(Orthoptera: Gryllidae) from The

ZOOSYSTEMATICA ROSSICA, 27(1): 40–76 25 JUNE 2018

Taxonomic studies on the cricket subfamilies Pteroplistinae, Phaloriinae and Cacoplistinae (Orthoptera: Gryllidae) from the Old World Таксономические исследования по cверчкам подсемейств Pteroplistinae, Phaloriinae и Cacoplistinae (Orthoptera: Gryllidae) из Старого Света

A.V. GOROCHOV

А.В. ГОРОХОВ

A.V. Gorochov, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia. E-mail: [email protected]

A new material on the subfamilies Pteroplistinae, Phaloriinae and Cacoplistinae from the Indo-Malayan and Afrotropical regions of the Old World is considered. Nineteen new taxa of these subfamilies are described: Pteroplistes malaccanus sp. nov. from Malacca, P. borneoensis sabahi subsp. nov. from Borneo, Tramlapiola? bugiamap sp. nov. from Vietnam [Pteroplisti- nae]; Trellius (Zatrellius) kinabalu sp. nov., T. (Z.) crocker sp. nov. and T. (Z.) tawau sp. nov. from Borneo, T. (Neotrellius) logunovi sp. nov. and T. (N.) simulator sp. nov. from Vietnam, Borneloria spinosa gen. et sp. nov. and B. moorei occidentalis subsp. nov. from Borneo, Pha- loria (Papuloria) multa sp. nov. from Sulawesi and a nearest island [Phaloriinae: Phaloriini]; Subtiloria semota sp. nov. and S. succinea korup subsp. nov. from Cameroon, Schizotrypus conradti nigericus subsp. nov. from Nigeria, Afrophaloria malawi sp. nov. from Malawi, A. dja sp. nov. from Cameroon [Phaloriinae: Subtiloriini]; Homoeogryllus ambo sp. nov. from Ethiopia, H. reticulatus limbe subsp. nov. from Cameroon [Cacoplistinae: Homoeogryllini]. New data on distribution of some other species are also given. Рассмотрен новый материал по подсемействам Pteroplistinae, Phaloriinae и Cacoplisti- nae из Индо-Малайской и Афротропической областей Старого Света. Описаны девят- надцать новых таксонов, относящихся к этим подсемействам: Pteroplistes malaccanus sp. nov. из Малакки, P. borneoensis sabahi subsp. nov. с Борнео, Tramlapiola? bugiamap sp. nov. из Вьетнама [Pteroplistinae]; Trellius (Zatrellius) kinabalu. sp., nov T. (Z.) crocker sp. nov. и T. (Z.) tawau sp. nov. с Борнео, T. (Neotrellius) logunovi sp. nov. и T. (N.) simulator sp. nov. из Вьетнама, Borneloria spinosa gen. et sp. nov. и B. moorei occidentalis subsp. nov. с Борнео, Phaloria (Papuloria) multa sp. nov. с Сулавеси и соседнего острова [Phaloriinae: Phalori- ini]; Subtiloria semota sp. nov. and S. succinea korup subsp. nov. из Камеруна, Schizotrypus conradti nigericus subsp. nov. из Нигерии, Afrophaloria malawi sp. nov. из Малави, A. dja sp. nov. из Камеруна [Phaloriinae: Subtiloriini]; Homoeogryllus ambo sp. nov. из Эфиопии, H. reticulatus limbe subsp. nov. из Камеруна [Cacoplistinae: Homoeogryllini]. Приведены также новые данные по распространению некоторых других видов. Key words: crickets, taxonomy, Old World, Orthoptera, Gryllidae, Pteroplistinae, Phalor- iinae, Cacoplistinae, new taxa, new localities Ключевые слова: сверчки, таксономия, Старый Свет, Orthoptera, Gryllidae, Pteroplisti- nae, Phaloriinae, Cacoplistinae, новые таксоны, новые местонахождения

INTRODUCTION subfamily group of Gryllidae. It is a continu- ation of some my previous papers containing This paper is devoted to three cricket information on new and little known repre- subfamilies (Pteroplistinae, Phaloriinae and sentatives of these taxa (Gorochov, 1985, Cacoplistinae) from the Phalangopsinae 1986, 1988a, b, 1990, 1992, 1996, 1999, 2001,

2003, 2004a, b, c, 2005, 2010, 2011a, b, 2014; Paratype. Female, same data as for holotype Gorochov & Kostia, 1999; Gorochov & Tan, (ZIN). 2012). The latter taxa are distributed in the Description. Male (holotype). Body Old World and unknown in the New World, rather large for this subfamily, clearly dor- but the Phalangopsinae, a fourth subfamily soventrally flattened and with following of this group, is distributed in the both Old colouration (Figs 1–3): epicranium dark World and New World. Representatives of brown with grey eyes, yellowish ocelli, this subfamily from the Old World are now light brown spot on rostral apex and spot considered by me in a special series of papers near (under) median ocellus (these spots (Gorochov, 2015a, b, 2017a, b, 2018). connected with each other by light brown The subfamilies studied are distributed median line), light yellowish brown trans- mainly in tropical regions. All species of verse stripe between middle parts of eyes Pteroplistinae live on and under the bark of (this stripe located slightly behind medi- forest trees. Most representatives of Pha- an ocellus and crossing lateral ocelli) and loriinae also live in forests (but usually near much wider transverse band between distal water) and may be found on leaves of trees parts of eyes (this band somewhat arcuate and shrubs, on tree trunks or on rocks near and with posterior edge convex), and these brooks; species of Cacoplistinae have been light parts separated from each other and found on leaves and branches of bushes in from clypeus by brown transverse stripes forests and savannahs or in some more spe- (stripe crossing median ocellus almost dark cialized conditions (on walls of termitaries; brown); antennae brown with almost dark Bhargava, 1982). Oviposition in Pteroplis- brown numerous and small spots on flagel- tinae is produced in the fissures of bark or lum (but these spots on proximal part of fla- wood, in Phaloriinae, in the bark but some- gellum indistinct); mouthparts brown with times possibly in the soil (some African light brown lateral parts of upper clypeal genera), and in Cacoplistinae, probably in half as well as small areas on maxillae and the soil. on labium, and with dark brown apical The material used in this paper is mainly segment of each maxillary palpus; prono- deposited at the Zoological Institute, Rus- tum also dark brown but with rather large sian Academy of Sciences, St Petersburg (wide) light brown median area on anterior (ZIN). However, two paratypes of one new half of disc; tegmina greyish brown with species are at the Museum of Natural His- barely darker venation in apical area of dor- tory, London (BMNH). All the specimens sal field and in lateral field; hind wings with are dry and pinned; photographs of their distal parts rather dark, greyish brown; morphological structures were made with a legs with light greyish brown femora hav- Leica M216 stereomicroscope. ing dark brown distal part of hind femur and intensively greyish brown reticular TAXONOMIC PART oblique pattern on dorsal half of rest part of this femur, with reddish brown tibiae hav- Subfamily PTEROPLISTINAE ing dorsal surface of hind tibia as well as Chopard, 1936 distal parts of its outer and inner surfaces Pteroplistes malaccanus sp. nov. dark, and with brown tarsi having proximal (Figs 1–6, 10–12) two thirds of hind basitarsus lighter (light greyish brown); venter of body light brown Holotype. Male, Malaysia, Pahang State (Ma- but with greyish brown genital plate; distal lacca = Malay Peninsula), Taman Negara National Park on Tembeling River, ~100 m, primary forest, part of abdominal dorsal half dark brown on bark of living tree under table attached to its with greyish brown cerci and dorsal areas trunck (at daytime), 28.XI–5.XII.2014, A. Goro- on tergites, whitish spot in central part of chov, M. Berezin, E. Tkatsheva (ZIN). anal plate and brown distal and lateral parts

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 42 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD of this plate. Structure of head typical of and labrum almost yellowish, pronotum this genus, but median ocellus much smaller with almost parallel lateral sides, tegmina than lateral ones, and space between anten- with narrower dorsal field having nine lon- nal cavities approximately 1.1 times as wide gitudinal veins (these veins shallowly S- as scape; pronotum distinctly wider than shaped, i.e. with more or less longitudinal long, somewhat widening to pterothorax, proximal and distal parts, and with slightly with slightly concave anterior and almost oblique and arcuate middle parts) and regu- straight posterior edges of disc (Fig. 1), and lar crossveins between them (Fig. 5), lateral with lateral lobes low (anterior half of these tegminal field with moderately narrow Sc-R lobes widely convex ventrally, but posterior and R-M areas having regular crossveins one very low, with oblique ventral edge and as well as with 13–14 Sc branches (these narrowly rounded posteroventral corner). branches obliquely longitudinal and with Tegmina long and rather wide, with vena- sparse crossveins), hind wings insignifi- tion of dorsal field as in Fig. 2; lateral teg- cantly protruding beyond tegminal apices, minal field with slightly widened R-M area colouraton of wings as in male but with (having regular crossveins) and lancet-like dark brown venation and darkened Sc-R cell, with narrow Sc-R area lacking cross- area (however, veins of dorsal field situated veins, and with numerous (24–26) moder- along its lateral edge lighter, almost as mem- ately long and strongly oblique branches branes of this field), fore and middle tibiae of Sc (crossveins between these branches barely darker, and anal plate brown and un- absent); hind wings distinctly protruding specialized. Genital plate light brown, small beyond tegminal apices. Legs rather short and more or less triangular but with distinct and stout; fore tibia with inner and outer rounded posteromedian notch; ovipositor tympana moderately small and not im- as in Fig. 6. mersed (inner tympanum oval, but outer Length in mm. Body: male 15.5, female one somewhat narrowed); hind femur al- 15; body with wings: male 22, female 20; most 1.45 times as long as hind tibia; latter pronotum: male 3.3, female 3.4; tegmina: tibia with six apical spurs, three short and male 15, female 14; hind femora: male 12.4, articulated distal spines on outer dorsal female 12.3; ovipositor 9.5. keel, two such spines on inner dorsal keel, Comparison. The new species is most re- and numerous small and unarticulated den- lated to P. sumatranus from Sumatra, but it ticles on both these keels in more proximal differs from the latter species in the teeth of parts of this tibia; hind basitarsus with six male anal plate distinctly larger, and in the outer and five inner dorsal denticles (except characters of male genitalia listed above, in for a pair of apical spurs). Anal plate with a the description. From P. acinaceus Sauss., pair of rather large teeth on anterior part of described from the same peninsula (Sau- its dorsum and with rather deep and round- ssure, 1877: “Malacca”), P. malaccanus is ly angular notch between them (Fig. 4); distinguished by the teeth of male anal plate genital plate rather large, somewhat wider located on the anterior (not posterior) half than long, with three short angular projec- of this plate and having an almost angular tions at apex and very shallow and rather (not round) notch between them, distal wide notches between them (Fig. 3); cerci epiphallic lobes with the apical parts al- very long. Genitalia most similar to that of most hook-like and directed aside as well as P. sumatranus Gor., but basal part of rachis with the proximal parts lacking very large with dorsomedian process (spine) clearly ventral lobes reaching the median part of bifurcated at apex and with a pair of small anterior epiphallic edge, and rachis with tubercles around this process (Figs 10–12). the distal part strongly curved forwards- Female. General appearance similar to upwards and with the proximal part hav- that of male, but most part of both clypeus ing a dorsomedian process bifurcated at

Figs 1–9. Pteroplistinae: 1–6, Pteroplistes malaccanus sp. nov.; 7, P. borneoensis borneoensis Gor.; 8, P. b. sabahi subsp. nov.; 9, Tramlapiola? bugiamap sp. nov. Head and pronotum with visible part of left fore leg from above (1); male (2) and female (5) tegmina in rest position with visible parts of hind wings from above; male genital plate from below (3); male anal plate with last tergite and bases of cerci from above (schematically) (4, 7, 8); ovipositor with some distal parts of body from side (6). [7, after Gorochov (2004), modified].

