39.1±5.4 m.a.s.l.; 9 January 2019, 19h11; on the leaf litter; coll. M.K. Tan, R.
	Japir, M. Binti & J.L. Yukang; SDK.19.20 (FRC)
	Additional specimens examined. EAST MALAYSIA • 1♂; Sabah State, Sandakan, Sepilok, Rainforest
	Discovery Centre; N5.87554, E117.94135, 54.5±6.1 m.a.s.l.; 11 May 2022, 20h58; on the leaf litter; coll. M.K.
	Tan & T. Robillard; SBH.22.19 (FRC) • 1♂; Sabah State, Sandakan, Sepilok, Rainforest Discovery Centre;
	N5.87416, E117.93786, 89.2±6.8 m.a.s.l.; 18 May 2022, 20h07; on the leaf litter; coll. M.K. Tan & T. Robillard;
	SBH.22.152 (MNHN) • 1♂; Sabah State, Sandakan, Sepilok, Rainforest Discovery Centre; N5.87328,
	E117.93749, 95.1±5.6 m.a.s.l.; 18 May 2022, 20h57; on the leaf litter; coll. M.K. Tan & T. Robillard; SBH.22.153
	(FRC) • 1♂; Sabah State, Sepagaya Waterfall, near Lahad Datu; N4.98354, E118.14498, 182.4±7.4 m.a.s.l.; 13
	May 2022, 11h39; on the leaf litter; coll. M.K. Tan & T. Robillard; SBH.22.50 (ZRC) • 1♂, 1♀; Sabah State,
	Mount Silam, lowland forest; N4.97589, E118.19060, 341 m.a.s.l.; 13 May 2022, 20h; on the leaf litter; coll. M.K.
	Tan & T. Robillard; TR22-10 (MNHN)
	Distribution. Borneo, Sabah State: Sepilok, Sepagaya near Lahad Datu [new locality record]
	Type locality. EAST MALAYSIA, Sabah State, Sepilok
	Calling

Figs 1A, 2) Cycloptiloides bimaculata Tan et al., 2021: 410 Cycloptiloides bimaculatus -Tan et al. 2022c: 600 Specimen examined. Holotype: EAST MALAYSIA • ♂; Sabah State, Sandakan, Sepilok,

song (1 ♂, in captivity, 26.4°C) (Fig.

.

  Borneo, BRUNEI DARUSSALAM: Belait and Temburong [new locality record] Districts Type locality. BRUNEI DARUSSALAM, Belait District, near Andulau Forest Reserve He et al. 2021: 88 [assigning to species group Ornebius (flori) Ingrisch, 1998]; Tan et al. 2021: EAST MALAYSIA • 1♂; Sabah State, Tabin Wildlife Reserve; N5.19496, E118.50326, 117.1±10.3 m.a.s.l.; 15 May 2022, 22h17; on the branch of a small tree; coll. M.K.Tan, T. Robillard & R. Japir; SBH.22.95 (FRC)

	Ornebius pullus Ingrisch, 2006
	(Figs 1B, 4, 5)
	Ornebius pullus Ingrisch, 2006: 157
	Ornebius pullus -

416 [notes on new locality and variations with Singapore specimens]; Tan et al. 2022a: under review [calling songs from Singapore] Ornebius cf. pullus -Tan 2012: 34; Tan & Ingrisch 2013: 25; Tan 2017: 55; Tan 2019: 335 Specimens examined.

Tan, sp. nov.

  Ornebius sp. -Tan et al. 2022a: under review [calling songs from Singapore]

	(Figs 1C, 1D, 6, 7)
	Specimens examined. Holotype: SINGAPORE • 1♂; Pasir Ris mangrove boardwalk; N1.37805, E103.95147; 19
	March 2022, night time; on foliage of mangrove tree; coll. M.K. Tan; ZRC
	Paratypes: SINGAPORE • 1♂; Pasir Ris mangrove boardwalk; N1.37805, E103.95147; 2 May 2022, night
	time; on foliage of mangrove tree; coll. M.K. Tan; MNHN-EO-ENSIF11343

