Taxonomy of Podoscirtinae (Orthoptera: Gryllidae). Part 9: the American Tribe Paroecanthini Таксономия Подсемейства Podoscirtinae (Orthoptera: Gryllidae)

ZOOSYSTEMATICA ROSSICA, 20(2): 216–270 25 DECEMBER 2011

Taxonomy of Podoscirtinae (Orthoptera: Gryllidae). Part 9: the American tribe Paroecanthini Таксономия подсемейства Podoscirtinae (Orthoptera: Gryllidae). Часть 9: американская триба Paroecanthini

A.V. GOROCHOV

A.В. ГОРОХОВ

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

Systematic position and composition of the endemic American tribe Paroecanthini are discussed. This tribe is divided into two subtribes: Paroecanthina stat. nov. (from Paroecanthini Gorochov, 1986) and Tafaliscina stat. nov. (from Tafaliscinae Desutter, 1988). Five new genera, 24 new species and 6 new subspecies are described. Systematic position and distribution of true and possible taxa of Paroecanthini are clarified, and some of these taxa are redescribed. Orocharis eclectos Otte, 2006, syn. nov. is synonymised with Paroecanthus mexicanus Sau- ssure, 1859 which is restored as type species of Paroecanthus Saussure, 1859 according to original monotypy of this genus. Orocharis signatus Walker, 1869 and Carsidava Walker, 1869 are excluded from synonymy of P. mexicanus and Paroecanthus, respectively. Orocharis signatus is considered to be a probable synonym of P. aztecus aztecus Saussure, 1874. Carsidava and Chremon Rehn, 1930, syn. nov. are considered possible and evident synonyms of Ectotrypa Saussure, 1874, respectively. Angustitrella vicina (Chopard, 1912), sp. resurr. and A. picipes (Bruner, 1916), sp. resurr. are restored from synonymy of A. podagrosa (Saussure, 1897) and Siccotrella niger (Saussure, 1874), respectively. Lectotype of Amblyrhethus brevipes (Saussure, 1878) and type species of Metrypa Brunner-Wattenwyl, 1873 (Tafalisca lurida Walker, 1869) are designated. Pseudogryllus Chopard, 1912, gen. resurr. is restored from synonymy of Ta - falisca Walker, 1869. Обсуждены cистематическое положение и состав эндемичной американской трибы Paroecanthini. Эта триба подразделяется на две подтрибы: Paroecanthina stat. nov. (от Paroecanthini Gorochov, 1986) и Tafaliscina stat. nov. (от Tafaliscinae Desutter, 1988). Опи- саны 5 новых родов, 24 новых вида и 6 новых подвидов. Выяснены систематическое положение и распространение настоящих и возможных таксонов Paroecanthini, и неко- торые из этих таксонов переописаны. Orocharis eclectos Otte, 2006, syn. nov. синонимизи- рован с Paroecanthus mexicanus Saussure, 1859, который восстанавливается как типовой вид Paroecanthus Saussure, 1859 в связи с оригинальной монотипией этого рода. Orocharis signatus Walker, 1869 и Carsidava Walker, 1869 исключаются из синонимии P. mexicanus и Paroecanthus соответственно. Orocharis signatus предлагается считать вероятным сино- нимом P. aztecus aztecus Saussure, 1874. Carsidava и Chremon Rehn, 1930, syn. nov. рассма- триваются как возможный и явный синонимы Ectotrypa Saussure, 1874 соответственно. Angustitrella vicina (Chopard, 1912), sp. resurr. и A. picipes (Bruner, 1916), sp. resurr. ис- ключаются из синонимии A. podagrosa (Saussure, 1897) и Siccotrella niger (Saussure, 1874) соответственнно. Лектотип Amblyrhethus brevipes (Saussure, 1878) и типовой вид для Me- trypa Brunner-Wattenwyl, 1873 (Tafalisca lurida Walker, 1869) обозначаются. Pseudogryllus Chopard, 1912, gen. resurr. восстанавливается из синонимии Tafalisca Walker, 1869. Key words: America, taxonomy, Orthoptera, Gryllidae, Podoscirtinae, Paroecanthini, new taxa Ключевые слова: Америка, таксономия, Orthoptera, Gryllidae, Podoscirtinae, Paroecan- thini, новые таксоны