Figs 10–19. Pteroplistinae, male: 10–12, Pteroplistes malaccanus sp. nov.; 13–15, P. borneoensis sabahi subsp. nov.; 16, P. b. borneoensis Gor.; 17–19, Tramlapiola? bugiamap sp. nov. (deutonymph collected during imaginal moulting). Genitalia from above (10, 13, 17), from below (11, 14, 18), and from side (12, 15, 19); rachis, left part of epiphallus and posterior parts of endoparameres from above (16).

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 45 the apex. From P. lagrecai Gor. and P. bor- Paratypes. Two males, same data as for ho- neoensis Gor. (both from Borneo), the new lotype (ZIN); male and female, same state, species differs in the rachis of male genitalia Tawau Hills National Park near Tawau City, more strongly curved in the profile, and in 200–400 m, primary-secondary forest, on trunks of living trees at night (female, 14–20.V.2013, the endoparameral sclerites having length A. Gorochov, M. Berezin, E. Tkatsheva; male, of these sclerites and width of their anterior 6–14.III.2014, M. Berezin) (ZIN). parts intermediate between those of the lat- Description. Male (holotype). Size, co- ter congeners; and from Indian representa- louraton and structure of body similar to tives, in the female tegmina and ovipositor those of P. malaccanus but with follow- clearly longer (from P.? platycleis Bol.), or ing differences: epicranium light yellowish in the posterolateral epiphallic lobes dis- brown with blackish eyes, a pair of small tinctly different in the shape (from P. kerva- brown spots on rostrum near medial edges sae Jaiswara and P. masinagudi Jaiswara; see of antennal cavities, three brown transverse Jaiswara & Desutter-Grandcolas, 2014). stripes on more posterior parts of dorsum Etymology. The new species is named af- (one narrow stripe crossing median ocel- ter the Malacca Peninsula. lus, one more posterior stripe widened in median two thirds and located immediately Pteroplistes borneoensis borneoensis behind lateral ocelli, and one band between Gorochov, 2004 posterior parts of eyes running along pos- (Figs 7, 16) terior edge of epicranial dorsum), slightly darkened posterior parts of genae and areas New material. Malaysia: male and 3 females, Sarawak State (Borneo), environs of Kuching under eyes, and greyish stripe along clypeal City, Kubah National Park on Matang Mt, 200– suture forming low triangle in median part; 500 m, primary forest, on trunks of living trees at antennae light brown with almost yellow- night, 10–17.III.2012, A. Gorochov, M. Berezin, ish scape, numerous (dense) small greyish E. Tkatsheva, I. Kamskov (ZIN). brown spots on proximal half of flagellum, Note. This subspecies was originally and greyish brown rest of flagellum; mouth- described as a species from Sarawak but parts also light yellowish brown but with without any more exact locality (Goro- dark brown margins of clypeus (including a chov, 2004). The male studied here is in ac- pair of stripes along transverse folds between cordance to the original description of this upper and lower clypeal halves), small spot species (including all the characters of male in median part of labrum as well as marks on genitalia) but has the anal plate with a bare- distal and lateral parts of mandibles; prono- ly wider notch between its dorsal teeth (this tum brown with almost reddish brown disc; notch is almost as in Fig. 8; in holotype, this wings distinguished from those of P. malac- notch is as in Fig. 7). Now this subspecies canus by dorsal tegminal field with mirror is recorded from the above-mentioned lo- slightly longer than chords (vs. slightly cality (situated in the western part of this shorter than chords), lateral tegminal field state) for the first time. with 21–23 Sc branches, and hind wings slightly shorter (insignificantly projected Pteroplistes borneoensis sabahi behind tegminal apices); legs somewhat subsp. nov. lighter than in this species; hind basitarsus (Figs 8, 13–15) with seven outer and six inner dorsal denticles; teeth of anal plate shorter, with shal- Holotype. Male, Malaysia, Sabah State (Bor- lower (roundly transverse) notch between neo), Sandakan Distr., environs of Sukau Vill. on Kinabatangan River (~35 km from sea), almost them (Fig. 8); genitalia almost as in Figs sea level, secondary / primary forest, on trunk of 13–15, very similar to those of nominotypi- living tree at night, 8–13.V.2013, A. Gorochov, cal subspecies in structure (see Gorochov, M. Berezin, E. Tkatsheva (ZIN). 2004: figs 28–30) but slightly distinguished

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 46 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD from them by narrow sclerotized posterior eus and labrum as well as most part of maxil- parts of distal epiphallic lobes somewhat lae and labium (including their palpi) light longer, and rachis with wider dorsoproxi- greyish brown; rest mouthparts greyish mal process and narrower subapical part brown with dark apices of mandibles; pro- (this part of rachis clearly narrower than its notum with brown disc having rather large nearest but more proximal part). light yellowish brown spots (Fig. 9) and with Variations. In male from Tawau Hills dark brown lateral lobes; wing pads brown National Park, colouration of head less con- with almost light brown “dorsal tegminal trasting, and pronotum almost uniformly field” (Fig. 9); legs greyish brown with a few brown; number of Sc branches in tegmina lighter but blurred spots on fore and middle and number of denticles on hind basitarsi in- femora as well as on all tibiae, and with light significantly varied; subapical part of rachis greyish brown hind femur having a few dark sometimes barely wider or barely narrower. brown distal areas, numerous brown oblique Female. General appearance practically lines on dorsolateral surface of rest part of as in female of P. malaccanus, but coloura- this femur and less regular brown marks on tion of head with slightly lighter rostral dorsomedial surface of this part; venter of part, and all sternites more darkened (geni- body light brown with almost brown pos- tal plate damaged). terior abdominal sternites and genital plate; Length in mm. Body: male 14.5–15; fe- thoracic tergites light greyish brown, but male 14.3; body with wings: male 20–21, abdominal ones brown with rather numer- female 20; pronotum: male 2.9–3.1, female ous light brown spots; other structures of 3.2; tegmina: male 13.4–13.8, female 14.3; abdominal apex (including cerci) more or hind femora: male 11.5–12, female 12; ovi- less light brown. Head slightly deformed, positor 9. but most of its structures kept in good con- Comparison. The new subspecies from dition: median ocellus much smaller than the Sabah State of Borneo is distinguished lateral ones; width of scape almost equal from P. b. borneoensis, distributed in the Sar- to distance between antennal cavities. Pro- awak State of this island, by the posterior notum slightly wider than long, somewhat (narrow) sclerotized parts of distal epiphal- narrowing to head (Fig. 9), and with lateral lic lobes longer, and by the rachis of male lobes similar to those of P. malaccanus but genitalia with the subapical part narrower having more widely rounded posteroven- and with the dorsoproximal process wider. tral corners; wing pads reaching middle part of third abdominal tergite, approximately Tramlapiola? bugiamap sp. nov. equal to each other in length, with visible (Figs 9, 17–19) rudiments of stridulatory apparatus on tegminal pads (Fig. 9); legs similar to those of Holotype. Male deutonymph collected dur- P. malaccanus in structure, but tympana un- ing imaginal moulting, Vietnam, Binh Phuoc Prov., 13 km NE of Bu Gia Map Vill., Bu Gia developed, hind femur almost 1.9 times as Map National Park on Dak A River, 12°12'20''N, long as hind tibia, and hind basitarsus with 107°12'15''E, 350 m, 18–31.V.2011, L. Anisyut- 6–7 outer and 4–6 inner dorsal denticles. kin, A. Anichkin (ZIN). Anal plate small, roundly triangular and un- Description. Male deutonymph (holo- specialized (i.e. without rudiments of teeth type). Body rather small for this subfamily or spines); genital plate short (almost twice and clearly dorsoventrally flattened. Co- as wide as long) and with strongly convex louration following: epicranium dark brown posterior edge (this plate also unspecial- with light greyish eyes and ocelli, and with ized: without rudiments of spines). Genita- seven light yellowish brown longitudinal lia completely formed, except for rami and stripes on dorsum behind lateral ocelli; an- formula (or these sclerotized structures ab- tennae more or less uniformly brown; clyp- sent): epiphallus divided into left and right

Figs 20–28. Phaloriinae (Phaloriini): 20, 21, Trellius (Zatrellius) kinabalu sp. nov.; 22, T. (Z.) crocker sp. nov.; 23, 24, T. (Z.) tawau sp. nov.; 25, T. (Neotrellius) logunovi sp. nov.; 26, T. (N.) simulator sp. nov.; 27, Borneloria sp.; 28, Phaloria (Papuloria) multa sp. nov. Male tegmina in rest position with visible parts of hind wings from above (20, 22, 23); same structures but with head and pronotum from above (25); distal part of ovipositor from below, reconstructed (21, 24) and original (26–28) views.

Figs 29–43. Phaloriinae (Phaloriini), male: 29–31, Trellius (Zatrellius) kinabalu sp. nov.; 32–34, T. (Z.) crocker sp. nov.; 35–37, T. (Z.) tawau sp. nov.; 38, 39, T. (Neotrellius) logunovi sp. nov.; 40, 41, T. (N.) tonkinensis (Chop.); 42, 43, T. (N.) simulator sp. nov. Genitalia from above (29, 32, 35, 39, 41), from below (30, 33, 36) and from side (31, 34, 37, 38, 40), as well as their distal half from side (42) and from above (43).

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 49 sclerotized plates by narrow median mem- leaf of bush at night, 26.IV–1.V.2013, A. Goro- branous stripe (as in Tramlapiola Gorochov, chov, M. Berezin, V. Gorochova, E. Tkatsheva 1990 and Crockeriola Gorochov et Kostia, (ZIN). 1999), but these plates without distinct Paratypes. Male and 2 females, same data as posteroventral processes, and their anterior for holotype. parts connected with each other by very Description. Male (holotype). Body narrow arcuate ribbon (as in Crockeriola); medium-sized for this genus. Colouration each distal lobe of these epiphallic plates not brownish grey with following pattern: epi- very long, slightly curved and with angular cranium (including eyes) almost uniformly projection on dorsal surface; endoparameres greyish brown with somewhat lighter ocelli; presented by a pair of rather wide transverse antennae uniformly light brown but with ribbons; rachis long, rather narrow and with slightly darker area on medial surface of thin dorsal process which somewhat shorter scapes; mouthparts light brown with up- than ventral rachial lobe (i.e. rachis more per half of clypeus and proximal part of similar to that of Tramlapiola than to that of mandibles brown, and with more or less Crockeriola) (Figs 17–19). whitish palpi having darkened areas on Imago unknown. proximal half of third and fourth segments Length in mm. Body 10.5; pronotum 2.4; of maxillary palpus as well as on distal part tegminal pads 2.8; hind femora 8.5. of fifth segment of this palpus; pronotum Comparison. The new species is prob- with almost uniformly dark brown lateral ably most related to the representatives of lobes and spotted (brown with a few light Tramlapiola and Crockeriola (each of these brown marks) disc; tegmina brownish grey, genera is with one species) but clearly dis- rather light and semitransparent but with tinguished from them by the male genital somewhat darker venation in basal and api- characters listed in the description. From cal areas of dorsal field as well as in lateral all the other representatives of Pteroplisti- field; hind wings with almost grey distal nae, the new species also distinctly differs part; legs spotted, i.e. with numerous light in the structure of male copulatory device: brown, brown and dark brown marks; rest male anal and genital plates evidently lack of body more or less uniformly brownish teeth and spines; genitalia are symmetrical, grey. Structure of head and pronotum typi- epiphallus is almost divided into two halves cal of genus Trellius Gor. Tegmina strongly by a rather long median membranous stripe; protruding beyond abdominal apex, rather rachis is long and rather narrow and with wide, with venation of dorsal field almost as thin dorsal process; ectoparameres are un- in Fig. 20, and with lateral field having most developed. Thus, this species may belong to part of costal area very wide and contain- a new genus, but for a more correct decision, ing 21–22 branches of Sc (these branches it is necessary to study additional material. oblique in proximal part and almost trans- Etymology. The new species is named af- verse in rest parts), Sc-R area very narrow, ter the Bu Gia Map National Park. and R-M area moderately wide and crossed by rather sparse and regular crossveins; Subfamily PHALORIINAE hind wings strongly projected behind teg- Gorochov, 1985 minal apices. Legs rather long and thin, Tribe PHALORIINI Gorochov, 1985 but hind femur with distinctly thickened proximal half; fore tibia somewhat widened Trellius (Zatrellius) kinabalu sp. nov. near base, with rather large, elongated and (Figs 20, 21, 29–31) barely immersed inner tympanum, and with Holotype. Male, Malaysia, Sabah State (Bor- slightly smaller, oval and not immersed neo), southern part of Kinabalu National Park on outer tympanum; hind tibia with rather Kinabalu Mt, 1500–2000 m, primary forest, on long and movable dorsal spines, very long