Acknowledgements

The authors thank Sigfrid Ingrisch for his comments to improve the manuscript; Huiqing Yeo (in Brunei Darussalam), Mohamad Azizan Bin Asidi, Momin Binti, John Lee Yukang and Saudi Bintang (in Sandakan) for field assistance. The permissions for collecting and exporting material in Brunei Darussalam were granted by the Forestry Department, Ministry of Primary Resources and Tourism, Brunei Darussalam (JPH/PDK/01 Pt 2 and BioRIC/HOB/TAD/51-80 respectively). The permission for research in Sabah was granted by the Sabah Biodiversity Centre (JKM/MBS.1000-2/3 JLD.3 (99) and JKM/MBS.1000-2/2 JLD.10 (179)); Osman Bangkong [the District Forestry Officer (DFO) of Lahad Datu]; Geungnong Ah Jing (the assistant DFO of Lahad Datu); and Jackly Ambrose (the DFO of Ulu Segama-Malua). The research in Tabin Wildlife Reserve was facilitated by Zainal Zahari Zainuddin and Bryan Wong Haoen. Permission for collecting material in Singapore was granted by the National Parks Board [NP/RP18-064-2a], Singapore. The work of MKT was supported by the Orthoptera Species File Grant 2019 under the taxonomic research project titled "Contribution to the species diversity and acoustic data on Orthoptera from Sandakan (Borneo, East Malaysia, Sabah)"; and Percy Sladen Memorial Fund (The Linnaean Society of London) under the project titled "Advancing biodiversity informatics of Orthoptera from Brunei Darussalam".


Ming Kai Tan

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

Cahyo Rahmadi
email: cahyo.rahmadi@lipi.go.id

Tony Robillard
email: tony.robillard@mnhn.fr

Cahyo Ramadi

A new species of Heminicsara (Orthoptera: Tettigoniidae: Agraeciini) from West Papua (New Guinea)

Keywords: Axylus, Conocephalinae, Indonesia, katydid, Southeast Asia

Introduction

Katydids of the tribe Agraeciini are highly diversified in species number and morphology. In the Southeast Asian and Oriental regions, the first major taxonomic revision of Agraeciini was done by [START_REF] Ingrisch | Monograph of the Oriental Agraeciini (Insecta: Ensifera: Tettigoniidae): Taxonomic revision, phylogeny, stridulation, and development[END_REF]. This led to further new species discoveries and taxonomic treatments of specific groups [START_REF] Gorochov | Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 2[END_REF][START_REF] Gorochov | New data on the genus Peracca (Orthoptera: Tettigoniidae: Conocephalinae) in Malaysia, with description of two new species[END_REF][START_REF] Gorochov | Taxonomy of the katydids (Orthoptera: Tettigoniidae) from East Asia and adjacent islands. Communication 11[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 | New taxa and notes on spineheaded katydids (Orthoptera: Conocephalinae: Agraeciini) from the Philippines[END_REF]. Many of these publications focus on species from areas west of the Wallace Line. To the east of Wallace Line, [START_REF] Willemse | Notes on the genus Salomona Blanchard (Orthoptera, Tettigonioidea, subfam. Agraecinae)[END_REF] provided a thorough revision of the genus Salomona which includes many species from, but not limited to New Guinea nor to the east of the Wallace Line. More recently, monographic contributions to the taxonomy of Agraeciini from New Guinea and surrounding regions also include studies by [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 | Revision of the genus Pseudonicsara Karny, 1912 (Orthoptera: Tettigoniidae: Conocephalinae: Agraeciini). Revision of the Indo-Australian Conocephalinae, part 2[END_REF][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][START_REF] Ingrisch | Revision of the genus Rhytidaspis Redtenbacher, 1891 including the description of a new genus Haudrhytidaspis gen. nov. (Orthoptera: Tettigoniidae: Conocephalinae)[END_REF]Ingrisch ( , 2020) ) and [START_REF] Rentz | Studies in Australian Tettigoniidae: Three New Species of Agraeciini From North-eastern Australia Orthoptera: Tettigoniidae; Conocephalinae; Agraeciini[END_REF], in addition to smaller contributions (Tan & Robillard,2020). These contributions, which include many new species and genera, demonstrate that a huge part of the species we of the Agraeciini from this biodiverse region remain unknown.

Heminicsara [START_REF] Karny | Orthoptera Fam. Locustidae Subfam[END_REF] is one of the Agraeciini genera limited to New Guinea and the surrounding archipelagos. It belongs to the Axylus genus group Ingrisch, 2015, characterised by deep lateral lobes of the pronotum, long prosternal spines, short and obtuse fastigium verticis and quadrangular cross-section in the anterior tibiae [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]. This genus currently comprises 62 species, 51 species of which were only described recently [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]. Even though recent new species were described from old museum specimens, more species can be expected with more intensive sampling in under-explored parts of the genus distribution. Recent fieldwork in West Papua, Indonesia led to the discovery of a new species, which is described here: Heminicsara incrassata sp. nov. This represents the first species of Heminicsara described and known the southwest of New Guinea.

