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ZOOSYSTEMATICA ROSSICA, 23(1): 7–88 25 JUNE 2014

Classification of the Phalangopsinae subfamily group, and new taxa from the subfamilies Phalangopsinae and Phaloriinae (Orthoptera: Gryllidae) Классификация группы подсемейств, родственных подсемейству Phalangopsinae, и новые таксоны из подсемейств Phalangopsinae и Phaloriinae(Orthoptera: Gryllidae)

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]

Preliminary classification of the Phalangopsinae subfamily group (= group “Phalangopsidae”: Phalangopsinae, Cacoplistinae, Phaloriinae and Pteroplistinae) are given. Some problems of evolution and taxonomy of this group are discussed. The former tribe Luzarini (= Luzarinae auct.) is synonymized with the tribe Phalangopsini divided into several subtribes (including the subtribe Luzarina). Sixty eight new taxa from Neotropical, Indo-Malayan and Papuan regions are described: Palpigera aluzara sp. nov., Asymmetracla subgen. nov. and Glandulacla subgen. nov. (in the genus Ochraperites Desutter-Grandcolas, 1993), O. (A.) asymmetricus sp. nov., O. (O.) cuyabeno sp. nov., O. (G.) aguarico sp. nov., Laozacla gen. nov., L. furca sp. nov., Eidmanacris longa sp. nov., Caribacusta gen. nov., C. antigua sp. nov., Noctivox orizaba sp. nov., Modestozarina subtrib. nov., Modestozara gen. nov., M. modesta sp. nov., M. troglophila sp. nov., M. satipo sp. nov., Notendecous subgen. nov. and Pedroecous subgen. nov. (in the genus Endecous Saussure, 1878), Daedalonotum gen. nov., D. daedalum sp. nov., Nemozarina subtrib. nov., Nemozara gen. nov., N. rio sp. nov., N. riorio sp. nov., N. pastaza sp. nov., N. vulcanica sp. nov., Anemozara gen. nov., Zacmozara subgen. nov. (in the genus Anemozara), A. (A.) vera sp. nov., A. (A.) propria sp. nov., A. (A.) umbrosa sp. nov., A. (Z.) eximia sp. nov., Lernecina subtrib. nov.; Lerneca sylvestris sp. nov., L. inalata amboro subsp. nov., L. inalata pantanal subsp. nov., Parendacustina subtrib. nov., ?Uvaroviella problematica sp. nov., Boli- vacla gen. nov., B. boliviana sp. nov., Brevizaclina subtrib. nov., Stridulacla subgen. nov. (in the genus Mikluchomaklaia Gorochov, 1986), M. (M.) enarotali sp. nov., Papuzacla subgen. nov. and Lobulacla subgen. nov. (in the genus Brevizacla Gorochov, 2003), B. (P.) fawi sp. nov., B. (L.) nabire sp. nov., B. (L.) halmahera sp. nov., Terrozacla gen. nov., T. jambi sp. nov., T. harau sp. nov., T. trusmadi sp. nov., T. borneo sp. nov., T. kubah sp. nov., T. gading sp. nov., Vescelia mulu sp. nov., Phaloria (Papuloria) latiuscula sp. nov., Ph. (P.) tristis sp. nov., Ph. (P.) paratristis sp. nov., Ph. (P.) tariku sp. nov., Ph. (P.) manifesta sp. nov., Ph. (P.) waena sp. nov., Ph. (P.) neorava sp. nov.; Ph. sulawesi sp. nov.; Tremellia timah orientalis subsp. nov.; Pseudotrigonidium borneo sp. nov.; P. gaponi sp. nov. One former genus is included in the genus Longuripes Desutter-Grandcolas et Hubbell, 1993 as its subgenus Prolonguripes Desutter- Grandcolas, 1993, stat. nov. Prosthacusta beliza (Otte et Perez-Gelabert, 2009), comb. nov. is transferred from the synonymic genus Doposia Otte et Perez-Gelabert, 2009 (= Lerneca Walker, 1869) to Prosthacusta Saussure, 1874. The genus Otteus Koçak et Kemal, 2009, gen. dist. is restored from synonymy with Cubacophus Ruiz-Baliú et Otte, 1997. Phaloria galoa Otte et Cowper, 2007, syn. nov. is synonymized with Ph. heterotrypoides Gorochov, 1999, and this species as well as Ph. insularis Bolivar, 1912 are transferred from the nominotypical subgenus to the subgenus Papuloria Gorochov, 1996.

Дана предварительная классификация группы подсемейств, родственых Phalangopsinae (группа “Phalangopsidae”: Phalangopsinae, Cacoplistinae, Phaloriinae and Pteroplistinae). Рассмотрены некоторые проблемы эволюции и таксономии этой группы. Бывшая триба Luzarini (= Luzarinae auct.) синонимизирована с трибой Phalangopsini, разделенной на несколько подтриб (включая подтрибу Luzarina). Шестьдесят восемь новых таксонов описаны из неотропической, индо-малайской и папуасской областей: Palpigera aluzara sp. nov., Asymmetracla subgen. nov. и Glandulacla subgen. nov. (в составе рода Ochraper- ites Desutter-Grandcolas, 1993), O. (A.) asymmetricus sp. nov., O. (O.) cuyabeno sp. nov., O. (G.) aguarico sp. nov., Laozacla gen. nov., L. furca sp. nov., Eidmanacris longa sp. nov., Caribacusta gen. nov., C. antigua sp. nov., Noctivox orizaba sp. nov., Modestozarina subtrib. nov., Modestozara gen. nov., M. modesta sp. nov., M. troglophila sp. nov., M. satipo sp. nov., Notendecous subgen. nov. и Pedroecous subgen. nov. (в составе рода Endecous Saussure, 1878), Daedalonotum gen. nov., D. daedalum sp. nov., Nemozarina subtrib. nov., Nemozara gen. nov., N. rio sp. nov., N. riorio sp. nov., N. pastaza sp. nov., N. vulcanica sp. nov., Anemo- zara gen. nov., Zacmozara subgen. nov. (в составе рода Anemozara), A. (A.) vera sp. nov., A. (A.) propria sp. nov., A. (A.) umbrosa sp. nov., A. (Z.) eximia sp. nov., Lernecina subtrib. nov.; Lerneca sylvestris sp. nov., L. inalata amboro subsp. nov., L. inalata pantanal subsp. nov., Par- endacustina subtrib. nov., ?Uvaroviella problematica sp. nov., Bolivacla gen. nov., B. boliviana sp. nov., Brevizaclina subtrib. nov., Stridulacla subgen. nov. (в составе рода Mikluchomaklaia Gorochov, 1986), M. (M.) enarotali sp. nov., Papuzacla subgen. nov. и Lobulacla subgen. nov. (в составе рода Brevizacla Gorochov, 2003), B. (P.) fawi sp. nov., B. (L.) nabire sp. nov., B. (L.) halmahera sp. nov., Terrozacla gen. nov., T. jambi sp. nov., T. harau sp. nov., T. trusmadi sp. nov., T. borneo sp. nov., T. kubah sp. nov., T. gading sp. nov., Vescelia mulu sp. nov., Pha- loria (Papuloria) latiuscula sp. nov., Ph. (P.) tristis sp. nov., Ph. (P.) paratristis sp. nov., Ph. (P.) tariku sp. nov., Ph. (P.) manifesta sp. nov., Ph. (P.) waena sp. nov., Ph. (P.) neorava sp. nov.; Ph. sulawesi sp. nov.; Tremellia timah orientalis subsp. nov.; Pseudotrigonidium borneo sp. nov.; P. gaponi sp. nov. Один бывший род включен в состав рода Longuripes Desutter-Grandcolas et Hubbell, 1993 как его подрод Prolonguripes Desutter-Grandcolas, 1993, stat. nov. Prost- hacusta beliza (Otte et Perez-Gelabert, 2009), comb. nov. перенесена из синонимического рода Doposia Otte et Perez-Gelabert, 2009 (= Lerneca Walker, 1869) в Prosthacusta Saussure, 1874. Род Otteus Koçak et Kemal, 2009, gen. dist. восстановлен из синонимии с Cuba- cophus Ruiz-Baliú et Otte, 1997. Phaloria galoa Otte et Cowper, 2007, syn. nov. сведена в синонимы к Ph. heterotrypoides Gorochov, 1999, и этот вид вместе с Ph. insularis Bolivar, 1912 перенесены из номинативного подрода в подрод Papuloria Gorochov, 1996. Key words: crickets, taxonomy, evolution, America, Indo-Malayan and Papuan regions, Or- thoptera, Gryllidae, Phalangopsinae, Phaloriinae, new taxa Ключевые слова: сверчки, таксономия, эволюция, Америка, индо-малайская и папуас- ская области, Orthoptera, Gryllidae, Phalangopsinae, Phaloriinae, новые таксоны

INTRODUCTION provisional hypotheses on its evolution and divergence: Classification of the Phalangopsinae 1) Branching of the Phalangopsinae sub- subfamily group (= the group “Phalangop- family group from a main stock of the fam- sidae”; Gorochov, 2001) is in need of an ily Gryllidae might be caused by a special- additional study, including revisions of nu- ization of its general ancestor to life on the merous insufficiently described taxa (with forest floor in warm forests or in ecotones of unclear taxonomical position) as well as dis- such forests. Its mode of life might be similar coveries of new taxa and of other new data to that of the genus Lerneca Walker, 1869: (on morphology, mode of life and so on). At many representatives of this genus have present, this group looks as holophyletic completely developed wings and live among one and allows us to ground the following dry leaves on the forest floor, among the

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 9 grass in forest glades, or even on the ground 2) Neotropical fauna of the Phalangop- in shrubby areas (with sparse groups of low sinae subfamily group is richest but seems trees) such as the Pantanal in Paraguay. to me less morphologically diverse than the Early differenciation of this group might Old World fauna. Possibly, evolution of this be connected with specializations to life in group in the Old World was more rapid (in the bark crevices of tree trunks (Ptero- connection with largest territory) and/or plistinae) or on the leaves of living bushes longer (paleontological data on this group, (Phaloriinae); origin of Cacoplistinae and formed probably before Caenozoic, are Phalangopsinae previously been associated known only since Middle Eocene; Goro- by me (Gorochov, 1995) with adaptation chov, 2012). In the both cases, extinction to life on the tree trunks, because almost inside this group in the Old World might all the genera with completely developed be more significant than in the New World. wings from different tribes of Phalangopsi- Thus, it is reasonable to consider the Old nae have such or similar mode of life. Now, World fauna more morphologically diverse this opinion is changed, as many of the (divided into a few distinct subfamilies) but phalangopsines live on the forest floor and somewhat impoverished, and to include the preserve a rather primitive habitus (semi- New World representatives in only a single globular head with a wide and almost not very rich subfamily (Phalangopsinae). projected rostrum, rather short legs). If it 3) Tribal composition of Phalangopsinae is correct, some adaptive characters for life in the Old World is also richer than in the on the tree trunks (angular shape of head New World: Luzaropsini and Endacustini in the profile, narrow and projected head are known only in the Old World; Phalan- rostrum, long legs) might be independently gopsini and Paragryllini are known in the developed in different tribes (Phalangop- Old World as well as in the New World. The sini, Paragryllini). former tribe Luzarini, often considered as a An opposite hypothesis (for example unique endemic American tribe (subfam- admitting that origin of this group was as- ily in some authors: Desutter, 1990; Mel- sociated with a certain adaptation to life lo, 1992; Mews et al., 2009), is probably a in caves or in similar stations) is much less probable. One of the important attributes synonym of Phalangopsini (see below). In of the latter adaptation is a partial (as mini- this connection, I can assume that the Pha- mum) reduction of the both pairs of wings, langopsinae penetrated America from the including diminution or loss of the stridula- Old World, and that two such penetrations tory apparatus. But in all subfamilies of this (as minimum) might have place. Richness group, there are recent taxa with a primitive of the recent Neotropical fauna of phalan- (not reduced) condition of wings: Paragryl- gopsines may be a result of rapid adaptive lus Guérin-Méneville, 1844 and many other radiation on the new enormous territory genera in Phalangopsinae; all or almost all practically lacking competitors and/or very genera in Pteroplistinae, Cacoplistinae and dangerous enemies. Phaloriinae. All these primitive genera do These hypotheses are supported by the not live in caves or in similar stations, but material on this subfamily group studied many of them live on the bark of tree trunks by me. Most part of this material contain- or on the forest floor. Transition to life in ing numerous new (unpublished) data is the caves might be independent in different collected by the Russian collectors and de- groups of Phalangopsini from life on the for- posited at the Zoological Institute, Russian est floor or/and on the tree trunk (through Academy of Sciences, St Petersburg (ZIN). intermediate stages connected with the But some specimens is from the Natural cavities among stones, with the rock crev- History Museum (former British Museum ices or with the tree hollows). of Natural History), London (NHM).

STATUS OF THE FORMER TRIBE (and slightly curved in the profile, judging LUZARINI AND PRELIMINARY by her figures 207 and 208); in “Phalangop- CLASSIFICATION OF THE sinae”, this segment is ostensibly apically PHALANGOPSINAE SUBFAMILY (almost transversally) truncate (see her fig- GROUP ure 206 for “Aclodes”, possibly genus Uva- roviella). However, this segment in Phalan- The former tribe Luzarini was originally gopsis (type genus of “Phalangopsinae”) is established by Hebard (1928) as the group clearly obliquely truncate and curved (Fig. Luzarae. He described this group on the I: 1), more similar to that of her Luzarinae base of external morphology only (without than to that of her “Aclodes”. study of stridulatory apparatus and genita- 2) According to Desutter, “first” inner lia in male). The characters found by him for apical spur of hind tibiae is longer than distinguishing of “Luzarae” from the other “first” outer one in “Luzarinae” and short- groups of Phalangopsinae are mainly con- er than latter spur in “Phalangopsinae” nected with different modes of life: in “Luz- (p. 49). But in Phalangopsis, all the inner arae” living mainly on the forest floor (among apical spurs of hind tibiae are longer than dry leaves, among grass-like plants, and so the respective outer spurs. on), body is not dorsoventrally depressed, 3) In “Luzarinae”, hind basitarsus is with rostrum of head is short and rather wide as two rows of denticles (Hebard’s character), well as obtuse in the profile, ocelli are situat- and hind tibia is with four inner and four ed in the shape of transverse triangle, prono- outer movable spines (excepting six apical tum is with weakly projected anteroventral spurs). But in Uvaroviella included by De- corners, legs are rather short (or moderately sutter in her Phalangopsinae, hind basitar- long) and more or less slender, hind basitarsus is similar in the structure; moreover, sus is with two distinct rows of denticles hind tibia in this genus and in Phalangop- on its dorsum; Uvaroviella Chopard, 1923, sis is usually with four inner and four outer Parendacustes Chopard, 1924 and many movable spines also. other phalangopsines, living mainly on tree 4) Reduction of stridulatory apparatus trunks, have opposite or partly opposite in the male tegmina is usual phenomenon characters. The subsequent authors erected for some representatives of her Luzarinae as rank of “Luzarae” up to tribal (Chopard, well as for some species of Uvaroviella (her 1968) and even subfamily (Desutter, 1990) Phalangopsinae), therefore this reduction level. The latter author (contra Hebard and as well as preservation of normal hind wings Chopard) included Amphiacusta Saussure, only in some genera of her Luzarinae do not 1874 and some other genera (for example, provide us any possibility for distinguishing Eidmanacris Chopard, 1956) in her subfam- of these “subfamilies” from each other on ily Luzarinae (Desutter-Grandcolas, 1993, the base of wing morphology. 1995). These actions violated Hebard’s un- 5) Male genitalia in “Luzarinae”, accord- derstanding of differences between the rela- ing to Desutter, have a tendency to regress tives of Luzara Walker, 1869 (type genus of of “lophi médians” (a pair of posteromedial “Luzarae”) and the relatives of Phalangopsis lobes of epiphallus) accompanied by length- Audinet-Serville, 1831. ening of the posterolateral lobes of epiphal- Desutter (1990) proposed a new list of lus. But in “Phalangopsinae” including differences between her Luzarinae and her Uvaroviella, these posteromedial lobes are Phalangopsinae; she included in this list ostensibly preserved. In reality, there is only some new characters of external morphol- one pair of sclerotized posterior epiphallic ogy as well as some data on male genitalia: lobes in Phalangopsis, in Uvaroviella, and in 1) In “Luzarinae”, apical segment of “Luzarinae” (see Figs II: 1–3, 6–8, 13–15). maxillary palpi is very obliquely truncate Posteromedian part of their epiphallus is

Figs I (1–6). Maxillary palpus from side: 1, Phalangopsis longipes A.-Serv.; 2, Eidmanacris longa sp. nov.; 3, Uvaroviella parantennalis Gor.; 4, Ochraperites cuabeno sp. nov.; 5, Luzara venado Gor.; 6, Daedalonotum daedalum sp. nov. membranous: in Phalangopsis and Luzara, Phalangopsis, but they are located rather far this part is large; in Uvaroviella, it is very from each other and from hind edge of this small (narrow) or practically absent. Thus, part in Eidmanacris; Figs II: 5, 12); epiphal- the structure of epiphallus in Phalangopsis lus of Luzara (Figs II: 9, 10) is practically and in Luzara is probably characterized by without membranous fold between epiphal- the synapomorphic features: median part of lic sclerite and rachis (possible autopomor- epiphallus is short and bridge-like, postero- phy of Luzara and its close-relatives), and lateral epiphallc arms are large (long), and without distinct upper lobes on the mem- membranous posteromediain epiphallic part branous posteromedian epiphallic part (ple- is very large (Figs II: 1–3, 6–8). However, siomorphy). So, the presence of characteris- epiphallus in Uvaroviella is more primitive: tic “lophi médians” (u) in the epiphallus of its median epiphallic part is rather long, pos- Phalangopsis is probably a unique specializa- terolateral epiphallic arms are rather small, tion but not primitive character; Eidmanac- and membranous part between these arms is ris shows an intermediate condition of these very small or absent (Figs II: 13–16). More- lobes between Luzara and Phalangopsis and over, epiphallus in Eidmanacris (included by may be a close relative of Phalangopsis (one Desutter in “Luzarinae”) is similar to that subtribe); Luzara is also more or less related of Phalangopsis in the presence of a charac- to this subtribe (the same tribe but another teristic membranous fold between the ra- subtribe); and Uvaroviella is somewhat less chis (= guiding rod) and sclerotized median related to these subtribes (another tribe or epiphallic part (Figs II: 4, 5, 11, 12) and in third subtribe). the presence of a pair of upper lobes (u) on 6) Desutter also mentions a tendency the membranous posteromedian epiphallic to the development of “l’apodème endopal- part (these lobes are situated near each other lique” (apodeme of formula = apodeme of and along hind edge of this epiphallic part in mold of spermatophore attachment plate)

Figs II (1–16). Male genitalia: 1–5, Phalangopsis longipes A.-Serv.; 6–10, Luzara venado Gor.; 11, 12, Eidmanacris longa sp. nov.; 13–16, Uvaroviella antennalis Gor. Genitalia from above (1, 6, 13), from below (2, 7, 14) and from side (3, 8, 15); sagittal section of genitalia, schematically (4, 9, 11, 16); distal half of genitalia from above, scheme (5, 10, 12). Abbreviations: a, apodeme of endoparamere; e, endoparamere fused with opposite endoparamere by median bridge; ec, ectoparamere; ep, epiphallus; epa, posterolateral epiphallic arm (lobe); f, formula (mold of spermatophore attachment plate); m, membranous posteromedian epiphallic part; r, rachis (guiding rod); ra, ramus; u, upper lobe of m; v, valvae.

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 13 in male genitalia of her Luzarinae. However, Grandcolas, 1993; Koilenoma Desutter-Grand- such apodeme is independently developed colas, 1993; Leptopsis Desutter-Grandcolas, or lost in numerous genera from different 1996; Peru Koçak et Kemal, 2008; Peruzara subfamilies of Gryllidae; for example, this Gorochov, 2011; Amazonacla Gorochov, 2011; apodeme is indistinct in Phalangopsis and Ucayacla Gorochov, 2011; Lecticusta Cadena- Castañeda et Garcia, 2012; possibly Gryllosoma low but distinct in Uvaroviella. Thus, this Hebard, 1928, Tairona Hebard, 1928, Rehniella character also cannot be used for clear dis- Hebard, 1928, Amusodes Hebard, 1928, Amusina tinguishing of these “subfamilies”. Hebard, 1928, Megalamusus Hebard, 1928 and So, Phalangopsis and Luzara are related Laozacla gen. nov.]; subtribe Phalangopsina genera and must be included in one tribe; Blanchard, 1845 [Phalangopsis Audinet-Serville, and Luzarinae and Luzarini are junior syn- 1831; Eidmanacris Chopard, 1956; Philippopsis onyms to Phalangopsinae and Phalangop- Desutter-Grandcolas, 1992; possibly Dysco- sini, respectively. The genus Uvaroviella is phogryllus Rehn, 1901 and Strinatia Chopard, somewhat more separated from majority 1970]; subtribe Amphiacustina Hubbell, 1938 of the studied American representatives of [Amphiacusta Saussure, 1878; Noctivox Desut- ter-Grandcolas et Hubbell, 1993; Longuripes De- Phalangopsini (including Phalangopsis and sutter-Grandcolas et Hubbell, 1993 (including Luzara) than these representatives from subgenus Prolonguripes Desutter-Grandcolas, each other, and this genus may be included 1993, stat. nov.); Nemoricantor Desutter-Grand- in a special tribe. However, such inclusion colas et Hubbell, 1993; Mayagryllus Desutter- must be grounded by an additional study Grandcolas et Hubbell, 1993; Arachnopsita De- of Phalangopsini from the New World as sutter-Grandcolas et Hubbell, 1993; Cantrallia well as from the Old World. At present, it Desutter-Grandcolas, 1994; Leptopedetes De- is reasonable to divide this rich and rather sutter-Grandcolas, 1994; Caribacusta gen. nov.; diverse tribe into several subtribes only. possibly fossil Araneagryllus Heads, 2010]; sub- Classification of the Phalangopsinae tribe Modestozarina subtrib. nov. [Endecous Saussure, 1878; Modestozara gen. nov.; possibly subfamily group from the Orthoptera Spe- Daedalonotum gen. nov.]; subtribe Nemozarina cies File (Eades et al., 2014) is somewhat subtrib. nov. [Lernecella Hebard, 1928; Nemo- eclectic: some parts of this classification are zara gen. nov.; Anemozara gen. nov.]; subtribe given in accordance to different views of Lernecina subtrib. nov. (= “Lernecae” Desut- different authors or has not any published ter, 1987, unavailable name published without ground. It is a reason that I put here (see description or diagnosis) [Lerneca Walker, 1869; below) a preliminary classification of this Prosthacusta Saussure, 1874; possibly Microler- group in accordance to my views (however, neca Mello, 1995]; subtribe Parendacustina the taxa of Phalangopsinae from Africa and subtrib. nov. [Arachnomimus Saussure, 1897; adjacent islands are not included, as they Parendacustes Chopard, 1924; Luzonogryllus Ya- masaki, 1978; Longizacla Gorochov, 2003]; sub- are in need of a special study): tribe ? Heterogryllina Hebard, 1928 (?= “Aclo- dae” Desutter-Grandcolas, 1992) [Uvaroviella PHALANGOPSINAE subfamily group Chopard, 1923; possibly Heterogryllus Saussure, (= group “PHALANGOPSIDAE”) 1874]; Phalangopsini incertae sedis (subtribal position unclear) [Miogryllodes Hebard, 1928; I. Subfamily PHALANGOPSINAE Blanchard, Prosthama Hebard, 1928; Anacusta Hebard, 1845 1928; Cophella Hebard, 1928; Paracophella He- 1) Tribe PHALANGOPSINI Blanchard, bard, 1928; Phalangopsina Chopard, 1933; Kem- 1845: subtribe Luzarina Hebard, 1928 [Luzara piola Uvarov, 1940; Endophallusia Mello, 1990; Walker, 1869; Luzarida Hebard, 1928; Palpi- Guabamima Mello, 1993; Aracamby Mello, 1993; gera Hebard, 1928; Niquirana Hebard, 1928; Cacruzia Mello, 1993; Izecksohniella Mello, 1993; Acantoluzarida Desutter-Grandcolas, 1992; Lu- Vanzoliniella Mello et Cezar dos Reis, 1994; Le- zaridella Desutter-Grandcolas, 1992; Melanotes rnecopsis Mello, 1995; Anophtalmotes Desutter- Desutter-Grandcolas, 1993; Ochraperites De- Grandcolas, 1995; Zacla Gorochov, 2003; Otte- sutter-Grandcolas, 1993; Allochrates Desutter- dana Mello et Andrade, 2003; Saopaoloa Koçak

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 14 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP et Kemal, 2008; Marliella Mews et Mol, 2009; II. Subfamily CACOPLISTINAE Saussure, Joadis Mews et Sperber, 2009; Grandcolasia 1877 (it is originally described as Cachoplistites, Koçak & Kemal, 2010; Mellopsis Mews et Sper- but this name is based on unjustified emenda- ber, 2010; Adenopygus Bolfarini et Mello, 2012; tion of generic name and must be corrected with Opiliosina Desutter-Grandcolas, 2012; Spelun- preservation of the same author and date; see casina Desutter-Grandcolas, 2012; Bambuina The Code, 1999: article 35.4.1 and example in it) Mello, Horta et Bolfarini, 2013] 1) Tribe CACOPLISTINI Saussure, 1877 2) Tribe PARAGRYLLINI Desutter, 1988: [Cacoplistes Brunner-Wattenwyl, 1873] subtribe Paragryllina Desutter, 1988 [Paragryl- 2) Tribe HOMOEOGRYLLINI Gorochov, lus Guerin-Meneville, 1844; Benoistella Uvarov, 1986 [Homoeogryllus Guérin-Méneville, 1847; 1939; Rumea Desutter, 1988; Silvastella Desut- Meloimorpha Walker, 1879] ter-Grandcolas, 1992; Bolivacla gen. nov.]; subtribe Neoaclina Desutter, 1988 [Neoacla De- III. Subfamily PHALORIINAE Gorochov, 1985 sutter, 1988; Kevanacla Desutter-Grandcolas, ú 1) Tribe PHALORIINI Gorochov, 1985 [Pha- 1992; Yoyuteris Ruiz-Bali et Otte, 1997; Aclella å å å Desutter-Grandcolas, 2000; Selvacla Otte, 2006; loria St l, 1877; Vescelia St l, 1877; Tremellia St l, possibly Escondacla Nischk et Otte, 2000 and Pe- 1877; Pseudotrigonidium Chopard, 1915; Strophi- ruacla Gorochov, 2011]; subtribe Strogulomor- ola Uvarov, 1940; Trellius Gorochov, 1988; Ceylo- phina Desutter, 1988 [Strogulomorpha Desutter, ria Gorochov, 1996; Sumatloria Gorochov, 2003; 1988; Loretana Desutter, 1991; possibly Eugryl- Gorochovius Xie et al., 2004; possibly fossil Elec- lina Hebard, 1928, Nigrothema Desutter-Grand- trogryllus Gorochov, 1992 (it contains one species colas, 1991, Unithema Desutter-Grandcolas, only, because “E. electrum Gorochov, 1992” does 1991, Anomaloterga Mello et Bolfarini, 2010 and not exist: it is a lapse probably based on incorrect Ecuadoracla Gorochov, 2011]; subtribe Mexi- translation of my sentence about etymology of aclina subtrib. nov. [Mexiacla Gorochov, 2007; the generic name; see Eades et al., 2014)] Oaxacla Gorochov, 2007]; subtribe Brevizaclina 2) Tribe SUBTILORIINI Gorochov, 2003 subtrib. nov. [Mikluchomaklaia Gorochov, 1986; (2011) [Heterotrypus Saussure, 1878; Schizotry- Brevizacla Gorochov, 2003]; Paragryllini incer- pus Chopard, 1954; Subtiloria Gorochov, 1999; tae sedis (subtribal position unclear) [Laranda Kameruloria Gorochov, 2003] Walker, 1869; Ectecous Saussure, 1878; Caltath- IV. Subfamily PTEROPLISTINAE Chopard, ra Otte, 1987; Adelosgryllus Mesa et Zefa, 2004; 1951 (Pteroplistinae Chopard, 1936 is unavail- Apteracla Gorochov, 2009] able name published without description or di- 3) Tribe ENDACUSTINI Gorochov, 1986 agnosis; see The Code, 1999: article 13.1) [Endacusta Brunner-Wattenwyl, 1873; Endo- taria Chopard, 1951; Tathra Otte et Alexander, [Pteroplistes Brunner-Wattenwyl, 1873; Tram- 1983; Nesitathra Otte et Rentz, 1985; Lucienia lapiola Gorochov, 1990; Crockeriola Gorochov et Corochov, 1986; Anendacusta Gorochov, 2003; Kostia, 1999; Kerinciola Gorochov, 2004; Chan- Itarotathra Gorochov, 2003; Pseudendacusta giola Gorochov, 2004; Tembelingiola Gorochov, Gorochov, 2003; Zaclotathra Gorochov, 2003; 2004; Pangrangiola Gorochov, 2004; Asymmet- possibly Discotathra Gorochov, 2003] riola Gorochov, 2010; Singapuriola Gorochov et 4) Tribe LUZAROPSINI Gorochov, 1986 Tan, 2012; possibly fossil Trichogryllus Chopard, [Luzaropsis Chopard, 1925; Terrozacla gen. nov.; 1936 and Eneopterotrypus Zeuner, 1937] possibly Larandopsis Chopard, 1924] 5) Tribe OTTEINI Koçak et Kemal, 2009 DESCRIPTIONS OF NEW TAXA [Cubacophus Ruiz-Baliú et Otte, 1997; Otteus Koçak et Kemal, 2009, gen. dist.] Subfamily PHALANGOPSINAE 6) PHALANGOPSINAE INCERTAE SE- Tribe PHALANGOPSINI DIS (tribal position unclear) [Hemicophus Sau- ssure, 1878; Aspidogryllus Chopard, 1933; Pro- Subtribe LUZARINA tathra Desutter-Grandcolas, 1997; Dambachia Nischk et Otte, 2000; Antilliclodes Otte et Perez- Note. The subtribe is characterized by Gelabert, 2009; Megacris Desutter-Grandcolas, an almost semiglobular head (its rostrum is 2012; fossil Eozacla Gorochov, 2012; possibly widely rounded in the profile, more or less fossil Eotrella Gorochov, 2012] wide, not very projected forwards, and not