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 50 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD inner dorsal spur reaching apex of second 2.5, female 2.3–2.6; tegmina: male 17–17.5, tarsal segment, comparatively short inner female 16.5–17; hind femora: male 11–11.7, ventral and outer middle spurs, and very female 11.4–11.7; ovipositor 4.9–5.1. short inner dorsal and inner ventral spurs; Comparison. The new species clearly dif- hind basitarsus without distinct denticles fers from T. (Z.) communis (Thailand and but with a pair of spurs (outer spur short; Cambodia) and T. (Z.) andamanensis (An- inner spur almost twice as long as outer daman Islands) in the characters of male one). Abdominal apex also typical of genus genitalia listed above, in the description. Trellius; genitalia similar to those of T. (Z.) Etymology. The new species is named af- communis Gor. and T. (Z.) andamanensis ter the Kinabalu Mount. Gor. in development of a pair of long and characteristically widened distal epiphal- Trellius (Zatrellius) crocker sp. nov. lic lobes (Figs 29, 30), but widened part of (Figs 22, 32–34) these lobes with widely rounded (not angular) proxilateral projection and without Holotype. Male, Malaysia, Sabah State (Bor- any distomedial hook, ectoparameres much neo), Crocker Range National Park not far from Keningau Town, 1000–1300 m, secondary/pri- longer, rachis (= guiding rod) less deeply mary forest, 2–6.V.2013, A.Gorochov, M. Bere- bifurcated, endoparameres larger and with zin, V. Gorochova, E. Tkatsheva (ZIN). much longer apodemes, and rami not fused Description. Male (holotype). Size, co- with each other in anterior part of genitalia louration and structure of body similar to (Figs 29–31). those of males of T. (Z.) kinabalu but with Variations. Second male with barely following differences: epicranium with dark lighter epicranium having slightly darker brown dorsal half (including rostral apex) spot on rostrum and a pair of longitudinal having brownish grey eyes and middle parts lines on posterior half of dorsum, and with of genae, and with light greyish brown ven- less distinctly spotted maxillary palpi. tral half having a pair of dark stripes along Female. General appearance similar to medial edges of antennal cavities and one that of male paratype, but: pronotum less rather small dark area under each eye; an- widened in posterior part; tegmina with tennae and mouthparts light brown with much narrower dorsal field having 12–13 yellowish labrum and almost whitish palpi longitudinal but partly oblique veins (these (but apex of apical segment of maxillary veins barely arcuate in proximal half of this palpus slightly darkened); pronotum dark field) and regular and numerous (but not brown with a pair of rather large light brown very dense) crossveins, with costal area spots on distal half of disc; tegmina without having 17–19 branches of Sc (these branch- darkened marks on basal area of dorsal field es somewhat more oblique than in male), and with lighter venation (all veins of dor- and with widened Sc-R area which almost sal field not darker than its membranes, and twice as wide as R-M area; colouration of veins of lateral field only barely darker than dorsal tegminal field somewhat darker than membranes); venation of dorsal tegminal in males (intensively greyish brown with field as in Fig. 22; tegminal Sc with 23–24 dark venation). Genital plate large, slightly branches; inner dorsal spur of hind tibia longer than wide, somewhat narrowing to slightly protruding beyond apex of second apex, with widely concave posterior edge; tarsal segment; genitalia in dorsal view ovipositor moderately short (hind femur with distal epiphallic lobes more arcuate in approximately 2.3 times as long as oviposi- proximal half and narrower in distal half, tor), with apical part as in Fig. 21. with much smaller (shorter) ectoparameres Length in mm. Body: male 10–11.5, fe- lacking distinct processes projected behind male 10.5–11; body with wings: male 24– bases of above-mentioned epiphallic lobes, 25, female 23.5–25; pronotum: male 2.3– with larger rachis which more deeply bi-

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 51 furcated and strongly curved upwards, and Female. General appearance similar to with somewhat longer rami (Figs 32–34). that of female of T. (Z.) kinabalu, but head Female unknown. dorsum and pronotal disc almost uniformly Length in mm. Body 11.3; body with brown, tegmina light brown with brown wings 23; pronotum 2.1; tegmina 11; hind venation, tegminal venation with 14–15 femora 10.5. branches of Sc, legs with slightly less con- Comparison. The new species is distin- trasting colouration, ovipositor somewhat guished from T. (Z.) kinabalu by the male shorter (hind femur almost 2.6 times as genital characters listed in the descrip- long as ovipositor) and with apical part as tion. From T. (Z.) communis and T. (Z.) in Fig. 24. andamanensis, the new species differs in Length in mm. Body: male 12.5, female the epiphallic distal lobes less widened in 11; body with wings: male 22, female 21.5; the distal half, ectoparameres lacking dis- pronotum: male 2.1, female 2.2; tegmina: tinct processes projected behind the bases male 16, female 15.5; hind femora: male 10, of these epiphallic lobes, rachis strongly female 10.5; ovipositor 4. curved upwards, endoparameres larger and Comparison. The new species is most with longer apodemes, and rami not fused similar to T. (Z.) crocker in the rachis of with each other in the anterior part. male genitalia strongly curved upwards, but Etymology. The new species is named af- it is distinguished from the latter species by ter the Crocker Range. the characters of these genitalia listed in the description. From T. (Z.) kinabalu, T. (Z.) Trellius (Zatrellius) tawau sp. nov. communis and T. (Z.) andamanensis, the (Figs 23, 24, 35–37) new species differs in the same characters as T. (Z.) crocker. Holotype. Male, Malaysia, Sabah State (Bor- Etymology. The new species is named af- neo), Tawau Hills National Park near Tawau ter the Tawau Hills National Park. City, 200–400 m, secondary/primary forest, on leaf of tree near brook at night, 14–20.V.2013, Trellius (Neotrellius) logunovi sp. nov. A. Gorochov, M. Berezin, E. Tkatsheva (ZIN). (Figs 25, 38, 39) Paratype. Female, same locality as in holotype, 6–14.III.2014, M. Berezin. Holotype. Male, Vietnam, Tuyen Quang Description. Male (holotype). Size, co- Prov., 5 km E of Na Hang, 22°20'59''N, 105°25' louration and structure of body very similar 36''E, 290 m, 4–13. XI.2015, D. Logunov (ZIN). to those of male of T. (Z.) crocker but with Description. Male (holotype). Body me- following differences: dorsum of epicranium dium-sized for this genus and with follow- behind lateral ocelli slightly lighter (brown) ing colouration: head light greyish brown but with four dark longitudinal lines; lower with a few greyish brown marks on rostrum, part of epicranium crossed by a pair of dark four longitudinal brown stripes on dorsum brown stripes, running from medial halves behind rostrum, two pairs of brown areas of antennal cavities to clypeal suture, and behind eyes and on lower parts of genae, with dark transverse band dividing lighter four vertical greyish brown stripes on epi- median part of lower rostral surface into cranium under eyes and antennal cavities, two areas (small dorsal area and longer ven- a few small darkened spots on mouthparts tral one); venation of dorsal tegminal field (including somewhat darkened apical parts as in Fig. 23; genitalia with distal epiphal- of maxillary palpi), and almost greyish lic lobes slightly different in shape (Figs 35, brown distal part of antennal flagellum; pro- 36), with rachis having distal processes dis- notum almost completely dark brown but tinctly wider in middle part, and with notch with poorly visible barely lighter marks on between proximal parts of these processes disc and lateral lobes; tegmina light brown much wider (almost round) (Figs 35–37). with brown venation; legs and visible dis-

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 52 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD tal parts of hind wings also light brown, Comparison. The new species is most re- but legs with barely darker spots; venter lated to T. (N.) tonkinensis described from of body light brown with partly darkened another province of Vietnam and belongs to genital plate; rest part of abdominal apex the same subgenus; T. (N.) logunovi differs with more or less darkened areas (including from this species (previously only a single slightly darkened cerci). Rostrum of head species of this subgenus) in the characters between antennal cavities almost equal to of male genitalia listed in the description. scape in width; all ocelli rather small (me- Etymology. The new species is named af- dian ocellus almost equal to lateral ones in ter its collector. size); pronotum similar to that of above- mentioned congeners in structure; venation Trellius (Neotrellius) simulator sp. nov. of tegminal dorsal field as in Fig. 25; tegmi- (Figs 26, 42, 43) nal lateral field with wide costal area having Holotype. Male, Vietnam, Ha Bac Prov., en- 21–22 almost transverse Sc branches, with virons of Chi Linh Vill., X.1997, N. Orlov (ZIN). very narrow Sc-R area (these areas without Paratype. Female, same data as for holotype crossveins), and with rather widened R-M (ZIN). area having regular crossveins; hind wings Description. Male (holotype). General barely projected behind tegminal apices; appearance very similar to that of T. (N.) legs with medium-sized and elongately oval logunovi but with following small differ- inner and outer tympana (inner tympanum ences: head with more distinct (wider) slightly longer than outer one), with dor- light vertical median stripe near rostral sal surface of hind tibia having four pairs of apex, with 18–19 branches of tegminal Sc, rather long and movable articulated spines and with slightly less transverse mirror in as well as moderately numerous and very dorsal tegminal field. Genitalia most simi- small unarticulated denticles mainly on lar to those of T. (N.) tonkinensis; however, dorsolateral edge, and with inner dorsoapi- posterolateral epiphallic lobes (processes) cal spur of this tibia reaching apex of second almost straight in profile, and posterome- segment of hind tarsus. Abdominal apex dial epiphallic lobules distinctly shorter as similar to that of other congeners; genitalia well as rounded (not almost rectangularly most similar to those of T. (N.) tonkinensis projected) posteriorly (for comparison see (Chop.), but distal epiphallic lobes more ar- Figs 40, 41 and 42, 43). cuate in posterior half, dorsoproximal lob- Female. Size, colouration and structure ules of these lobes narrower and thinner in of body similar to those of male, but: pro- profile, posteromedial sclerites of epiphallus notum slightly less widening to prothorax; (ectoparameres?) between bases of above- wings somewhat shorter, but hind wings mentioned epiphallic lobes clearly narrower slightly more protruding beyond tegminal in posterior part, endoparameres (which apices; dorsal tegminal field darker (almost located almost as in this species, i.e. much greyish brown), with convex longitudinal before all posteromedial sclerotized parts vein parallel to lateral edge and located of genitalia) with larger plate-like lateral near this edge (area between these vein and parts, formula with distinctly shorter and edge without crossveins), with 7–8 slightly narrower apodeme, and rami clearly longer oblique longitudinal branches, and with than rest structures of genitalia but much rather numerous (but more or less oblique) shorter that those of T. (N.) tonkinensis (for crossveins between these branches; lateral comparison see Figs 38, 39 and 40, 41). tegminal field light greyish brown with Female unknown. brown venation, with Sc–R area moder- Length in mm. Body 15.3; body with ately wide and having rather sparse cross- wings 21; pronotum 3.2; tegmina 15.7; hind veins, and with 10–11 oblique branches of femora 14.5. Sc (crossveins between these branches ab-

Figs 44–50. Phaloriinae (Phaloriini): 44–46, Phaloria (Papuloria) multa sp. nov.; 47, 48, Borneloria spinosa sp. nov.; 49, B. moorei occidentalis subsp. nov.; 50, B. sp. Male head, pronotum and tegmina with one or some legs, as well as with distal parts of hind wings and cerci in rest position from above (44, 47, 49); female tegmina with distal parts of hind wings in rest position from above (45); female genital plate from below (46, 50); male head with pronotum and fore leg from side as well as slightly in front and slightly from above (48).