Materials and Methods

The katydids were collected during the "Lengguru 2014" expedition in Papua, Indonesia and will be deposited in the collections of the Muséum national d'Histoire naturelle Paris, France (MNHN) and in the Museum Zoologicum Bogoriense, Bogor, Indonesia (MZB). Square brackets in the list of specimens are used for additional information not mentioned on the specimen labels or translated from French labels.

Male titillators were dissected in specimens, softened using KOH, by cutting the membranes between the paraprocts and the subgenital plate. They were then cleaned with cold KOH, and subsequently kept in glycerine. Close-up images of habitus and morphological features were done using a Canon EOS 6D digital SLR camera with a macro photo lens MP-E 65 mm f/2.8 USM (1-5×). Imaging stacking was done using Helicon Remote version 9.3.1. W and Helicon Focus 6.8.0. Photographs of male titillators were done with a binocular microscope Leica MZ16 with AmScope Microscope Eyepiece Camera (MU1000, 10 MP Aptina Color, CMO50) attached via an AmScope FMA050 fixed microscope adaptor and the montage software AMScope version ×64, 3.7. Image editing was accomplished using Adobe Photoshop CC2014.

Measurements of specimens were made using ImageJ 1.51j8 (Wayne Rasband, Research Services Branch, National Institute of Mental Health, Bethesda, MD, USA). In the measurements, the following abbreviations are used: BL = body length; PL = pronotum length; PW = pronotum width; TL = tegmen length; FIIIL = posterior femur length; FIIIW = posterior femur width; TIIIL = posterior tibia length.

Family Tettigoniidae Krauss, 1902 Subfamily Conocephalinae Burmeister, 1838

Tribe Agraeciini Redtenbacher, 1891

Genus group Axylus Ingrisch, 2015

Genus Heminicsara [START_REF] Karny | Orthoptera Fam. Locustidae Subfam[END_REF]Heminicsara Karny, 1912: 9 -1926: 200;Ingrisch, 2015: 166 (with Etymology. The species name refers to the enlarged and thickened tenth abdominal tergite, the most distinct character of this species from the congeners (from Latin, incrassata = thickened).

Diagnosis. This new species is different from species from the Axylus genus group by the combination of these characters: colouration generally pale green, mottled dark; face dark between the eyes, frons grey-brown; pronotum disc with two transverse lateral spots at anterior third and an X-shaped dark pattern behind anterior third, male tenth abdominal tergite forming a large shield-shaped plate with two long apical lobes pointing ventrad, male cercus conical with a strongly curved inner-basal process and titillators fused and elongated.

This new species is similar to fully-winged species Heminicsara bilobata Ingrisch, 2015 from north-central New Guinea, Heminicsara mamberamo Ingrisch, 2015 from Mamberamo in West Papua and Heminicsara sica Ingrisch, 2015 from East Sepik in Papua New Guinea by the slightly prolonged tenth abdominal tergite, but it differs by the presence of two long apical lobes pointing ventrad. It also differs from these congeners by face and pronotum colouration, titillators being fused and elongated (instead of separated) and male cercus with only one internal basal process, which is strongly curved (instead of long with apical spines) and male cercus without distal internal process. It also differs from H. bilobata and H. mamberamo by the fused titillators (instead of separated or free). It is also similar to H. sica by the fused titillators but differs by the male cercus missing a semi-circular bulge at apex, and the male subgenital plate being missing long, laterally compressed projections curved dorsad and fused with styli.

This new species has a somewhat similar colouration as long-winged Heminicsara despecta Naskrecki & Ingrisch, 2015 from Muller Range in Papua New Guinea and Heminicsara schlaginhaufeni [START_REF] Karny | Orthoptera Fam. Locustidae Subfam[END_REF] occurring along the northern coast of New Guinea but differs by less black mottled patterns, the male abdominal apex (including tenth abdominal tergite and cercus) and titillators.