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 15 separated from the rest of vertex by a dor- edges of latter field, greyish brown to al- sal concavity), the median ocellus located most light brown areas on metanotum and at the rostral apex or near it, lateral ocelli on first abdominal tergite, moderately dark situated on lateral parts of rostral base or reddish brown area on middle tibiae, light near rostrum and eyes (but not on rostral brown most part of spines and spurs of legs, dorsum), maxillary palpi with a moderately and light brown small marks on basal part of long and clearly widened apical segment outer surface of hind femora as well as larg- having an oblique cutting and slightly arcu- er areas on proximal half of inner surface of ately curved distal part in the profile (Figs these femora. Head with apical segment of I: 4, 5; Fig. III: 5), legs usually not very long maxillary palpi strongly widened in profile and not very thin, hind tibiae with the dor- (almost semicircular in shape; Fig. III: 5) sal inner apical spur longest, and rachis of and lamellar, with rostral apex barely nar- male genitalia usually more or less membra- rower than scape, and without concavity nous and very small but sometimes almost on epicranial dorsum in profile. Pronotum absent (Figs II: 6–10) or rather short and slightly wider than head and with widely with the apical lobe directed upwards (Figs rounded (not distinctly projected) antero- III: 6–8). However, distribution of these ventral corners. Notal gland developed, characters among many included genera is consisting of rather wide and rounded me- partly studied only, and the genera listed dian lobe of hind part of mesonotum (this above as members of Luzarina may belong lobe curved upwards) as well as of two pairs to more than one subtribe. Majority of of rounded tubercles on metanotum (lat- the Luzarina representatives live on forest eral tubercles distinctly larger than medial floor; often they may be collected on soil or ones; Fig. III: 3). Tegmina rather wide but low grass-like plants but sometimes on tree distinctly shortened (reaching apex of fifth trunks near ground during stridulation. abdominal tergite); tegminal lateral field with six almost straight and parallel longi- Palpigera aluzara sp. nov. tudinal veins (including vein separating lat- (Figs III: 1–13) eral field from dorsal one) and a few small additional branches (crossveins almost Holotype. Male; Bolivia, Santa Cruz De- lost); tegminal dorsal field with developed partment, 23 km SW of Santa Cruz City, El Soc stridulatory apparatus having numerous Natural Park (small private area with secondary oblique veins (sinuate in shape) and not forest), about 600 m, on bark of lower part of living tree trunk at night, 14–16 Feb. 2014, A. large obliquely triangular mirror; dorsal Gorochov (ZIN). field of right (upper) tegmen slightly more Paratypes. Five males and 4 females, same thickened than that of left (lower) tegmen data as for holotype, but some females collected (Figs III: 1, 2). Hind wings absent. Legs not on small forest road at night (ZIN). very long and not very thin, with only in- Description. Male (holotype). Coloura- ner medium-sized open tympana and mod- tion dark brown, rather uniform but with erately thickened (for jumping) hind fem- white apical and subapical segments of ora. Anal plate almost square, with barely maxillary palpi, reddish brown both small notched hind edge and tubercle-like pos- area between ocelli and transverse band on terolateral corners; genital plate distinctly dorsum between hind halves of eyes, dark longer than previous plate and with barely reddish brown disc and almost blackish concave apex (Fig. III: 4). Genitalia with lateral lobes in pronotum, brown tegmina epiphallus typical of Luzarina and having having dorsal field of right tegmen barely membranous lobules on apices of postero- lighter and that of left tegmen almost trans- lateral epiphallic arms, very characteristic parent with light brown apical area and rachis having rather large sac-like process with very light narrow areas along other directed upwards and apical spine curved

Figs III (1–13). Palpigera aluzara sp. nov.: 1, 2, dorsal field of lower (1) and upper (2) male tegmina; 3, male metanotal gland from above; 4, male abdominal apex from above; 5, maxillary palpus from side; 6–8, male genitalia from above (6), from below (7) and from side (8); 9, head, pronotum and tegmina of female from above; 10, female genital plate from below; 11, 12, female copulatory papilla from above (11) and from side (12); 13, distal part of ovipositor from side.

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 17 upwards also, most part of ectoparameres shorter; and from P. fratercula Hebard, heavily sclerotized and compact but situat- 1928, by a much wider apical segment of the ed near distal part of above-mentioned arms maxillary palpi and distinctly concave (not and partly fused with them (medial part of convex) hind edge of the male anal plate. ectoparameres with strongly S-shaped thin Etymology. Name of this species consists process having apex directed to apex of ra- of the Greek prefix meaning “not” and ge- chis), endoparameres H-shaped and rather neric name Luzara. long (their posterior arms very long, reaching distal half of ectoparameres), rami very Genus Ochraperites Desutter-Grandcolas, long, and formula located distinctly before 1993 rami and with rather large apodeme (Figs III: 6–8). Note. This genus includes crickets with Variations. Some males with uniformly a medium-sized to rather small body, almost dark brown middle tibiae, slightly darker or semiglobular and moderately high head lighter tegmina, or somewhat more concave having the rostrum almost as wide as scape, hind edge of anal plate. almost square pronotum with a slightly nar- Female. General appearance as in males, rowing anterior part (Figs IV: 1, 5, 9) and but tegmina different (both tegmina prac- widely rounded anteroventral corners, the tically identical in structure, somewhat presence of metanotal gland in male, sig- smaller but not very short, slightly over- nificantly shortened male tegmina having lapping each other, with thickened dorsal a widely rounded distal part of the dorsal field, with 8–9 convex and parallel longi- field and very asymmetrical structure of the tudinal veins in dorsal field as well as with upper tegmen in comparison to lower one 5–6 almost parallel ones in lateral field, (dorsal field in the upper tegmen is semi- with weakly developed crossveins, and with sclerotized and almost without venation, colouration of dorsal field in both tegmina but such field in the lower tegmen is almost similar to that of right male tegmen; Fig. completely or partly membranous and with III: 9), notal gland absent, and anal plate a thin stridulatory vein dividing the mem- slightly narrowing to rather widely round- branous space into two parts almost equal ed apex; genital plate rather short and with to each other in the size; Figs IV: 2, 3, 6, 7, small rounded posteromedian notch (Fig. 10, 11), almost scale-like female tegmina III: 10); ovipositor normal and with simple (hind wings are absent in both sexes), mod- apex (Fig. III: 13); copulatory papilla as in erately short and thin legs (but the hind Figs III: 11, 12. femora are distinctly thickened, adapted to Length in mm. Body: male 19–22, fe- jumping) having inner (oval) tympana only male 18–21; pronotum: male 3–3.3, female and numerous denticles between and before 3.2–3.4; tegmina: male 9–10, female 5.5–6; four outer and four inner dorsal articulated hind femora: male 15–16, female 15.5–16.5; spines of the hind tibiae, a short male anal ovipositor 11.5–12. plate having a widely truncate hind part Comparison. Maxillary palpi of the new and slightly projected posterolateral cor- species are with the apical segment inter- ners (Figs IV: 4, 8, 12), the male genital mediate between those of P. borellii (Giglio- plate approximately twice longer than anal Tos, 1897) and P. boliviana (Bruner, 1916) plate, and the male genitalia rather diverse. in the shape; it is slightly narrower than in These genitalia are with only a few general the first species and clearly wider than in characters: the epiphallus strongly curved P. boliviana. From P. borellii, the new spe- in the profile and with a pair of postero- cies is also distinguished by more angular lateral arms fused or partly fused with the posterolateral projections of the male anal ectoparameres; each ectoparamere is with plate; from P. boliviana, by these projections an angular or lobule-like short anterodorsal

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 18 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP projection situated very near such projec- [Included species: O. (A.) asymmetricus sp. tion of the opposite ectoparamere (possibly nov. (type species) only. Etymology: the sub- this character is one of the unique synapo- generic name originates from the Latin word morphies of the genus Ochraperites); the en- of Greek origin “asymmetricus” (asymmetri- doparameres are H-shaped and with rather cal) and genus Acla.] – Male genitalia narrow; epiphallus with an- long posterior arms; formula is semitubular terior part weakly widened and curved up- and without long apodeme; rami are well wards, as well as with posterolateral arms developed (Figs V: 1–9). long and sclerotized or partly sclerotized; ec- This genus includes a few species with toparameres small; endoparameral apodemes somewhat different structure of the male narrow (Figs V: 7–9) ...... 4 tegmina and male genitalia. These differ- 4. Epiphallus with long anterior (curved) part, ences allow me to divide it into three sub- with narrowed middle part (between ante- genera. I cannot exclude that these subgen- rior and posterior parts), and with compara- era may be considered as separate genera, or tively wide basal part of posterolateral arms that some other genera described previous- (near middle part of ectoparameres); rachis of male genitalia not protruding upwards; ly (Allochrates and Peru) may be included endoparameral apodemes much shorter than in the genus Ochraperites as its subgenera rami (Figs V: 7–9, 11). Male tegmina with (see a key for subgenera of this genus and distinct tegminal gland in distal part of lat- for close-related genera below). eral field and without mirror in stridulatory 1. Male genitalia moderately wide; epiphallus apparatus (Figs IV: 10; V: 10) ...... with anterior part comparatively small and ...... subgen. Glandulacla subgen. nov. curved upwards and partly backwards, as [Included species: O. (G.) aguarico sp. nov. well as with distal half of posterolateral arms (type species) only. Etymology: the subge- membranous; endoparameral apodemes long neric name originates from the Latin word and narrow (Figs V: 4–6) ...... 2 “glandulosus” (glandular) and genus Acla.)] – Male genitalia wide or narrow; epiphallus – Epiphallus with short anterior (curved) part, with anterior part diverse and with distal with middle part not narrower than more dis- half of posterolateral arms sclerotized or tal parts, and with rather narrow basal part partly sclerotized; endoparameral apodemes of posterolateral arms (near middle part of diverse (Figs V: 1–3, 7–9) ...... 3 ectoparameres); rachis of male genitalia dis- 2. Anterior part of epiphallus without any tinctly protruding upwards (clearly visible median tubercle; lateral part of ectoparam- above dorsal edge of middle part of epiphal- eres in male genitalia sclerotized and almost lus in profile); endoparameral apodemes spine-like; endoparameral apodemes very slightly shorter than rami. Structure of lat- long, projected before anterior part of rami eral field of male tegmina unknown; mirror in ...... tegminal stridulatory apparatus of male de- gen. Allochrates Desutter-Grandcolas, 1993 veloped . . . gen. Peru Koçak et Kemal, 2008 – Anterior part of epiphallus with distinct median tubercle; lateral part of ectoparameres in male genitalia membranous and lobe-like; Ochraperites (Asymmetracla) endoparameral apodemes not very long, not asymmetricus sp. nov. projected before anterior part of rami (Figs (Figs IV: 1–4; V: 1–3) V: 4–6) ...... subgen. Ochraperites s. str. [Included species: Ochraperites ottei Desut- Holotype. Male; Ecuador, western slopes of ter-Grandcolas, 1993 (type species); O. (O.) Andes not far from Pacific Coast, 10 km E of cuyabeno sp. nov.] Agua Blanca Vill. (environs of Puerto Lopez 3. Male genitalia wide; epiphallus with ante- Town), San Sebastian Camp, about 700 m, misty rior part strongly widened and curved back- primary forest, among dry leaves on forest floor wards, as well as with posterolateral arms at night, 26–29 Oct. 2005, A. Gorochov, A. Ovt- short and sclerotized; ectoparameres large; shinnikov (ZIN). endoparameral apodemes wide (Figs V: 1–3) Paratypes. Male and 2 females, same data as ...... subgen. Asymmetracla subgen. nov. for holotype (ZIN).

Figs IV (1–12). Ochraperites, male: 1–4, O. asymmetricus sp. nov.; 5–8, O. cuyabeno sp. nov.; 9–12, O. aguarico sp. nov. Head and pronotum from above (1, 5, 9); dorsal field of lower (2, 6, 10) and upper (3, 7, 11) tegmina; abdominal apex from above (4, 8, 12).

Figs V (1–11). Ochraperites, male: 1–3, O. asymmetricus sp. nov.; 4–6, O. cuyabeno sp. nov.; 7–11, O. aguarico sp. nov. Genitalia from above (1, 4, 7), from below (2, 5, 8) and from side (3, 6, 9); lateral field of upper tegmen (10); scheme of sagittal section of formula (coloured area) with membranous rachis, and schematic picture of genitalia from side (11). Abbreviations: a, anterodorsal projection of ectoparamere; ec, main body of ectoparamere; en, endoparamere with apodeme; f, formula; paep, posterolateral arm of epiphallus; ra, rachis.

Description. Male (holotype). Body dominal tergite and almost contacted with rather large for this genus. Coloration dark each other (dorsal field in both tegmina brown with four yellowish longitudinal semisclerotized and lacking both vena- lines on dorsum of head, light brown labrum, tion and light stripe along posterior edge); brown antennae having yellowish proximal metanotal gland absent; dorsum of abdomi- part and a few light brown small spots near nal tergites more or less spotted, consisting this part, whitish palpi having darkened of brown and light brown marks; anal plate (almost brown) three proximal segments simple and slightly narrowing to obtusely of maxillary palpi and areas on two proxi- angular apex. Genital plate almost as long as mal segments of labial palpi, pronotum hav- wide and with rounded but slightly concave ing a pair of short light brown longitudinal apex; hind femur approximately 1.5 times as stripes on hind half of disc (Fig. IV: 1) and long as ovipositor; apical part of ovipositor small yellow spot in anteroventral corner narrow and acute. of lateral lobes, tegmina having lateral field Length in mm. Body: male 11–11.5, fe- dark brown and dorsal field differently co- male 13.5–14; pronotum: male 2.2–2.4, fe- loured in upper and lower tegmina (in upper male 2.9–3.1; tegmina: male 2.9–3.2, female tegmen, latter field brown with light brown 2–2.2; hind femora: male 8.6–9, female 10– stripes along lateral and posterior edges; in 10.5; ovipositor 6.5–6.9. lower tegmen, it light greyish, semitrans- Etymology. The species name is the parent, having light brown stripe along lat- Latin word of Greek origin “asymmetricus” eral edge and yellowish one along posterior (asymmetrical) given in connection with a edge; Figs IV: 2, 3), pterothoracic tergites strong asymmetry of the male tegmina. partly and abdominal ones completely (excepting cerci) dark brown to blackish, legs and venter of thorax light brown but having Ochraperites (Ochraperites) cuyabeno numerous oblique brown stripes on outer sp. nov. surface of hind femora, and cerci brown to (Figs I: 4; IV: 5–8; V: 4–6) light brown. Head and pronotum shinning; Holotype. Male; Ecuador, eastern plain, 80– tegmina reaching distal third of second ab- 85 km E of Lago Agrio Town, environs of Lago dominal tergite, with long and thin stridula- Grande Lake on Rio Cuyabeno, very lowlying tory vein of upper tegmen clearly visible on primary forest, among dry leaves on forest floor ventral surface and arcuate in shape, with at night, 2–9 Nov. 2005, A. Gorochov, A. Ovt- almost completely membranous dorsal field shinnikov (ZIN). of lower tegmen having small but distinct Paratypes. Three females, same data as for mirror (Figs IV: 2), and with narrow lateral holotype (ZIN). field (in both tegmina) having three longi- Description. Male (holotype). Body tudinal and almost parallel veins only; hind medium-sized for this genus. Colouration basitarsus with two rows of dorsal denticles; (Figs IV: 5–8) light brown with slightly genital plate with rounded and slightly bi- darker dorsum of head, a pair of large al- lobate apex; anal plate and genitalia as in most dark brown areas on face contacting figures (Figs IV: 4; V: 1–3). with ventral parts of eyes and antennal Variations. Second male with a pair of cavities, brown both spot behind each eye additional light brown spots on anterior and most part of antennae (only their small part of disc and weakly distinct brown areas proximal part light), whitish maxillary pal- on femora (brown oblique stripes on hind pi having brownish areas on proximal half, femora also developed). dark brown upper half of pronotal lateral Female. General appearance as in male lobes, rather small brown spots on pronotal but with following differences: tegmina disc, greyish brown lateral field of tegmina, shorter, reaching middle part of first ab- greyish light brown dorsal field of upper

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 22 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP tegmen having yellowish stripes along lat- Ochraperites (Glandulacla) aguarico eral and posterior edges, almost complete- sp. nov. ly transparent most part of dorsal field of (Figs IV: 9–12; V: 7–11) lower tegmen, spotted (with light brown Holotype. Male; Ecuador, eastern plain, and slightly darker small marks) abdomi- about 70 km SE of Lago Agrio Town, environs nal tergites, and rather numerous brown of San Pablo de Kantesiya Vill. on Rio Aguarico, marks on legs. Structure of body similar to lowlying partly primary / partly secondary for- that of O. asymmetricus, but tegmina longer est, on forest floor at night, 10–17 Nov. 2005, and with narrower lateral field having one A. Gorochov, A. Ovtshinnikov (ZIN). longitudinal vein only, upper tegmen with Paratype. Female, same data as for holotype straight and short stridulatory vein (situat- (ZIN). ed in medial half of dorsal field) on ventral Description. Male (holotype). Body surface, lower tegmen with distinctly larger rather small for this genus. Colouration mirror (Figs IV: 6, 7), hind basitarsus with (Figs IV: 9–12) dark brown with head dor- inner row of dorsal denticles consisting of sum and pronotal disc brown, yellowish line one apical denticle only, anal plate with along dorsal edge of each eye, a pair of small slightly convex hind edge between postero- yellowish marks on rostral apex, antennae lateral corners, genital plate with roundly and maxillary palpi as in O. asymmetricus truncate apex (Fig. IV: 8), and genitalia as but with slightly darker scape, pronotum in Figs V: 4–6. with a pair of light brown longitudinal lat- Female. General appearance similar to eral stripes on disc (these stripes distinct in that of male, however dorsum of head with posterior half and less distinct in anterior more or less distinct light longitudinal one), dorsal field of upper tegmen with yel- stripes, colouration of legs and abdominal lowish grey lateral band and wider yellow tergites sometimes slightly darker (with band along posterior edge (Fig. IV: 11), more contrast spots and marks), tegmina dorsal field of lower tegmen yellowish with similar to those of females of O. asymmetri- semitransparent membranous part (Fig. IV: cus but somewhat shorter and not contact- 10), colouration of legs and abdomen ap- ing with each other (their colouration with proximately as in holotype of O. cuyabeno light part of dorsal field larger, occupying but slightly darker, venter of thorax and third or half of this field), other bodyparts pleurites light brown. Structure of body also similar to those of these females but similar to that of O. asymmetricus, but teg- with ovipositor slightly longer (hind femur mina somewhat different in size and with 1.3–1.4 times as long as ovipositor). lateral field modified (only one short lon- Length in mm. Body: male 10, female gitudinal vein in proximal half of this field 9–11; pronotum: male 1.8, female 1.9–2.2; preserved, and tegminal gland with hairs on tegmina: male 3.5–3.8, female 1.5–2; hind ventral surface of special fold in distal half femora: male 7.2, female 7.5–8.7; ovipositor of this field developed; Fig. V: 10), dorsal 5.3–6.7. field of upper tegmen distinctly widened in Comparison. The new species differs distal half (Fig. IV: 11) and with less dis- from O. ottei (another species of this sub- tinct stridulatory vein on ventral surface, genus) in the male genitalia with somewhat dorsal field of lower tegmen with membra- longer membranous parts of the posterolat- nous part occupying about half of surface eral epiphallic arms, with shorter ectopara- of this field (this part slightly immersed meres and with much wider membranous and without mirror) and with almost semi- lateral lobes of the ectoparameres. sclerotized rest part lacking distinct vena- Etymology. The new species is named af- tion (Fig. IV: 10), hind basitarsus and geni- ter the Cuyabeno River. tal plate as in O. cuyabeno, anal plate with

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 23 slightly less angular posterolateral corners, simple, with rounded apex. Male genital and genitalia as in Figs V: 7–9, 11. plate short and with three small angular Female. General appearance as in male, apical lobules located rather far from each however body clearly larger, colouration other (Fig. XVI: 3); female genital plate dis- of pronotal disc less contrast, colouration tinctly smaller, almost triangular but with of abdominal tergites more contrast, other widely rounded and slightly notched apex bodyparts similar to those of females of O. (Fig. XVI: 8). Male genitalia: epiphallus cuyabeno but with ovipositor insignificantly strongly curved in profile; posterolateral longer (hind femur approximately 1.2 times epiphallic arms long and distally bifurcate; as long as ovipositor). ectoparameres also long, partly fused with Length in mm. Body: male 8.5, female epiphallus and articulated with endoparam- 11; pronotum: male 1.6, female 2; male up- eres (distal part of ectoparameres well vis- per tegmen 2.8; male lower tegmen 2; female ible above posterolateral epiphallic arms in tegmina 1.4; hind femora: male 6, female 7.5; profile); endoparameres short, connected ovipositor 6.3. with each other by narrow sclerotized rib- Etymology. The new species is named af- bon; endoparameral apodemes short; poste- ter the Aguarico River. rior endoparameral arms practically absent; formula small and with not large anterior Genus Laozacla gen. nov. apodeme (Figs XVI: 5–7). Ovipositor normal, with slightly higher subapical part as Type species L. furca sp. nov. well as with narrower and acute apical part Diagnosis. Body medium-sized. Head (Fig. XVI: 9). almost semiglobular, distinctly (in male) Included species. Type species only. or slightly (in female) narrower than pro- Comparison. The new genus is more or notum and with rather thin apical segment less similar to Luzarina in the structure of of maxillary palpi (its shape intermediate head and of male genitalia (including size between those of Luzara and Uvaroviella; of rachis), but it is distinguished by the see Figs I: 3, 5). Pronotum transverse, dis- following combination of characters: both tinctly (in male) or slightly (in female) nar- male tegmina with normal membranes in rowing to head and having rather widely the stridulatory apparatus, with a rounded rounded anteroventral corners; male meta- mirror having two distinct dividing veins, notal gland developed (Figs XVI: 1, 4). and practically without apical area of the Male tegmina rather wide but somewhat dorsal field; male anal plate with a pair of shortened (practically without apical area), rather long and thin posterolateral process- with stridulatory apparatus distinct, with es, and male genitalia with long ectopara- venation similar in both tegmina, and with meres situated mainly above (not under) large and almost rounded mirror crossed the posterolateral epiphallic arms and with by two dividing veins (Fig. XVI: 1); hind strongly reduced posterior endoparameral wings in male and all wings in female ab- arms. sent. Legs thin (but hind femora thickened Etymology. The generic name originates in proximal half, adapted for jumping) and from the Laos Country and genus Zacla. moderately long; fore tibiae with only inner tympana; hind tibiae with denticles Laozacla furca sp. nov. (excepting four pairs of articulated spines) (Figs XVI: 1–10) situated mainly on outer dorsal edge and Holotype. Male; Laos, Champasak Prov., “Bo- with inner dorsal apical spur longer than laven Plateau, 14 km SE of Muang Paxong, Ban all other apical spurs. Male anal plate with Houayteuay”, 15°4.655´N, 106°16.848´E, 1200 m, a pair of rather long and thin posterolateral disturbed mountain forest, “pitfall traps”, 6 Dec. processes (Fig. XVI: 2); female anal plate 2007 – 6 May 2008, S. Tarasov (ZIN).

Paratypes. Thirty two males and 38 females, pronotum somewhat lighter (almost light same data as for holotype (ZIN); 2 males, same brown in head, and with light brown me- data but “carrion trap”, 6 May – 14 June 2008 dian and hind parts in pronotal disc), ptero- (ZIN). thoracic and abdominal tergites light brown Description. Male (holotype). Coloura- with numerous darkish marks, abdominal tion light brown but with following marks: venter also somewhat lighter than in male, head with areas behind eyes dark brown, anal plate uniformly light brown, and meta- with apical segment of maxillary palpi notum without gland; structure of genital mainly greyish brown, and with spots on plate and of ovipositor as in Figs XVI: 8, 9. mandibles and under eyes as well as not Length in mm. Body: male 10–11, fe- very distinct longitudinal stripes on dor- male 10.5; pronotum: male 1.9–2.2, female sum and mark on third segment of maxil- 2.3; tegmina, male 6–7; hind femora: male lary palpi brown; pronotum brown with al- 7.5–8.5, female 8; ovipositor 7.5. most light brown posterior part of disc and Etymology. The species name is the Latin small lightish spot on anteroventral part of word “furca” (fork). each lateral lobe; tegmina with brown upper half of lateral field, slightly lighter (greyish Subtribe PHALANGOPSINA brown) lower half of this field, yellowish venation in this field, and almost whitish Note. This subtribe has a characteristic veins in dorsal field of lower tegmen; legs shape of the head: it is with a rather nar- with not very distinct darkish spots on fore row, distinctly projected and more or less and middle femora as well as somewhat angular (in the profile) rostrum separated more distinct brown spots on tibiae and from the rest of vertex by a dorsal concav- tarsi; abdomen with dark brown tergites ity (sometimes almost indistinct) and hav- as well as with greyish brown genital plate ing the ocelli located almost on the rostral and nearest sternites; anal plate with brown dorsum (near its apex and base); and max- processes and a pair of small spots near cer- illary palpi with a very long and thin apical bases (Figs XVI: 1–3). Ocelli well de- cal segment having the distal part more or veloped, not large; rostrum between anten- less similar to that of Luzarina (Figs I: 1, nal cavities approximately twice wider than 2; VI: 1, 2). Also this subtribe has long and scape; general shape of head and pronotum relatively thin legs (such structure of the (dorsal view) as well as structure of dorsal head and legs are usual for representatives field of tegmina as in Fig. XVI: 1. Metanotal of Phalangopsinae living on tree trunks), gland as in Fig. XVI: 4. Tegmina reaching the hind tibiae with the middle inner api- hind part of ninth abdominal tergite (Fig. cal spur distinctly longest, and rachis of the XVI: 1), with lateral area having moder- male genitalia elongate but thin or almost ately widened R-M area and rather narrow finger-like (Figs II: 1–5, 11, 12; VI: 5–8). (low) area of Sc branches (this area almost equal to Sc-M area in width) as well as al- Eidmanacris longa sp. nov. most lacking crossveins. Anal and genital (Figs I: 2; VI: 1–12) plates as in Figs XVI: 2, 3. Genitalia as in Figs XVI: 5–7, 10. Holotype. Male; Bolivia, Santa Cruz De- partment, 23 km SW of Santa Cruz City, El Soc Variations. Some males with dorsal field Natural Park (small private area with secondary of upper tegmen somewhat darker (almost forest), about 600 m, on wet soil in dry brook at greyish brown) as well as with lateral teg- night, 14–16 Feb. 2014, A. Gorochov (ZIN). minal field having R-M area and large spot Paratypes. Three males and 2 females, same before this area dark brown. data as for holotype (ZIN). Female. General appearance more or less Description. Male (holotype). Body similar to male, but dorsum of head and of rather large for this genus. Colouration

Figs VI (1–16). Eidmanacris: 1–12, E. longa sp. nov.; 13–16, E. ?marmorata (Bruner). Head, pronotum and tegmina of male from above (1) and from side (2); male abdominal apex from above (3); male metanotal gland from above (4); male genitalia from above (5), from below (6), from side and slightly above (7), and from side (8); female copulatory papilla from side (9, 14) and from above (10, 15); distal part of ovipositor from side (11); female genital plate from below (12, 16); dorsal field of male tegmen (13), after Desutter-Grandcolas (1995).