Figs 51–62. Phaloriinae (Phaloriini), male: 51–53, Borneloria spinosa sp. nov.; 54–56, B. moorei occidentalis subsp. nov.; 57–61, Phaloria (Papuloria) multa sp. nov. ; 62, B. spinosa. Genitalia from above (51, 54, 57), from below (52, 55, 58) and from side (53, 56, 59); distal part of genitalia from above, schematically (60); left ectoparamere from below (61); genital plate, ventral and lateral views (62). sent). Genital plate almost square but with Length in mm. Body: male 12.5, female narrowing distal part and rather shallow 11; body with wings: male 22, female 21.5; and widely rounded apical notch; distal part pronotum: male 2.1, female 2.2; tegmina: of ovipositor as in Fig. 26. male 16, female 15.5; hind femora: male 10,

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 55 female 10.5; ovipositor 4. Trellius (Protrellius) perbonus Comparison. The new species is most Gorochov, 1999 similar to T. (N.) tonkinensis in the structure and length of its male genitalia but New material. Indonesia: male, Western Java, Cinumpang in environs of Sukabumi City, distinguished from the latter species by the 800 m, forest, 28.III.2003, M. Berezin (ZIN). characters of these genitalia listed in the Note. This species originally described description. From T. (N.) logunovi, T. (N.) from one male with the label “Java” (Goro- simulator differs in the same characters as chov, 1999) is here indicated for a more ex- well as in the male genitalia longer and hav- act locality. ing the epiphallus less distinctly narrowed near the bases of posterolateral epiphallic Trellius (Protrellius) duplicatus lobes (see Figs 39 and 43). Gorochov, 1999 Etymology. This species name is the Lat- in word “simulator” (imitator, shammer). New material. Indonesia: 2 males, Western Java, 20–25 km SE of Bogor City, environs of Trellius (Trellius) vitalisi (Chopard, 1925) Cemande Vill. in Pangrango Mts, 1000 m, secondary forest, on leaves of trees near small river, New material. Vietnam: male, Vinh Phu 27.XI–7.XII.1999, A. Gorochov (ZIN). Prov., environs of Tam Dao Vill., 800–900 m, Note. This species originally described primary/secondary forest, on leaf of tree near also from Java without other data (Goro- small river at night, 1–11.VI.1995, A. Gorochov chov, 1999) is here recorded from a more (ZIN); 2 male and female, Cao Bang Prov., Nguen exact locality. Binh Distr., Quang Thanh Vill., 4–13.V.1998, N. Orlov (ZIN); male, Ha Tinh Prov., Huong Son Vill. on Rao An River, 18°21'N, 105°13'E, pri- Genus Borneloria gen. nov. mary forest, IV.2000, N. Orlov (ZIN). Type species Borneloria spinosa sp. nov. Note. These specimens are very similar Diagnosis. Body rather small and slen- to each other and to the original description der. Colouration characteristic, contrasting: of this species, but they have very small dif- dorsum of head, pronotal disc and tegmina ferences in the body colouration and shape more or less greyish brown, but head with of some male genital structures. I cannot almost yellowish line separating this greyish exclude that they may be divided into a few brown part from nearest parts; lower half of subspecies; however, this question is in need head and band on pronotal lateral lobe along of additional data. This North Vietnam- its ventral edge very light; areas on both ese species was described from “Hoa Binh” head and pronotum situated between grey- (Chopard, 1925) and recorded from the Bak ish brown and very light parts dark brown Thai Province (Gorochov, 1988a). Here, it or blackish; hind tibiae and tarsi also dark or is indicated for three other localities of the very dark but with three whitish transverse North Vietnam. bands (two of them on tibia, and one band on basitarsus). Rostrum of head rather long Trellius (Protrellius) siveci (longer than in most genera of this tribe), in Gorochov, 1996 profile with more or less angular apex and New material. Malaysia: male, Penang I. be- oblique anterior edge between rostral apex tween Malacca and Sumatra, Penang Mt, 800 m, and lower part of epicranium (Figs 47–49); secondary forest, on leaf of bush near brook at pronotum distinctly widening to pterotho- night, 20–24.XI.2014, A. Gorochov, M. Berezin, rax, with sinuate ventral edges of lateral E. Tkatsheva (ZIN). lobes (Fig. 48); wings completely developed, Note. This species originally described typical of this tribe (Figs 47, 49); legs long from Malacca (Perak State) is here record- and very thin but with somewhat thickened ed from the Penang Island for the first time. proximal two thirds of hind femur; fore tibia

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 56 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD with two oval tympana on both sides; hind much lower. From Pseudotrigonidium tibia with four pairs of rather short and mov- Chop., the new genus differs in the presence able dorsal spines, more numerous and very of both tympana (not only inner one) and in small dorsal denticles, and six apical spurs the formula of male genitalia having the un- very inequal in length; abdominal apex also paired apodeme directed backwards but not typical of Phaloriini, but male genital plate forwards or upwards; from Phaloria Stål and elongate and with almost truncated apex Strophiola Uv., in the endoparameres fused having one small median angular lobule with each other; from Trellius and Ceyloria curved upwards/forwards (almost hook- Gor., in the epiphallus undivided into two like; Fig. 62), and female genital plate mod- or more isolated sclerites; from Vescelia Stål erately large and slightly wider than long as and Tremellia Stål, in the ectoparameres well as with shallow rounded posteromedian much shorter and in the formula of male notch and short rounded lobes around it genitalia more strongly notched; and from (Fig. 50). Male genitalia peculiar: epiphallus all these genera, in the head rostrum more almost plate-like and semimembranous but or less longer and with the anteroventral with a pair of moderately long and sclero- edge more oblique in the profile. tized distal lobes somewhat curved upwards Etymology. The new generic name con- and having slightly widened (in profile) dis- sists of parts of the following names: Borneo tal parts; ectoparameres small, plate-like and (geographic name) and Phaloria (generic almost not separated from bases of epiphal- name). lic distal lobes; endoparameres fused with each other, with rather long posterior arms Borneloria spinosa sp. nov. and anterior apodemes; rachis rather large (Figs 47, 48, 51–53, 62) (long), gradually narrowing to almost spine- Holotype. Male, Malaysia, Sabah State (Bor- like apex and more or less fused with endo- neo), Tawau Hills National Park near Tawau parameral median lobe near rachial base; City, 200–400 m, secondary/primary forest, on formula deeply notched in anterior part and leaf of bush near brook at night, 14–20.V.2013, with rather thin unpaired apodeme directed A. Gorochov, M. Berezin, E. Tkatsheva (ZIN). backwards; rami long and thin (Figs 51–56). Description. Male (holotype). Antennae Ovipositor moderately long (but distinctly with scape light greyish but having large shorter than hind femur) and with clearly greyish brown dorsomedial area, with pedi- drilling apex (Fig. 27). cel also greyish brown, and with flagellum Included species. Type species and Hete- uniformly light yellowish grey (Figs 47, 48). rotrypus moorei Chopard, 1940 (Borneo). Tegmina distinctly longer than body; their Comparison. The new genus is most sim- dorsal field light greyish brown but semi- ilar to the genus Sumatloria Gor. in the gen- transparent, with venation as in Fig. 47; eral appearance, but it is distinguished from tegminal lateral field with band along dor- the latter by the epiphallus larger (longer), sal edge from intensively brown at base to ectoparameres much shorter, distal half of light brownish grey in distal part, with rest rachis distinctly narrower, median endo- areas of this field whitish, and with vena- parameral part clearly larger, formula differ- tion similar to that of T. (Z.) kinabalu; hind ent in the shape, and its unpaired apodeme wings distinctly protruding beyond tegmi-

Figs 63–72. Phaloriinae (Subtiloriini): 63, Subtiloria semota sp. nov.; 64, 65, S. succinea korup subsp. nov.; 66, Schizotrypus conradti nigericus subsp. nov.; 67, Sch. c. conradti (Gor.); 68–70, Afrophaloria malawi sp. nov.; 71, 72, A. dja sp. nov. Male body in rest position from above, without some legs (63, 64, 66, 68) and without head, pronotum and all legs (67); male anal plate without lateral parts from behind (65); anterior half of body from side, male (69) and female (72); female genital plate from below (70, 71).

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 59 nal apices and with almost dark grey distal Description. Male (holotype). General parts (Fig. 47). Fore and middle legs light appearance very similar to that of B. spi- yellowish grey but with slightly darkened nosa, but with following differences: grey- rings around tympana and two elongate ish brown areas on dorsal parts of both head dorsal spots on tibiae (on their middle and and pronotum slightly lighter and with distal parts), with greyish ventral mark on barely wider yellowish line separating such distal part of middle femur and dorsal mark area on head from darker parts (Fig. 49); an- on proximal part of middle tibia, and with tennal scape completely brown; distal half barely darkened distal two thirds of fore and of antennal flagellum somewhat darkened; middle tarsi; hind femur yellowish with two middle femur with almost complete brown- darkish areas in middle third and almost ish grey ring near apex; tegmina with dorsal dark brown distal femoral portion (Fig. 47); field yellowish grey and venation as in Fig. outer tympanum oval but not very long; 49; genitalia with anterior edge of epiphal- inner tympanum slightly larger (longer) lus having slightly deeper and almost an- and almost not immersed. Cerci and ven- gular median notch, with distal epiphallic ter of body yellowish grey, light; distal part lobes lower and having less widened distal of abdominal dorsum somewhat darkened. part, and without spine on ventromedial Genitalia with anterior edge of epiphallus edge of each distal epiphallic lobe (however, having shallow and rounded median notch, apodemes of endoparameres and of formula with distal epiphallic lobes rather high and almost as long as in B. spinosa, and apex of having distinctly widened (in profile) distal rachis not bifurcated, similar to that of this parts, and with one distinct spine on ven- species) (Figs 54–56). tromedial edge of distal part of each distal Female unknown. epiphalic lobe (Figs 51–53). Length in mm. Body 10; body with Female unknown. wings 17; pronotum 2.1; tegmina 12; hind Length in mm. Body 10; body with femora 9.5. wings 17; pronotum 2.1; tegmina 12; hind Comparison. The new subspecies is de- femora 9.5. scribed from the western part of Sarawak Etymology. This new species is named by and differs from the nominotypical subspe- the Latin word “spinosa” (spinose, spiked) cies, known only from the eastern part of in connection with the presence of charac- Sarawak (Dulit Mt), in the apical area of teristic spines on the distal epiphallic lobes male tegminal dorsal field somewhat longer in male. [in B. m. occidentalis, this area is approximately 1.1 times as long as the mirror, but Borneloria moorei occidentalis in B. m. moorei (Chop.), such ratio is about subsp. nov. 0.8], distal epiphallic lobes more strongly (Figs 49, 54–56) curved upwards, apodemes of endoparameres and of formula longer, and apex of rachis Holotype. Male, Malaysia, Sarawak State acute, spine-like (in B. m. moorei, this apex (Borneo), environs of Kuching City, Kubah National Park on Matang Mt, 200–500 m, pri- is barely bifurcated; see Gorochov, 1999: mary forest, on leaf of bush near brook, 27.XI–1. figs 39–45). From B. spinosa, the new sub- XII.2016, A. Gorochov, M. Berezin, E. Tkatshe- species is distinguished by the characters va, I. Kamskov (ZIN). listed in the description.