This new species also differs from fully winged species (with only females known) by dark patterns on the pronotal lateral lobe limited to the anterior and posterior ends (instead of being connected or only present at posterior end) and FW anterior part of costal area distinctly narrow and widening posteriorly: Heminicsara cingima Ingrisch, 2015 from central mountain range of New Guinea, Heminicsara coriformis Ingrisch, 2015 from Nabire, Heminicsara excisa [START_REF] Karny | Beiträge zur malayischen Orthopterenfauna XV. Die Conocephalinen (s.l.)[END_REF], Heminicsara scutula Ingrisch, 2015 from Central Province of Papua New Guinea, Heminicsara siwi Ingrisch, 2015 from Arfak Mountains, Heminicsara tabtab Ingrisch, 2015 and Heminicsara wanuma Ingrisch, 2015 from Adelbert Range in Papua New Guinea. It is similar to H. coriformis provided in the pre-subcostal area with network of white veinlets but differs from it by larger eyes. It also differs from H. excisa by the absence of vertical black band widening ventrad on the face.

Descriptions. Fully-winged, medium-sized species with habitus typical of the genus, as shown in Fig. 1. Fastigium verticis conical, fairly stout, not surpassing scapus, apex obtuse (Fig. 2A). Frons flat, oblique, subsmooth in the middle and very weakly rugose on the lateral parts; without lateral carinae (Fig. 2B). Maxillary palps elongated and slender; apical segment longest with apex truncated, subapical segment slightly longer than third segment. Eyes oval, weakly protruding (Figs. 2A,2B); median ocellus reduced, indicated only by light oval spot (Fig. 2B); lateral ocelli between fastigium of vertex and scapus, small and oblong (Fig. 2A). Pronotum smooth, lateral lobe slightly more rugose; disc distinctly longer than wide, broadly rounded into paranota (Figs. 2A,2C); anterior margin very faintly emarginated in middle; posterior area not raised, projecting posteriorly and covering stridulatory file and base of mirror, margin rounded (Fig. 2A). Lateral lobe about 3.2 times as long as high; ventral margin very faintly concave, ventro-posterior angle slightly projecting, posterior margin broadly rounded (Fig. 2C). Thoracic auditory spiracle completely covered by paranota (Fig. 2C). Prosternum with two fairly long spines (Fig. 2D). Mesosternal and metasternal lobes conical with blunt apices; median plate less broad than meso-and meta-sterna (Fig. 2D). Anterior coxa with a long dorsal spine pointing anteriorly, apex acute (Fig. 2A). Anterior tibia with both tympana slit-shaped covered by slightly swollen conchate covering (Fig. 2B). Femora with the following number of robust spines on ventral margins: anterior femur 7 external (slightly smaller than internal ones), 7 internal of increasing size distally; middle femur 7 external, 3 small internal at base; posterior femur 11 external of increasing size distally, 6 barely visible small internal (all anterior of the most basal external spines). All knee lobes long spinose externally and internally, except for external knee lobe of middle femora. Tibiae with the following number of spines on ventral margins: anterior tibia 9 external, 9 internal; middle tibia 10 external, 9 internal.

Male. Tegmen macropterous, surpassing hind femora, apex rounded (Fig. 1). Mirror fully developed, somewhat triangular, 1.2 times wider than long, anterior margin wider than posterior margin (Fig. 3A). Stridulatory file straight; about 2.2 mm long; with teeth evenly distributed; with circa 78 teeth (Fig. 3B). Hind wings concealed under tegmina (Fig. 1). Tenth abdominal tergite highly modified, forms a large shield-shaped plate; setose, especially in the medial area; apical half bent ventrally; apical margin producing two elongated lobes in the middle, pointing ventrad, apex oblique, rounded between the two apical lobes; apical lobe carinate along external margin with apex subacute (Figs. 3D-E). Epiproct concealed under the prolongation of tenth abdominal tergite. Cercus elongate and conical with apex rounded; ventro-inner margin with a stout and strongly curved basal lobe with apex rounded and pointing ventrad (Fig. 3F). Subgenital plate rectangular, slightly longer than broad, posterior margin narrowly and angularly excised in middle, lateral lobes stout; styli cylindrical, long, apex obtuse (Fig. 3G). Phallus with fused titillators (Figs. 3H,3I). Titillator slender and elongated; dorso-lateral margins carinate and diverging posteriorly into laminate lobules (Fig. 3H); posterior end truncated (Fig. 3I).