INTRODUCTION us to transfer this tribe from Podoscirtinae to Eneopterinae. This paper is the ninth communication in the series of publications on taxonomy of Tribe PAROECANTHINI Gorochov, 1986 Podoscirtinae. In the previous communi- cations, the tribes Podoscirtini and Apho- A most characteristic feature of this tribe noidini from Old World (Gorochov, 2002, is the unique structure of ovipositor which 2003, 2004, 2005, 2006, 2007, 2008) as well is dorsoventrally flattened and with the dis- as Podoscirtini from New World (Goro- tal part strongly flattened and having small chov, 2010) were discussed. denticles on the lateral edges of valves (Figs Most part of the material examined (in- I: 103; II: 3, 7, 10; IX: 4, 9, 17, 18, 21; XV: cluding the type series of all new species) 7, 8, 10, 11, 13, 14, 17, 18, 21–24; XVIII: is deposited at the Zoological Institute, 7–9; XIX: 7, 8). It is somewhat other type Russian Academy of Sciences, St Peters- of drilling ovipositor than in all the other burg (ZIN). But some specimens are from taxa of Podoscirtinae; this type is a result the following institutions: Museum für of independent specialisation to the ovipo- Naturkunde der Humboldt-Universität, sition into plant tissues (general ancestor of Berlin (MNH); Museum and Institute of Podoscirtinae possibly had the ovipositor Zoology, Polish Academy of Sciences, War- weakly specialised to such oviposition, but saw (MIZ). all the members of Eneopterinae have the Majority of specimens studied were col- ovipositor not drilling, adapted to the ovi- lected in tropical forests at night, on leaves position into soil or similar loose substrate). of trees and bushes or at light. However Usually small tubercles and low transverse some species were collected in less usual ridges are additionally present on the ven- conditions which will be briefly character- tral and /or dorsal surfaces of the ovipositor ised for each such species. distal part. However sometimes all drilling denticles and ridges on the ovipositor are TAXONOMIC PART partly reduced or practically absent (Figs The brief characteristics of Podoscirti- XVI: 10, 11); such structure may be a result nae and differences of this subfamily from of a return transition from the oviposition Eneopterinae are given in the preceding into plants to the oviposition into moulder- communication of this series (Gorochov, ing wood, soil or cracks of bark. 2010). Here it is necessary to mention only a All other external characters of Paroe- problem with the tribe Paroecanthini Goro- canthini are typical of Podoscirtinae: head chov, 1986. This tribe was originally includ- somewhat dorsoventrally depressed and ed in Podoscirtinae (Gorochov, 1986), but with more or less flattened dorsal area (from Desutter (1987, 1988) put Paroecanthini rostrum to hind part of vertex), angular or in Eneopterinae (=Eneopteridae sensu De- roundly angular rostrum of head in profile, sutter). She considered that Podoscirtinae and not very large eyes usually situated and Eneopterinae are families, and created near dorsal surface of head; thorax usually the subfamily Tafaliscinae Desutter, 1988 with low or moderately low pronotum and with Paroecanthini as one of her tribes of often with male metanotal gland typical of Tafaliscinae. These her actions are not very Podoscirtinae in structure; all tarsi with understandable, because Podoscirtinae and very wide second segment; fore and middle Eneopterinae are very different groups hav- legs with very short basitarsus; apical spines ing only weak convergent similarity con- (spurs) of hind basitarsi large; stridulatory nected with the adaptation to life on leaves apparatus of male tegmina (if it well devel- of plants (Gorochov, 2010), and Paroecan- oped) with rather numerous oblique veins, thini does not have any characters allowing oval or round mirror, and very long arcuate