Figs 73–85. Phaloriinae (Subtiloriini): 73–77, Subtiloria semota sp. nov.; 78–81, S. succinea korup subsp. nov.; 82–84, S. subtilis (Gor.); 85, Schizotrypus conradti nigericus subsp. nov. Male genitalia from above (73, 78, 82), from below (74, 79, 83) and from side (75, 80, 84); male (76, 81, 85) and female (77) genital plates from below.

Etymology. The new subspecies name is 122°34'12''E, secondary/primary forest on hills, the Latin word “occidentalis” (western). on leaf of bush near brook at night, 21.IX– 2.X.2015, A. Gorochov, M. Berezin, I. Kamskov, E. Tkatsheva (ZIN). Borneloria sp. Paratypes. Indonesia: 6 males and 4 fe- (Figs 27, 50) males, same data as for holotype (ZIN); 2 males, Southeast Sulawesi, ~30 km SE of Kendari New material. Indonesia: female, East Kali- City, environs of Moramo Waterfall, 4°13'33''S, mantan (Borneo), ~20 km N of Balikpapan City, 122°44'9''E, primary forest on hill, on leaves Bukit Bangkirai Park, 1°1'43''S, 116°51'49''E, of bushes at night, 29.IX.2015, A. Gorochov, secondary forest on hills, on leaf of bush at night, M. Berezin, I. Kamskov, E. Tkatsheva (ZIN); 4–8.X.2015, A. Gorochov, M. Berezin, I. Kams- 2 males and 2 females, Buton I. near Southeast kov, E. Tkatsheva (ZIN). Sulawesi, environs of Baubau Town, 5°28'55''S, Note. This female is very similar to the 122°38'18''E, secondary forest on hills, on other congeners, described only from males, leaves of bushes near water at night, 25–26. and cannot be determined more exactly. It XI.2015, A. Gorochov, M. Berezin, I. Kamskov, is the first record of this genus from the In- E. Tkatsheva (ZIN). donesian part of Borneo. Description. Male (holotype). General appearance more or less similar to that Phaloria (Papuloria) tristis of Trellius representatives previously dis- Gorochov, 2014 cussed here: colouration of body light greyish brown with characteristic brown marks New material. Malaysia: 3 males and female, on head dorsum (Fig. 44), a pair of brown Sabah State (Borneo), Tawau Hills National Park near Tawau City, 200–400 m, primary/sec- spots behind eyes, six greyish brown ver- ondary forest, on leaves of bushes at night, 14– tical stripes on lower part of epicranium 20.V.2013, A. Gorochov, M. Berezin, E. Tkatshe- (a pair of medial stripes running to rostral va (ZIN). apex), two pairs of slightly darkened spots Note. This species, originally described on upper half of clypeus and on subgenae, from two localities in Sarawak and one lo- greyish lateral parts of labrum, small brown cality in Sabah (Gorochov, 2014), is here mark on outer surface of third segment of recorded from another locality of the Sabah maxillary palpus, greyish brown distal two State. thirds of antennal flagellum, dark brown pronotum having three small lighter spots Phaloria (Papuloria) paratristis on each lateral lobe, distinctly spotted (with Gorochov, 2014 brown and yellowish spots) dorsal tegminal field (Fig. 44), almost transparent lateral New material. Malaysia: male, Sarawak State tegminal field having brown to light brown (Borneo), 80–90 km WWN of Kuching City, venation, brownish grey distal parts of hind Gunung Gading National Park, 100–300 m, pri- wings visible behind tegminal apices, dis- mary/secondary forest, on leaves of bushes at tinctly spotted legs, and barely darkened night, 22–26.X.2016, A. Gorochov, M. Berezin, E. Tkatsheva, I. Kamskov (ZIN). dorsum of abdominal apex (but cerci light); Note. The species was originally de- scape approximately as wide as rostrum scribed from three localities in Sarawak between antennal cavities; mediam ocellus (Gorochov, 2014). Now it is indicated for much smaller than lateral ones; pronotum another locality of the same state. less widening to pterothorax (Fig. 44); tegmina with somewhat narrower dorsal field (its venation as in Fig. 44) and with lateral Phaloria (Papuloria) multa sp. nov. field having 25–26 Sc branches; hind wings (Figs 28, 44–46, 57–61) distinctly protruding beyond tegminal api- Holotype. Male, Indonesia, Southeast Su- ces; tympana oval, but outer tympanum lawesi, environs of Kendari City, 3°57'28''S, distinctly shorter than inner one, i.e. less

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 61 elongated than in above-mentioned repre- er widely truncated apex and having very sentatives of Trellius; armament of hind tibia small (narrow) posteromedian notch (Fig. similar to that of T. (N.) logunovi. Genitalia 46); ovipositor rather short (hind femur characteristic of this subgenus of Phaloria approximately twice as long as ovipositor) but with following features: epiphallus nar- and with distal part as in Fig. 28. rowing to apex, with long and rather wide Length in mm. Body: male 16–17, fe- (high) anterolateral parts, with short (but male 13.5–15; body with wings: male 21– not very short) median part, with moder- 23, female 19–22; pronotum: male 3.2–3.4, ately short posterolateral (distal) lobes hav- female 3–3.2; tegmina: male 18–19, female ing slightly concave proximedial edges and 17.5–18.5; hind femora: male 14.5–15.5, fe- convex distomedial edges as well as almost male 13–14; ovipositor 6.5–7. straight (oblique) lateral edges and more Comparison. The new species is most or less angular apical parts; each ectopara- similar to Ph. (P.) aspersa Gor. widely dis- mere rather small and in shape of oblique tributed in the Papuan Region (New Gunea ribbon having four small distal tubercles and nearest islands), but it differs from the (two apical tubercles situated very near latter species in the distal epiphallic lobes each other, and two subapical ones slight- shorter, rachis smaller, and ectoparameres ly larger than apical tubercles and located almost straight (not slightly S-shaped). dorsally; Fig. 61); each endoparamere with From some other species of this subgenus two rather short and curved posterior arms having similar shape of the ectoparameres, connected with base of ectoparamere and Ph. (P.) multa is distinguished by the ab- with base of rachis (but not with analogous sence of large apical spine on the ectopara- arms of opposite endoparamere), and with mere (from Ph. solita Gor. distributed in wide but not very long anterior apodeme; the same region), by the presence of apical rachis moderately large, slightly elongated ectoparameral tubercles (from Ph. vulgata (almost subsquare in shape), with rather Gor. with similar distribution), by the dis- deep and moderately wide posteromedian tal epiphallic lobes shorter (from Ph. tariku notch, with slightly longer and wider an- Gor.; New Guinea) and not bilobate (from teromedian notch, and with lamellar and Ph. verecunda Gor.; New Guinea), by the almost acute apical lobules; formula rather apical lobules of rachis not very long and large, in shape of roundly square semisclero- not curved (from Ph. pulchra Gor., Ph. rava tized plate having posteromedian part more Gor., Ph. pararava Gor., Ph. neorava Gor. sclerotized; rami fused with epiphallus (but and Ph. gilva Gor.; New Guinea), and by the not with each other) and distinctly shorter apical area of male dorsal tegminal field long than this structure (Figs 57–61). (from Ph. curta Gor. having this area signifi- Variations. Dark marks on head some- cantly shortened; New Guinea). From Ph. times barely larger or smaller; pronotal disc sulawesi Gor. (Northeast Sulawesi) with in some males with traces of small lightish unclear subgeneric position, the new spe- spots; size of tegminal mirror, number of Sc cies differs in the pronotum darker, tegmina branches in tegmina and armament of hind with the dorsal field spotted (not unicolor- legs insignificantly varied. ous) and with the venation of lateral field Female. Size, colouration and structure darker, female genital plate truncated and of body similar to those of male, but struc- with a small (narrow) posteromedian notch ture of tegmina more similar to that of fe- (vs. this plate strongly and widely concave males of T. (Z.) kinabalu and T. (Z.) tawau at the apex), and ovipositor longer. From (Sc having 15–17 branches, and dorsal field the other similar congeners, Ph. (P.) multa with 10–12 longitudinal veins; Fig. 45); is distinguished by the following combina- genital plate moderately large, almost as tion of characters: ectoparameres ribbon- long as wide, distinctly narrowing to rath- like and obliquely situated, distal epiphal-

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 62 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD lic lobes not very long and not bifurcated, a. Sepik”; “Supiori I. not far from northern endoparameral apodemes not very wide and coast of New Guinea, environs of vill. Ko- not very long. rido”). Etymology. The new species name is the Latin word “multa” (numerous). Phaloria (Papuloria) ostensa Gorochov, 1999 Phaloria (Papuloria) vulgata New material. Indonesia: 8 males and 5 fe- Gorochov, 1996 males, Papua Prov. (New Guinea), environs New material. Indonesia, Papua Prov. (New of Fawi [= Faowi] Vill. in upper part of Tariku Guinea): male and female, Bagusa Vill. near River (tributary of Mamberamo River), partly mouth of Mamberamo River, lowlying for- lowlying forest / partly forest on hills, on leaves est, 22–24.I.2012, A. Gorochov (ZIN); 2 males of bushes and trees at night, 29.I–17.II.2012, and female, Kasonaweja Vill. on Mamberamo A. Gorochov (ZIN). River near Van Rees Range, forest on hills, 25– Note. This species, originally described 27.I.2012, A. Gorochov (ZIN); 3 males and 7 fe- from “Torecella Mt., Afua” in New Guinea males, environs of Fawi [= Faowi] Vill. in upper (Gorochov, 1999), is here recorded from an- part of Tariku River (tributary of Mamberamo other locality. River), partly lowlying forest / partly forest on hills, 29.I–17.II.2012, A. Gorochov (ZIN). Phaloria (Papuloria) harzi Note. This species, widely distributed Gorochov, 1999 in Papuan Region (New Guinea, Solomon Islands and some other nearest islands; New material. Indonesia: male and 2 fe- Gorochov, 1996, 1999, 2005), is here re- males, Papua Prov. (New Guinea), Waena Vill. corded from three new localities. These near Jayapura City, 9–13.VIII.2012, N. Kluge, specimens have very small differences L. Sheyko (ZIN). Note. Here, this species is recorded from in the body colouration as well as in the a new locality situated not far from its shape of rachial apex; possibly this species type locality (Gorochov, 1999: “Kyklop Be. may be divided into a few subspecies, but [= Cyclops Mts]”). The male studied has their description is now premature due to its genitalia also barely distinguished from the presence of specimens with intermedi- those of Ph. (P.) harzi holotype, but pos- ate characters. sibly these differences are connected with insufficient sclerotization of the first geni- Phaloria (Papuloria) aspersa talia. Gorochov, 1996

New material. Indonesia, Papua Prov. (New Tribe SUBTILORIINI Gorochov, 2003 Guinea): 9 males and 6 females, environs of Fawi (2011) [= Faowi] Vill. in upper part of Tariku River (tributary of Mamberamo River), partly lowly- Subtiloria semota sp. nov. ing forest, on walls of small cave among rocks (Figs 63, 73–77, 102, 103) along river at daytime, 29.I–17.II.2012, A. Goro- chov (ZIN); 1 male and 1 female, environs of Holotype. Male, Cameroon, Southwest Re- Nabire Town, secondary forest on hill, on rocks gion, Korup National Park not far from Nigeria, along brook at night, 28.II–2.III.2012, A. Goro- ~300 m, primary forest, on leaf of bush at night, chov (ZIN). 1–8.II.2016, A. Gorochov (ZIN). Paratypes. Cameroon: male and 2 females, Note. This species is here recorded from same data as for holotype (ZIN); female, bor- two localities different from those indicated der of South and East Regions, Dja Reserve on previously (Gorochov, 1996, 2005: “Hol. Dja River, ~600 m, primary / secondary forest, N. Guinea, Pauwi-Samberi”; “D.N. Guinea on leaves of bushes at night, 15–22.II.2016, i. Lager 29 km unterh. d. Mäanderberges A. Gorochov (ZIN).