(Figs VI: 1–4) greyish brown with dark pi; antennae dark brown with light brown brown, light brown and yellowish spots: scape (having a few darkish marks), rather head (without antennae) yellowish with sparse and small whitish spots on proximal more or less dark dorsum, rostrum, three part of flagellum, moderately long whitish vertical stripes on lower half of head, a pair spot on flagellum near this part, and very of spots on genae, and darkish areas on pal- sparse and small whitish spots on rest of

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 26 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP flagellum; pronotum dark brown with nu- Ovipositor rather long (hind femur 1.2 merous small and not very distinct lighter times as long as ovipositor) and with apex marks; tegmina brown (rather dark) but as in Fig. VI: 11; copulatory papilla as in with yellow stripe in distolateral part; legs Fig. VI: 9, 10. with dark and light spots more or less equal Length in mm. Body: male 19–22, fe- to each other in size; abdomen less distinct- male 18–20; pronotum: male 4.5–4.7, fe- ly spotted (darkish) but with long median male 4.4–4.6; tegmina, male 3.5–3.8; hind yellowish line on dorsum, yellowish venter, femora: male 20–22, female 20–21; oviposi- and light greyish brown anal plate and cer- tor 17–17.5. ci; other parts of body yellowish with more Comparison. The new species is most or less distinct darkish marks on pleurites similar to E. marmorata from Sara Province and tergites. Head typical of Eidmanacris, of Bolivia and to E. corumbatai Garcia, 1998 however its dorsum in profile with shallow from Brazil (“Cerrado de Corumbatai”). It but distinct concavity near lateral ocelli differs from E. marmorata in the dorsum of (Fig. VI: 2). Thorax, wings and legs also head with a slightly concave rostral base in similar to those of other congeners (legs the profile (this part of rostrum is practical- very long, tympana and hind wings absent) ly straight in E. marmorata), male tegminal but with some peculiarities: metanotum apex less narrow (for comparison see Figs with gland consisting of characteristically VI: 1, 13), male anal plate with less thin and curved anteromedian process having apical not curved posterolateral processes as well part widened and with hairs (Fig. VI: 4); as with a shorter posteromedian projection, tegmina semisclerotized, short (reaching the characters of male genitalia mentioned base of first abdominal tergite), narrowing above, female genital plate with a somewhat to almost angular apex, almost without dis- wider notch (see Figs VI: 12, 16), female tinct venation excepting one longitudinal copulatory papilla larger and with a clearly vein between dorsal and lateral fields (Figs wider proximal notch (see Figs VI: 9, 10, VI: 1, 2); longest (middle inner) apical spur 14, 15), as well as in a distinctly longer ovi- of hind tibiae slightly longer than half of positor (in E. marmorata, hind femur is 1.45 hind basitarsus. Anal plate with a pair of times as long as ovipositor). From E. corum- elongate posterolateral processes and very batai, the new species differs in the median short, rounded and slightly bilobate median edge of the tegminal proximal half lacking projection (Fig. VI: 3); genital plate mod- any light stripe, and lateral epiphallic lobes erately elongate and with roundly bilobate without long finger-like distal processes; apex (Fig. VI: 3); genitalia similar to those and from all the other congeners, in the of E. marmorata (Bruner, 1916) but with shape of some structures: epiphallus, male posterolateral lobes of epiphallus somewhat anal plate, male tegmina, or female copula- different in shape (their hook-like apical tory papilla. spines thinner and longer), dorsal sclerite of Etymology. The species name is the Latin rachis longer and more deeply notched, and word “longa” (long). anteromedian notch of epiphallus distinctly deeper and narrower (Figs VI: 5–8). Eidmanacris ?marmorata (Bruner, 1916) Variations. Dorsal sclerite of rachis well (Figs VI: 13–16) sclerotized in one male (paratype) and semimembranous in all other males. Material examined. Female; Bolivia, Female. General appearance similar to Santa Cruz Department, near 80 km NWW that of male, but tegmina and metanotal of Santa Cruz City, Amboro National Park, gland absent, anal plate rounded (without Mataracu Camp, about 800 m, primary for- processes), genital plate with not very nar- est, on leaf of very low bush at night, 8–19 row posteromedian notch (Fig. VI: 12). Feb. 2014, A. Gorochov (ZIN).

Note. This specimen is in accordance to viding veins in mirror; tegmina in female the original description and redescription and hind wings in both sexes absent. Legs by Desutter-Grandcolas (1995) including with open oval inner tympanum only, and length of its ovipositor, but its copulatory longest apical spur (middle inner) of hind papilla slightly higher (it is a reason that my tibia clearly longer than half of hind basitar- determination of this female is under ques- sus. Metanotal gland absent: only traces of tion). Possibly it was collected in a locality it (looking as a pair of slight convexities on situated less far from the type locality of E. metanotum) visible. Male anal plate with a marmorata than from that of E. longa, and pair of rather long and thin posterolateral I cannot exclude that these species may be processes (Fig. XV: 1); male genital plate two geographical subspecies of the same slightly longer and with roundly truncate species only. apex. Male genitalia rather long and distinctly narrowed before distal part; epiphal- Subtribe AMPHIACUSTINA lus elongate; inner ectoparameres (articulated with endoparameres) long and nar- Note. The subtribe is similar to Pha- row; outer ectoparameres (articulated with langopsina in the general appearance (in- posterolateral epiphalliuc arms) more or cluding shape of head, length of legs, and less ovally widened and almost horizontally structure of tibial spurs) and structure of situated; rachis large (long) and directed the male genitalia (its rachis may be rath- backwards; formula small; endoparameral er large and partly sclerotized). However apodeme rather short and directed laterally often this rachis is short and directed up- and forwards; rami very narrow (almost in- wards (almost as in Luzarina) but distinctly distinct; Figs XIII: 13, 14). sclerotized. From Phalangopsina (as well Included species. Type species; Amphia- as from Luzarina and Lernecina), this sub- custes caraibea Saussure, 1897; Amphia- tribe is not very clearly distinguished by custa saba Desutter-Grandcolas et Otte, the presence of inner and outer ectopara- 1997; A. dominica Otte et Perez-Gelabert, meres (outer ectoparameres are formed by 2009; A. iviei Otte et Perez-Gelabert, 2009; articulated or partly articulated distal parts A. renodis Otte et Perez-Gelabert, 2009; A. of the posterolateral epiphallic arms, and in- kittsia Otte et Perez-Gelabert, 2009; A. ca- ner ectoparameres sometimes may be partly icosensis Otte et Perez-Gelabert, 2009; A. fused with lateral ones or deeply divided cavicola Otte et Perez-Gelabert, 2009. into two sclerotized lobes; Figs XIII: 13, 14; Comparison. The new genus is distin- XV: 5–7). guished from Amphiacusta and Noctivox by a strong reduction of the male abdomi- Genus Caribacusta gen.nov. nal gland and a few characters of the male genitalia: epiphallus elongate; lateral ecto- Type species: C. antigua sp. nov. parameres almost oval and partly horizon- Diagnosis. Scape approximately 1.5 tal; rachis large and straight. From all the times as wide as head rostrum between other genera of Amphiacusta, it differs in the antennal cavities. Ocelli distinct: median male genitalia distinctly narrowed before ocellus situated almost at apex of rostrum; distal part and some additional characters: lateral ocelli located on head dorsum near from Longuripes (including the subgenus base of rostrum, not very near each other. Prolonguripes Desutter-Grandcolas, 1993, Pronotal lateral lobes with anteroventral stat. nov.) by not bifurcated outer ecto- corner strongly projected downwards but parameres, from Nemoricantor by the medial rounded. Tegmina in male distinctly (but ectoparameres not divided into two sepa- not very strongly) shortened and with nor- rate sclerites, from Mayagryllus by a much mal stridulatory apparatus having 1–2 di- longer anterior part of epiphallus (curved

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 28 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP upwards and backwards), from Cantrallia Comparison. The new species is most by a distinctly longer epiphallus, and from similar to C. iviei comb. nov. in the head Arachnopsita and Leptopedetes by the pres- colouration and structure of the male geni- ence of tympana and male tegmina (in Lep- talia, however its head dorsum is with a topedetes, tegmina are developed but very narrower transverse band (not reaching the small, scale-like; however in the new genus, lateral ocelli), and its outer ectoparameres male tegmina are much larger and with a are with a more angular apical part. From C. normal stridulatory apparatus). renodis comb. nov., the new species differs Etymology. The generic name originates in somewhat narrower outer ectoparameres from the Caribbean Sea (Mare Caribaeum having a more angular apical part; and from in Latin) and genus Amphiacusta. C. caraibea comb. nov., C. saba comb. nov., C. dominica comb. nov., C. kittsia comb. Caribacusta antigua sp. nov. nov., C. caicosensis comb. nov. and C. cavi- (Figs XIII: 13, 14; XV: 1) cola comb. nov., in significantly less protruding lateroproximal edges of the lateral Holotype. Male; The Lesser Antilles, Anti- ectoparameres. gua I., “W. Indies: Antigua, 1932, A.D. Torlesse, Etymology. This species is named after B.M. 1932-148” (NHM). the Antigua Island. Description. Male (holotype). Body colouration brown (rather dark) with follow- Noctivox orizaba sp. nov. ing marks: head with yellowish face (from (Figs XV: 2–7) median ocellus to labrum) having four brown vertical bands on epicranium and Holotype. Male; Mexico, Veracrus State, two such bands on mouthparts, with light “Magdalena, Orizaba”, 18°48.2´N, 97°03.7´W, stripe along hind edge of each eye, with 1198 m, 9 Aug 2011, V. Sinjaev (ZIN). light brown transverse band on dorsum be- Description. Male (holotype). Body tween hind part of eyes, with more or less rather small for this genus and more or less yellowish longitudinal stripes on palpi, and dark. Head with black dorsum having light with almost light brown scapes; pronotum brown transverse line between lateral ocel- with yellowish stripe along all edges and a lus and eye as well as very short longitu- pair of light brown spots on disc; tegmina dinal lines on hind part, with dark brown light greyish brown; legs almost reddish face having light brown transverse stripe brown but slightly lighter than most part on rostral apex (including median ocellus) of body and with weakly distinct darkish and three short vertical stripes near clyp- and lightish spots on distal half of fore and eus as well as a few light spots on mouth- middle femora as well as on middle tibiae; parts, with greyish brown genae having meso- and metanotum as well as venter of light brown lower parts and lateral surfaces body almost light brown. Tibia with oval of mandibles as well as yellowish subgenae, open tympanum on inner side of both fore with greyish brown palpi and antennae hav- legs and with smaller one on outer side of ing slightly lighter apical segment of palpi only left fore leg; hind tibiae with four inner as well as partly yellowish scape and small and four outer articulated dorsal spines sit- sparse light brown spots on antennal fla- uated not near each other. Tegmina reach- gellum; pronotum black with small barely ing middle of abdomen as minimum (their lighter (dark reddish brown) mark on each distal part missing). Anal plate and genita- anterolateral corner; legs dark brown with lia as in figures (Figs XIII: 13, 14; XV: 1). rather small and not numerous light brown Female unknown. spots on fore and middle legs as well as with Length in mm. Body 14.5; pronotum 2.8; light greyish brown proximal half of hind hind femora 14.5. femora having longitudinal dark median

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 29 stripe on lateral surface (this stripe fused Subtribe MODESTOZARINA with dark distal half); tegmina greyish subtrib. nov. brown with somewhat lighter (almost light greyish brown) dorsal field; venter of body Type genus Modestozara gen. nov. (gen- and rest of thorax greyish brown; abdomi- der feminine) nal tergites and anal plate dark brown; cerci Diagnosis. Structure of head similar to brown. Rostrum of head between antennal that of Luzarina, but maxillary palpi thin- cavities almost as wide as scape; maxillary ner and longer, intermediate between those palpi approximately intermediate between of Phalangopsina and Luzarina (sometimes those of Luzara (Fig. I: 5) and Uvaroviella possibly more or less similar to that of Lu- zara; Fig. I: 6). Hind tibia with dorsal inner (Fig. I: 3). Structure of metanotal gland and apical spur longest. Rachis of male genitalia dorsal field of tegmina as in figures (Figs long or moderately long and having finger- XV: 3, 4); stridulatory vein of upper tegmen like or angular apex (Figs VII: 4–6, 12–14; with 114 teeth; hind wings absent. Legs VIII: 2–4, 6–8, 10–12; IX: 5–7). with inner tympanum open and oval, with Note. Included genera are listed in the outer tympanum almost twice smaller, and classification of Phalangopsinae subfamily with longest (inner median) apical spur of group given above. From the other subtribes hind tibiae slightly longer than half of hind of Phalangopsini, the new subtribe differs in basitarsus. Anal plate with characteristic the head outlines and apical spurs of hind lateroproximal tubercles (Fig. XV: 2); geni- tibiae as in Luzarina and Lernecina, and in talia as in Figs XV: 5–7. the rachis of male genitalia as in Phalangop- Female unknown. sina (from Amphiacustina, the new subtribe Length in mm. Body 13.5; pronotum 2.3; additionally differs in the same characters tegmina 7.8; hind femora 13. of ectoparameres as Phalangopsina; see the Comparison. The new species is similar notes on Amphiacustina above). to N. bolivari (Chopard, 1947) described also from Veracruz State (Atoyac) but dis- Genus Modestozara gen. nov. tinguished by the male genitalia having a wider lateral ectoparameres in the profile, Type species M. modesta sp. nov. higher (longer) arcuate sclerites articulated Diagnosis. Body medium-sized, with with the posterolateral epiphallic arms as short pubescence, with very small (scale- well as with the median ectoparameres, a like) male tegmina lacking venation and longer (wider) proximal part of these scler- stridulatory apparatus, without tegmina in ites, and somewhat different shape of the female, and with very long thin legs (Figs median ectoparameres in the profile (they VII: 1, 2, 10) lacking tympana and having have a distinctly narrower distal part and only a few very small dorsal denticles on are situated very near the proximal epiphal- distal part of hind basitarsus (excepting a lic part; Fig. XV: 7). From N. chopardi De- pair of larger apical spurs). Anal plate sim- sutter-Grandcolas, 1995 poorly described ple: somewhat elongate, with flat or slightly from environs of Cordoba City (another concave dorsum, and with rounded apex locality in Veracruz State), the new species (Figs VII: 3, 11); male genital plate distinct- differs in the epiphallus not longer than in ly longer, with truncate or slightly bilobate N. bolivari, endoparameral apodemes (their but more or less rounded apex (Figs VII: 3, proximal part as minimum) not larger than 11); female genital plate short, weakly and in this species, and stridulatory vein with roundly bilobate at apex (Fig. VII: 9). Male 114 (not 253–291) teeth. genitalia (Figs VII: 4–6, 12–14) similar to Etymology. The new species is named af- those of Endecous but distinguished by pos- ter its type locality. terolateral epiphallic arms rather thin and

Figs VII (1–18). Modestozara: 1–9, M. modesta sp. nov.; 10–16, M. troglophila sp. nov.; 17, 18, M. satipo sp. nov. Head and thorax of male from above (1) and from side (2); male abdominal apex from above (3, 11); male genitalia from above (4, 12), from below (5, 13) and from side (6, 14); female copulatory papilla from above (7, 15, 17) and from side (8, 16, 18); female genital plate from below (9); male pterothorax from above (10).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 31 not bifurcate, rachis moderately long (ap- yellow median line, yellowish lateral parts, proximately finger-like) and almost com- and brown to light brown marks along hind pletely membranous (not semisclerotized edge of dorsum; abdominal tergites similar dorsally) as well as with small semisclero- to it in colouration but with very dark lat- tized ventral lobule (absent in Endecous), eral parts and without light median line in and ectoparameres wide (not elongate) and hind half of abdomen; venter of body light distinctly not reaching apices of posterolat- brown to yellowish, but genital plate as well eral epiphallic arms (in Endecous, ectopara- as anal plate darker (greyish brown); cerci meres narrower, more or less elongate and light greyish brown. Head large, high, with reaching or almost reaching apices of the rostrum slightly narrower than scape; lat- latter arms). eral lobes of pronotum with anteroventral Included species. Type species; M. tro- corner distinctly angularly projected down- glophila sp. nov.; M. satipo sp. nov. wards; metanotum without distinct traces Comparison. The new genus is distin- of gland; scale-like tegmina partly covered guished from Endecous by the above-men- by pronotum, not contacted with each oth- tioned characters of hind tarsi and of male er, and having rounded posterior edge (Fig. genitalia. VII: 1, 2); hind wings absent; anal plate with Etymology. The generic name originates rather strongly widened base and slightly from the Latin word “modestus” (modest) concave dorsum; genital plate with roundly and a part of the generic name Luzara. truncate apex (Fig. VII: 3); genitalia as in Figs VII: 4–6. Modestozara modesta sp. nov. Female. General appearance similar to (Figs VII: 1–9) that of male, but dorsum of head and rostrum almost completely darkened, tegmina Holotype. Male; Ecuador, western slopes of absent, and anal plate slightly shorter and Andes not far from Pacific Coast, 10 km E of gradually narrowing to apex; genital plate Agua Blanca Vill. (environs of Puerto Lopez short and weakly roundly bilobate (Fig. Town), San Sebastian Camp, about 700 m, misty primary forest, on wall of small and widely open VII: 9); hind femur approximately 1.5 times cavity between stones very near brook, at night, as long as ovipositor; copulatory papilla as 26–29 Oct. 2005, A. Gorochov, A. Ovtshinnikov in Figs VII: 7, 8. (ZIN). Length in mm. Body: male 10.5, female Paratype. Female, same data as for holotype 11; pronotum: male 2.5, female 3.1; exposed (ZIN). part of tegmina, male 0.5; hind femora: male Description. Male (holotype). Body 13, female 14.5; ovipositor 9.5. rather large for this genus. Coloration grey- Etymology. Name of the new species is ish brown (more or less light) but with nu- the Latin word “modesta” (modest). merous and usually not very contrast lighter and darker spots (Figs VII: 1, 2): head Modestozara troglophila sp. nov. almost yellowish with three brown verti- (Figs VII: 10–16) cal stripe on face, brown areas along dorsal edges of antennal cavities, light brown to Holotype. Male; Ecuador, Los Tayos, 3°10´S, brown mandibles and marks on hind half of 78°12´W, “on wall of main cave at bottom of sec- dorsum, light brown palpi, and dark brown ond shaft”, 12 July 1976, Edinburg University antennae (but scape, pedicel and distinct Expedition “B.M. 1977-243” (NHM). Paratypes. Male, same data as for holotype spots on flagellum light, yellowish to whit- but “in main cave silty ledge above climb down ish); pronotum with somewhat darker (al- to stream passage” (ZIN); 2 females, one of them most uniformly brown) lateral lobes; teg- with data as for male paratype, and second female mina almost uniformly light brown; meta- with same data but having label “main shaft, on notum dark brown (almost blackish) with wall of N. E. passage” (NHM, ZIN).

Description. Male (holotype). Size, co- 800 m, partly primary / partly secondary forest louration and structure of body similar to near waterfall, among stones on bank of brook at those of M. modesta but with following dif- night, 4–5 Nov. 2008, A. Gorochov, M. Berezin, ferences: colouration almost uniformly light L. Anisyutkin, E. Tkatsheva, V. Izersky (ZIN). Paratypes. Three females, same data as for brown with brown lower half of head (but holotype (ZIN). labrum and palpi yellowish, and mandibles Description. Female (holotype). Body dark brown), light greyish brown antennae distinctly smaller than in both previous having yellowish scape and whitish spots congeners. Colouration similar to that of M. on flagellum, brown pronotum having disc modesta but lighter: head (including man- slightly lighter and anteroventral corners dibles) yellowish with three brown vertical lightish, brown to light brown other tergites stripes on lower half, a pair of brown marks (proximal ones darker), legs with barely on genae, and brown antennal flagellum spotted femora, and venter of body (except- having rather sparse and very small whitish ing genital plate) almost yellowish; tegmina spots; pronotum with light brown disc and with almost truncate distal part (Fig. VII: brown lateral lobes having light stripe along 10); anal plate more gradually narrowing to lower half of anterior edge; metanotum and apex (Fig. VII: 11); genital plate with slight- legs approximately as in M. modesta in co- ly roundly bilobate apex (Fig. VII: 11); geni- louration; abdomen light brown (almost yel- talia slightly wider and with distinctly more lowish) with brown areas on lateral parts of curved (sinuate in profile) posterolateral first, fourth, fifth and tenth tergites, as well epiphallic arms (Figs VII: 12–14). as with brown and brownish dots on rest of Variations. Paratype with scape and ros- tergites. Structure of body also similar to trum somewhat darker (almost brown) and that of female of M. modesta, but copula- tibiae barely spotted (almost as femora). tory papilla (Figs VII: 17, 18) and oviposi- Female. General appearance similar to tor somewhat shorter (hind femur approxi- that of male, but structure of tegmina and mately 1.7 times as long as ovipositor). of anal and genital plates as in female of Variations. Sometimes colouration slight- M. modesta; hind femur approximately 1.8 ly darker (general colour light brown) and times as long as ovipositor; copulatory pa- with almost blackish spots on metanotum pilla distinctly wider than in this species and hind femora. (Figs VII: 15, 16). Male unknown. Length in mm. Body: male 11.5–12, fe- Length in mm. Body 9–11; pronotum male 12–14.5; pronotum: male 2.3–2.5, fe- 2–2.2; hind femora 10.5–11.5; ovipositor male 2.6–3; exposed part of tegmina, male 6.2–6.7. 0.4–0.6; hind femora: male 15–16, female Comparison. The new species differs from 16–17.8; ovipositor 9–9.6. all the other congeners in a smaller body Comparison. The new species differs size; from M. troglophila, in a more spotted from M. modesta in a less spotted coloura- colouration, less convex ventral part of the tion, dark most part of lower half of head, female copulatory papilla (see in the pro- more curved posterolateral epiphallic arms file), and barely longer ovipositor; and from in the male genitalia, and shorter ovipositor. M. modesta, in the characters listed above Etymology. The species name originates (in the description). from the Greek roots meaning “cave-lover”. Etymology. The species is named after Satipo Province. Modestozara satipo sp. nov. (Figs VII: 17, 18) Genus Endecous Saussure, 1878

Holotype. Female, Peru, Junin Department, Note. This genus is rather diverse in the Satipo Prov., environs of Satipo Town, about structure of body and male genitalia and

Figs VIII (1–12). Endecous, male: 1–4, E. lizeri Rehn; 5–8, E. arachnopsis Sauss.; 9–12, E. onthophagus (Berg). Dorsal field of lower (1) and upper (5, 9) tegmina; genitalia from above (2, 7, 11), from below (3, 8, 12) and from side (4, 6, 10).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 34 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP must be divided into three subgenera as Sperber, 2008: fig. 5, C) ...... minimum. Descriptions of these subgenera ...... subgen. Endecous s. str. are given below, in a key for subgenera of [Included species: E. arachnopsis Saussure, the genus Endecous: 1878 (type species) from Brazil (Figs VIII: 5–8) and some of the other congeners from 1. Male genitalia: rachis very large (signifi- Brazil and possibly from Bolivia listed as be- cantly protruding behind epiphallic postero- longing to the genus Endecous in the Orthop- lateral arms), lamellar (strongly laterally tera Species File (Eades et al., 2014).] compressed), and almost completely membranous; formula very long, forming dorsal Genus Daedalonotum gen. nov. part of distinctly elongate membranous cavity sometimes looking as sacculus of Grylli- Type species D. daedalum sp. nov. nae (Figs VIII: 2–4, 10–12) ...... subgen. Notendecous subgen. nov. Diagnosis. Body medium-sized for Pha- [Included species: Dyscophus onthophagus langopsini. Head not large, almost equal to Berg, 1891 (type species) from Uruguay pronotum in width, with apical segment of (Figs VIII: 9–12; E. lizeri Rehn, 1918 from maxillary palpi having oblique distal part Argentina (Figs VIII: 1–4); possibly E. hub- rather short and barely arcuate (Fig. I: belli Liebermann, 1965 from Argentina. Ety- 6); pronotum with anteroventral corners mology: the subgeneric name originates from rather widely rounded (not distinctly pro- the Greek prefix meaning “southern” and the jected downwards); male metanotum with generic name Endecous.] gland consisting of very large transverse – Male genitalia: rachis not very large (not protruding behind epiphallic posterolat- inflation having additional transverse lobe eral arms), more or less finger-like but not with hairs, transverse lamellar fold situated lamellar, and partly membranous or semi- behind (near) this inflation and provided sclerotized; formula moderately long or short with vertical median process, and second (elongate membranous cavity weakly devel- transverse lamellar fold formed by hind oped and not looking as sacculus, or undevel- part of metanotum curved upwards (Fig. oped; Figs VIII: 6–8) ...... 2 IX: 2). Male tegmina distinctly shortened 2. Body apterous in both sexes; fore tibiae (reaching middle part of abdomen), semi- without tympana. Epiphallus in male genitalia short (its width much greater than its sclerotized, lacking stridulatory apparatus, length), with posterolateral arms divided without venation in somewhat inflate dor- into lateral and medial processes or projec- sal field of upper tegmen, and with traces of tions (but lacking distinct ventral projection venation in less inflate dorsal field of lower or even tubercle; see Souza-Dias et al., 2014: tegmen (Figs IX: 1, 3); female tegmina al- figs 3, A–D) ...... most scale-like; hind wings absent. Legs not ...... subgen. Pedroecous subgen. nov. very long, moderately thin but with clearly [Included species: E. apterus Bolfarini et Sou- thickened (for jumping) hind femora; tym- za-Dias, 2014 (type species) from Brazil only. Etymology: the subgeneric name includes the pana absent; dorsal inner apical spur of hind first name of Pedro G.B. Souza-Dias (one of tibiae longest; hind basitarsus with two the authors of E. apterus) and a part of the rows of dorsal denticles (excepting a pair of generic name Endecous.] apical spurs). Male anal plate short but with – Male tegmina well developed, with normal a pair of rather long and narrow posterolat- stridulatory apparatus having distinct mir- eral lobes; male genital plate moderately ror (Fig. VIII: 5); fore tibiae with rather elongate (Fig. IX: 4); female genital plate small inner tympana. Epiphallus in male approximately as long as wide and with pos- genitalia elongate (its width slightly greater teromedian notch (Fig. IX: 9). Epiphallus than its length), with posterolateral arms divided into dorsal and ventral processes or in male genitalia very strongly transversally projections (ventral one from rather large curved (“anterolateral” epiphallic lobes di- and roundly lobe-like to small, almost tu- rected backwards, although in taxa with bercle-like; Figs VIII: 6–8; see also Mews & less curved epiphallus, latter lobes directed