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 218 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI cell completely or almost completely bor- Note. This genus was originally described dering hind half of mirror (Figs II: 1, 4, 5; as Paroccanthus for only a single species – P. VII: 1–5; X; XIV: 9; XV: 3, 9, 12). But in mexicanus described (judging by its name, difference from the other taxa of Podoscir- because geographical data were omitted) tinae, Paroecanthini with such stridulatory from Mexico (Saussure, 1859). So, type spe- apparatus has the lateral part of stridula- cies of this genus is P. mexicanus by mono- tory vein clearly S-shaped and lacking any typy (!), but not P. olmecus Saussure, 1897 stridulatory teeth, and oblique veins normal which was incorrectly indicated as type of (long, weakly sinuate or almost straight, Paroecanthus by Kirby (1906) and subse- but not zigzag-shaped; Figs I: 4–7). quent authors (Chopard, 1968; Otte, 1994; Male genitalia in this tribe are with Otte, Gelabert, 2009; Eades et al, 2011). some elements reduced. In one group of In one of his later publications, Saussure genera, several genital sclerites are reduced (1874) changed the name Paroccanthus for or absent (Figs I: 8–10; IV; XII); in second Paroecanthus, because he produced it from group of genera, all these sclerites are well the genus Oecanthus Audinet-Serville, 1831. developed, but another sclerite (guiding The following words present in the first de- rod) may be reduced (Figs XV: 4–6; XVIII: scription of Saussure’s genus: “Generibus 1–8). These two groups are considered here Occantho et Trigonodio affinis”. However as subtribes. there are only two most similar and older generic names of crickets: Oecanthus and Subtribe PAROECANTHINA Gorochov, Trigonidium. This is an indication that Sau- 1986, stat. nov. ssure’s generic name is based on Oecanthus, but he or (more probably) typography made This subtribal name originates from some mistakes in the generic names which the tribe Paroecanthini described earlier were corrected by the same author in 1874. (Gorochov, 1986). The subtribe is charac- Also the name Paroecanthus have been used terised by the lateral part of male stridula- by all the authors since 1874 as a valid ge- tory vein (lacking stridulatory teeth) long neric name proposed in 1859. So, the above- and strongly S-shaped (Figs I: 4–6), male mentioned correction is in accordance to the genitalia with several sclerites reduced Code of Zoological Nomenclature: articles and/or absent (ectoparameres are often not 32.5.1 or 33.2.3.1 (International Commis- separated from the epiphallus or practically sion on Zoological Nomenclature, 1999) (as absent; endoparameres are absent or pos- “the Code” elsewhere below). sibly very small, lacking distinct apodemes The genus Paroecanthus is characterised and fused with guiding rod; mold of sper- by the following characters (Figs I: 4, 8–10; matophore attachment plate completely II; III: 1–13; IV; V; VI: 1–9): head is rather membranous; but guding rod is not reduced; short, with the roundly angular (in profile) Figs I: 8–10; IV–VI; VIII; XI: 1–18; XII), and distal part of ovipositor with the acute rostrum narrower than scape, and with or narrowly rounded apex (Figs I: 1–3; II: 3, the comparatively large lateral ocelli and 7, 10; IX: 4, 9, 17, 18, 21). smaller median ocellus (three these ocelli Paroecanthina includes the following gen- situated in shape of triangle); pronotum era: Paroecanthus Saussure, 1859; Siccotrella is not very long (distinctly wider than its gen. nov.; Angustitrella gen. nov.; Ectotrypa length) and distinctly or strongly narrow- Saussure, 1874; Selvagryllus Otte, 2006. ing to head; male metanotal gland is with the middle process transverse and more or Genus Paroecanthus Saussure, 1859 less high (Figs III: 1, 5, 7, 9); fore tibiae are hardly or moderately inflated in the tympa- Type species: Paroccanthus mexicanus nal region, with the inner tympanum long, Saussure, 1859. deeply immersed and slit-like, and with the

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 219 outer tympanum distinctly shorter (almost 1869; P. fallax Saussure, 1874; P. hwinanus round or slightly longitudinal), opened, Otte et Perez-Gelabert, 2009; P. otaros Otte and weakly immersed; tegmina are long or et Perez-Gelabert, 2009; P. scinax Otte et with the weakly shortened apical area; hind Perez-Gelabert, 2009; P. sporadicos Otte et wings are with the rather long or short (but Perez-Gelabert, 2009; P. strongylops Otte distinct) part exposed behind tegmina; anal et Perez-Gelabert, 2009; P. valens Otte et and genital plates of both sexes are simple Perez-Gelabert, 2009) and Selvagryllus (P. (anal plate is rather small and with the foraminatus Saussure, 1878; P. olmecus Sau- rounded apex; male genital plate is much ssure, 1897; P. tibialis Saussure, 1897), (2) longer, and its apex similar to previous one; must be included in the tribe Diatrypini female genital plate is not longer than anal (P. desumptus Otte, 2006; P. eudoxos Otte, plate, transverse, with the roundly truncate 2006; P. exaetos Otte, 2006; P. optivus Otte, apex); male genitalia are with the epiphallus 2006; P. pipizon Otte, 2006), and (3) are partly membranous and with the ectopara- with unknown tribal and generic positions meres developed or strongly reduced (Figs (O. affinis Walker, 1869 from Brazil; P. litu- I: 8–10; IV–VI); distal part of ovipositor is ratus Walker, 1871 from Nicaragua). almost triangular and with the numerous denticles on the lateral edges, numerous Paroecanthus mexicanus Saussure, 1859 small transverse ridges on the hind half of (Figs I: 4, 8–10; II: 1–3; III: 1, 2; V: 1–3) its ventral surface and on the lateral parts of basal half of this surface, and a few larger Orocharis scitulus Walker, 1869 (synonymised by transverse ridges at the middle of this ven- Saussure, 1874). tral surface and at base of this surface (Figs Orocharis eclectos Otte, 2006, syn. nov. II: 3, 7, 10 ), but dorsal surface of ovipositor Material examined. Two males, 1 female; distal part more smooth (only rather long Mexico, Chiapas, environs of Palenque Town and weak transverse ridges at base distinct). near Maya Archaeological Centre, ~ 200 m, sec- Paroecanthus contains the following spe- ondary forest, on leaves, 18–20 Nov. 2006 (1 cies (named in original binomen): P. mexi- male collected as nymph of middle age, imago canus (type species); P. aztecus Saussure, Feb. 2007); coll. A. Gorochov & A. Ovtshin- nikov; ZIN. One male, 3 females; Mexico, Ve- 1874 from Mexico; Orocharis aeschyntelos racruz, 15–20 km NE of Catemaco Town, Los Otte, 2006 and O. habros Otte, 2006 from Tuxtlas (biostation of Mexico University) in 2 Costa Rica; P. oaxaca sp. nov.; P. simplex km from coast of Mexican Gulf, partly primary sp. nov.; P. varius sp. nov.; P. minor sp. nov.; and partly secondary forest on hills, on leaves, P. jalisco sp. nov.; P. selva sp. nov.; possibly 6–17 Nov. 2006 (1 female collected as nymph of also O. signatus Walker, 1869 from Mexico middle age, imago Jan. 2007); coll. A. Gorochov (see Notes after the redescriptions of P. & A. Ovtshinnikov; ZIN. mexicanus and P. aztecus aztecus). Redescription. Male. Colouration light All the other species, included in Paroe- (yellowish), but with following marks: canthus in the electronic catalogue (Eades head with brownish transverse stripe along et al., 2011), evidently or possibly (1) be- clypeal suture, small brown spots under long to the genera Siccotrella gen. nov. eyes, and blackish or dark brown both most (P. guatemalae Saussure, 1874; P. niger part of antennae (scape and sparse spots on Saussure, 1874; P. toltecus Saussure, 1878; flagellum light) and three distal segments P. sulcatus Saussure, 1897), Angustitrella of maxillary palpi; pronotum with very dis- gen. nov. (P. podagrosus Saussure, 1897; tinct black or dark brown border along all P. vicinus Chopard, 1912; P. picipes Bruner, edges (in one male from Chiapas, fore half 1916; P. roosevelti Rehn, 1917; P. hespe- of ventral edge of lateral lobes with yel- rus Hebard, 1924; P. versutus Otte, 2006), lowish border, and in both males from Chi- Ectotrypa (Carsidava cinerascens Walker, apas, pronotum with additional large light