Description. Male (holotype). Body solateral denticles only (except for a pair of rather large for this genus. Colouration apical spurs); anal plate without spines but following: head light yellowish (almost with posterior part almost roundly trun- whitish) with light yellowish grey dorsum cated at apex; genital plate almost rectan- having slightly darker (brownish grey) dor- gular but somewhat semitubular and with sal part of rostrum and four longitudinal slightly concave posterior edge (Fig. 76). stripes on dorsum behind rostrum, with a Genitalia characteristic: epiphallus short, pair of greyish stripes on anterior surface transverse, partly membranous (with weak- of rostrum (these stripes running from me- ly sclerotized anterior, median and poste- dian ocellus to almost clypeal suture), with rior parts) and with small angular postero- a few light greyish marks under eye and an- median notch; ectoparameres absent; endo- tennal cavity, with brownish grey antennal parameres rather short (their posterolateral flagellum (except for its small basal part), arms distinctly not reaching posterior edge as well as with dark brown eyes and whit- of epiphallus), widely separated from each ish ocelli; pronotum brownish grey (but other but having short medial arms fused rather light) with very light (yellowish) with rachis, and having small anterolateral band on each lateral lobe along its ventral apodemes; rachis very large, having a pair of edge and with several very light marks on long posterolateral processes (located not disc; tegmina almost brownish grey with near each other) and much shorter postero- yellowish venation, slightly darker spots on median lobe (this lobe clearly wider than some membranes of dorsal field, and almost these processes) as well as low longitudinal transparent membranes of lateral field; ventromedian keel, and fused with rather distal parts of hind wings darker (greyish short formula (anterior part of formula brown) with whitish venation; legs light barely bilobated); rami similar to those of with brownish grey spots and stripes; rest other congeners but distinctly shorter than of body light brown with yellowish thoracic in S. stena Gor. (Figs 73–75). sternites and distal part of genital plate, as Variations. One male paratype with well as with greyish tinge on anal plate and darkened parts of body slightly darker, but cerci. Structure of body more or less simi- second one with light marks larger; dorsum lar to that of other congeners, but: scape of both pterothorax and abdomen partly or almost 1.2 times as wide as space between almost completely darkened. antennal cavities; ocelli large and almost Female. General appearance as in males, equal to each other in size; eyes very large but tegmina barely darker and without (high); pronotum transverse and moderate- stridulatory apparatus (dorsal field with ly widening to pterothorax; tegmina long 9–10 longitudinal but slightly arcuate and and with venation as in Fig. 63; hind wings oblique veins, and with rather sparse cross- strongly protruding beyond tegminal api- veins), tegminal areas located near dorsal ces; legs very slender but with hind femora edge of lateral field moderately narrow and somewhat thickened in proximal half; inner with sparse crossveins, tegminal Sc with tympanum distinct (shortly oval, opened); 10–11 oblique veins (crossveins between outer tympanum larger (longer) and partly them absent), spots on legs slightly more covered with large and inflated lobe of fore distinct, and anal plate shorter; genital plate tibia; hind tibia with four pairs of long mov- longer than wide, slightly narrowing to able spines, six apical spurs (inner spines apex, and with rather wide and moderately and spurs clearly longer than outer ones; in- deep apical notch (Fig. 77); ovipositor with ner dorsal spur longest, protruding slightly distal part as in Figs 102 and 103. beyond second tarsal segment), and 4–5 Length in mm. Body: male 10–12, female small dorsolateral denticles more proximal 10–11; body with wings: male 17–20.5, than spines; hind basitarsus with 6–7 dor- female 17–19; pronotum: male 1.9–2.1, fe-

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 64 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD male 2–2.1; tegmina: male 13–14, female and rather large (distances between them 12–12.5; hind femora: male 8.5–9.3, female almost equal their diameters); scape more 8.5–8.8; ovipositor 7–7.5. or less equal to space between antennal Comparison. The new species is clearly cavities in width; venation of dorsal tegmi- distinguished from all the congeners by the nal field as in Fig. 64; lateral tegminal field absence of spines on the male anal plate, and with very narrow Sc-R area lacking cross- by the rachis of male genitalia rather wide veins, and with 19–20 oblique branches and having posterolateral processes located of Sc (areas between these branches also not near each other. without crossveins); hind wings strongly Etymology. The new species is named by protruding beyond tegminal apices; inner the Latin word “semota” (distant, dissimi- tympanum opened, rather small and round; lar). outer tympanum with membrane larger and oval as well as with strongly inflated Subtiloria succinea korup subsp. nov. lobe near this membrane (this tympanum (Figs 64, 65, 78–81) partly opened and with strongly immersed membrane); hind tibia with four pairs of Holotype. Male, Cameroon, Southwest Re- rather long and movable spines, six apical gion, Korup National Park not far from Nigeria, spurs, a few very small dorsolateral and ~300 m, primary forest, on leaf of bush at night, dorsomedial denticles proximal in relation 1–8.II.2016, A. Gorochov (ZIN). to these spines, and inner dorsal (longest) Description. Male (holotype). Body spur reaching base of second tarsal seg- rather small for this genus, slender. Colou- ment; hind tarsus with 4–5 dorsal denticles ration following: head yellowish with grey- located almost on median line of basitarsus ish brown eyes, thin transverse stripe along and with a pair of moderately long apical upper edge of median ocellus, and small spurs on this basitarsus (inner spur almost spot on upper part of membrane of each an- twice as long as outer one); anal plate weak- tennal cavity, with barely darkened dorsal ly elongate, slightly narrowing to widely surface of rostrum, four short longitudinal truncated apex, and with one row of small stripes on posterior part of epicranial dor- and densely situated spinules at this apex sum, and a pair of oblique bands running (Fig. 65); genital plate slightly elongate, from lower parts of eyes to middle part of barely narrowing to widely notched apex clypeus, and with light greyish brown to (apical notch very wide, roundly trapezoi- greyish brown most part of antennal fla- dal and moderately deep; Fig. 81). Genita- gellum (short basal part of this flagellum lia most similar to those of nominotypical lighter); pronotum also barely darkened subspecies, but posterior part of epiphal- (greyish), but longitudinal ventral band lus somewhat narrower, ectoparameral on each lateral lobe as well as a few marks sclerotizations clearly not reaching apices on disc and between disc and lateral lobes of posterolateral membranous lobes, and lighter (yellowish); tegmina greyish with posteroventral processes of rachis slightly light yellowish (almost whitish) venation S-shaped (not almost straight) in profile and almost transparent areas between Sc (Figs 78–80). branches; hind wings with distal parts al- Female unknown. most brownish grey and having whitish ve- Length in mm. Body 10.5; body with nation (Fig. 64); venter of body (including wings 16.5; pronotum 1.6; tegmina 11; hind genital plate) and cerci yellowish; posterior femora 8. part of abdominal dorsum barely darkened Comparison. The new subspecies is dis- and with apical part of anal plate greyish tinguished from S. s. succinea (Bol.) (Equa- brown. Structure of body typical of Sub- torial Guinea) by the mirror of male tegmi- tiloria, but: all ocelli almost equal in size na slightly larger (longer), and by the male

Figs 86–103. Phaloriini (Subtiloriini): 86–89, Schizotrypus conradti nigericus subsp. nov.; 90–93, Sch. c. conradti (Gor.); 94–98, Afrophaloria malawi sp. nov.; 99–101, A. dja sp. nov.; 102, 103, Sub- tiloria semota sp. nov. Male genitalia from above (86, 90, 94), from below (87, 91, 95) and from side (88, 92, 96); posteromedian projection of rachis from side and slightly above (89, 93); distal part of ovipositor from above (97, 100, 102) and from side (98, 101, 103); distal half of female copulatory papilla from below, schematically (99).

Figs 104–111. Cacoplistinae (Homoeogryllini): 104, Homoeogryllus xanthographus Guér.-Mén.; 105, H. ambo sp. nov.; 106, H. reticulatus limbe subsp. nov.; 107, H. r. reticulatus (F.); 108–111, H. gabonensis Des. Dorsal field of right male tegmen (104–108); female body from above (109); distal part of ovipositor, reconstructed view from above (110) and original view from side (111).

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 67 genital characters listed above (in its de- nominotypical subspecies, but: tegminal scription). From the other congeners with mirror somewhat shorter (for comparison the male genitalia similar, S. s. korup differs see Figs 66 and 67); anal plate with very in the ocelli larger, ectoparameral sclero- shallow and asymmetrical apical notch and tizations shorter, and rachis longer [from without any armament; genital plate with S. subtilis (Gor.), Nigeria and Cameroon; see somewhat deeper apical notch (Fig. 85); Figs 78–80 and 82–84], in the posterolat- and genitalia with anteromedian epiphallic eral lobes of male genitalia clearly shorter plate having posteromedian part very nar- [from S. villosa (Chop.), Congo], and in the row, with lateral epiphallic plates (located apical notch of male genital plate less shal- around latter part) having lateral arms nar- low, proximal part of tegminal mirror (be- row (short), and with posteromedian angu- fore dividing veins) shorter, and distal part lar process of rachis almost acute in profile of this mirror (behind dividing veins) lon- and slightly shorter (Figs 86–89). ger [from S. angusta (Chop.), San Tome I.]. Variations and female. Differences of all Etymology. This subspecies is named af- above-mentioned males in external struc- ter the Korup National Park. ture from holotype of nominotypical subspecies as well as description of female para- Schizotrypus conradti nigericus type given in one of previous publications subsp. nov. (Gorochov, 1999: Sch. conradti). (Figs 66, 85, 86–89) Length in mm. Body: male 13.5–16.5, Holotype. Male, Nigeria, Cross River State, female 15; body with wings: male 23–24, “900–1100 km Bueoro”, 8.XI.1983 (ZIN). female 25; pronotum: male 2.2–2.3, female Paratypes. Nigeria, same state: male, “2000, 2.6; tegmina: male 17–18, female 17.7; hind 27 km NE Calabar”, 22.XI.1983 (BMNH); fe- femora: male 11–11.3, female 11.2; oviposi- male, “80 km E of Calabar”, 3.V.1982 (BMNH). tor 11. Description. Male (holotype). General Comparison. The new subspecies differs appearance very similar to that of nomi- from nominotypical one in the male tegmi- notypical subspecies. Body medium-sized; nal mirror shorter, male anal plate without colouration yellowish with brown eyes, any armament, and male genitalia with the four slightly brownish longitudinal lines on differences listed above (especially in the posterior half of head dorsum, light brown posteromedian rachial projection more median longitudinal stripe on rostral dor- acute; see Figs 86–89 and 90–93). sum and small area between lateral ocelli, a Etymology. This subspecies is named af- few barely distinct darkish marks on lower ter the Nigeria Country where it was col- half of anterior part of epicranium, greyish lected. brown antennal flagellum, almost transparent membranes in tegminal mirror and Schizotrypus conradti conradti between tegminal branches of Sc, barely (Gorochov, 1996) distinct darkish spots on femora and hind (Figs 67, 90–93) tibia (but on hind leg, some of these spots slightly more distinct), brownish grey dis- New material. Cameroon: male and 2 females, tal half of fore and middle tibiae as well as Southwest Region, Korup National Park not far dorsal part of rest half of fore tibia (except from Nigeria, ~300 m, primary forest, at light, for small basal part which light) and distal 1–8.II.2016, A. Gorochov (ZIN). half of fore and middle tarsi (middle tibia Note. This subspecies originally de- also with slightly darkened spot on dorsal scribed as a species from “N. Kamerun, Job- surface of its proximal half), and barely Albrechtshöhe” (Gorochov, 1996: “Hetero- darkened posterolateral edges of anal plate. trypus conradti”) is here recorded from an- Structure of body also similar to that of other locality of this country.