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 35 forwards or upwards), having posterolat- approximately 300 m, primary forest, among eral (lower) arms distally articulated with dry leaves on forest floor at night, 26–31 Oct. thin stick-like sclerites (directed more or 2008, A. Gorochov, M. Berezin, L. Anisyutkin, less medially) as well as basal parts of these E. Tkatsheva, V. Izersky (ZIN). Paratypes. Five males and 4 females, same arms partly fused with two pairs of long and data as for holotype (ZIN). more or less narrow ectoparameres (lateral Description. Male (holotype). Body col- ectoparameres possibly formed by separat- oration uniformly brown but with barely ed parts of posterolateral epiphallic arms); lighter antennae and pronotum, yellowish rachis long, membranous (not with small white distal half of maxillary palpi and dis- and narrow ventral sclerite), and having tal part of tegminal dorsal field (this field in almost lamellar distal part; formula long, upper tegmen with lightish medial area cov- anteriorly bifurcate, with high characteris- ered by upper tegmen, and ventral surface tic apodeme situated near middle of formula of both tegmina greyish white), contrast and directed upwards; rami developed (Figs colouration of metanotal gland (Fig. IX: 1, IX: 5–7). Ovipositor rather long and with 2, 3), light brown legs with darkish apical simple, acute apex. part of hind femora and sparse and weakly Included species. Type species only. darkened (almost indistinct) spots on rest Comparison. The new genus differs from of legs, almost dark brown and distinctly all the other genera of Phalangopsini in a pubescent abdominal tergites having large very large and rather highly projected male lightish areas on first and ninth tergites as metanotal gland, the absence of tegminal well as light brown anal plate (with a few stridulatory apparatus, a somewhat inflate small darker marks; Fig. IX: 4), and yel- semisclerotized dorsal field in the upper lowish to light brown cerci and venter of male tegmen, very strongly curved epiphal- body (including abdominal sternites and lus, the presence of two pairs of long ecto- genital plate). Dorsum of head convex in parameres, and a very large formula in the profile; head rostrum almost equal to scape male genitalia (this formula has a very char- in width; pronotum almost square in dor- acteristic vertical apodeme dissimilar to sal view; metanotal gland as in Fig. IX: 2. that of all the other congeners of this tribe). Tegmina reaching base of fourth additional Etymology. The generic name originates tergite; tegminal dorsal field with obliquely from the Latin word of Greek origin “dae- and roundly truncate hind edge (Figs IX: dalus” (intricate) and morphological term 1); tegminal lateral field with three parallel “notum” (dorsum of thorax). longitudinal veins and without crossveins. Remarks. Belonging of this genus to Eight and ninth abdominal tergites with Modestozarina is not evident, as its teg- wide concavity in hind part covered with mina are more similar to those of Luza- dense yellowish hairs (additional gland?); rina, its male anal plate similar to that of anal plate with strongly curved lobule on Amphiacustina and Phalangopsina (but apex of each posterolateral lobe and with more or less projected lateroproximal cor- small median keel (between bases of latter ners of this plate may be developed in some lobes) having one row of almost spine-like Luzarina also), and its male genitalia are but very small denticles (Fig. IX: 4); genital rather different from all the other genera of plate with rounded and very slightly bilo- Modestozarina. bate apex (Fig. IX: 4); genitalia as in Figs IX: 5–7. Daedalonotum daedalum sp. nov. Variations. Sometimes hind femora with (Figs I: 6; IX: 1–9) more distinct (darker) spots, and coloura- Holotype. Male; Peru, Ucayali Department, tion of pubescence on most part of abdomi- Atalaya Prov., about 35 km NWW of Atalaya nal tergites varied: from dark brown to Town on Rio Ucayali, environs of Sapani Vill., whitish grey.

Figs IX (1–9). Daedalonotum daedalum sp. nov.: 1, head, pronotum and tegmina of male from above; 2, male metanotal gland from above; 3, dorsal field of male lower tegmen; 4, male abdominal apex from above; 5–7, male genitalia from above (5), from below (6) and from side (7); 8, female pterothorax from above; 9, female genital plate from below.

Female. General appearance similar to anal plate more simple in shape (almost that of male, but tegmina reaching base or square with very short angular posteromedi- middle part of first abdominal tergite and an projection); genital plate as in Fig. IX: 9. with roundly angular apex, distance be- Length in mm. Body: male 11–12, fe- tween medial tegminal edges rather wide, male 10–11.5; pronotum: male 2–2.3, fe- metanotum without gland (Fig. IX: 8), and male 2.2–2.5; tegmina: male 3–3.4, female

1.3–1.5; hind femora: male 8–9, female 8.2– absent in both sexes. Hind tibiae with four 8.8; ovipositor 6.2–6.5. inner and four outer moderately long and Etymology. Name of the new species is articulated dorsal spines, without distinct the Latin word of Greek origin “daedalum” denticles between them (only more proxi- (intricate). mal denticles well developed), and with inner dorsal apical spur longest (as compared Subtribe NEMOZARINA subtrib. nov. with five other apical spurs). Median part of third and fourth tergites of abdomen in Type genus Nemozara gen. nov. (gender male with abdominal gland (Figs X: 1, 5, 9; feminine) XI: 1, 5); anal plate simple, looking as not Diagnosis. Head more or less semi- large and not long (in male) or somewhat globular (rostrum rather weakly projected elongate (in female) lobe with more or less forwards and almost not separated from rounded or almost truncate apex; genital other parts of head by a pair of anterolat- plate of male slightly longer than its anal eral concavities between eyes); maxillary plate; genital plate of female approximately palpi with apical segment short and distally twice shorter than its anal plate and widely wide (length of this segment 2–3 times as truncate (but very shallowly concave) at great as its width; XII: 2). Legs rather short apex (Fig. XI: 9). Structure of male genita- but not thick (excepting hind femora); hind lia rather primitive: epiphallus not curved, tibia with dorsal inner apical spur longest. with a pair of large posterolateral lobes di- Epiphallus not transversally curved or rected backwards; ectoparameres not large, weakly transversally curved, with differ- elongate, partly or almost completely sepa- ently developed ectoparameres, and with rated from epiphallus and articulated with rather diverse rachis (Figs X: 2–4, 6–8; XI: posterior arms of endoparameres; rachis 2–4, 6–8; XIII: 1–3, 9–11). rather large (elongate or moderately short), Note. Included genera are listed in the partly or almost completely membranous; classification of Phalangopsinae subfam- formula small but with rather long unpaired ily group given above. From the other sub- apodeme directed forwards; rami developed tribes of Phalangopsini, this subtribe dif- (Figs X: 2–4, 6–8; XI: 2–4, 6–8). Oviposi- fers in the general appearance more or less tor rather short, barely curved upwards, and similar to that of Nemobiinae, a short and with acute apical part and slightly higher wide apical segment of the maxillary palpi, (than nearest parts) subapical part (Figs and rather primitive (not distinctly curved XI: 10); width of ovipositor (without distal in the profile) male genitalia. part) more or less greater than its height. Included species. Type species; N. riorio Genus Nemozara gen. nov. sp. nov.; N. pastaza sp. nov.; N. vulcanica sp. nov. Type species N. rio sp. nov. Comparison. The new genus is distin- Diagnosis. General appearance more guished from all the other genera of Pha- or less similar to that of Nemobiinae: body langopsini by the characters given in the small and with rather large setae (especial- diagnosis of Nemozarina as well as by some ly distinct on legs), legs short and thin (as additional features: the size and shape of compared with other bodyparts) but with body more similar to those of Nemobiinae significantly thickened hind femora. Head than to those of the other taxa of Phalan- almost semiglobular (almost roundly tri- gopsini; the absence of wings, tympana and angular in front), as wide as pronotum and male notal gland; the presence of abdominal with rostrum barely wider than scape; pro- gland on third and/or fourth tergites of the notum transverse and with more or less par- male abdomen; a rather primitive structure allel sides; notal gland, wings and tympana of the male genitalia but with an elongate

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 38 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP and membranous rachis; and a comparative- usually with several or numerous brown to ly short ovipositor with the subapical part light brown marks, and anal plate somewhat slightly higher than nearest parts. different (see generic diagnosis); genital plate and distal part of ovipositor as in Figs Nemozara rio sp. nov. XI: 9, 10. (Figs X: 1–4; XI: 9, 10) Length in mm. Body: male 6.5–7, fe- Holotype. Male; Peru, Junin Department, male 7.5–8; pronotum: male 1.2–1.4, female Satipo Prov., about 25 km SE of Satipo Town, en- 1.3–1.5; hind femora: male 4.7–5.2, female virons of Rio Venado Vill., approximately 1200 m, 5.5–6; ovipositor 3.2–3.4. partly primary / partly secondary forest, on large Etymology. The species name originates leaf of grass-like plant near brook at night, 20–23 from the Spanish word “rio” meaning one Oct. 2008, A. Gorochov, M. Berezin, L. Anisyut- river (Rio Venado). kin, E. Tkatsheva, V. Izersky (ZIN). Paratypes. Two males and 6 females, same data as for holotype, but some specimens collect- Nemozara riorio sp. nov. ed among dry leaves on forest floor (ZIN). (Figs X: 5–9) Description. Male (holotype). Body rather small for this genus; colouration uni- Holotype. Male; Ecuador, eastern slopes of formly brown but with almost dark brown Andes, about 95 km E of Quito City, environs of epicranium, upper parts of mandibles, pro- San Rafael Waterfall on Rio Coca, approximate- notum, dorsum and pleurites of both meso- ly 1300 m, primary forest, on large leaf of grass- like plant near river at night, 23–26 Nov. 2005, thorax and metathorax, abdomen (except- A. Gorochov, A. Ovtshinnikov (ZIN). ing rather large whitish area of abdominal Paratypes. Male, same data as for holotype gland on fourth tergite) and apical seg- (ZIN); male and female, same data but about ments of maxillary palpi, as well as with al- 75 km SEE of Quito City, environs of El Chaco most light brown ventral parts of hind fem- Vill. on Rio Quijos, approximately 1500 m, sec- ora and small areas on all tarsi. Dorsum of ondary forest, 18–22 Nov. 2005, A. Gorochov, head convex in profile; ocelli small but dis- A. Ovtshinnikov (ZIN). tinct; pronotum approximately 1.4 times as Description. Male (holotype). Colou- wide as long and with widely rounded (not ration and structure of body very similar projected) anteroventral corners. Abdomen to those of holotype of N. rio, but maxil- with weak median concavity on third ter- lary palpi almost completely light (greyish gite and flat whitish median area on fourth white), scape light brown, third abdominal tergite (this area outlined by low and thin tergite with small (low and rather weakly keel along anterior and lateral edges; Fig. X: distinct) transverse keel situated behind 1); genital plate with roundly truncate and median concavity of this tergite (this keel very slightly concave apex; genitalia with light, yellowish; Fig. X: 5), genital plate distal part of posterolateral epiphallic arms without distinct apical concavity, and geni- more or less membranous, straight in pro- talia distinguished from those of N. rio by file and divided into very small dorsal lob- distal part of posterolateral epiphallic arms ule (almost indistinct in profile) and larger somewhat curved upwards in profile and ventral lobe, as well as with almost straight not bifurcate as well as by strongly curved rachis (Figs X: 2–4). rachis with distal part directed upwards and Variations. Other males with mesono- well visible above dorsal surface of epiphal- tum having a pair of small lightish spots or lus in profile (Figs X: 6–8). with antennal flagellum almost light brown. Variations. Second male from Rio Coca Female. General appearance as in male, with third abdominal tergite having trans- but head sometimes with weakly distinct verse keel behind median concavity slightly lighter (brown) longitudinal stripes on dor- more distinct, and male from Rio Quijos sum, pterothoracic and abdominal dorsum with this tergite having a pair of additional

Figs X (1–7). Nemozara, male: 1–4, N. rio sp. nov.; 5–9, N. riorio sp. nov., holotype (5–8) and paratype (9). Abdominal gland from above (1, 5, 9); genitalia from above (2, 6), from below (3, 7) and from side (4, 8). yellowish stripes running laterally from Comparison. The new species differs above-mentioned keel (these stripes and from N. rio in the presence of a weak light keel together forming sinuate and rather transverse keel on the third abdominal ter- long transverse yellowish line; Fig. X: 9). gite (behind median concavity of abdomi- Female. General appearance as in male, nal gland) and the above-mentioned (in de- but with weakly distinct lightish longi- scription) characters of male genitalia. tudinal lines on hind part of head dorsum Etymology. The species name originates and marks on pronotal disc, with somewhat from the Spanish word “rio” twice repeated more distinct small lightish spots on ptero- and meaning two rivers (Rio Coca and Rio thoracic and abdominal tergites, and with Quijos). anal and genital plates as well as with ovipositor similar to those of female of N. rio. Nemozara pastaza sp. nov. Length in mm. Body: male 7–8, female (Figs XI: 1–4) 8.5; pronotum: male 1.3–1.5, female 1.5; hind femora: male 5.5–6, female 6.5; ovi- Holotype. Male; Ecuador, Pastaza Prov., positor 3.8. about 10 km W of Puyo City, environs of Shell

Figs XI (1–10). Nemozara: 1–4, N. pastaza sp. nov.; 5–8, N. vulcanica sp. nov.; 9, 10, N. rio sp. nov. Male abdominal gland from above (1, 5); male genitalia from above (2, 6), from below (3, 7) and from side (4, 8); female genital plate from below (9); distal part of ovipositor from side (10).

Town, 1000–1500 m, secondary forest, on bark yellowish labrum, greyish brown antennae of tree near soil, 1–3 Jan. 2010, A. Gorochov having light brown proximal part as well (ZIN). as sparse and very small lightish spots on Paratypes. Three females, same data as for holotype (ZIN). rest of flagellum, dark brown pronotum and Description. Male (holotype). Body other tergites but having a few small light medium-sized for this genus. Colouration brown spots on pronotal disc and pterotho- brown with light brown areas on epicrani- racic dorsum as well as whitish grey abdom- um near dorsal edge of eyes, yellowish palpi inal gland and very small lightish marks on having darkish marks on proximal half, fifth and sixth abdominal tergites, yellow-

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 41 ish legs with distinct brown spots on femora marks on proximal half of maxillary palpi), and tibiae as well as on hind tarsi, yellow- labrum, coxae, and proximal part of femora, ish thoracic venter and pleurites, and light with brown lower half of mandibles, thorac- brown cerci. Structure of body similar to ic pleurites, hind tibiae (excepting lighter that of both previous species, but abdomi- spines and spurs), abdominal sternites, and nal gland consisting of two small obtusely genital plate, as well as with whitish grey angled median keels (one of them located abdominal gland (Fig. XI: 5). Structure of on third abdominal tergite and directed body very similar to that of N. pastaza (in- somewhat backwards, but second keel lo- cluding abdominal gland: Fig. XI: 5), how- cated on fourth abdominal tergite and di- ever genital plate distinctly but not deeply rected almost upwards; Fig. XI: 1), male notched at apex and with small postero- genital plate rounded but with very small median lobule, and genitalia with epiphal- posteromedian notch, and male genitalia lus lacking posteromedian lobe and having with clear anterolateral epiphallic lobes and longer posterolateral arms as well as with posteromedian lobe of epiphallus as well as somewhat longer and basally narrower ra- with rather short rachis and posterolateral chis (Figs XI: 6–8). epiphallic arms (Figs XI: 2–4). Female unknown. Female. General appearance as in male, Length in mm. Body 10.5; pronotum 2; but head somewhat lighter (with larger hind femora 9. light parts) or sometimes almost completely Comparison. From N. pastaza, the new light brown, abdomen without gland, anal species differs in a larger body size and the and genital plates as well as ovipositor simi- characters of male genitalia listed in its lar to those of female of N. rio. description; from N. rio and N. riorio, the Length in mm. Body: male 8, female new species differs in a much shorter (nar- 8.5–9.5; pronotum: male 1.5, female 1.5– rower) part of gland located on the fourth 1.6; hind femora: male 7.5, female 6.5–7.5; abdominal segment, clearly longer epiphal- ovipositor 3.5–4. lus having distinct anterolateral lobes, and Comparison. The new species is dis- the endoparameres with distinctly wider tinguished from N. rio and N. riorio by the apodemes. absence of wide flat and whitish area on Etymology. This species name is the fourth abdominal tergite in male and the Latin word “vulcanica” (volcanic) given in characters of male genitalia listed above (in connection with the type locality (Reman- description). tador Volcano). Etymology. The species is named after the Pastaza Province. Anemozara gen. nov. Type species A. vera sp. nov. Nemozara vulcanica sp. nov. Diagnosis. General appearance similar to (Figs XI: 5–8) that of Nemozara. Body small, with rather Holotype. Male; Ecuador, eastern slopes of large setae on head, pronotum and legs; Andes, about 85 km E of Quito City, Reman- head with slightly oblique face in profile; tador Volcano, primary forest, 1800–2000 m, 2 legs with fore tibiae having open oval inner Dec. 2005, A. Ovtshinnikov (ZIN). tympanum only and with hind tibiae hav- Description. Male (holotype). Body ing middle inner apical spur longest (barely rather large for this genus. Colouration dark longer than dorsal inner apical spur) and brown with light brown antennae (having slightly thickened; male tegmina (Figs XII: small sparse yellowish spots on flagellum), 1, 3, 6) extending behind middle of abdo- most part of legs, cerci, and thoracic venter, men, with normal stridulatory apparatus with yellowish palpi (having slight greyish having short stridulatory vein, with rath-

Figs XII (1–8). Anemozara: 1–5, A. vera sp. nov.; 6, 7, A. eximia sp. nov.; 8, A. propria sp. nov. Male body without cerci and distal half of hind legs from above (1); distal half of maxillary palpi from side (2); head in front and dorsal field of lower tegmen in male (3); male metanotal gland from above (4, 7); distal part of ovipositor from side (5); head in front and dorsal fields of both tegmina in male (6); head and pronotum from side and somewhat in front (8).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 43 er numerous and almost straight oblique these genera, in the posteromedian lobe of veins, with short and somewhat arcuate epiphallus longer than its posterolateral diagonal vein (its proximal part almost par- lobes as well as in the sclerotized epiphallic allel to that of lateral chord), with more or lateral parts fused with the rami and more less triangular mirror having one more or or less isolated from the median epiphallic less angularly curved dividing vein, with sclerite. very short apical area, with a few approxi- Composition. Four species grouped in mately parallel longitudinal veins in lateral two subgenera (see list of species in key for field, and without crossveins between them; subgenera of this genus below). tegmina in female and hind wings in both Etymology. Name of this genus consists sexes absent; male metanotal gland clearly of the Greek prefix meaning “not” and ge- developed (Figs XII: 4, 7); ninth abdomi- neric name Nemozara. nal tergite of male with very short median 1. Apical segment of maxillary palpi whitish. lobe; anal plate in both sexes almost equal Concavity of male metanotal gland (cov- in size and shape (rather short, with nar- ered with hairs) partly divided into a pair rowed and rounded hind part); male genital of almost round lateral concavities by low plate longer than anal plate; female geni- posteromedian convexity (Fig. XII: 4). Male tal plate with widely rounded apex having genitalia with rather short and wide epiphal- barely distinct posteromedian concavity. lus having moderately narrow and bilobate Male genitalia more or less similar to those apical lobule of posteromedian lobe directed of Nemozara but: they with a pair of partly upwards, with short and more or less simple ectoparameres, and with short rachis; Figs membranous posterolateral epiphallic lobes XIII: 1–3) ...... subgen. Anemozara s. str. (these lobes modified and more or less look- [Included species: type species; A. propria sp. ing as containing rather long posterior pro- nov.; A. umbrosa sp. nov.] jections of rami articulated at base with – Apical segment of maxillary palpi darkened anterolateral parts of epiphallus) and with or partly darkened. Concavity of male meta- larger posteromedian epiphallic lobe articu- notal gland (covered with hairs) not divided lated with ectoparameres (in one subgenus, into a pair of lateral concavities (Fig. XII: 7). ectoparameres short and looking as sepa- Male genitalia with rather long and narrow rated lateroproximal parts of this lobe, but epiphallus having simple posteromedian lobe widely truncated at apex, with elongate ec- in second subgenus, ectoparameres elongate toparameres divided into two sclerites, and and divided into two isolated sclerites); en- with elongate rachis (Figs XIII: 9–11) ...... doparameres articulated with ectoparamer- ...... subgen. Zacmozara subgen. nov. es and with long apodemes; rachis membra- [Included species: A. eximia sp. nov. (type nous, short or elongate; formula rather long species). Etymology: the generic name con- but without distinct apodeme (Figs XIII: sists of parts of the generic names Zacla and 1–5, 9–11). Ovipositor with distal part nar- Nemozara.] row (not dorsoventrally flattened) and having movable structure of upper valvae (Fig. Anemozara (Anemozara) vera sp. nov. XII: 5). (Figs XII: 1–5; XIII: 1–6) Comparison. The new genus differs from Nemozara in the presence of inner tympana Holotype. Male; Malaysia, Borneo I., Sabah State, Trus Madi Mount, about 1000 m, partly and of male tegmina; from Lernecella prob- primary / partly secondary forest, among dry ably also belonging to Nemozarina and hav- leaves on forest floor at night, 13–25 May 2007, ing approximately similar shape of dividing A. Gorochov (ZIN). veins in the male tegminal mirror, in a some- Paratypes. Five males and 3 female, same da- what wider apical segment of maxillary pal- ta as for holotype (ZIN). pi and the presence of one (not two) divid- Description. Male (holotype). Coloura- ing veins in the male mirror; and from both tion (Figs XII: 1–4) light brown, but head

Figs XIII (1–14). Anemozara and Caribacusta: 1–6, A. vera sp. nov.; 7, A. umbrosa sp. nov.; 8, A. propria sp. nov.; 9–12, A. eximia sp. nov. ; 13, 14, C. antigua sp. nov. Male genitalia from above (1, 9, 13), from below (2, 10, 14) and from side (3, 11); posteromedian epiphallic lobe with ectoparameres from behind (4, 5); outlines of female copulatory papilla from above (6–8, 12).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 45 dorsum with a few brown spots and stripes, Etymology. The species name is the Latin face (excepting labrum, areas on mandibles, word “vera” (true). and median spot on frons near clypeus) and genae brown, antennal flagellum grey- Anemozara (Anemozara) propria sp. nov. ish brown with a few small and very sparse (Figs XII: 8; XIII: 8) lightish spots, distal half of maxillary palpi whitish, pronotum with dark brown most Holotype. Female; Malaysia, Borneo I., Sa- part of lateral lobes and large area on disc, rawak State, environs of Miri Town, Lambir Hills National Park, 100–300 m, primary forest, both tegmina with almost dark brown lat- among dry leaves on forest floor at daytime, 29 eral tegminal field and yellowish humeral March – 1 Apr. 2012, A. Gorochov, M. Berezin, stripe, dorsal tegminal field brown in up- E. Tkatsheva, I. Kamskov (ZIN). per tegmen and transparent in lower teg- Description. Female (holotype). Gen- men (but its apical area also brown), other eral appearance (Fig. XII: 8) similar to bodyparts with yellowish white hairs of that of female of A. vera, but colouration metanotal gland and weakly distinct dark- and structure of body somewhat different: ish marks and lines on legs as well as dis- epicranium almost uniformly brown (with tinct dark brown spots on lateral parts of ab- only barely distinct lightish marks on dor- dominal tergites and brown venter of body sum); labrum and proximal part of anten- (including genital plate and fore coxae but nae light brown with probably somewhat not including other coxae). Scape almost darker middle part of antennae (most part 1.3 times as wide as head rostrum between of antennae missing); other mouthparts antennal cavities; pronotum and metanotal from brown to almost light brown but with gland as in Figs XII: 1, 4. Tegmina reaching whitish distal half of palpi; pronotum light base of seventh abdominal tergite, with 3–4 greyish brown with dark brown posterior longitudinal veins in lateral field, and with two thirds of lateral lobes; legs and venter of venation of dorsal field as in Figs XII: 1, 3. body light brown (but not very light) with Ninth abdominal tergite with rounded and brown spots on upper half of hind femora not very wide posteromedian lobe; genital and slightly darkish proximal areas on ab- plate approximately twice longer than anal dominal sternites; other parts of thorax and plate, almost square but with slightly nar- abdomen light greyish brown with a pair of rowed and roundly truncate hind part; gen- dark spots on lateral parts of mesonotum, italia as in Figs XIII: 1–4. large greyish brown area on metanotum, Variations. Sometimes face of head, pro- and several small dark and darkish marks notal disc and/or dorsal tegminal field al- on abdominal tergites and on anal plate; most completely dark brown; apical lobule scape approximately 1.2 times as wide as (directed upwards) of epiphallic postero- rostrum between antennal cavities; genital median lobe somewhat varied in width and plate weakly transverse; and copulatory pa- size of apical notch (Figs XIII: 4, 5). pilla rather large, strongly sclerotized, and Female. General appearance similar to distinctly bilobate (Fig. XIII: 8). that of males, but metanotal gland absent, Male unknown. and ninth abdominal tergite simple; geni- Length in mm. Body 9; pronotum 1.9; tal plate distinctly transverse, almost twice hind femora 6.4; ovipositor 5.3. shorter than male genital plate; copulatory Comparison. The new species differs papilla and ovipositor as in figures (Figs from A. vera in a distinctly wider head ros- XII: 5; XIII: 6). trum, larger and strongly bilobate female Length in mm. Body: male 9–9.5, female copulatory papilla, and some peculiarities 8.5–9; pronotum: male 1.8–1.9, female 1.8– of body colouration named above. 1.9; tegmina, male 5–5.2; hind femora: male Etymology. The species name is the Latin 6.4–6.6, female 6.5–6.7; ovipositor 5–5.2. word “propria” (separate, isolate).