Female. Unknown. Colouration. Pale green (yellow when preserved and discoloured) with mottled dark patterns (Fig. 1). Head dorsum pale green with some faint grey-brown strips, fastigium verticis grey-brown (Fig. 2A). Scapus dark brown and pedicel yellow brown (Fig. 2B); antennal segments red brown to dark brown (Fig. 2A). Face dark between the eyes, frons grey-brown. Clypeus black in dorsal parts, yellow brown on ventral half; maxillary palpus and labrum pale yellow; mandible black (Fig. 2B). Pronotum generally pale green. Dorsal disc with anterior part with lateral transverse black spots, anterior third dark brown in middle, between the lateral spots; posterior of the anterior third with X-shaped dark pattern, posterior arms of X-shaped pattern larger and extending to dorsal part of lateral lobe (Fig. 2B). Lateral lobe of pronotum green to yellow green (Fig. 2C). Tegmen generally with cells black; veins white (Figs. 1B, 2C, 3A). Coxae pale yellow. Femora pale green with a faint dark dorsal band at apical end just before knees (most obvious in posterior femora); knees yellow green; spines brown with black apices (Figs. 1A,2C). Anterior tibia yellow brown, with dark spots between and posterior of tympana (Fig. 2B). Abdominal tergites, sternites and apex pale green (Figs. 1, 2D, 3D-E).

Measurements (mm).

Holotype BL = 27.0, PL = 6.0, PW = 7.0, TL = 28. [START_REF] Tan | New taxa and notes of some described species of Agraeciini (Orthoptera: Tettigoniidae: Conocephalinae) from Malay Peninsula[END_REF]FIIIL = 17.8,FIIIW = 4.6,TIIIL = 17.0;paratype BL = 28.5,PL = 9.9,PW = 7.2,TL = 27.9,FIIIL = 18.7,FIIIW = 4.6 TIIIL = 17.3 (slightly deformed).

Habitats. The two known specimens were collected in primary forest, one on a Pandanus tree, one on a tree trunk at about 2.5 m high.

  key to species) Type species: Heminicsara jacobii Karny, by original monotypy Heminicsara incrassata Tan, Ingrisch & Robillard, sp. nov. Material examined. Holotype: INDONESIA • ♂; Nouvelle-Guinée [New Guinea], Lobo [village in the Bay of Kaimana], S3.7033056, E134.071444 (LOBO6), 221 m.a.s.l., forêt proche [forest near] PK8/ route Lobo-Kaimana, crête [ridge] prox. camp, forêt primaire [primary forest], 21-30.X.2014, jour [day], on Pandanus, T. Robillard leg., LEN2014-TR346 (MZB). Paratype. INDONESIA • 1♂; Nouvelle-Guinée [New Guinea], Lobo [village in the Bay of Kaimana], S3.764667, E134.100111 (LOBO1), 23 m.a.s.l., forêt littorale sur pente, autour du camp [littoral forest on slope, around the camp], sur tronc (height = 2.5 m) [on tree trunk]; 21-30.X.2014, nuit [night], T. Robillard leg., LEN2014-TR66 (MNHN-EO-ENSIF11334). Type locality. Indonesia, West Papua: Lobo (a coastal village in the Bay of Kaimana)

Figure captions FIGURE 1 .

 captions1 Figure captions

FIGURE 2 .

 2 FIGURE 2. Heminicsara incrassata sp. nov. head and pronotum in dorsal view (A), head and anterior legs in anterior view (B), body in lateral view (C), thorax in ventral view (D). Scale bar: 10 mm.

FIGURE 3 .

 3 FIGURE 3. Heminicsara incrassata sp. nov. base of tegmen in dorsal view (A), stridulatory file in ventral view (B), abdominal apex without titillators in lateral (C) and posterior (D) views, abdominal apex with titillators in lateral view (E), cercus in dorsal view (F), titillators in lateral (H) and dorsal (I) views. Scale bars: 1 mm (A, C-G), 0.5 mm (B, H, I).

Acknowledgments

Fieldwork was supported by the Project Lengguru 2014 (www.lengguru.org), conducted by the French Institut de Recherche pour le Développement (IRD), the Indonesian Institute of Sciences (LIPI), the University of Papua (UNIPA), the University of Cendrawasih (UNCEN), the University of Musamus (UNMUS) and the Sorong Fisheries Academy (APSOR) with corporate sponsorship from COLAS Group, Veolia Water and the Total Foundation. The work of MKT was supported by the Fyssen Foundation Postdoctoral Fellowship.