Schizotrypus planus Gorochov, 1999 and middle femora as well as on fore and middle tibiae, with dense and barely dis- New material. Cameroon: male, Southwest tinct darkish oblique stripes on dorsal and Region, Korup National Park not far from Ni- outer surfaces of hind femur, with slightly geria, ~300 m, primary forest, on leaf of bush at night, 1–8.II.2016, A. Gorochov (ZIN). darkened distal part of this femur, with Note. This male is very similar to the ho- more spotted hind tibia having distal part lotype of Sch. planus from Nigeria, but its darkened (proximal two thirds of its spines rostrum has less contrasting spots, and its also darkened), and with all basitarsi partly male genitalia are slightly narrower. How- darkened; pleurites and sternites yellow- ever, there are only two males of this species ish to light brown; anal and genital plates from different localities, i.e. this material is as well as cerci almost greyish brown but insufficient for understanding their subspe- with light cercal bases (Figs 68, 69). Head cies position. typical of Subtiloriini, with scape slightly wider than space between antennal cavities, with ocelli medium-sized and almost Heterotrypus laqueatus Karsch, 1893 equal in size, and with maxillary palpus New material. Cameroon: male, Southwest rather long and thin (its fifth segment with Region, Korup National Park not far from Ni- oblique subapical part and almost trans- geria, ~300 m, primary forest, on leaf of bush versally truncated apical part; these palpi at night, 1–8.II.2016, A. Gorochov (ZIN); 2 fe- more or less similar to those of A. hempae males, border of South and East Regions, Dja Des.-Grand.). Pronotum barely narrowing Reserve on Dja River, ~600 m, primary / second- to head, with almost square disc and moder- ary forest, on leaves of bushes at night, 15–22. ately low lateral lobes; these lobes with ven- II.2016, A. Gorochov (ZIN). tral edge weakly oblique and having poste- Note. This species is here recorded from rior corner distinctly more widely rounded two new localities (it was previously known than anterior one (Figs 68, 69). Legs mod- from Ghana, Equatorial Guinea and “Barom- erately long, with inner tympanum small bi-Stat.” of Cameroon; Gorochov, 1996). and rounded, without outer tympanum, with distinctly thickened proximal half of Afrophaloria malawi sp. nov. hind femur, with four pairs of moderately (Figs 68–70, 94–98) long and movable spines on hind tibia, with Holotype. Male, Malawi, northern part, “Mu- a few spinule-like denticles on this tibia be- ghese FR”, 28–29.XII.2010, A. Sochivko (ZIN). fore above-mentioned spines, with longest Paratype. Female, same data as for holotype (inner dorsal) spur of this tibia distinctly (ZIN). not reaching second tarsal segment, with Desription. Male (holotype). Body com- six spinule-like dorsolateral and one simi- pletely apterous, slightly robust. Upper half lar dorsomedial denticle on hind basitarsus of head, entire pronotum and all other ter- (latter denticle located near apical spurs of gites uniformly brown (almost dark brown); this basitarsus), and with second segment of eyes and ocelli greyish; rest of head yellow- all tarsi almost not widened (approximately ish with a pair of brown vertical stripes on as in genus Kameruloria Gor.). Anal plate anterior surface of rostrum running from simple, rather small and with rounded apex; median ocellus to almost clypeal suture, genital plate elongate (almost twice as long with brown genae having lower parts yel- as previous plate), semitubular, somewhat lowish, with light brown to greyish brown narrowing to almost truncated apex; geni- antennal flagellum (but its short proximal talia as in Figs 94–96. part yellowish), and with darkened distal Female. General appearance very simi- parts of mandibles; legs light brown with lar to that of male, but vertical stripes on slightly darker spots on distal half of fore anterior surface of epicranium less distinct;

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 69 genital plate short (transverse), narrowing yellowish but having a pair of darkish dots to rounded distal part having moderately in lateral corners of clypeus and darkened wide and rather deep apical notch (Fig. 70); apical parts of mandibles, and with middle ovipositor slightly shorter than hind femur, and distal parts of antennal flagellum grey- thin but with distal part as in Figs 97 and 98. ish brown; pronotum dark brown with a pair Length in mm. Body: male 13.5, female of reddish brown marks on disc; other ter- 15.5; pronotum: male 2.9, female 3.6; hind gites also dark brown but with lightish ar- femora: male 13.4, female 13.6; ovipositor eas on anterolateral parts of metanotum and 12.5. on lateral parts of ninth abdominal tergite; Comparison. The male genitalia of A. ma- legs with more contrasting spots on fore lawi are similar to those of A. hempae and and middle femora as well as on all tibiae A. amani Des.-Grand. from Tanzania but (hind tibia with spotted but not completely distinguished by the presence of inner tym- darkened distal part and with light bases of panum, by the bases of hook-like parts of spines), and with lightish basal parts of all posterior epiphallic lobes (processes) locat- basitarsi; sternites more or less light greyish ed less far from the anteromedian epiphal- brown; pleurites and genital plate almost lic sclerite, by these hook-like parts longer, yellowish but with a few small darkened by the distal half of rachis clearly wider in marks on pterothoracic pleurites; ocelli dis- its distal part or distinctly narrower in its tinctly smaller; scape and space between subdistal part in the profile (in A. amani, antennal cavities approximately equal in this half is narrower in its distal part, and in width; maxillary palpus obliquely truncated A. hempae, it is wider in its subdistal part), at apex (similar to those of A. apiariensis); and by the formula of male genitalia having pronotum somewhat shorter and slightly a pair of acute-angled anterior projections lower (Fig. 72), with almost parallel lateral (in A. hempae, structure of this formula is sides; legs also somewhat shorter, without unknown). From A. apiariensis Des.-Grand. both inner and outer tympana, with lon- (Gabon), the new species differs in the gest (inner dorsal) spur of hind tibia almost epiphallic lobes and rachis very different in reaching base of second tarsal segment, and the shape (see Desutter-Grandcolas, 2015). with 3–4 dorsolateral denticles of hind ba- Etymology. The new species is named af- sitarsus. Genital plate also similar to that ter the Malawi Country. of female of A. malawi, but almost as long as wide and with distinctly narrower apical Afrophaloria dja sp. nov. notch (Fig. 71); ovipositor somewhat lon- (Figs 71, 72, 99–101) ger than hind femur, with distal part as in Figs 100 and 101; copulatory papilla with a Holotype. Female, Cameroon, border of pair of rather long lobules at apex and deep South and East Regions, Dja Reserve on Dja Ri- notch between them (as in Fig. 99). ver, ~600 m, primary/secondary forest, on leaves of bushes at night, 15–22.II.2016, A. Gorochov Variations. Second female with slightly (ZIN). darker head having less distinct pattern, Paratype. Female, same data as for holotype with darker anal and genital plates (almost (ZIN). dark brown and light brown, respectively), Description. Female (holotype). General and with pronotal disc barely widening to appearance similar to that of A. malawi but pterothorax. with following differences: body slightly Male unknown. smaller; head light brown with brown dor- Length in mm. Body 13.5–14; pronotum sal area between eyes (anterior edge of this 2.5–2.7; hind femora 10.8–11.2; ovipositor area situated near lateral ocelli) and a pair 12–12.5. of spots behind eyes, with rostral apex (un- Comparison. The new species is similar der median ocellus) and mouthparts almost to A. hempae, A. amani and A. apiariensis

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 70 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD in the absence of all tympana, but it is dis- blackish genital plate and cerci (but cercal tinguished from A. amani and A. hempae by bases greyish brown). Body structure typi- the female genital plate less transverse and cal of this genus: scape approximately 2.5 having a distinctly narrower apical notch, times as wide as rostrum between antennal and from A. apiariensis, by the female copu- cavities; ocelli almost indistinct; head dor- latory papilla having a pair of clearly longer sum with distinct transverse and fold-like apical lobules. From A. malawi, A. dja dif- concavity behind (near) antennal cavities; fers in the absence of all tympana and in the pronotum strongly widening to ptero- characters listed in its description. thorax, with disc having almost angularly Etymology. This new species is named af- concave anterior edge and straight poste- ter the Dja Reserve. rior edge as well as distinct transverse fold (concavity) in middle part, and with lat- Subfamily CACOPLISTINAE eral lobe not very flat but rather high and Saussure, 1877 with rounded antero- and posteroventral corners; tegmina with dorsal field as in Fig. Tribe HOMOEOGRYLLINI 105; lateral tegminal field with Sc-R and Gorochov, 1986 R-M areas weakly widened and more or less Homoeogryllus ambo sp. nov. equal in width as well as having moderately (Figs 105, 112–115) sparse crossveins, and with 16–17 partly S-shaped branches of Sc (crossveins be- Holotype. Male, Ethiopia, ~100 km W of tween these branches absent); hind wings Addis Ababa City, environs of Ambo Town, invisible in rest position; fore leg with inner XI.2011, A. Zagorinskij (ZIN). and outer tympana narrowly oval, not large Description. Male (holotype). Body mo- and with acute-angled distal parts; fore and derately large for this genus. Colouration middle tibiae with a pair of ventral spurs mostly very dark: head light brown with only; hind tibia with three inner and two black eyes, rostral dorsum, transverse stripe outer movable dorsal spines in distal third between middle parts of eyes running along as well as with six apical spurs (middle in- posterior edges of antennal cavities and ner spur longest and reaching middle part crossing lateral ocelli, a pair of large spots of basitarsus); anal plate widely rounded behind eyes, a pair of narrow longitudinal at apex; genital plate rather large, some- stripes on dorsum behind lateral ocelli, and what elongated and with rather narrowly most part of antennae (scape with light rounded apex. Genitalia most similar to brown dorsomedian area and brown ven- those of H. xanthographus Guér.-Mén. but tromedian area; proximal half of flagellum with posteromedian (ventromedian) edge with sparse whitish and yellowish spots; of epiphallus more oblique in caudal view, and rest of flagellum probably greyish with right posterolateral epiphallic lobe brown), as well as with brown spot under directed more aside and having apical part each eye and dark brown to greyish brown more curved upwards, with left postero- palpi; pronotum black with a few brown lateral epiphallic lobe barely shorter and and light brown marks on disc as well as less inflated in distal half as well as hav- with brown anteroventral part of each lat- ing apical hook strongly curved upwards/ eral lobe; tegmina dark greyish brown with backwards (anterior surface of this lobe some veins reddish brown (Fig. 105); legs almost without longitudinal keel-like ridge also dark brown but with greyish brown near this hook), and with left (longer) ec- coxae and bases of femora, with blackish toparamere having apical part truncated tibiae and basitarsi, and with light brown (not almost acute) (for comparison see Figs movable spines and spurs of hind tibia; rest 112–115 and 116–120). of body from brown to dark brown and with Female unknown.

Figs 112–120. Cacoplistinae (Homoeogryllini), male: 112–115, Homoeogryllus ambo sp. nov.; 116– 120, H. xanthographus Guér.-Mén. Genitalia from below, but their movable distal part in somewhat different positions (112, 117); genitalia from behind (113, 118) and from side (114, 115, 119, 120); distal part of genitalia from below and slightly in front (116), i.e. in same position as in Fig. 112.

Figs 121–130. Cacoplistinae (Homoeogryllini), male: 121–123, Homoeogryllus reticulatus limbe subsp. nov.; 124–126, H. r. retuculatus (F.); 127–130, H. gabonensis Des. Genitalia from below (121, 124, 127), from side (122, 125, 129, 130), from behind and slightly below (123, 126), and from below (128).