Anemozara (Anemozara) umbrosa Hills National Park, 100–300 m, primary forest, sp. nov. among dry leaves on forest floor at daytime, 29 (Figs XIII: 7) March – 1 Apr. 2012, A. Gorochov, M. Berezin, E. Tkatsheva, I. Kamskov (ZIN). Holotype. Female; Malaysia, Borneo I., Sar- Paratype. Female, same data as for holotype awak State, environs of Kuching City, Kubah (ZIN). National Park, Matang Mount, 200–500 m, pri- Description. Male (holotype). General mary forest, among dry leaves on forest floor at appearance similar to that of male of A. night, 10–17 March 2012, A. Gorochov, M. Ber- vera. Body colouration (Fig. XII: 6, 7) light ezin, E. Tkatsheva, I. Kamskov (ZIN). brown but with following marks: head ros- Paratype. Two females, same data as for ho- trum slightly darker; genae and mouthparts lotype (ZIN). (excepting upper half of clypeus) yellow- Description. Female (holotype). General ish; palpi whitish with greyish brown apex appearance similar to that of female of A. of subapical segment and distal half of api- propria, but colouration and shape of some cal segment; pronotum brown with barely structures somewhat different: body practi- lighter disc; lateral tegminal field brown cally uniformly brown but with almost dark with yellowish humeral stripe; dorsal field brown hind half of head dorsum, pronotal in upper tegmen and distal part of this field disc, upper part of pronotal lateral lobes, in lower tegmen greyish brown (slightly and abdominal tergites, with greyish white lighter than lateral field); rest of latter field distal half of maxillary palpi, with lightish transparent; pterothoracic and abdomi- stripe along hind edge of metanotum and nal tergites brown with light brown areas several small marks on other parts of ptero- on metanotum and not very distinct spots thoracic and abdominal tergites, and with on abdominal tergites as well as yellowish dark brown dorsal surface of hind femora hairs on metanotal gland; abdominal ster- (having a few small lightish marks); scape nites, genital plate and hind tibiae slightly approximately 1.3 times as wide as head ros- darkened; and base of cerci yellowish. Scape trum between antennal cavities; and copu- approximately 1.4 times as wide as head ros- latory papilla very small, rather narrow, and trum between antennal cavities; metanotal practically not bilobate (Fig. XIII: 7). gland as in Figs XII: 7. Tegmina similar Variations. Second female with slightly to those of A. vera but somewhat smaller, lighter both posterior part of pronotal lat- with comparatively larger mirror, shorter eral lobes (greyish brown) and hind femur chords, and distinctly longer and narrow- (reddish brown). er apical area of dorsal field (Figs XII: 6). Male unknown. Ninth abdominal tergite with clearly wider Length in mm. Body 8.5–9.5; prono- and truncate posteromedian lobe; genital tum 1.8–1.9; hind femora 6–6.5; ovipositor plate elongate, almost thrice longer than 5–5.2. anal plate, with almost rounded apex hav- Comparison. The new species differs ing small median part semimembranous and from A. vera and A. propria in a distinctly slightly folded in rest position; genitalia as smaller female copulatory papilla as well as in Figs XIII: 9–11. darker body colouration. Female. Body colouration similar to Etymology. The species name is the Latin that of male, but metanotum with darker word “umbrosa” (dark, shady). (almost dark brown) anterior part and lighter (almost yellowish) stripe along pos- Anemozara (Zacmozara) eximia sp. nov. terior edge, tergites of abdomen brown with (Figs XII: 6, 7; XIII: 9–12) slightly darker lateral parts and narrow median stripe, hind femora with slightly dark- Holotype. Male; Malaysia, Borneo I., Sar- ened dorsal surface, and genital plate light awak State, environs of Miri Town, Lambir brown. Structure of body similar to that of

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 47 female of A. umbrosa, but copulatory pa- like apical part and slightly curved subapi- pilla larger, almost round but with slightly cally (in P. circumcincta, these epiphallic truncate one edge and concave opposite one arms are somewhat widened distally, pro- (Fig. XIII: 12). vided with a rather wide and rounded sub- Length in mm. Body: male 9.5, female apical lobe, and having a medial apical spine 9; pronotum: male 1.8, female 1.9; tegmina, separated from the latter lobe by a small but male 4; hind femora: male 6.2, female 6.4; distinct notch; see Figs XIV: 13, 14). ovipositor 5.1. Etymology. The species name is the Latin Lerneca sylvestris sp. nov. word “eximia” (peculiar, particular). (Figs XIV: 1–6)

Subtribe LERNECINA subtrib. nov. Holotype. Male; Bolivia, northern part of Santa Cruz Department (near Brazil), Noel = Lernecae Desutter, 1987 (unavailable name Kempff Mercado National Park, Los Fierros published without description or diagnosis) Camp, about 300 m, primary forest, among dry leaves on forest floor at daytime, 23–28 Jan. Type genus Lerneca Walker, 1869 (gen- 2014, A. Gorochov (ZIN). der feminine). Paratypes. Male and 3 females, same data as Diagnosis. This subtribe is similar to the for holotype (ZIN). subtribe Luzarina in the structure of palpi, Description. Male (holotype). General hind tibial spurs and male genitalia (hav- appearance typical of this genus. Coloura- ing small rachis) but distinguished from it tion (Fig. XIV: 1, 2) dark brown (almost by the presence of additional sclerites con- blackish) but with proximal part of antennal necting semisclerotized ectoparameres with flagellum light brown and having two small bases of the posterolateral epiphallic arms whitish spots, other parts of this flagel- (Figs XIV: 3–5, 10, 12). lum having longer whitish spots, pronotal Included species. Lernecina includes two disc and tegmina greyish brown (tegminal or three genera listed above (see a prelimi- membranes with small barely lighter spots; nary classification of the Phalangopsinae tegminal veins slightly lighter, almost light subfamily group). greyish brown), dorsum of pterothorax and Comparison. Differences from Luzarina of first abdominal tergite almost yellowish, are given above. From all the other sub- venter of body from greyish brown to light tribes of Phalangopsini, it differs in the brown, and cerci greyish brown in distal same characters as Luzarina. half. Metanotal gland consisting of a pair of rather small inflations situated near each Prosthacusta beliza other (Fig. XIV: 2). Tegmina insignificant- (Otte et Perez-Gelabert, 2009), comb. nov. ly extending behind abdominal apex, with dorsal field as in Fig. XIV: 1. Hind wings = Doposia beliza Otte et Perez-Gelabert, 2009 strongly shortened (Fig. XIV: 2). Fore tibi- Note. Judging by the photographs pub- ae with outer and inner tympana open, oval. lished by Otte & Perez-Gelabert (2009: fig. Anal plate not long and somewhat narrow- 608), this almost undescribed species from ing to widely rounded apex; genital plate ap- Belize was erroneously included by its au- proximately twice longer and with notched thors in the related genus Lerneca (= Dopo- apex. Genitalia similar to those of L. ornata sia). In reality P. beliza is close-related and Desutter-Grandcolas, 1992 but with inner very similar to the Mexican species P. cir- (upper) posterolateral epiphallic arms thin- cumcincta (Scudder, 1869) (type species of ner and having denticles on rounded api- the genus Prosthacusta), but differs from the cal part only as well as with outer (lower) latter in the posterolateral epiphallic arms posterolateral epiphallic arms narrower in (lobes) gradually narrowing to the spine- proximal part and wider in distal part: these

Figs XIV (1–14). Lerneca and Prosthacusta, male: 1–6, L. sylvestris sp. nov.; 7, L. ornata Desutter- Grandcolas; 8–10, L. inalata amboro subsp. nov.; 11, 12, L. inalata pantanal subsp. nov.; 13, P. circumcincta (Scud.), Mexico; 14, P. beliza (Otte et Perez-Gelabert), comb. nov., after Otte & Perez- Gelabert (2009). Body (with upper tegmen) from above (1, 8, 11); metanotal gland from above (2, 9); genitalia from above (3), from below (4) and from side (5, 10, 12); lower posterolateral arm of epiphallus from side (6, 7); distal part of posterolateral epiphallic arm (lobe) from below (13, 14).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 49 parts almost equal to each other in width specimens of L. sylvestris (head and pro- (height), but distal part with clearly shorter notum are blackish with yellowish lines apical spine (see Figs XIV: 3–7). along dorsomedial edges of the eyes and Variations. Second male with dorsum of with light brown stripes along lateral edges head having weakly distinct lightish lines of the pronotal disc). Male from Guyana is along dorsomedial edges of eyes and with lighter, with the pronotal disc and tegmina light brown areas on pronotal disc along its uniformly light brown, and with the mir- lateral edges. ror of upper tegmen having one dividing Female. Colouration and structure of vein only (aberration?). Genitalia in all body similar to those of males, but tegmina the males studied are practically identical slightly not reaching abdominal apex and to those pictured by Desutter-Grandcolas with rather narrow dorsal and lateral fields (1992a) for L. ornata from French Guiana having five almost parallel longitudinal (type locality). veins in both fields (crossvenation well developed in dorsal field and strongly reduced Lerneca inalata amboro subsp. nov. in lateral one), colouration of tegmina dark (Figs XIV: 8–10) brown to black with brown venation of dorsal field and light brown humeral stripe, Holotype. Male; Bolivia, Santa Cruz Depart- metanotal gland absent, and anal plate ment, about 70 km NWW of Santa Cruz City, smaller and with somewhat narrower apex; Amboro National Park, Mataracu Camp, approximately 800 m, on grass near small house genital plate short (not longer than anal in primary forest, at daytime, 8–13 Feb. 2014, plate) and with widely truncate hind part. A. Gorochov (ZIN). Length in mm. Body: male 8.5–9.5, fe- Paratypes. Male and female, same data as for male 8–10; body with wings, male 10–11; holotype (ZIN). pronotum: male 1.5–1.7, female 1.6–1.8; Description. Male (holotype). General tegmina: male 7.5–8, female 5.5–6; hind appearance similar to that of L. sylvestris, femora: male 7–7.5, female 6.5–7; oviposi- but epicranium with light greyish brown tor 6–6.5. dorsum having darkish longitudinal stripes, Comparison. The new species is most antennae greyish brown (almost uniformly similar to L. ornata but distinguished from it dark) and with light brown proximal part, by a less contrast body colouration, clearly maxillary palpi with two distal segments less transverse mirror in the male tegmina, white, pronotum with light greyish brown and the above-mentioned features of male disc having small darker marks, tegmina genitalia (see description). slightly lighter and with venation prac- Etymology. The species name is the Latin tically of same colour as tegminal mem- word “sylvestris” (applying to forest). branes, tergites of both pterothorax and abdomen completely dark brown, anal plate Lerneca ornata Desutter-Grandcolas, 1992 moderately brown, cerci light brown, head (Fig. XIV: 7) smaller, and structure of tegminal dorsal field somewhat different (Fig. XIV: 8, 9). Material examined. Five males and 5 females; All these features as well as structure of Ecuador, eastern plain, 80–85 km E of Lago genitalia very similar to those of L. inalata Agrio Town, environs of Lago Grande Lake on (Saussure, 1874), but this new subspecies Rio Cuyabeno, very lowlying primary forest, among dry leaves on forest floor at night, 2–9 distinguished from all other subspecies of Nov. 2005, A. Gorochov, A. Ovtshinnikov (ZIN). latter species (L. i. inalata distributed from Male; Guyana, “British Guiana: New River, 750 Surinam to Panama; L. i. mexicana Goro- ft”, 4–7 March 1938, C. Hudson (NHM). chov, 2007 from Chiapas State in Mexico; Note. The specimens from Ecuador have Doposia tobago Otte et Perez-Gelabert, a contrast colouration almost as in some 2009 originally described as species from

Tobago I. but possibly belonging to L. in- Description. Male (holotype). Size, co- alata as separate subspecies or synonym of louration and structure of body very similar its nominotypical subspecies) by slightly to those of L. i. amboro, but tegmina (Fig. narrower or smaller tegmina with clearly XIV: 11) with somewhat longer mirror smaller mirror (Fig. XIV: 8) and distinctly (mirror of upper tegmen almost as wide as longer ventral spines in middle part of lower long, but in L. i. amboro, it is clearly trans- posterolateral arms of epiphallus as well as verse) and with apical area of dorsal field almost truncate (not round) apical part of distinctly shorter (mirror of upper tegmen these arms in profile (Fig. XIV: 10). approximately 3.5 times as long as this area, Variations. Tibiae and tarsi of paratype but in L. i. amboro, this ratio 2.1–2.4); geni- with more distinct lightish spots, and dark talia also very similar to those of this sub- marks on its pronotal disc more numerous. species, but semimembranous ectoparamer- Female. General appearance very simi- es somewhat longer and directed backwards lar to that of male paratype, but coloura- more than upwards (Fig. XIV: 12). tion of pronotal disc as in holotype; struc- Variations. Lower posterolateral epi- ture and colouration of tegmina, of ptero- phallic arms in male genitalia with some- thorax and of abdomen almost as in female what different length of distal part (from of L. sylvestris. apex to most distal denticle of these arms): Length in mm. Body: male 8–9.5, female from rather short (as in Fig. XIV: 12) to al- 10; body with wings, male 9–11; pronotum: most as in L. i. amboro (Fig. XIV: 10) male 1.5–1.7, female 1.9; tegmina: male 7–8, Female. General appearance very similar female 7; hind femora: male 7–8, female 8; to that of male; structure and colouration of ovipositor 6.8. tegmina, of pterothorax and of abdomen al- Comparison. Differences from the other most as in female of L. sylvestris. subspecies (or possible subspecies) of this Length in mm. Body: male 8–10, female species are given above. Lerneca digrediens 9–11; body with wings, male 9.5–11.5; pro- (Otte, 2006), originally described from Cos- notum: male 1.6–1.8, female 1.8–2.2; tegmi- ta Rica as a member of the genus Amphiacus- na: male 7.5–8.5, female 7–8; hind femora: ta, and L. occidentalis Gorochov, 2007 (Oax- male 7.5–8.5, female 8–9; ovipositor 6.5–7. aca State in Mexico) are close-related to L. Comparison. Differences of the new sub- inalata and differ from the new subspecies in species from L. i. amboro are given above. a distinctly less concave middle part of ven- From all the other subspecies of this species, tral edge of lower posterolateral epiphallic the new subspecies differs in clearly longer arms (having also much shorter spines) or ventral spines in middle part of the lower clearly greater length of these arms (espe- posterolateral epiphallic arms; and from cially their distal part), respectively. close-related L. digrediens and L. occiden- Etymology. The species is named after talis, in the same characters as L. i. amboro the Amboro National Park. (see its comparison above). Etymology. The species is named after Lerneca inalata pantanal subsp. nov. “Reserva Pantanal Raraguayo”. (Figs XIV: 11, 12) Subtribe PARENDACUSTINA Holotype. Male; Paraguay, “Reserva Panta- nal Raraguayo” near Bolivia, Los Tres Gigantes subtrib. nov. Biological Station on Rio Negro (basin of Parana Type genus Parendacustes Chopard, 1924 River), open landscape (without forest) with bushes and high grass, on road at night, 31 Jan. – (gender masculine). 4 Feb. 2014, A. Gorochov (ZIN). Diagnosis. General appearance as in Pha- Paratypes. Six males and 4 females, same data langopsina, Amphiacustina and Uvaroviella as for holotype (ZIN). (? Heterogryllina), but head rostrum usu-

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 51 ally somewhat less projected and slightly tablished for a few “genera” by Desutter- less angular (but more projected and an- Grandcolas (1992b); but the latter taxa are gular than in other subtribes of Phalangop- considered as subgenera of Uvaroviella by sini). Hind tibiae with middle inner apical me (Gorochov, 2007). Thus, this group may spur longest (as in Phalangopsina and Am- be a synonym of Heterogryllina. It is char- phiacustina) or with middle and dorsal in- acterized by the shape of head and general ner apical spurs longest but almost equal appearance more or less similar to those of to each other in length (as in Uvaroviella). Phalangopsina, by a rather long and mod- Male genitalia: epiphallus with very short erately thin apical segment of the maxillary (almost transversally ribbon-like) proximal palpi slightly widened to the apex and hav- part divided into two lateral halves by small ing a rather short oblique apical truncation membranous median area (sometimes these (Fig. I: 3), by two (dorsal and median) inner halves almost contacting with each other), apical spurs of the hind tibiae longest and with large posterolateral arms (almost as almost equal to each other in the length, by in Luzara; Fig. II: 6), and with posterome- the inner ventral apical spur of these tibiae dian part shorter and membranous but usu- strongly reduced or absent, and by the fol- ally with a pair of isolated sclerites (these lowing characters of male genitalia: epiphal- sclerites sometimes more numerous or fused lus is almost not curved in the profile and with each other, very small or rather large, with a not very large posteromedian notch; semimembranous and almost indistinct or ectoparameres are not moved to the apical heavily sclerotized); rachis membranous, or subapical epiphallic parts; and rachis is more or less short; ectoparameres developed more or less small and not protruding or but sometimes not very distinctly separated almost not protruding behind the median from epiphallus [see Gorochov, 1996 (figs epiphallic part (Figs II: 13–16). 113–128), 2003a (figs 2 and 3), 2003b (fig. 3), 2006 (figs 1–3)]. ?Uvaroviella problematica sp. nov. Note. This subtribe is distinguished from (Figs XV: 8, 9) Phalangopsina, Amphiacustina and Uva- Holotype. Female; Ecuador, Morona Santia- roviella by a Luzara-like epiphallus but with go Prov., bank of Rio Morona near border with the membranous posteromedian epiphallic Peru, environs of Puerto Morona Vill., about part having one or a few sclerites. From the 300 m, primary forest, 5–15 Jan. 2010, A. Goro- other subtribes of Phalangopsini, Parenda- chov (ZIN). custina differs in not semiglobular head and Description. Female (holotype). Body in the presence of posteromedian membra- medium-sized and spotted. Head light grey- nous epiphallic part with isolated sclerites. ish with a pair of brown areas between an- Generic composition of this subtribe are tennal cavities under median ocellus (these given above, in the classification of Phalan- areas separated from each other by narrow gopsinae subfamily group. light triangle but almost contacting with each other ventrally), small median brown ? Subtribe HETEROGRYLLINA spot on epicranium near clypeus, a pair of similar spots on genae behind eyes, brown Note. This subtribe possibly includes two most part of mandibles (excepting proximal genera. The genus Heterogryllus is insuffi- parts), and greyish brown (slightly lighter) ciently studied but may be close-related to antennal flagellum and marks on scape; pro- Uvaroviella. Judging by Saussure (1874), it notum also light greyish brown but with is more or less similar to Uvaroviella species brown upper halves of lateral lobes and lacking wings in female but distinguished most part of disc (excepting light stripes by somewhat longer legs. The genus Uva- along all edges of disc); tegmina greyish roviella is equal to the group Aclodae es- brown with lighter humeral part; legs light

Figs XV (1–9). Caribacusta , Noctivox and ?Uvaroviella: 1, C. antigua sp. nov.; 2–7, N. orizaba sp. nov. ; 8, 9 , ?U. problematica sp. nov. Male anal plate from above (1, 2); male metanotal gland from above (3); dorsal field of male upper tegmen (4); male genitalia from above (5), from below (6) and from side (7); female copulatory papilla from above (8) and from side (9). greyish brown with dark spots on fore and strongly oblique ventral edge (their an- middle legs as well as on hind tibiae and teroventral corner protruding downwards, tarsi, and with dark two transverse bands roundly angular; their posterior part low) on distal part of hind femora and numer- and slightly oblique anterior edge. Tegmina ous oblique stripes on rest of these femora; reaching posterior third of first abdominal other thoracic parts light greyish brown tergite, with oblique and slightly rounded with slightly darker sternites and most pat posterior edge of dorsal field (lateral part of of metanotum; abdomen greyish brown this field distinctly longer than medial one) with slightly darker and slightly lighter having 5–6 short and more or less oblique small marks on tergites as well as with longitudinal veins as well as numerous ir- moderately light greyish brown cerci and regular crossveins between them, and with ovipositor. Scape almost 1.5 times as wide 3–4 longitudinal veins and not numerous as head rostrum between antennal cavities; (and less distinct) crossveins in lateral field. maxillary palpi similar to those pictured Legs with rather small and moderately nar- in Fig. I: 3. Pronotum with anterior part row inner tympana only as well as with two slightly narrowing to head; disc almost 1.25 inner apical spurs of hind tibiae practically times as wide as long; pronotal lobes with equal to each other in length (ventral inner

Figs XVI (1–10). Laozacla furca sp. nov.: 1, body of male from above; 2, 3, male abdominal apex from above (2) and from below (3); 4, male metanotal gland from above; 5–7, male genitalia from above (5), from below (6) and from side (7); 8, female abdominal apex from below; 9, distal part of ovipositor from side; 10, scheme of rachis with formula (formula looking as median sclerite located on ventral surface of rachis) from below. apical spur of hind tibiae almost lost). Geni- with much larger opposite notch (distinct tal plate slightly transverse, narrowing to but short hook-like process located between widely truncate apex; ovipositor with nar- these notches; Fig. XV: 8, 9). row apical part typical of genus Uvaroviella Male unknown. in structure; copulatory papilla large, al- Length in mm. Body 10.5; pronotum most round but dorsoventrally depressed, 2.5; tegmina 2; hind femora 11.5; ovipositor with small but distinct median notch, and 10.5.

Comparison. The new species is distin- cepting short basal part); hind femora some- guished from all the other congeners by what thickened; hind tibiae with apical spurs more oblique ventral edges of the pronotal not inflate and with middle inner apical spur lateral lobes and a very characteristic shape longest but distinctly not reaching middle of the female copulatory papilla (this papil- of hind basitarsus. Male metanotum with la has a short hook-like process distinct in weakly developed gland (Fig. XVIII: 3); the profile). I cannot exclude that this spe- anal plate simple, almost triangular and with cies may belong to a close-related genus of rounded apex; male genital plate somewhat the same subtribe. From the congeners with longer and narrowing to narrowly rounded an unknown copulatory papilla, the new apex; female genital plate slightly smaller species differs also in the female tegmina and with distinctly angular and rather wide short (approximately 2 mm in the length) posteromedian notch. Male genitalia similar but slightly covering each other, and hind to those of Benoistella, but epiphallus fused femur approximately 1.1 times as long as with rami and having posterolateral lobes ovipositor. separated from main epiphallic body by rather deep transverse fold (distinctly visible in profile), and rachis with clearly wider Tribe PARAGRYLLINI apical part (Figs XVIII: 4–6). Ovipositor Subtribe PARAGRYLLINA with ventral part of subapical lobule of dor- Genus Bolivacla gen. nov. sal valve truncate (as in Paragryllus; in Ben- oistella, this part rounded) and with ventral Type species B. boliviana sp. nov. edge of apical part of ventral valve lacking Diagnosis. General appearance similar to denticles (as in Benoistella; in Paragryllus, that of genus Paragryllus. Head depressed this edge denticulate) (Fig. XVIII: 7). in front (face with rather wide but not deep Included species. Type species only. concavity between eyes), with narrow and Comparison. The new genus differs from projected rostrum having small median ocel- Paragryllus and Rumea in the male geni- lus on dorsal surface near apex and similar talia more similar to those of Benoistella (in size) lateral ocelli situated at base of this (epiphallus is less curved in profile, with a surface very near each other, and with rather clearly longer median part, or rachis is dis- short maxillary palpi (Fig. XVIII: 2). Pro- tinctly shorter), and from Benoistella and notum clearly narrowing to head, weakly Silvastella, in the male tegmina more similar transverse (its width slightly greater than its to those of Paragryllus (mirror is larger and length), with rather low pronotal lobes hav- with much more numerous dividing veins). ing rounded and not projected anteroventral Etymology. The generic name originates corners. Tegmina long, distinctly protruding from the Bolivia Country and genus Acla. behind abdominal apex; male tegmina (Fig. XVIII: 1) with wide dorsal field having Bolivacla boliviana sp. nov. large stridulatory apparatus (this apparatus (Figs XVIII: 1–7) characterized by long and somewhat arcuate stridulatory vein, numerous and partly Holotype. Male; Bolivia, northern part of transverse oblique veins, and large oval mir- Santa Cruz Department (not far from Brazil), ror with numerous and strongly curved di- environs of Florida Vill. on Rio Paragua (Ama- zon Basin), almost 300 m, secondary forest, on viding veins), with rather long apical area, trunk of large tree rather high above ground at and with distinctly widened middle part of night, 22–29 Jan. 2014, A. Gorochov (ZIN). R–M area; hind wings slightly protruding Paratypes. Male and female, same data as for behind tegmina. Fore tibiae with small oval holotype (ZIN). tympana on both sides; middle tibiae barely Description. Male (holotype). Body inflate in middle and proximal thirds (ex- rather large. Colouration dark brown with

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 55 light greyish spots on lateral sides of la- mina: male 18–19, female 19.5; hind femora: brum, brown antennae and maxillary palpi, male 12–14, female 13; ovipositor 11.5. brown hind half of pronotum and dorsal Etymology. The species is named after part of pterothorax as well as areas on tibiae the Bolivia Country. and tarsi of middle and hind legs, light grey large areas on dorsal field of upper tegmen Subtribe MEXIACLINA subtrib. nov. (Fig. XVIII: 1) and most part of lateral field in both tegmina (but Sc–R area dark Type genus Mexiacla Gorochov, 2007 brown, and most part of R–M area greyish (gender feminine) brown), greyish brown apical area of both Diagnosis. General appearance (includ- tegmina and some marks on other parts of ing structure of both head and pronotum dorsal field in upper tegmen, almost trans- as well as length of legs) similar to that in parent membranes of stridulatory apparatus Phalangopsina, Amphiacustina and Neoac- in lower tegmen, greyish hind wings, and lina (probable convergence caused by simi- brown coxae and venter of body (but geni- lar adaptation to life on bark of living trees; tal plate almost dark brown). Scape almost Figs XVII: 1, 5). Maxillary palpi similar to twice wider than head rostrum between those of Uvaroviella (see Fig. I: 3). Tegmina antennal cavities; metanotal gland consist- strongly shortened in male and absent in ing of anterior fold widened laterally and female; hind wings absent in both sexes. partly covered with short hairs, and darker Hind tibiae with inner middle apical spur (blackish) posterior convexity (Fig. XVIII: longest. Male abdomen (Figs XVII: 1, 5) 3); tegmina with dorsal field having 6–7 di- with specialized tubercles on dorsum (ab- viding veins in mirror (Fig. XVIII: 1) and dominal gland); male genitalia (Figs XVII: with lateral field having 19–20 branches of 2, 3, 6) with rather small epiphallus having Sc and without crossveins between these transverse anterior part and strongly or branches. Genitalia as in Figs XVIII: 4–6. very strongly bifurcate posteromedian part Variations. Second male with complete- (sometimes epiphallus divided into a few ly dark brown labrum and reddish brown isolated sclerites; Fig. 6), with large (lon- antennal flagellum. ger than epiphallus or not shorter than it) Female. General appearance (includ- and partly membranous or semisclerotized ing colouration of labrum) similar to that ectoparameres originating from posterolat- of holotype, but antennal flagellum as in eral epiphallic lobes (arms) and separated second male, tegmina dark greyish brown from epiphallus near proximal epiphallic with light brown marks on proximal half part, and endoparameres articulated mainly of dorsal field and with greyish brown to with ectoparameres. Ovipositor with rath- light brown ventroproximal part of lateral er primitive structure of distal part (Fig. field, dorsal tegminal field narrower than in XVII: 4). male and with 9–10 not very regular longi- Note. This subtribe is most similar to tudinal veins and comparatively numerous Neoaclina, but it differs from the latter in a crossveins, and lateral tegminal field with strongly bifurcate posteromedian epiphallic distinctly narrower R–M area and wider part, larger and more distinctly articulated Sc–R area as well as with 10–12 branches posterolateral epiphallic lobes separated of Sc and not very numerous crossveins be- from the epiphallus near its proximal part tween these branches; genital plate almost (but not near its distal or middle parts), and dark brown, and ovipositor with distal part the endoparameres connected mainly with as in Fig. XVIII: 7. the ectoparameres (but not with rachis). Length in mm. Body: male 17–23, fe- From Paragryllina, the new subtribe differs male 18; body with wings: male 24–27, fe- in a more primitive structure of the oviposi- male 26; pronotum: male 4, female 4.2; teg- tor: its apical part is non-denticulate, and

Figs XVII (1–6). Mexiacla and Oaxacla, after Gorochov (2007): 1–4, M. ecosuri Gor.; 5, 6, O. squa- miptera Gor. Male body from above (1, 5); male genitalia from above (2, 6) and from below (3); distal part of ovipositor from side (4). Abbreviations: en.a, endoparameral apodemes; ep.a, ectoparameres (almost articulated posterolateral epiphallic arms); ep.b, anterior (basal) epiphallic parts; ep.l, partly or very strongly bifurcate posteromedian lobe of epiphallus; m, formula (mold of spermatophore attachment plate); s.p, semisclerotized part of ectoparameres (thickened areas of membranes around formula fused with ectoparameres). ventral edge of the subapical part of dor- Diagnosis. General appearance similar sal valves is rounded (but not angular) in to that of Phalangopsina, Amphiacustina, the profile; and from Strogulomorphina, in Neoaclina and Mexiaclina. Male tegmina a much larger body as well as in the male from distinctly shortened to scale-like; fe- genitalia with the median part of epiphal- male apterous. Male genitalia consisting lus clearly shorter than its posterolateral of more or less short epiphallus having di- lobes (ectoparameres) and with the formula verse but narrower posteromedian process, distinctly smaller. Composition of this sub- a pair of distinctly sclerotized ectoparameres clearly separated from epiphallus and tribe is given above, in the classification of strongly projected behind its posterolateral Phalangopsinae subfamily group. parts, and endoparameres articulated with ectoparameres. Subtribe BREVIZACLINA subtrib. nov. Note. Included genera are listed in the classification of Phalangopsinae subfam- Type genus Brevizacla Gorochov, 2003 ily group given above. From the similar (gender feminine) subtribes Neoaclina and Mexiaclina, this

Figs XVIII (1–7). Bolivacla boliviana sp. nov.: 1, dorsal field of male upper tegmen with distal parts of hind wings and parts of cerci from above; 2, distal half of maxillary palpus from side; 3, male metanotal gland from above; 4–6, male genitalia from side (4), from above (5) and from below (6); 7, distal part of ovipositor from side. subtribe differs in the following characters: sclerotized structures); from Mexiaclina, in from Neoaclina, in the male genitalia with- a not or almost not bifurcate posteromedian out membranous, semimembranous or scler- epiphallic part and the presence of distinctly otized lobules around the posteromedian sclerotized and clearly articulated ecto- epiphallic process and with clearly developed parameres (instead less sclerotized and less sclerotized ectoparameres (in Neoaclina, articulated posterolateral epiphallic lobes). these lobules are often almost not separat- In Brevizaclina, formation of ectoparameres ed from this process, or sometimes they are is probably finished, but in Neoaclina and more or less turned into ectoparamere-like Mexiaclina, convergent formation of ecto-

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 58 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP parameres is in initial stage. From the other [Included species: M. papuana Gorochov, subtribes of Paragryllini, Brevizaclina dif- 1986 (type species); M. buergersi Gorochov, fers in a characteristic general appearance, 1996; M. enarotali sp. nov.] the presence of strongly shortened tegmina – Tympana developed. Both tegmina in male with more or less normally developed stridu- in male, a short epiphallus lacking deep pos- latory apparatus (see Otte, 2007: figs 65–67) teromedian notch, and usually rather small ...... subgen. Stridulacla subgen. nov. formula in the male genitalia. [Included species: M. gorochovi Otte, 2007 (type species); M. biaru Otte, 2007; M. chu- Genus Mikluchomaklaia Gorochov, 1986 aye Otte, 2007; possibly M. longicerca Goro- chov, 1986 and M. korido Gorochov, 2006. Etymology: the subgeneric name originates Note. This genus is characterized by the from the Latin word “stridulus” (stridula- male tegmina strongly shortened but not tory) and genus Acla.] scale-like, dorsal field membranous (with strongly reduced venation) in the lower Mikluchomaklaia (Mikluchomaklaia) tegmen and sclerotized (changed or partly enarotali sp. nov. changed into an attractive gland) in the up- (Figs XIX: 1–4, 18) per tegmen or with a more or less normally developed stridulatory apparatus, and male Holotype. Male; Indonesia, New Guinea I., genitalia with the endoparameres lacking Papua Prov., about 120 km SEE of Nabire Town, any ribbon or plate-like structure connected Central Range, environs of Enarotali [Enarato- the left endoparamere with right one or hav- li] Vill. on Paniai Lake, approximately 2000 m, ing such ribbon interrupted in the median mountain primary forest, on tree trunk near soil part (these endoparameres are also with at night, 22–27 Feb. 2012, A. Gorochov (ZIN). long apodemes directed forwards; Figs XIX: Paratype. Male and 3 females, same data as for holotype, but male collected on rock in forest 2–4; Gorochov, 1996: figs 100–105; Goro- at night (ZIN). chov, 2003: figs 1, 1–3, 11–13). The genus Description. Male (holotype). Body com- is known from New Guinea and nearest is- paratively small. Colouration dark brown lands and divided into three subgenera di- with lightish spots on mouthparts, very agnosed below, in a key for subgenera of the sparse and small lightish marks on anten- genus Mikluchomaklaia. nal flagellum, a few light brown longitu- 1. Male head with deep concavity having hairs dinal lines on dorsum of head, brown both (attractive gland) on dorsum between eyes. keel around tegminal dorsal field and ante- Tympana very small. Male upper tegmen rior widening of this keel, light brown spots sclerotized, with rather large concavity in on legs, weakly distinct lightish marks on dorsal field lacking hairs, and without vena- pterothoracic dorsum and on abdominal tion in lateral field (see Gorochov, 2003b: figs 1, 4, 5, 7) ...... tergites, and light greyish brown venter of ...... subgen. Phantazacla Gorochov, 2003 body (excepting dark genital plate). Exter- [Included species: M. phantastica Gorochov, nal structure of body typical of this subge- 1996 (type species); M. mystica Gorochov, nus: scape approximately 1.5 times as wide 2003.] as head rostrum between antennal cavities; – Male head without any concavity on dorsum. pronotum slightly transverse, not wider Tympana developed or absent. Male upper than head, with almost parallel lateral sides, tegmen diverse: sclerotized but with hairs, and with anteroventral corners roundly or not sclerotized and with more or less nor- projected downwards; upper tegmen with mally developed stridulatory apparatus . . . . 2 2. Tympana absent. Dorsal tegminal field of up- almost round dorsal field reaching middle per tegmen in male sclerotized, with group of part of second abdominal tergite, with nar- distinct hairs at centre (Fig. XIX: 1) ...... row and low keel around this field having al- ...... subgen. Mikluchomaklaia s. str. most angular widening in anterior part, with

Figs XIX (1–18). Mikluchomaklaia and Brevizacla, male: 1–4, M. enarotali sp. nov.; 5–9, B. fawi sp. nov.; 10–13, B. nabire sp. nov.; 14–17, B. halmahera sp. nov.; 18, M. enarotali. Anterior part of body from above (1, 10) and same but with right tegmen erected (5, 14); genitalia from above (2, 7, 12, 16), from below (3, 8, 13, 17) and from side (4, 9, 11, 15); 6, right tegmen; 18, ectoparamere from below. not large convexity at centre of very large widely and roundly truncate apex; genital but not deep concavity of this field, and plate distinctly larger, slightly elongate, and with group of distinct light brown hairs on with widely rounded apex having weakly these widening and convexity (Fig. XIX: 1); distinct shallow median notch. Genitalia middle and dorsal inner apical spurs of hind with ectoparameres having two long and tibiae almost equal to each other in length, narrow posterior processes; both medial ec- clearly longer than other apical spurs, and toparameral processes covered by postero- slightly not reaching middle of hind basitar- median epiphallic process (in dorsal view; sus; anal plate short, distinctly narrowed to Figs XIX: 2–4, 18).

Variations. Second male with coloura- 2003b: fig. 2, 4). Male genitalia with very tion of head lighter (brown to light brown) long ectoparameres, with wide plate con- and having less distinct marks on dorsum nected left and right endoparameres, and and mouthparts. with short endoparameral apodemes (see Gorochov, 2003b: figs 2, 1–3) ...... Female. General appearance similar to ...... subgen. Brevizacla s. str. that of males of this species, but body co- [Included species: Mikluchomaklaia curta louration as in second male, wings absent, Gorochov, 2003 (type species) only.] and anal plate only barely smaller (shape – Tympana absent or very small. Male tegmina of this plate almost indistinguished from lobule-like (i. e. with dorsal fields transversal- that of male); genital plate almost equal ly elongate, directed more or less medially or to anal plate in length, slightly transverse, somewhat overlapping each other; Figs XIX: somewhat narrowing to widely and roundly 5, 6, 10, 14). Male abdomen without attrac- truncate apex having barely concave medi- tive gland. Male genitalia with short or moderately long ectoparameres as well as with an edge; ovipositor with distal part typical diverse endoparameres and their apodemes of this genus. (Figs XIX: 7–9, 11–13, 15–17) ...... 2 Length in mm. Body: male 11–12.5, 2. Fore tibiae with very small inner tympanum female 10–11.5; pronotum: male 1.8–2, fe- only. Male tegmina partly overlapping each male 2–2.2; upper tegmen, male 2.6–2.8; other; lower tegmen somewhat smaller than hind femora: male 9.5–10.8, female 10–11; upper one and slightly different in shape ovipositor 7.3–7.7. (Figs XIX: 5, 6). Male genitalia with distinct Comparison. The new species differs sacculus (spermatophore sac) between formula and rachis (Figs XIX: 7–9) ...... from M. papuana and M. buergersi (see ...... subgen. Papuzacla subgen. nov. Gorochov, 1996: figs 100–105) in the male [Included species: B. fawi sp. nov. (type spe- genitalia having long and strongly bifurcate cies) only. Etymology: the subgeneric name ectoparameres. originates from the Papua Province and ge- Etymology. The species is named after nus Zacla.] the Enarotali Village. – Tympana absent. Male tegmina not overlapping each other (i. e. contacting or not contacting with each other), equal in shape and Genus Brevizacla Gorochov, 2003 size (Figs XIX: 10, 14). Male genitalia more Note. This genus contains several rather primitive, without distinct sacculus between formula and rachis (Figs XIX: 11–13, 15– diverse species characterized by scale-like or 17) ...... subgen. Lobulacla subgen. nov. almost scale-like tegmina lacking stridulato- [Included species: B. nabire sp. nov. (type spe- ry apparatus in male; probably ventral mem- cies); Opilionacris chandani Bhowmik, 1982; branes of these tegmina are glandular, but Arachnomimus ranjani Bhowmik, 1982; M. sometimes tegmina are with a small attrac- discoptila Gorochov, 1996; B. speranda Goro- tive gland (looking as a concavity with short chov, 2006; possibly B. halmahera sp. nov. hairs) on the dorsal surface of each tegmen Etymology: the subgeneric name originates (see Gorochov, 2003b: figs 2, 13, 17). Male from the Latin word “lobulus” (lobule) and genitalia in Brevizacla are rather diverse, genus Acla.] and this genus must be divided into a few subgenera or related genera (see a key for Brevizacla (Papuzacla) fawi sp. nov. subgenera of the genus Brevizacla below). (Figs XIX: 5–9) 1. Tympana absent. Male tegmina scale-like Holotype. Male; Indonesia, New Guinea I., (i.e. rounded, with dorsal fields not transver- Papua Prov., Fawi [Faowi] Vill. in upper part of sally elongate and not contacting with each Tariku River (tributary of Mamberamo River), other; see Gorochov, 2003b: fig. 2, 4). Male primary forest on low hill, on tree trunk not abdomen with attractive gland on third and far from soil at night, 29 Jan. – 17 Feb. 2012, fourth abdominal tergites (see Gorochov, A. Gorochov (ZIN).

Description. Male (holotype). Body with a pair of long and very narrow ribbons rather large. Colouration variegate (Fig. running from apex of rachis to anterior part XIX: 5): head yellow with light greyish of sacculus, and with very large formula brown dorsum, three dark brown vertical situated around lateral and anterior sides stripes on face (from brown rostrum to la- of ventral part of sacculus (Figs XIX: 7–9). brum and from eyes to mandibles), a pair Female unknown. of dark brown spots near dorsal edges of Length in mm. Body 17.5; pronotum 3.5; antennal cavities, two small dark marks on upper tegmen 1.9; lower tegmen 1.6; hind membrane of each this cavity, brown areas femora 17.3. and small marks on genae, light brown palpi Etymology. The species is named after and antennae having darkened distal part of the Fawi Village. apical segment of maxillary palpi and darkish spot on scape; pronotum dark brown Brevizacla (Lobulacla) nabire sp. nov. with greyish brown disc, two yellowish (Figs XIX: 10–13) spots on anterior half of pronotal lobes, and lightish mark on each anterolateral corner Holotype. Male; Indonesia, New Guinea I., Papua Prov., environs of Nabire Town, second- of these lobes; tegmina almost uniformly ary forest on hill not far from sea, on tree trunk light brown; legs, pterothorax and abdomen near soil at night, 28 Feb. – 2 March 2012, A. dark brown with light brown venter (in- Gorochov (ZIN). cluding genital plate), brown cerci having Description. Male (holotype). Body lighter basal part, and numerous yellowish rather small. Colouration variegate but to light brown marks on rest parts. Scape al- somewhat darker than in B. fawi (Fig. XIX: most twice as wide as head rostrum between 10): head yellowish with three vertical dark antennal cavities; ocelli small and situated brown bands on face (almost as in B. fawi), near apex and at base of rostral dorsum. with a few dark brown spots on dorsum and Pronotum approximately as wide as head, behind eyes, with brown areas on maxil- narrowing to head and to pterothorax (Fig. lary palpi, and with dark brown antennae XIX: 5), with low hind part of lateral lobes, having yellowish scape marked by brown and with roundly angular anteroventral medial spot; pronotum dark brown with corners directed downwards. Tegmina (Figs a few yellowish and light brown marks on XIX: 5, 6) reaching middle third of meta- disc; other bodyparts more or less similar notum; upper tegmen semisclerotized, al- in colouration to those of B. fawi but with most oval, with very transversally elongate abdominal tergites darker and genital plate dorsal field lacking dorsal gland and vena- slightly darkened. Scape approximately tion; lower tegmen similar to upper one but 1.5 times as wide as head rostrum between slightly smaller, with less rounded medial antennal cavities; ocelli approximately as part and almost membranous basal area of in B. fawi. Pronotum also similar to that dorsal field. Hind tibiae with middle inner of this species but with somewhat less dis- apical spur longest but slightly not reaching tinct anterior and posterior narrowings middle of hind basitarsus. Anal and genital (Fig. XIX: 10). Tegmina reaching middle of plates more or less similar to those of M. en- metanotum, weakly transversally elongate arotali; genitalia with rather long epiphal- (almost oval), contacting with each other, lus and comparatively short ectoparameres, and without distinct gland on dorsal surface with a pair of narrow ventral additional (Fig. XIX: 10). Hind tibiae with middle and ribbons fused with base of rami (also fused dorsal inner apical spurs equal to each other with epiphallus), with long endoparameres in length, longer than all other apical spurs, connected with each other by narrow trans- and practically reaching middle of hind ba- verse ribbon as well as having rather short sitarsus. Anal and genital plate somewhat and strongly laterally curved apodemes, similar to those of B. fawi but with rounded

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 62 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP apices; genitalia more or less similar to those longer and more bifurcate ectoparameres as of M. ranjani (see Gorochov, 2003b: figs 2, well as with larger posteromedian process 10–12) but with distinctly larger (higher) of epiphallus and endoparameres not con- ectoparameres (Figs XIX: 11–13). nected with each other (Figs XIX: 15–17). Female unknown. Female unknown. Length in mm. Body 7.5; pronotum 1.7; Length in mm. Body 11; pronotum 2.5; tegmina 0.7; hind femora 8.2. tegmina 0.8; hind femora 14. Comparison. The new species dif- Comparison. The new species is distin- fers from B. discoptila and B. speranda in guished from all the other species of Lobu- larger male tegmina contacting with each lacla by the ectoparameres of male genitalia other and in the posteromedian process of having two posterior processes, by lateral epiphallus lacking ventrolateral teeth or one of these processes long and rather thin denticles; and from B. chandani and B. ran- (in the other species, ectoparameres shorter jani, in the absence of dorsal tegminal gland or wider), and by male tegmina without as well as much shorter (only from B. chan- distinct dorsal gland. It is necessary to note dani) or higher (only from B. ranjani) ecto- that male genitalia of the new species are parameres. most similar to those of M. papuana belong- Etymology. The species is named after ing to another genus (Mikluchomaklaia); it the Nabire Town. is a reason that inclusion of the new species in Lobulacla and Brevizacla is problematic, Brevizacla (Lobulacla) halmahera as this species may have convergent similar- sp. nov. ity to M. papuana in the genital structure or (Figs XIX: 14–17) to L. nabire in the tegminal shape. Etymology. The species is named after Holotype. Male; Indonesia, Maluku Utara the Halmahera Island. Prov., Halmahera I., environs of Subaim Vill. (not far from Lolobata Vill.) near Wasile Bay, disturbed primary forest, on tree trunk not far Tribe LUZAROPSINI from ground at night, 27 Jan. – 1 Feb. 2011, A. Gorochov (ZIN). Note. This tribe is characterized by an Description. Male (holotype). Body me- almost semiglobular head, the development dium-sized. Colouration (Fig. XIX: 14) and of a moderately large sacculus (= spermato- structure of bodyparts more or less similar phore sac) in the male genitalia, and absence to those of B. nabire, however dorsum of of characteristic folds before the formula of head with two narrow dark brown trans- these genitalia (the latter character distin- verse stripes between eyes above antennal guishes this tribe from the Otteini which cavities and greyish brown area behind has a similar sacculus in one of its genera; these stripes, labrum almost completely this partial similarity shows that such sac- dark brown, palpi yellowish, scape and prox- culus is convergently developed in these imal part of antennal flagellum light brown, tribes). Generic composition of Luzaropsini tegmina yellow with light brown proximal is given above, in the classification of Pha- area, cerci and genital plate almost com- langopsinae subfamily group. pletely light greyish brown, tegmina reaching middle third of metanotum and not con- Luzaropsis ferruginea (Walker, 1869) tacting with each other, apical spurs of hind tibiae more similar to those of B. fawi, anal Note. Gorochov (2003) supposed that plate with clearly truncate hind part, geni- L. confusa Chopard, 1969 from Sri Lanka tal plate with narrowly rounded apex, and may be a synonym of L. ferruginea (also genitalia similar to those of M. papuana (see described from Sri Lanka); he led some Gorochov, 1996: figs 100–102) but with arguments showing that the male genita-

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 63 lia, pictured by Chopard (1969: fig. 157) shorter than hind basitarsus). Anal plate in as belonging to L. confusa, may belong to both sexes almost equal in size; its hind part another species (i.e. genitalia of Chopard’s widely truncate, concave or notched in male specimens or pictures of these genitalia may (Figs XX: 3, 6–10), as well as slightly nar- be confused). Desutter-Grandcolas (2014) rower and rounded in female; male genital did not agree with this synonymy; however, plate rather large, not long (its width and she did not publish any explanation for her length almost equal), and with widely trun- opinion or comments for Gorochov’s argu- cate, slightly concave or notched hind part. ments. Thus, it is reasonable to consider Male genitalia: epiphallus fused with rami, these species names as probable synonyms having transverse fold (distinct in profile), up to checking of the above-mentioned ar- and divided into two H-shaped sclerites guments. (proximal and distal ones) articulated with each other in two points; endoparameres Genus Terrozacla gen. nov. connected with ectoparameres but not with each other, lacking medial projections, and Type species T. jambi sp. nov. having rather long apodemes; rachis moder- Diagnosis. Body medium-sized. Head ately large and with large elongate sclerite almost semiglobular but rather high (its not connected with other sclerites; mem- height slightly greater than width) and brane between rachis and formula forming barely narrower than pronotum; apical seg- distinct but not large sacculus (Figs XXII: ment of maxillary palpi distinctly (but not 1–3, 6–11, 13–18). Ovipositor with acute strongly) widened to apex (similar to that movable distal structure of upper valves of Luzara but with less oblique apical trun- (Fig. XXII: 4). cation). Pronotum almost square, with an- Included species. Type species; T. harau terior part barely narrower than middle or sp. nov.; T. trusmadi sp. nov.; T. borneo sp. posterior parts (Figs XX: 1, 4), and with an- nov.; T. kubah sp. nov.; T. gading sp. nov.; teroventral corners widely rounded; meta- possibly Luzaropsis mjobergi Chopard, 1930 notum with clearly developed gland consist- and L. omissa Gorochov, 2003. ing of rather deep transverse concavity and Comparison. The new genus is most sim- median lobe located behind this concavity ilar to the genus Luzaropsis from Sri Lanka and directed forwards (Figs XX: 2, 5). Teg- but distinguished by the epiphallus divided mina strongly shortened but contacting into two sclerites (in majority of Luzaropsis with each other or barely overlapping, with- species, it is undeveloped; see illustrations out stridulatory apparatus or with traces of in Gorochov, 2003b: figs 3, 4–16), its proxi- it in male, and with partly reduced venation mal (anterior) sclerite H-shaped and with (Figs XX: 1, 4; XXI); hind wings looking posterolateral parts supporting a pair of as very small lateral lobules reaching hind large membranous lobes situated around the edge of metanotum (Figs XX: 2, 5). Legs not distal (posterior) epiphallic sclerite, and ra- long and moderately thin (but hind femora chis distinctly projected backwards behind clearly thickened, well adapted to jumping); the posteromedian epiphallic part. One of fore tibiae without tympana or with inner species of Luzaropsis (L. omissa) has simi- tympanum only; hind tibiae with denticles lar structure of the epiphallus (but with- (except for four pair of articulated spines) out deep posteromedian notch and with situated mainly on outer dorsal edge as well the rachis invisible from above) and may as with dorsal and middle inner apical spurs belong to this new genus or to a separate almost equal in length and longer than all group of Luzaropsis having a parallel evolu- other apical spurs or with dorsal inner api- tion of the male genitalia. From Larandop- cal spur slightly longer than middle inner sis (India) known from a few nymphs of the apical spur (these longest spurs distinctly same species only, the new genus differs in

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 64 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP a clearly shorter inner dorsal apical spur of tic: elongate, distally widened, with apical the hind tibiae (it is distinctly shorter than lobule curved upwards and forwards and hind basitarsus in the new genus and not with small apical hook directed downwards shorter than this basitarsus in Larandopsis; and laterally). Chopard, 1969). Variations. Sometimes short basal part Etymology. This generic name originates of antennae and most part of pronotal disc from the Latin word “terra” (ground) and light brown, as well as lighter spots on legs genus Zacla. somewhat more distinct. Female. General appearance as in males, Terrozacla jambi sp. nov. but metanotal gland absent, tegmina with (Figs XX: 1–3, XXI: 1, 2; XXII: 1–5) four longitudinal veins in dorsal field and 3–4 ones in lateral field, and anal plate Holotype. Male; Indonesia, Sumatra I., Jambi brown or greyish brown; genital plate and Prov., about 35 km N of Sungaipenuh Town, en- distal part of ovipositor as in Figs XXII: 4, 5. virons of Kerinci-Seblat National Park, Kerinci Length in mm. Body: male 10–12, fe- Mount, 1500–2000 m, primary forest, among dry leaves on forest floor, 18–22 Nov. 1999, A. Goro- male 11–14; pronotum: male 2–2.3, female chov (ZIN). 2.3–2.7; tegmina: male 2.7–2.9, female Paratypes. Five males and 8 females, same 2.8–3.2; hind femora: male 9–10, female data as for holotype (ZIN). 10–11.5; ovipositor 10.5–11.5. Description. Male (holotype). Coloura- Comparison. The new species differs tion of body (Figs XX: 1–3) greyish brown from L. mjobergi and L. omissa (which may with dark brown lower part of head (ex- be members of this genus) in the absence of cepting whitish labrum), areas on head tympana (from the both species), a smaller behind eyes, pronotal lateral lobes, and body size and not very transverse female tegminal lateral field, with a pair of yellow- genital plate (from L. mjobergi), as well as ish narrow stripes on dorsum of head (from shorter male tegmina, a deeply bifurcate lateral parts of rostrum to hind part of ver- posterior sclerite of the epiphallus, and more tex and along dorsomedial edges of eyes), protruding rachis clearly visible behind the with light brown small spots on proximal median epiphallic part (from L. omissa). part of antennae, stripes on pronotal disc Etymology. The species is named after (along lateral edges) and on dorsal tegmi- the Jambi Province. nal fields (along lateral edge of each dorsal field), and weakly distinct spots on legs, as Terrozacla harau sp. nov. well as with light greyish brown anal plate (Figs XXI: 3, 4; XXII: 6–8) and cercal bases. Scape approximately 1.5 times as wide as rostrum between anten- Holotype. Male; Indonesia, Sumatra I., West nal cavities; median ocellus well developed Sumatra Prov., about 20 km E of Sasak Town, but distinctly smaller than lateral ocelli. environs of Harau Valley National Park, equator, approximately 600 m, partly primary / partly Pronotum and metanotal gland as in Figs secondary forest, among dry leaves on forest XX: 1, 2. Tegmina reaching base of third floor at daytime, 24–26 Nov. 1999, A. Gorochov abdominal tergite, with rounded apex, 4–5 (ZIN). irregular longitudinal veins in dorsal field Paratypes. Two females, same data as for ho- (i. e. without traces of stridulatory appara- lotype (ZIN). tus), four longitudinal veins in lateral field, Description. Male (holotype). General and without distinct crossveins (Figs XX: appearance very similar to that of T. jambi 1; XXI: 1, 2). Fore tibiae without tympana. but tegmina slightly larger (reaching mid- Anal and genital plates with barely concave dle part of third abdominal tergite) as well hind part (Fig. XX: 3). Genitalia as in Figs as with weakly distinct traces of stridula- XXII: 1–3 (ectoparameres very charactetis- tory apparatus in both dorsal fields (Figs

Figs XX (1–10). Terrozacla, male: 1–3, T. jambi sp. nov.; 4–6, T. trusmadi sp. nov.; 7, 8, T. borneo sp. nov.; 9, T. kubah sp. nov.; 10, T. gading sp. nov. Body of male without distal part from above (1, 4); metanotal gland from above (2, 5); abdominal apex from above (3, 6, 8–10) and from above/ behind (7).

XXI: 3, 4) and five longitudinal veins in Paratypes. Three males and 2 females, same lateral fields, fore tibiae with small and oval data as for holotype (ZIN). inner tympanum only, metanotal gland with Description. Male (holotype). General slightly larger hind median lobe directed appearance similar to that of T. jambi (Fig. forwards, and anal plate roundly truncate XX: 4) but colouration of head with less dis- in hind part. Genitalia also very similar to tinct yellowish stripes on dorsum as well as those of T. jambi, but posterolateral projec- slightly lighter (almost light brown) lower tions of distal H-shaped epiphallic sclerite part of clypeus and area on mandibles, pro- with ventrolateral lobules more laterally notum and tegmina almost uniformly grey- projected and with distal part (behind these ish brown (with only lighter small basal lobules) narrower, posteromedian projec- humeral spot on tegmina), lightish spots tion of this epiphallic sclerite distinctly on fore and middle legs indistinct and on smaller, ectoparameres more strongly wid- hind legs almost absent, lateral ocelli less ened in distal part (almost laterally angu- distinct and barely larger than median ocel- lar), rachis with somewhat narrower apical lus, metanotal gland with clearly larger hind part as well as with longer median sclerite median lobe directed forwards (Fig. XX: 5), having proximal (narrower) part distinctly tegminal venation with very weak traces of longer, and formula with more distinct and stridulatory apparatus in both dorsal fields longer apodeme (Figs XXII: 6–8). (Fig. XXI: 5, 6) and with five longitudinal Female. Colouration and structure of veins in lateral field, fore tibiae with small body similar to those of male, but tegmina and oval inner tympanum only, anal plate reaching base of third abdominal tergite and with rather deep angular posteromedian with 5–6 parallel longitudinal veins in dor- notch and strongly angularly projected pos- sal field, latter field almost completely light terolateral lobes, and genital plate distinctly greyish brown, metanotum slightly lighter (but not deeply) notched at apex (Fig. XX: and without gland, and rest parts of abdo- 6). Genitalia: distal H-shaped epiphallic men approximately as in female of T. jambi. sclerite longer (especially its median part) Length in mm. Body: male 13, female and narrower, with somewhat shorter pos- 14–16; pronotum: male 2.5, female 2.9–3.1; terolateral projections, with much narrower tegmina: male 3.5, female 4–4.2; hind fem- notch between them, and with long and nar- ora: male 10.7, female 12–12.5; ovipositor rowly angular ventral processes directed 13.5–14.3. backwards and slightly downwards; ecto- Comparison. Differences of the new spe- parameres semisclerotized and with apical cies from T. jambi are given above. From L. part not bifurcate; rachis with somewhat mjobergi, the new species differs in a less narrower apical part; formula distinctly lon- transverse female genital plate; and from L. ger (Figs XXII: 9–11). omissa, in a very different shape of the distal Variations. Sometimes dorsum of head epiphallic sclerite (see the Terrozacla com- with more distinct yellowish stripes (al- parison above). most as in T. jambi), and anal plate with Etymology. The species is named after somewhat less deep and more rounded pos- the Harau Valley. teromedian notch. Female. Colouration and structure of Terrozacla trusmadi sp. nov. body as in holotype, but dorsal tegminal field (Figs XX: 4–6; XXI: 5, 6; XXII: 9–12) with 4–5 more or less distinct longitudinal veins, metanotal gland absent, and abdomen Holotype. Male; Malaysia, Borneo I., Sabah State, Trus Madi Mount, about 1000 m, partly similar to that of female of T. jambi and T. primary / partly secondary forest, among dry harau but with more transverse genital plate leaves on forest floor at night, 13–25 May 2007, having distinct posteromedian notch (this A. Gorochov (ZIN). notch slightly varied in size; Fig. XXII: 12).