Length in mm. Body 19; body with ter species in the body colouration darker, wings 21; pronotum 3.2; tegmina 17; hind mirror of male tegmina less transverse (see femora 11.7. Figs 104 and 105), and male genital char- Comparison. The new species is most acters listed in the description (in H. xan- similar to H. xanthographus (see below) in thographus, left posterolateral lobe has the the male genitalia, but it differs from the lat- apical hook curved mainly upwards, and an-

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 73 terior surface of this lobe is with a distinct nal Sc with 14–15 branches. Genitalia most keel-like ridge near this hook; Figs 118, similar to those of nominotypical subspecies 119). From all the other congeners, H. ambo of H. reticulatus (F.), but distance between is distinguished by the same characters of bases of posterolateral lobes of epiphallus male genitalia as H. xanthographus. clearly smaller (shorter) than in H. r. reticulatus, right posterolateral epiphallic lobe Homoeogryllus xanthographus with wider distal part in profile (i.e. with- Guérin-Méneville, 1847 out shallow ventrodistal notch), and left (Figs 104, 116–120) posterolateral epiphallic lobe having low dorsolateral convexity in middle part (this New material. Ethiopia: male, Illubabor Prov., convexity almost conical and looking as an- 25 km W of Abobo Vill. (near Alwero River), at gular projection in dorsal and ventral views; light, 10.XII.1986, L. Rybalov (ZIN). see Figs 121–123 and 124–126). Note. This species was originally de- Female unknown. scribed from Ethiopia, but later, its type Length in mm. Body 15; body with locality was changed by Desutter (1985) wings 18.5; pronotum 2.7; tegmina 15.5; in connection with the loss of type material hind femora 10.9. and the designation of a male from Kenya Comparison. The new subspecies is dis- as a neotype of H. xanthographus. This au- tinguished from nominotypical one (dis- thor and Gorochov (1988b) recorded this tributed probably in more western countries species also from Sudan, Chad, Egypt and of Africa: from Senegal to Burkina Faso) by some other localities of Ethiopia. Here, this the male genital characters listed above and species is indicated for a new locality in possibly by more numerous dividing veins Ethiopia. Also it is necessary to note that in the male tegminal mirror (see Figs 106 H. lyristes Gor. (Sudan) might be described and 107). from a young male of H. xanthographus hav- Etymology. The new subspecies is named ing insufficiently coloured tegmina and not after the Limbe Town situated near its type fully sclerotized genitalia. locality. Remark. My understanding of H. reticu- Homoeogryllus reticulatus limbe latus is in accordance to the proposal by subsp. nov. Desutter (1985) based on several specimens (Figs 106, 121–123) from different African countries. However, its type is a female from Senegal. It is a rea- Holotype. Male, Cameroon, Southwest Re- son that my understanding of H. reticulatus gion, environs of Limbe Town on sea coast, edge nominotypical subspecies is based on a male of field, on leaf of bush at night, 13–14.II.2016, A. Gorochov (ZIN). from Burkina Faso located near Senegal Desciption. Male (holotype). General ap- (Gorochov, 1988b) but not on Desutter’s pearance very similar to that of H. ambo, but “neoallotype” from Congo located very far body somewhat smaller (medium-sized), from Senegal, because the latter specimen colouration slightly lighter (lower part of may belong to a new subspecies of this spe- head with rather large yellowish areas; light cies. brown marks on pronotal disc larger; each lateral lobe of pronotum with whitish grey Homoeogryllus gabonensis Desutter, 1985 anterior half of lower third; majority of teg- (Figs 108–111, 127–130) minal veins light yellowish to intensively New material. Cameroon: male and 2 females, yellow; cercal bases light brown), longest border of South and East Regions, Dja Reserve spur of hind tibia slightly not reaching mid- on Dja River, ~600 m, primary/secondary forest, dle part of basitarsus, tegminal mirror with on forest floor at night, 15–22.II.2016, A. Goro- three dividing veins (Fig. 106), and tegmi- chov (ZIN).

Description. Female (nov.). General ap- (but not strongly) smaller than outer ones; pearance (Fig. 109) similar to that of male, genitalia very similar to those pictured by but: pronotum somewhat less widening to Desutter (1985: figs 48–52) but with pos- pterothorax and with roundly angular pos- terodorsal semimembranous lobules of terior edge of disc; tegmina without black- epiphallus smaller (Figs 127, 129, 130), left ish veins, insignificantly protruding beyond posterolateral epiphallic lobe slightly differ- abdominal apex, with 7–8 slightly oblique ent in shape (Figs 129), and sclerotized part longitudinal veins in dorsal field and rather of rachis having small notch at apex (Fig. regular and moderately numerous cross- 127, 128). veins between them, with 10–11 oblique Length in mm. Body: male 16, female branches of Sc and sparse crossveins be- 15–17; body with wings: male 19, female tween them, with Sc-R area moderately 18–19.5; pronotum: male 2.7, female 3–3.3; narrow, with R-M area clearly narrower tegmina: male 14.5, female 13–13.5; hind than previous area in proximal half and ap- femora: male 14, female 15–16; ovipositor proximately equal to it (in width) in distal 15.5–16. half, and with rather numerous crossveins Remark. This species was indicated for in both these areas. Genital plate short, several other African countries (Gabon, clearly narrowing to widely truncated or Congo, Zaire, Central African Republic) as widely notched apex (in latter case, this well as for some other localities of Camer- notch very shallow); ovipositor with distal oon (Desutter, 1985). However, the above- part as in Figs 110, 111. mentioned very small differences in the Male almost completely in accordance male genitalia suppose that this species may to very brief original description of this be represented on this enormous territory species, i.e. similar to all congeners de- by a few subspecies. scribed here but with following characteristic features: body medium-sized; coloura- Meloimorpha albicornis (Walker, 1869) tion slightly lighter and less spotted (head brown with dark brown marks on head New material. India: male, Himachal Pradesh similar to those of these congeners in shape, State, environs of Nagar [= Naggar ?] Vill., and with blackish antenna having sparse 1800 m, 10.X.2011, A. Lvovsky (ZIN). whitish and yellowish spots on flagellum; Note. This species was originally de- pronotum also brown but with a few rather scribed from North India, synonymized large light brown spots on disc and almost with M. japonica (Haan) by Chopard dark brown anterior two thirds of each lat- (1968), incorrectly indicated for Vietnam eral lobe; tegmina brown to light brown, (as a subspecies of M. japonica; Gorochov, with vein along distolateral part of mirror 1996), and restored as a distinct species on blackish, and other venation from yellow to the base of its holotype study (Gorochov, yellowish; legs brown with dark brown dis- 2003). In the latter publication, M. albicor- tal parts of femora, tibiae and basitarsi, as nis was recorded from another locality of well as with darkened bases of tibiae; rest of India (Delhi City); here, it is indicated for a body from brown to light brown, but bases more concrete locality in North India. of cerci almost yellowish); pronotum with anterior edge of disc roundly concave and AKCNOWLEDGEMENTS with its posterior edge hardly angularly I thank all my coleagues and friends help- convex; tegminal dorsal field as in Fig. 108; ing me during my trips in tropical regions and tegminal lateral field with 17–18 branches collecting some useful specimens for this study: of Sc and with R-M area almost twice as M.V. Berezin, E.Ju. Tkatsheva, L.N. Anisyutkin, wide as Sc-R area; hind wings invisible, legs D.V. Logunov, G.S. Irisov and others. This study as in H. r. limbe, but inner tympana distinctly was performed in the frames of the state research

© 2018 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 27(1): 40–76 A.V. GOROCHOV. PTEROPLISTINAE, PHALORIINAE AND CACOPLISTINAE FROM OLD WORLD 75 project No. AAAA-A17-117030310210-3 (Rus- Gorochov A.V. 1992. On some new and little sian Federation) and supported by the Russian known crickets (Orthoptera, Gryllidae) Foundation for Basic Research (grant No. 16- from Vietnam. Proceedings of the Zoological 04-01143). Institute RAS, 245: 3–16. (In Russian). Gorochov A.V. 1996. New and little known REFERENCES crickets from the collection of the Humboldt University and some other collections (Or- Bhargava R.N. 1982. A note on the ecology of thoptera: Grylloidea). Part 2. Zoosystematica the cricket, Homeogryllus cincticornis (Walk- rossica, 5(1): 29–90. er) with extension of its range. Geobios New Gorochov A.V. 1999. New and little known Reports, 1: 42–43. Phaloriinae (Orthoptera: Gryllidae). Zoosys- Chopard L. 1925. Descriptions de Gryllides tematica rossica, 8(1): 27–60. nouveaux [Orthoptères]. Annales de la So- Gorochov A.V. 2001. Preliminary notes on the ciété entomologique de France, 94: 291–332. history of South American Ensifera (Orthop- Chopard L. 1968. Gryllides. Orthopterorum cat- tera). Acta geologica leopoldensia, 24(52/53): alogus, 12: 213–500. 81–86. Desutter L. 1985. Etude préliminaire des es- Gorochov A.V. 2003. New and little known Ca- pèces africaines du genre Homoeogryllus choplistinae and Phaloriinae (Orthoptera: Guérin-Méneville (Grylloidea, Phalangopsi- Gryllidae). Zoosystematica rossica, 12(1): dae). Annales de la Société entomologique de 79–92. France (Nouvelle Série), 21(2): 189–206. Gorochov A.V. 2004a. Review of the subfam- Desutter-Grandcolas L. 2015. Phalangopsidae ily Pteroplistinae (Orthoptera Gryllidae). crickets from Tropical Africa (Orthoptera, Memorie della Societa entomologica italiana, Grylloidea), with descriptions of new taxa 82(2): 379–396. and an identification key for African genera. Gorochov A.V. 2004b. A new species of Trellius Zootaxa, 3948(3): 451–496. from the Philippines (Orthoptera: Grylli- Gorochov A.V. 1985. To the fauna of the crick- dae: Phaloriinae). Zoosystematica rossica, et subfamilies Itarinae, Podoscirtinae, and 13(1): 22. Nemobiinae (Orthoptera, Gryllidae) from Gorochov A.V. 2004c. New data on Tremellia Eastern Indochina. In: Medvedev L.N. (Orthoptera: Gryllidae: Phaloriinae). Zoo- (Ed.). Nasekomye Vietnama [Insects of Viet- systematica rossica, 13(1): 46. nam]: 17–25. Moscow: Nauka. (In Russian). Gorochov A.V. 2005. New and little known Gorochov A.V. 1986. On system and morpho- crickets of the subfamilies Phaloriinae, Pha- logical evolution of the cricket family Gryl- langopsinae and Landrevinae (Orthoptera, lidae (Orthoptera) with description of new Gryllidae) from Indonesia and South Africa. taxa. Communication 2. Zoologicheskii zhur- Proceedings of the Russian Entomological So- nal, 65(6): 851–858. (In Russian). ciety, 76: 25–46. (In Russian). Gorochov A.V. 1988a. New and little known Gorochov A.V. 2010. New cricket taxa of the crickets of the subfamilies Landrevinae and subfamilies Phaloriinae and Pteroplistinae Podoscirtinae (Orthoptera, Gryllidae) from (Orthoptera: Gryllidae) from South-East Vietnam and some other territories. In: Med- Asia. Proceedings of the Zoological Institute vedev L.N. & Striganova B.R. (Eds). Fau- RAS, 314(2): 184–190. na i ekologiya nasekomykh Vietnama [Fauna Gorochov A.V. 2011a. Crickets of the subfamily and ecology of Vietnamese insects]: 5–21. Pteroplistinae (Orthoptera: Gryllidae) from Moscow: Nauka. (In Russian). Malacca. Proceedings of the Zoological Insti- Gorochov A.V. 1988b. New and little known tute RAS, 315(2): 176–180. tropical Grylloidea (Orthoptera). Proceed- Gorochov A.V. 2011b. Crickets of the subfam- ings of the Zoological Institute, USSR Acad- ily Phaloriinae (Orthoptera: Gryllidae) from emy of Sciences, 178: 3–31. (In Russian). Malacca, Sulawesi and Moluccas. Proceed- Gorochov A.V. 1990. New and little known ings of the Zoological Institute RAS, 315(3): crickets (Orthoptera, Gryllidae) from Viet- 227–241. nam and some other territories. Proceedings Gorochov A.V. 2014. Classification of the Pha- of the Zoological Institute, USSR Academy of langopsinae subfamily group, and new taxa Sciences, 209: 3–28. (In Russian). from the subfamilies Phalangopsinae and

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