Figs XXI (1–12). Terrozacla, male: 1, 2, T. jambi sp. nov.; 3, 4, T. harau sp. nov.; 5, 6, T. trusmadi sp. nov.; 7, 8, T. borneo sp. nov.; 9, 10, T. kubah sp. nov.; 11, 12, T. gading sp. nov. Left tegmen (1); dorsal field of right (2, 4, 6, 8, 10, 12) and left (3, 5, 7, 9, 11) tegmina.

Length in mm. Body: male 13–15, fe- upper half of pronotal lobes and spots on male 16–18; pronotum: male 2.6–3, female fore and middle legs, and greyish brown 3–3.2; tegmina: male 4–4.5, female 5.3–5.5; small marks on inner surface of hind femora. hind femora: male 12–13, female 13–14; Female. General appearance as in holo- ovipositor 15–16. type, but tegmina with slightly narrower Comparison. The new species differs distal half and without traces of stridula- from T. jambi and T. harau in the characters tory apparatus in dorsal field (this field of male genitalia listed above. From L. mjo- somewhat wider than in female of T. trus- bergi, the new species differs in a notched madi, with brown medial and light brown (not straight) hind edge of the female geni- lateral halves), metanotum and abdomen tal plate; and from L. omissa, in a strongly practically identical to those of female of T. bifurcate distal epiphallic sclerite. trusmadi. Etymology. The species is named after Length in mm. Body: male 14–15, female the Trus Madi Mount. 15–17; pronotum: male 2.6–3, female 3–3.2; tegmina: male 4.8–5.2, female 5.4–5.7; hind Terrozacla borneo sp. nov. femora: male 12–12.5, female 13–14; ovi- (Figs XX: 7, 8; XXI: 7, 8; XXII: 13, 14) positor 15–16. Comparison. Differences from T. trusma- Holotype. Male; Malaysia, Borneo I., Sabah di are given above. From all the other true State, Trus Madi Mount, about 1000 m, partly and possible congeners, the new species dif- primary / partly secondary forest, among dry fers in the same characters as T. trusmadi. leaves on forest floor at night, 13–25 May 2007, Etymology. The species is named after A. Gorochov (ZIN). the Borneo Island. Paratypes. Malaysia, Borneo I.: male and 3 females, same data as for holotype (ZIN); 2 males, Sarawak State, environs of Miri Town, Terrozacla kubah sp. nov. Lambir Hills National Park, 200–300 m, primary (Figs XX: 9; XXI: 9, 10; XXII: 15, 16) forest, on forest road among dry leaves at daytime, 29 March – 1 Apr. 2012, A. Gorochov, M. Holotype. Male; Malaysia, Borneo I., Sar- Berezin, E. Tkatsheva, I. Kamskov (ZIN). awak State, environs of Kuching City, Kubah National Park on Matang Mount, 200–500 m, Description. Male (holotype). Coloura- primary forest, among dry leaves on forest floor tion and structure of body very similar to at night, 10–17 March 2012, A. Gorochov, those of holotype of T. trusmadi, but teg- M. Berezin, E. Tkatsheva, I. Kamskov (ZIN). mina with dorsal field distinctly wider and Description. Male (holotype). Coloura- having more distinct traces of stridulatory tion and structure of body similar to those of apparatus (Figs XXI: 7, 8), stripe along lat- holotypes of T. trusmadi and T. borneo, how- eral edge of this field yellowish, some veins ever head dorsum and pronotal disc light of basal area of dorsal field in upper tegmen greyish brown with barely distinct darkish almost yellowish, medial half of this field marks, antennae light brown with almost behind basal area in lower tegmen light yellowish proximal part and small sparse greyish (semitransparent), lateral tegminal whitish spots on rest of flagellum, lateral field with 5–6 longitudinal veins, hind part lobes of pronotum with light brown stripe of anal plate almost not concave (rather along ventral edge, tegmina intermediate in sharply truncate; Fig. XX: 7, 8), and geni- structure between those of T. trusmadi and talia with narrower and distinctly longer T. borneo (Figs XXI: 9, 10) as well as almost posterolateral projections of distal epiphal- light brown with yellowish stripe on dorsal lic sclerite (Figs XXII: 13, 14). field along its lateral edge (lateral tegminal Variations. Males from Sarawak slightly field with 5–6 longitudinal veins), rest of lighter: almost light greyish brown with thorax and abdominal tergites also almost brown areas on face and behind eyes, brown light brown, anal plate intermediate in shape

Figs XXII (1–12). Terrozacla: 1–5, T. jambi sp. nov.; 6–8, T. harau sp. nov.; 9–12, T. trusmadi sp. nov.; 13, 14, T. borneo sp. nov.; 15, 16, T. kubah sp. nov.; 17, 18, T. gading sp. nov. Male genitalia from above (1, 6, 9, 13, 15, 17), from below (2, 7, 10) and from side (3, 8, 11, 14, 16, 18); distal part of ovipositor from side (4); female genital plate from below (5, 12). between those of above-mentioned repre- by angularly narrowed apical part slightly sentatives (Fig. XX: 9), and genitalia with shorter and directed mainly backwards (not posterolateral epiphallic projections similar partly laterally; Figs XXII: 15, 16). to those of T. trusmadi but distinguished Female unknown.

Length in mm. Body 16; pronotum 2.6; Length in mm. Body 15; pronotum 2.8; tegmina 4.7; hind femora 12. tegmina 5; hind femora 12.5. Comparison. The new species differs Comparison. Differences from T. trus- from T. trusmadi in the characters of male madi, T. borneo and T. kubah in the male genitalia listed above; from T. borneo, in genitalia are given above, and differences a narrower dorsal field of the male teg- from L. mjobergi are also unclear. From all mina and wider and shorter posterolateral the other true and possible congeners, the epiphallic projections in the male genitalia; new species differs in the same characters as from T. jambi and T. harau, in a much nar- T. trusmadi. rower notch between the above-mentioned Etymology. The species is named after epiphallic projections; and from L. omissa, the Gading Mount (Gunung Gading in Ma- in the same characters as T. trusmadi. The lay language). differences of the new species from L. mjobergi (described from the same mount) are Tribe OTTEINI Koçak et Kemal, 2009 unclear, as these species are known from one male and one female, respectively; however, Note. This tribe includes a few apterous, the male is significantly smaller (length of troglophilous crickets with an almost semi- its pronotum 2.6 mm; vs. 3.5 mm) and with globular head, long and thin apical segment an almost uniformly light (not distinctly of maxillary palpi, long and rather thin legs banded) head dorsum; I cannot exclude lacking tympana, and characteristic male that these specimens may be two colour genitalia having a series of folds before the variants of the same species. formula. Ruiz-Baliú & Otte (1997) consider Etymology. The species is named after that these folds belong to “spermatophore the Kubah National Park. tube mold”, but spermatophore tube is formed by a sacculus (= spermatophore sac) Terrozacla gading sp. nov. behind the formula (=mold of spermato- (Figs XX: 10; XXI: 11, 12; XXII: 17, 18) phore ancora, = mold of spermatophore attachment plate), i. e. between formula and Holotype. Male; Malaysia, Borneo I., Sar- rachis. Probably in Otteini, spermatophore awak State, 80–90 km WWN of Kuching City, is with a rather long collum (= neck, nar- Gunung Gading National Park, 100–300 m, primary forest, among dry leaves on forest floor at row part between ampulla and ancora or daytime, 8–9 March 2012, A. Gorochov, M. Ber- attachment plate of spermatophore) which ezin, E. Tkatsheva, I. Kamskov (ZIN). is formed in these folds. If it is correct, this Description. Male (holotype). Coloura- character is unique for Phalangopsinae and tion and structure of body similar to those other cricket taxa. of holotype of T. trusmadi, but tegmina and This tribe was originally described as anal plate more similar to those of T. kubah Cophusini for the genera Cophus Saussure, in structure (Figs XX: 10; XXI: 11, 12), 1874 and Cubacophus Ruiz-Baliú et Otte, and genitalia distinguished from those of 1997. But the name of its type genus (Co- T. trusmadi by wider both proximal part of phus) is twice preoccupied: in Coleoptera posteromedian epiphallic notch and basal and in Hymenoptera (Koçak & Kemal, part of ventral processes of posterolateral 2009; Eades et al., 2014). Thus, the names epiphallic projections (medial parts of these Cophus and Cophusini are unavailable, and ventral processes clearly visible from above the names Otteus Koçak et Kemal, 2009 and between bases of dorsal processes of pos- Otteini were proposed instead them. At the terolateral epiphallic projections; vs. these same year, Otte & Perez-Gelabert (2009) parts invisible from above in T. trusmadi, T. synonymized Cophus and Cubacophus. All borneo and T. kubah; Figs XXII: 17, 18). these data were summarized in the Orthop- Female unknown. tera Species File (Eades et al., 2014), and its

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 71 authors supposed: if the name Otteus (=Co- dark) with following marks: dorsum of head phus) is a junior synonym of Cubacophus, and face under median ocellus with a few the new name Cubacophusini must be pro- small and slightly lighter spots; mouthparts posed instead Otteini. and antennae light brown with greyish However, these genera are clearly dis- brown most part of antennal flagellum; pro- tinguished from each other by the structure notum with a pair of barely lighter spots on of male genitalia: in the genus Otteus nom. disc; tegmina brown with semitransparent dist. including type species (Cophus tho- membranes between Sc branches; exposed racicus Saussure, 1874) only, epiphallus is parts of hind wings somewhar darker; legs short and widely notched at the apex, rami light greyish brown with dark distal half are large (much longer than epiphallus), ec- of femora as well as dark tibiae and tarsi toparameres are moderately wide (ventral (proximal half of fore and middle femora view), rachis is rather short (not protruding with somewhat darkened area in basal part; behind median part of epiphallus) and not this half in hind femora with numerous strongly arcuate in the profile, formula is greyish brown oblique stripes on dorsal sur- rather simple (consisting of a pair of sclero- face; dark parts with a few almost whitish tized stripes posteriorly fused with the base transverse bands: one band on femora, 2–3 of rachis), and endoparameral apodemes are bands on fore and middle tibiae, 4–5 bands rather large and normal in the structure; in on hind tibiae, and one band at base of ba- Cubacophus including four other species sitarsus); venter of body greyish brown (but (type species: Cubacophus gibaraensis Ruiz- not dark); cerci slightly lighter. Rostrum of Baliú et Otte, 1997), epiphallus is long and head between antennal cavities barely wid- with a deep and narrow notch at the apex, er than scape; pronotum 1.8 times as wide rami are small (not longer or almost not lon- as long, distinctly narrowing to head; dor- ger than epiphallus), ectoparameres are very sal field of tegmina moderately wide, with narrow (ventral view), rachis is very long slightly shortened apical area and barely (strongly protruding behind median part of transverse mirror (Fig. XXIII: 1); lateral epiphallus) and strongly arcuate in the pro- field of tegmina with 25–27 branches of file, structure of the formula and endoparam- Sc situated more or less transversally (ver- eres is not very clear but possibly somewhat tically) and with moderately wide R-M complicated (see pictures in: Ruiz-Baliú & area; hind wings significantly longer than Otte, 1997; Otte & Perez-Gelabert, 2009). tegmina; legs with distinct subproximal widening of fore tibiae having large oval inner tympanum and slightly smaller (but Subfamily PHALORIINAE similar in shape) outer tympanum (both Tribe PHALORIINI tympana open), with rather long spines of Vescelia mulu sp. nov. hind tibiae, and with longest (inner dorsal) (Figs XXIII: 1; XXIV: 1–3; XXVII: 4) apical spur of these tibiae reaching middle of third tarsal segment; anal plate simple, Holotype. Male; Malaysia, Borneo I., Sar- with narrowed but almost truncate distal awak State, Mulu National Park (not far from part; genital plate distinctly larger, almost borders with Brunei and Indonesia), 100–300 m, primary forest, on leaf of small tree at night, not narrowed to roundly angular apical part 24–27 March 2012, A. Gorochov, M. Berezin, having deep median fold; genitalia as in Figs E. Tkatsheva, I. Kamskov (ZIN). XXIV: 1–3. Paratype. Male and female, same data as for Variations. Second male with somewhat holotype (ZIN). lighter R-M area in tegmina. Description. Male (holotype). General Female. General appearance similar to appearance typical of this genus. Body rath- that of male but with following differences: er small. Colouration greyish brown (rather pronotum less strongly narrowing to head,

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 72 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP approximately 1.4 times as wide as long; Paratype. Two males and female, same data as tegmina typical of female of Phaloriinae, for holotype (ZIN). with 9–10 more or less oblique longitudinal Description. Male (holotype). Body me- veins in dorsal field and rather sparse cross- dium-sized. Colouration yellowish with fol- veins between them, with 11–12 oblique lowing marks: head dorsum with a few nar- branches of Sc and almost without cross- row longitudinal stripes and a pair of dis- veins between them, and with Sc-R area tinctly wider bands behind eyes brown; face much wider than R-M area (in male, latter of head with a pair of narrow brown verti- area much wider than Sc-R area) and with cal stripes running from rostral apex (and sparse crossveins. Genital plate weakly fused with medial stripes of head dorsum) elongate, almost triangular but with rather to clypeus and with a pair of similar stripes deep and rounded posteromedian notch; under lateral parts of antennal cavities as ovipositor not long, with drilling apical part well as with small brownish spot on upper typical of Vescelia (Fig. XXVII: 4). part of mandibles; each gena with rather Length in mm. Body: male 9–10, female small brownish spot near subgena; antennae 8.5; body with wings: male 19–19.7, female with small light brown marks on scape and 19.5; pronotum: male 1.9–2, female 2.2; teg- numerous brownish spots on flagellum; pro- mina: male 13–13.5, female 13; hind femora: notum with dark brown disc having a few male 8.7–9, female 9; ovipositor 4. light brown spots, with dark brown upper Comparison. The new species differs part of lateral lobes (lower edge of this part from V. variegata (Chopard, 1937) in the oblique, running from anterodorsal corner rachis of male genitalia lacking strong cur- of this lobe to its posteroventral corner), vature (visible in the profile) near the base and with brownish area in anteroventral and in much longer ectoparameres; from V. corner of these lobes; tegmina semitrans- picta (Chopard, 1931), in much longer male parent with yellowish tinge as well as with genitalia and ectoparameres as well as in the rather large light brown areas and brown rachis having a rather deep posteromedian marks on dorsal field (Fig. XXIII: 2); legs notch; from V. pieli (Chopard, 1939), in with rather small brown spots on distal much wider both rachis and posteromedian half of fore and middle femora, brown and epiphallic notch as well as a much narrower brownish ventral keels of these femora, nu- apical part of the posterolateral epiphal- merous brown oblique stripes on hind fem- lic lobes; from V. moorei Chopard, 1940, in ora (excepting distal part of these femora much longer ectoparameres and narrower having three brownish transverse bands), posterolateral epiphallic lobes; and from and rather wide brown transverse bands on V. infumata Stål, 1877 known only from a tibiae and tarsi; abdominal sternites with female (Philippines), in distinctly spotted large light greyish brown area on median hind femora. part. Scape almost 1.2 times as wide as ros- Etymology. This species is named after trum between antennal cavities; pronotum the Mulu National Park. 1.45 times as wide as long, clearly narrowing to head; tegmina rather wide, with strongly Phaloria (Papuloria) latiuscula sp. nov. shortened apical area, strongly transverse (Figs XXIII: 2; XXIV: 4–6; XXV: 1; mirror (Fig. XXIII: 2), 26–28 Sc branches XXVII: 1) similar to those of V. mulu in shape, and R-M area also similar to those of this spe- Holotype. Male; Malaysia, Borneo I., Sarawak cies but with distinctly less numerous cross- State, environs of Kuching City, Kubah National Park, Matang Mount, 200–500 m, primary for- veins; hind wings shortened, reaching distal est, on leaf of bush at night, 10–17 March 2012, quarter of tegmina; structure of legs distin- A. Gorochov, M. Berezin, E. Tkatsheva, I. Kam- guished from that of V. mulu only by some- skov (ZIN). what less widened fore tibiae having inner

Figs XXIII (1–8). Vescelia and Phaloria, dorsal field of male upper tegmen: 1, V. mulu sp. nov.; 2, Ph. latiuscula sp. nov.; 3, Ph. paratristis sp. nov.; 4, Ph. tristis sp. nov.; 5, Ph. tariku sp. nov.; 6, Ph. waena sp. nov.; 7, Ph. manifesta sp. nov.; 8, Ph. neorava sp. nov. and outer tympana medium-sized and oval Variations. Sometimes upper tegmen (but outer tympanum open, and inner one with slightly lighter dorsal field and genita- barely immersed), by rather short spines of lia with somewhat less deep posteromedian hind tibiae, and by longest (inner dorsal) notch of rachis. apical spur of these tibiae reaching base of Female. General appearance similar to second tarsal segment; abdomen more or that of male, but structure of pronotum and less similar to that of V. mulu but with geni- of tegmina more similar to that of female tal plate roundly truncate at apex and hav- of V. mulu excepting a few differences: pro- ing less distinct posteromedian fold as well notum approximately 1.3 times as wide as as with genitalia as in fugures (Figs XXIV: long; tegmina and hind wings reaching ab- 4–6; XXV: 1). dominal apex (tegmen clearly shorter than

Figs XXIV (1–14). Vescelia and Phaloria, male: 1–3, V. mulu sp. nov.; 4–6, Ph. latiuscula sp. nov.; 7–10, Ph. paratristis sp. nov.; 11–14, Ph. tristis sp. nov. Genitalia from above (1, 4, 7, 11), from below (2, 5, 8, 12) and from side (3, 6, 9, 13); ectoparamere from below (10, 14).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 75 in male); dorsal tegminal field with 6–7 lon- darker longitudinal marks on dorsum and gitudinal veins; lateral tegminal field with brown spot behind each eye, four very small 12–13 branches of Sc. Genital plate almost brownish spots under antennal cavities, and as in V. mulu but slightly shorter (its length numerous small and very slightly darkened and width almost equal) and with slightly spots on middle and distal parts of anten- less deep posteromedian notch; oviposi- nal flagellum; pronotum with colouration as tor rather long, with apical part as in Fig. in Ph. latiuscula, but disc almost uniformly XXVII: 1. brown and lateral lobes with slightly lighter Length in mm. Body: male 10.5–11, fe- brownish area in each anteroventral cor- male 11; body with wings: male 14–14.7, ner; tegmina with brown dorsal field and female 11.5; pronotum: male 2.1–2.3, female yellowish (semitransparent) membranes 2.3; tegmina: male 10.5–11, female 8.3; hind in lateral field; legs with pattern similar to femora: male 10.7–11.3, female 10.5; ovi- that of Ph. latiuscula, but darkened marks positor 9. smaller and less distinct (upper half of prox- Comparison. The new species is simi- imal two thirds of outer side of hind femora lar in the general appearance to Ph. hob- without darkenings); abdominal sternites byi (Chopard, 1940) described also from also similar to those of Ph. latiuscula in co- Sarawak State (male genitalia of the latter louration. Rostrum of head between anten- species is unknown; Gorochov, 1999); Ph. nal cavities approximately 1.2 times as wide latiuscula is distinguished from it by a dis- as scape; pronotum almost 1.6 times as wide tinctly wider mirror in the male tegmina. as long, distinctly narrowing to head; dorsal Etymology. Name of this species is the tegminal field similar to that of Ph. latius- Latin word “latiuscula” (widish). cula but somewhat narrower and with less transverse mirror (Fig. XXIII: 4); lateral Phaloria (Papuloria) tristis sp. nov. tegminal field and legs also similar to those (Figs XXIII: 4; XXIV: 11–14; XXV: 2) of this species, but with 28–29 Sc branches, with both tympana open (not immersed), Holotype. Male; Malaysia, Borneo I., Sar- and with outer tympanum somewhat longi- awak State, Mulu National Park (not far from tudinally narrowed (clearly narrower than borders with Brunei and Indonesia), 100–300 m, oval inner one); abdomen similar to that of primary forest, on leaf of small tree at night, 24–27 March 2012, A. Gorochov, M. Berezin, Ph. latiuscula but with genitalia having pos- E. Tkatsheva, I. Kamskov (ZIN). teromedian epiphallic notch wider, trans- Paratypes. Three males and female, same parent lobules located in this notch longer, data as for holotype (ZIN); male, same state but rachis with truncate apex, medial arms of environs of Miri Town, Lambir Hills National endoparameres longer, formula clearly de- Park, 100–300 m, primary forest, on leaf of bush veloped (Figs XXIV: 11–13; XXV: 2), and at night, 29 March – 1 Apr. 2012, A. Gorochov, ectoparameres as in Fig. XXIV: 14. M. Berezin, E. Tkatsheva, I. Kamskov (ZIN); 3 Variations. Sometimes dorsum of head males and 3 females, same country and island, lighter (light brown with brown marks), Sabah State, Sandakan Division, environs of Sukau Vill. on Kinabatangan River (about 35 pronotal disc with light brown spots, and km from sea), almost sea level, partly second- genitalia with rachis and formula slightly ary / partly primary forest, on leaves of bushes at varied in width. night; 8–13 May 2013, A. Gorochov, M. Berezin, Female. General appearance as in male, E. Tkatsheva (ZIN). but head dorsum as in lighter males, teg- Description. Male (holotype). Body mina with brown dorsal field having light medium-sized. Colouration light brown brown elongate spot near middle of lateral with following pattern: head with slightly edge and with almost yellowish lateral field darker (intermediate between light brown having semitransparent membranes in Sc-R and brown) dorsum having a few barely area and between Sc branches, and structure

Figs XXV (1–7). Phaloria, male (schematically): 1, Ph. latiuscula sp. nov.; 2, Ph. tristis sp. nov.; 3, Ph. paratristis sp. nov.; 4, Ph. tariku sp. nov.; 5, Ph. manifesta sp. nov.; 6, Ph. waena sp. nov.; 7, Ph. neorava sp. nov. Epiphallus (proximal part not pictured) with posterior part of rachis (1, 4, 7) and without it (2, 3, 5, 6) from above. Abbreviations: m, membranous medial epiphallic lobules; r, rachis. of tegmina and hind wings similar to that of National Park, 100–300 m, primary forest, on Ph. latiuscula. Genital plate and ovipositor leaf of bush at night, 29 March – 1 Apr. 2012, also similar to those of female of this species, A. Gorochov, M. Berezin, E. Tkatsheva, I. Kam- but ovipositor distinctly shorter. skov (ZIN). Length in mm. Body: male 10–11.5, fe- Paratypes. Two males and 2 females, same data as for holotype (ZIN); female, same state male 8–10; body with wings: male 13.5–14, but Mulu National Park (not far from bor- female 10.5–11.5; pronotum: male 2–2.2, fe- ders with Brunei and Indonesia), 100–300 m, male 2–2.3; tegmina: male 10–10.7, female primary forest, on leaf of small tree at night, 7.3–8; hind femora: male 10–10.5, female 24–27 March 2012, A. Gorochov, M. Berezin, 9.5–11; ovipositor 6.6–7. E. Tkatsheva, I. Kamskov (ZIN); 3 males, same Comparison. The new species is most state but environs of Kuching City, Kubah Na- similar to Ph. hobbyi and Ph. latiuscula but tional Park, Matang Mount, 200–500 m, pri- distinguished from them by a somewhat mary forest, among dry leaves on forest floor at wider mirror in the male tegmina (from Ph. night, 10–17 March 2012, A. Gorochov, M. Ber- hobbyi) and by narrower both dorsal teg- ezin, E. Tkatsheva, I. Kamskov (ZIN). Description. Male (holotype). Structu- minal field and its mirror as well as by the characters of male genitalia listed above re of body and colouration very similar to (from Ph. latiuscula). those of Ph. tristis, but dorsum of head and Etymology. Name of this species is the pronotal disc uniformly dark brown, face Latin word “tristis” (gloomy). under rostral apex very light brown but with a pair of anterolateral brownish dots on clypeus, lower part of pronotal lateral Phaloria (Papuloria) paratristis sp. nov. lobes yellowish, tegmina with somewhat (Figs XXIII: 3; XXIV: 7–10; XXV: 3) smaller dorsal field and 26–27 Sc branches Holotype. Male; Malaysia, Borneo I., Sar- in lateral field (Fig. XXIII: 3), and genitalia awak State, environs of Miri Town, Lambir Hills with somewhat different shape of postero-

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 77 lateral epiphallic lobes (Figs XXIV: 7–9; Description. Male (holotype). Body me- XXV: 3) and with more massive distal half dium-sized. Colouration uniformly yellow- of ectoparameres undivided into two medial ish but with two basal segments of antennal lobes by deep notch (Fig. XXIV: 10). flagellum light brown, 14–15 nearest seg- Variations. Head dorsum and prono- ments of this flagellum dark brown, its rest tal disc sometimes barely lighter and with part brown to light brown and having nu- traces of lighter spots, respectively; shape merous small dark brown spots, pronotum of ectoparameres in genitalia insignificantly having light brown narrow stripes along varied. ventral edges as well as along anterior and Female. General appearance similar to posterior edges, tegmina with almost light that of male and very similar to that of fe- brownish dorsal field, fore and middle fem- male of Ph. tristis but distinguished from ora with a few short brownish grey longitu- latter female in head dorsum without dis- dinal lines on outer surface and along both tinct pattern and in pronotal disc uniformly ventral keels, hind femora with numerous brown or with slight traces of lighter spots. oblique brownish grey lines on outer sur- Length in mm. Body: male 9–10.5, fe- face and a few irregular marks on inner sur- male 8.5–10; body with wings: male 12–13, face as well as with a pair of brownish lon- female 10–10.5; pronotum: male 1.8–2, fe- gitudinal lines between ventral keels, fore male 1.9–2.1; tegmina: male 9.5–10.5, fe- tibiae with greyish stripe along dorsal sur- male 6.8–7.2; hind femora: male 9–10, fe- face, middle tibiae with two brownish spots male 9–9.5; ovipositor 6.3–6.6. on distal half, hind tibiae with six barely Comparison. The new species is distin- darker (light brown) spots, all tarsi with guished from Ph. hobbyi by practically uni- two darkish spots. Scape almost 1.3 times as formly brown both head dorsum and pro- wide as rostrum between antennal cavities; notal disc. From Ph. tristis, the new species pronotum approximately 1.4 times as wide differs in the characters listed above (in the as long, distinctly narrowing to head; struc- description); and from Ph. latiuscula, in dis- ture of dorsal tegminal field as in figure tinctly narrower both dorsal field of the male (Fig. XXIII: 5); lateral tegminal field hav- tegmen and its mirror, as well as in the same ing 25–26 Sc branches and distinctly wid- characters of male genitalia as Ph. tristis. ened R-M area (crossveins absent between Etymology. Name of this species consists Sc branches and sparse in R-M area); hind of the Greek prefix “para” (near) and spe- wings slightly not reaching tegminal apex; cies name Ph. tristis. inner and outer tympana not very large, open, oval and almost equal to each other in Phaloria (Papuloria) tariku sp. nov. size; spines of hind tibiae not very long; lon- (Figs XXIII: 5; XXV: 4; XXVI: 1–4; gest (inner dorsal) apical spur reaching base XXVII: 2) of second tarsal segment; abdomen similar to that of Ph. latiuscula but with shallow Holotype. Male; Indonesia, New Guinea I., angular notch at apex of genital plate and Papua Prov., Fawi [Faowi] Vill. in upper part of with characteristic genitalia (see Figs XXV: Tariku River (tributary of Mamberamo River), 4; XXVI: 1–4). lowlying secondary forest near river, on leaf of Variations. Sometimes antennal flagel- bush at night, 29 Jan. – 17 Feb. 2012, A. Goro- lum somewhat lighter, darkened marks on chov (ZIN). femora less distinct, and darkish stripe on Paratypes. Male and four females, same data as for holotype (ZIN); male, same island and fore tibiae divided into a few shorter parts. province but environs of Kasonaweja Vill. on Female. Geneal appearance as in male Mamberamo River near Van Rees Range, second- but stripes along edges of pronotum almost ary forest on not high hill, on leaf of small tree at brown, pronotum less distinctly narrow- night, 25–27 Jan. 2012, A. Gorochov (ZIN). ing to head, dorsal tegminal field yellowish

Figs XXVI (1–16). Phaloria, male: 1–4, Ph. tariku sp. nov.; 5–8, Ph. manifesta sp. nov.; 9–12, Ph. waena sp. nov.; 13–16, Ph. neorava sp. nov. Genitalia from above (1, 5, 10, 14), from below (2, 6, 11, 15) and from side (3, 7, 9, 13); ectoparamere from below (4, 8, 12, 16). and with 10–11 oblique longitudinal veins Length in mm. Body: male 11.5–13, fe- (crossveins between them very sparse), lat- male 11–13; body with wings: male 15–16, eral tegminal field similar to that of previ- female 14.5–16; pronotum: male 2.6–2.8, ous representatives of Phaloriinae but with female 2.6–2.9; tegmina: male 12–12.5, fe- 15–16 Sc branches (wings slightly shorter male 11–12.5; hind femora: male 11.5–12, than in male). Abdominal apex more or less female 11–12.5; ovipositor 7.7–8.3. similar to that of Ph. latiuscula but with dis- Comparison. The new species is similar tal part of ovipositor as in Fig. XXVII: 2. to Ph. solomonica Gorochov, 1996, Ph. vul-

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 79 gata Gorochov, 1996, Ph. solita Gorochov, and slightly transverse mirror (Fig. XXIII: 1996, Ph. gilva Gorochov, 1996, Ph. aphana 7); lateral field of tegmina with 24–25 more Gorochov, 1999 and Ph. verecunda Goro- or less vertical (transverse) branches of Sc chov, 2005 in the general appearance in- and with clearly widened R-M area hav- cluding colouration, but it is distinguished ing several crossveins; hind wings reaching from all these species by the male genitalia tegminal apex; outer and inner tympana with rather long and almost straight pos- not very large, almost equal to each other, terolateral epiphallic lobes and with small and oval (outer tympanum open; inner one oblique ectoparameres situated near base barely immersed); dorsal spines of hind of these lobes and having distomedial part tibiae not very long; longest (inner dorsal) with five distinct denticles (Fig. XXVI: apical spur of hind tibiae reaching base of 4). The differences from some other more second tarsal segment; abdomen as in Ph. or less similar species, known only from tariku, but genitalia (Figs XXV: 5; XXVI: females or insufficiently described, are less 5–7) with posterolateral lobes of epiphallus clear; the new species may differ from them apically bilobate and having oblique medial in a smaller body and/or longer ovipositor. spine, with rachis having a pair of long and Etymology. The new species is named af- thin posterolateral lobules, and with ecto- ter the Tariku River. parameres as in Fig. XXVI: 8. Female unknown. Length in mm. Body 15.5; body with Phaloria (Papuloria) manifesta sp. nov. wings: 20; pronotum 3.7; tegmina 15.5; hind (Figs XXIII: 7; XXV: 5; XXVI: 5–8) femora 14.3. Holotype. Male; Indonesia, New Guinea I., Comparison. The new species is more or Papua Prov., Fawi [Faowi] Vill. in upper part of less similar to Ph. tariku and to the conge- Tariku River (tributary of Mamberamo River), ners listed above (see comparison for Ph. lowlying secondary forest near river, on leaf of tariku). However, the new species differs bush at night, 29 Jan. – 17 Feb. 2012, A. Goro- from them in the male genitalia having chov (ZIN). slightly bilobate posterolateral epiphallic Description. Male (holotype). Body lobes and a medial oblique spine on each of rather large. Colouration almost uniform- these lobes, in the rachis with long and thin ly light: head light brown with slightly posterolateral lobules, and a characteris- darker dorsum and antennal flagellum as tic shape of the ectoparameres. From some well as with a pair of brown stripes along other similar species, it may differ in a larger medial edges of antennal cavities and grey- body and/or longer tegmina. ish brown medial part of scape; pronotum Etymology. Name of this species is the with light brown lower half of lateral lobes Latin word “manifesta” (obvious). and slightly darker all other parts; tegmina yellowish with greyish brown Sc branches and light brownish rest venation; legs light Phaloria (Papuloria) waena sp. nov. brown with yellowish coxae, a pair of longi- (Figs XXIII: 6; XXV: 6; XXVI: 9–12) tudinal greyish brown lines on outer surface Holotype. Male; Indonesia, New Guinea of hind femora, and a few barely darkened I., Papua Prov., Waena Vill. not far from Jaya- spots on hind tibiae; other body parts yel- pura City, 9–13 Aug. 2012, N. Kluge, L. Sheyko lowish with light brown middle and distal (ZIN). parts of cerci. Scape 1.15 times as wide as Description. Male (holotype). Body rostrum between antennal cavities; prono- medium-sized. Colouration yellowish with tum approximately 1.3 times as wide as long, following marks: head with a few longitudi- distinctly narrowing to head; dorsal field of nal stripes on dorsum, spot behind each eye, tegmina with moderately long apical area marks on rostral apex and on membranes

Figs XXVII (1–7). Phaloria, Vescelia, Tremellia and Pseudotrigonidium, distal part of ovipositor from below: 1, Ph. latiuscula sp. nov.; 2, Ph. tariku sp. nov.; 3, Ph. sulawesi sp. nov. (in dry specimen, left and right distal parts of ovipositor slightly twisted: their lateral surfaces directed downwards but not aside); 4, V. mulu sp. nov.; 5, T. timah orientalis subsp. nov.; 6, P. borneo sp. nov.; 7, P. gaponi sp. nov. of antennal cavities, stripes along dorsal (genae with small and lighter, greyish spot; edges of antennal cavities, vertical stripes clypeus with a pair of very small brownish under these cavities, and middle and distal marks); pronotum greyish brown with dis- parts of antennal flagellum greyish brown tinct yellowish spots on disc and on lateral

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 81 lobes as well as with light brown distal part Phaloria (Papuloria) neorava sp. nov. of disc; tegmina with light greyish brown (Figs XXIII: 8; XXV: 7; XXVI: 13–16) dorsal field having whitish area on basal part, with whitish R-M area having brown Holotype. Male; Indonesia, New Guinea I., Papua Prov., Fawi [Faowi] Vill. in upper part of crossveins, and with semitransparent (light Tariku River (tributary of Mamberamo River), greyish) membranes between light brown lowlying secondary forest near river, on leaf Sc branches; exposed part of hind wings of small tree at night, 29 Jan. – 17 Feb. 2012, greyish brown; legs spotted. Scape approxi- A. Gorochov (ZIN). mately 1.2 times as wide as rostrum be- Description. Male (holotype). Body tween antennal cavities; pronotum almost moderately large. Colouration light grey- 1.9 times as wide as long, clearly narrowing ish brown with following marks: head with to head; dorsal tegminal field rather narrow barely darker dorsum, a pair of brown dots and with rather long apical area as well as on superolateral parts of clypeus, very with almost round mirror (Fig. XXIII: 6); light mouthparts (but apical part of man- lateral tegminal field with 20–21 almost dibles and maxillary palpi darkened) and vertical (transverse) Sc branches and with proximal part of antennae (other antennal R-M area similar to that of Ph. manifesta parts missing); pronotum with light greyish but insignificantly narrower; hind wings brown lower half of lateral lobes and slight- strongly protruding behind tegminal apex; ly darker (insignificantly darker than head tympana also similar to those of Ph. mani- dorsum) rest parts; tegmina with very light festa but with slightly longer and mod- membranes between Sc branches; exposed erately narrow inner tympanum; dorsal part of hind wings similar to pronotal disc spines of hind tibiae long; longest (inner in colouration; fore and middle femora with dorsal) apical spur of these tibiae reach- several small darkish spots; hind femora ing base of third tarsal segment; abdomen with two larger darkish spots on distal half more or less similar to that of Ph. tariku of inner surface only; hind tibiae slightly and Ph. manifesta, but genital plate with spotted; middle part of all tarsi darkened; truncately concave hind edge and genitalia venter of body (including genital plate and with following peculiarities: posterolateral cerci) very light, yellowish. Scape approxi- epiphallic lobes lancet-like and medially mately 1.6 times as wide as rostrum between curved; rachis large, with narrowed and antennal cavities; pronotum 1.15 times as slightly bilobate apical part reaching apex wide as long, weakly narrowing to head; of above-mentioned lobes; and ectoparam- tegmina long, not wide, with almost round eres consisting of plate-like dorsal lobe and mirror, with long apical area (Fig. XXIII: semitubular ventral lobe directed medially 8), with 24–25 almost vertical (transverse) and partly forwards (Figs XXV: 6; XXVI: branches of Sc, and with R-M area similar to 9–12). that of Ph. manifesta; inner tympana slight- Female unknown. ly immersed, oval, medium-sized; outer Length in mm. Body 11.8; body with tympana open, almost round and approxi- wings 22; pronotum 2.1; tegmina 14; hind mately twice smaller (shorter) than inner femora 9.8. ones; longest (inner dorsal) apical spur of Comparison. The new species differs hind tibiae reaching middle of second tar- from all the other similar congeners in sal segment. Abdomen similar to that of Ph. medially curved and acute posterolateral tariku and Ph. manifesta, but genitalia with epiphallic lobes as well as in a characteristic ectoparameres of characteristic shape (Fig. shape of the rachis and in the ectoparameres XXVI: 16) and with rachis having long and divided into two lobes. thin posterolateral lobules not protruding Etymology. This species is named after behind posterolateral epiphallic lobes (Figs the Waena Village. XXV: 7; XXVI: 13–15).

Figs XXVIII (1–9). Tremellia and Phaloria: 1–4, T. timah orientalis subsp. nov.; 5–7, T. t. timah Gor. et Tan; 8, 9, Ph. sulawesi sp. nov. Dorsal field of male upper tegmen (1); male genitalia from above (2, 5); distal part of rachis (3, 6) and of epiphallic posterolateral lobes (4, 7) from below; female body from above (8) and from side (9).

Female unknown. Phaloria neorava is additionally distin- Length in mm. Body 15.8; body with guished from Ph. rava and Ph. pararava by wings 24.5; pronotum 3.5; tegmina 16; hind wider ectoparameres of somewhat differ- femora 14. ent shape (Fig. XXVI: 16); from Ph. gilva, Comparison. The new species is similar by shorter posterolateral lobules of the to Ph. rava Gorochov, 1996, Ph. pararava rachis and also another shape of the ecto- Gorochov, 1999, Ph. gilva Gorochov, 1996 parameres; and from Ph. chopardi, by the and Ph. chopardi (Willemse, 1950) in the posterolateral epiphallic lobes not curved rachis of male genitalia with long and thin medially. posterolateral lobules. But it differs from Etymology. The new species name con- them in these lobules not protruding be- sists of the Greek prefix meaning “new” and hind the posterolateral epiphallic lobes. species Ph. rava.

Phaloria (Papuloria) heterotrypoides following pattern: epicranium with slightly Gorochov, 1999 darker most part of genae and a pair of spots under eyes, with distinctly darker (brown) = Phaloria galoa Otte et Cowper, 2007, syn. nov. rostrum and four longitudinal lines on dor- Note. The species was described from Fiji sum, and with almost yellowish lower part (Vity Levu I.) as a member of the nomino- of face; mouthparts also yellowish but with typical subgenus (Gorochov, 1999). Later whitish labrum; antennae with light brown this species was repeatedly described as Ph. proximal part and brown middle part (distal galoa from the same island: judging by its part missing); pronotum brown with slight- description and photographs (Otte & Cow- ly lighter posterior part and a pair of spots per, 2007), Ph. galoa is a junioir synonym on disc before this part; tegmina with semi- of Ph. heterotrypoides. Male genitalia of Ph. transparent membranes in lateral field (Figs heterotrypoides have distinct elongate me- XXVIII: 8, 9); exposed parts of hind wings dial membranous lobules on the posterior slightly darker than tegmina (almost grey- epiphallic edge (presence of a pair of mem- ish brown); legs with brown, light brown branous lobules or one membranous lobe in and yellowish spots (but on femora, these the posterior epiphallic notch is a distinct spots not very contrast); cerci and oviposi- diagnostic character of the subgenus Papu- tor almost yellowish but with reddish brown loria Gorochov, 1996); thus, this species marks in drilling part of ovipositor (Fig. must be transferred from the subgenus Pha- XXVII: 3). Rostrum of head between anten- loria to Papuloria. nal cavities almost equal to scape in width; pronotum 1.35 times as wide as long, mod- Phaloria (Papuloria) insularis erately narrowing to head; tegmina (Figs (Bolivar, 1912) XXVIII: 8, 9) long, distinctly protruding behind abdominal apex, with 11–12 weakly Note. This species, consisting of two sub- oblique longitudinal veins in dorsal field and species [Ph. i. insularis from Seychelles; Ph. rather numerous crossveins between them, i. karnyi (Chopard, 1929) from Java and Su- and with 14–15 oblique branches of Sc and matra], was previously placed by me in the rather numerous crossveins in Sc-R area nominotypical subgenus (Gorochov, 1996). (this area much wider than R-M area); hind However, its epiphallus has a pair of mem- wings significantly protruding behind teg- branous lobules in the posterior notch (a minal apex; fore tibiae with clearly widened diagnostic character for the subgenus Papu- part near their base, with large, oval and loria; see above); thus, Ph. insularis must be moderately immersed inner tympana, and also transferred from the subgenus Phaloria with distinctly smaller, oval and open outer to Papuloria. tympana (inner tympanum approximately 1.7 times as long as outer one); hind tibiae Phaloria sulawesi sp. nov. with not very long dorsal spines and longest (Figs XXVII: 3; XXVIII: 8, 9) (inner dorsal) apical spur reaching apex of second tarsal segment; genital plate approx- Holotype. Female: Indonesia, Sulawesi I., imately as long as wide, weakly narrowing Sulawesi Utara Prov., about 40 km NE of Ma- backwards, and with rather large (but not nado City, Tangkoko National Park on eastern coast of Minahassa Peninsula, environs of Tang- very deep) apical notch and short rounded koko Lodge, partly primary / partly secondary lobes around it; ovipositor short, with apical forest, on leaf of bush near brook at night, 3–6 (drilling) part as in Fig. XXVII: 3. Feb. 2011, A. Gorochov (ZIN). Male unknown. Description. Female (holotype). General Length in mm. Body 12.5; body with appearance typical of genus Phaloria. Body wings 24.5; pronotum 2.8; tegmina 17.2; medium-sized. Colouration light brown with hind femora 12.7; ovipositor 4.5.

Comparison. The new species is most Female. General appearance as in male, similar to Ph. doloduo Gorochov, 2011 (Su- but dorsal tegminal field and genital plate lawesi) in the structure of inner tympana as in female of nominotypical subspecies but distinguished from it by a smaller and (Gorochov & Tan, 2012), lateral tegminal lighter body with more uniformly coloured field with 14–16 Sc branches (vs. 11 in T. tegmina as well as with less contrast spots t. timah), and apical part of ovipositor as in on the legs and with a somewhat shorter Fig. XXVII: 5. ovipositor. From Ph. lindu Gorochov, 2011 Length in mm. Body: male 11–13.5, (also from Sulawesi), the new species dif- female 11–13; body with wings: male 16– fers in the same characters as Ph. doloduo 18.5, female 17–18.5; pronotum: male 2.4– (except for the ovipositor structure, as fe- 3.1, female 2.8–3.2; tegmina: male 11–13.5, male of the latter species is unknown), in female 11–13; hind femora: male 11–13, fe- more uniformly coloured genae and lateral male 13–13.5; ovipositor 5.6–6. pronotal lobes, and in the hind wings more Comparison. Differences of the new sub- protruding behind the tegminal apex. Sub- species from nominotypical one are given generic position of Ph. sulawesi is unclear. above, in the description. Etymology. The new species is named af- Etymology. The subspecies name is the ter the Sulawesi Island. Latin word “orientalis” (eastern).

Tremellia timah orientalis subsp. nov. Pseudotrigonidium borneo sp. nov. (Figs XXVII: 5; XXVIII: 1–4) (Figs XXVII: 6; XXIX: 1, 2)

Holotype. Male; Malaysia, Borneo I., Sabah Holotype. Female; Malaysia, Borneo I., Sabah State, Sandakan Division, environs of Sukau Vill. State, Sandakan Division, environs of Sukau Vill. on Kinabatangan River (about 35 km from sea), on Kinabatangan River (about 35 km from sea), almost sea level, partly secondary / partly primary almost sea level, partly secondary / partly primary forest, on leaves of bushes at night; 8–13 May 2013, forest, on leaves of bushes at night; 8–13 May 2013, A. Gorochov, M. Berezin, E. Tkatsheva (ZIN). A. Gorochov, M. Berezin, E. Tkatsheva (ZIN). Description. Male (holotype). Size, co- Description. Female (holotype). Body louration and structure of body very simi- small and thin, with long and thin legs. Co- lar to those of less spotted males of nomi- louration variegate: head yellowish with notypical subspecies (see Gorochov & Tan, greyish brown both large area on dorsum 2012), but head and pronotum with slightly and short longitudinal band behind each darker upper half (dorsum of head greyish eye, X-shaped dark brown spot on face, a brown, most part of pronotal disc and up- pair of brownish oblique stripes under eyes, per part of lateral lobes almost dark grey- a pair of darkish dots near superolateral ish brown), tegmina with somewhat more corners of clypeus, small darkened marks on arcuate stridulatory vein and narrower area scape, light brown proximal part of anten- between proximal parts of 1A and 2A (Fig. nal flagellum, and brown rest of flagellum XXVIII: 1), genital plate with slightly con- (but with rather numerous small lightish cave apical edge, and genitalia with thinner spots); pronotum greyish brown with sev- (narrower) subapical part of long postero- eral light spots on disc (Fig. XXIX: 1) and lateral epiphallic lobes and with slightly two yellowish longitudinal stripes on lateral notched apex of rachis (in T. t. timah, this lobes (Fig. XXIX: 2); tegmina light greyish epiphallic part clearly widened, and rachis brown with brown to dark brown (almost with truncate apex; for comparison see Figs black) venation; exposed part of hind wings XXVIII: 2–4 and 5–7). greyish brown; legs rather contrastingly Variations. One male with genital plate spotted but with almost uniformly light having barely convex apical edge (practi- greyish brown hind tibiae (having darkened cally as in nominotypical subspecies). apical part); venter of thorax and abdominal

Figs XXIX (1–4). Pseudotrigonidium, female: 1, 2, P. borneo sp. nov.; 3, 4, P. gaponi sp. nov. Body from above (1, 3) and from side (2, 4).

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 86 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP sternites light greyish brown; genital plate Pseudotrigonidium gaponi sp. nov. lighter; and cerci also light greyish brown (Figs XXVII: 7; XXIX: 3, 4) with greyish brown dorsal and lateral surfaces of proximal part. Head rather low Holotype. Female; Indonesia, Sulawesi I., Sulawesi Tenggara Prov. (Southeast Sulawesi), (somewhat dorsoventrally compressed) and environs of Kendari City, 9–12 Nov. 2011, D. with shallow dorsal concavity in region of Gapon (ZIN). rostral base; scape approximately 1.7 times Description. Female (holotype). General as wide as rostrum between antennal cavi- appearance somewhat similar to that of P. ties. Pronotum almost 1.4 times as wide as borneo, but body clearly larger, lighter and long, with more or less parallel lateral sides with different pattern. Head whitish with and low lateral lobes (Figs XXIX: 1, 2). Teg- a pair of greyish brown spots on dorsum, a mina weakly protruding behind abdominal few oblique greyish stripes on gena and be- apex, with 8–9 longitudinal veins in dor- hind eye, almost X-shaped greyish brown sal field (these veins slightly oblique and median spot on face, a pair of greyish curved somewhat irregular, curved), with 13–14 lines around it, sparse darkish marks along oblique branches of Sc, with R-M area much subgenae and between antennal cavities as narrower than Sc-R area, and with not nu- well as on clypeus and maxillary palpi, and merous crossveins in Sc-R area and in dorsal light brown antennal flagellum having nu- field (Figs XXIX: 1, 2); hind wings slightly merous lightish dots in middle part (distal protruding behind tegminal apex. Fore tibi- part missing); pronotum with brown disc ae slightly widened neare base, having rath- having a few light brown and yellowish er long (almost oval) and slightly immersed spots as well as with whitish lateral lobes inner tympanum only; hind tibiae with having greyish brown both upper half (with four pairs of rather short dorsal spines and small light brown to yellowish anterior and longest (inner dorsal) apical spur reaching posterior areas) and longitudinal stripe in middle part of hind basitarsus. Genital plate lower half (this stripe posteriorly fused with transverse, with rounded hind part having previous darkened half); tegmina whitish small shallow posteromedian notch; ovipos- with greyish medial two thirds of dorsal itor with apex as in Fig. XXVII: 6. field and distal part of lateral field, with Male unknown. greyish brown venation and a few spots Length in mm. Body 9.5; body with in dorsal field, with two blackish areas in wings 11.2; pronotum 1.9; tegmina 8; hind lateral field and almost transparent mem- femora 11.6; ovipositor 5.7. branes between distal halves of proximal Comparison. The new species is most Sc branches, and with whitish and greyish similar to P. javanicum (Chopard, 1954) in brown as well as blackish venation in lat- the colouration and shape of lateral prono- eral field (Figs XXIX: 3, 4); exposed parts tal lobes, but it is distinguished from the lat- of hind wings greyish brown; legs contrast- ter species by a lower head and a character- ingly spotted, but hind tibiae with light and istic pattern on the face of head (with a dark weakly spotted proximal and middle parts X-shaped spot and a pair of the darkened and with brown distal part; rest of body yel- oblique stripes under eyes, and without any lowish to whitish with greyish most part of dark stripe along clypeal suture). From all sternites (genital plate lighter) and partly the other congeners, P. borneo differs in the darkened ovipositor (Fig. XXVII: 7). Scape presence of two light longitudinal stripes on approximately 1.2 times as wide as rostrum the pronotal lobes and in the above-men- between antennal cavities; pronotum al- tioned pattern of face. Subgeneric position most 1.3 times as wide as long; tegmina with of the new species is unclear. 9–10 not very regular longitudinal veins in Etymology. This species is named after dorsal field as well as with irregular and not the Borneo Island. very numerous crossveins between them,

© 2014 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 23(1): 7–88 A.V. GOROCHOV. CLASSIFICATION AND NEW TAXA OF PHALANGOPSINAE SUBFAMILY GROUP 87 with 15–16 Sc branches and rather numer- de Doctorat, Universite de Paris, 1247: ous crossveins in Sc-R area (Figs XXIX: 3, 1–347. 4); hind wings barely protruding behind Desutter-Grandcolas L. 1992a. Les Phalan- ç è tegminal apex; inner tympanum distin- gopsidae de Guyane fran aise (Orthopt res, Grylloidea): systématique, éléments de phy- guished from that of P. borneo by narrower logénie et de biologie. Bulletin du Muséum distal part (outer tympanum absent); hind national d’Histoire naturelle (4e série), 14 tibiae with spines and spurs also similar to (section A, 1): 93–177. those of P. borneo; genital plate with widely Desutter-Grandcolas L. 1992b. Les Phalangop- truncate apex; ovipositor with apex as in sidae Neotropicaux (Orthoptera: Grylloi- Fig. XXVII: 7. dea). II. Le groupe des Aclodae. Annales de Male unknown. la Société entomologique de France (Nouvelle Length in mm. Body 13; body with série), 28(2): 171–199. wings 15.8; pronotum 2.7; tegmina 11.7; Desutter-Grandcolas L. 1993. The cricket fa- hind femora 14.5; ovipositor 6.5. u na of Chiapanecan caves (Mexico): sys- tematics, phylogeny and the evolution of Comparison. The new species is similar troglobitic life (Orthoptera, Grylloidea, to P. javanicum and P. borneo in the coloura- Phalangopsidae, Luzarinae). International tion and shape of lateral pronotal lobes. It Journal of Speleology, 22(1–4): 1–82. differs from P. javanicum in a more spotted Desutter-Grandcolas L. 1995. Le genre Eid- dorsum of the head, lighter pronotal disc, manacris Chopard, 1956 (Orthoptera, Gryl- and all the ocelli obliterated (in P. javani- loidea, Phalangopsidae, Luzarinae): habitat, cum, the median ocellus is developed); from répartition et espèces nouvelles. Bulletin du P. borneo, in a larger and lighter body as well Muséum national d’Histoire naturelle (4e sé- as almost truncate apex of the genital plate; rie), 16 (section A, 2–4), 1994: 453–474. and from all the other congeners, in the pres- Desutter-Grandcolas L. & Jaiswara R. 2012. Phalangopsidae crickets from the Indian Re- ence of two light longitudinal stripes on the gion (Orthoptera, Grylloidea), with the de- pronotal lobes and in the above-mentioned scriptions of new taxa, diagnoses for genera, pattern of face. Subgeneric position of this and a key to Indian genera. Zootaxa, 3444: new species is also unclear. 1–39. Etymology. The new species is named in Eades D.C., Otte D., Cigliano M.M. & Bra- honor of its collector. un H. 2014. Orthoptera Species File Online. Version 5.0/5.0. Visited 3 April 2014. Avail- ACKNOWLEDGEMENTS able from: < http://Orthoptera.SpeciesFile. org> The author is thankful to the collectors of Gorochov A.V. 1995. System and evolution of these insects as well as to Dr. J. Beccaloni and the suborder Ensifera (Orthoptera). Part 2. Mrs. J. Marshall (NHM) for supplying the ad- Proceedings of the Zoological Institute of RAS, ditional material. The study is supported by the 260: 1–213. Presidium of the Russian Academy of Sciences Gorochov A.V. 1996. New and little known (Program “Biosphere Origin and Evolution of crickets from the collection of the Humboldt Geo-biological Systems”). University and some other collections (Or- thoptera: Grylloidea). Part 2. Zoosystematica REFERENCES Rossica, 5(1): 29–90. Gorochov A.V. 1999. New and little known Chopard L. 1968. Gryllides. Orthopterorum Cat- Phaloriinae (Orthoptera: Gryllidae). Zoosys- alogus, 12: 213–500. tematica Rossica, 8(1): 27–60. Chopard L. 1969. Grylloidea. The fauna of India Gorochov A.V. 2001. Preliminary notes on the and the adjacent countries. Orthoptera, 2. Cal- history of South American Ensifera (Or- cutta: Baptist Mission Press. 421 p. thoptera). Acta Geologica Leopoldensia, Desutter L. 1990. Etude phylogenetique, bio- 24(52/53): 81–86. geographique et ecologique des Grylloidea Gorochov A.V. 2003a. New and little known neotropicaux (Insectes, Orthoptères). These crickets of the subfamily Phalangopsinae

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