Figs I (1–10). Different structures of body; schematic. 1–3, Siccotrella modesta modesta subsp. nov.; 4, Paroecanthus mexicanus; 5, Angustitrella trivialis sp. nov.; 6, S. mutabilis sp. nov.; 7, Perutrella originalis sp. nov.; 8–10, Paroecanthus mexicanus. Distal part of ovipositor from side (1), and its left half from above (2) and from below (3); regions of stridulatory vein and of oblique veins in stridulatory apparatus of male tegmen (4–7); male genitalia from above (8), from below (9) and from side (10). Abbreviations: e, ectoparamere; ep, epiphallus; g, giding rod; m, membranous processes on both sides of epiphallus or near its apex; r, ramus.

Figs II (1–10). Paroecanthus. 1–3, P. mexicanus; 4, P. aztecus aztecus; 5, P. aztecus tuxtla subsp. nov.; 6, 7, P. minor sp. nov.; 8–10, P. selva sp. nov. Male upper tegmen (1); dorsal field of female tegmina (2) and of male upper tegmen (4, 5); dorsal part of ovipositor from below (3, 7, 10); head with pronotum (upper photo) and tegmina with hind legs (lower photo) from above (6); head in front (8); hind tibia and tarsus from side (9).

Figs III (1–17). Paroecanthus and Siccotrella. 1, 2, P. mexicanus; 3, P. aztecus tuxtla subsp. nov.; 4, P. aztecus verapaz subsp. nov.; 5, 6, P. oaxaca sp. nov.; 7, 8, P. simplex simplex subsp. nov.; 9, P. simplex chamela subsp. nov.; 10, S. varius sp. nov.; 11, P. minor sp. nov.; 12, P. jalisco sp. nov.; 13, P. selva sp. nov.; 14, S. mutabilis sp. nov.; 15, S. sympatrica sp. nov.; 16, S. modesta modesta subsp. nov.; 17, S. managua sp. nov. Male metanotal gland (1, 5, 7, 9, 14–17); outer side of fore tibia (2–4, 6, 8, 10–13).

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 223 brown or reddish brown spot on dorsal photograph (Fig. II: 1), and venation of lat- two-thirds of lateral lobes and light brown eral field with 16–19 branches of Sc (some median area on disc); legs with rather of them more or less oblique and other wide reddish brown subapical (transverse ones somewhat S-shaped) and without and incomplete) band on fore and middle crossveins; hind wings shortly exposed be- femora (fore femora of male from Veracruz hind tegmina. Genitalia with rather short practically without such band), brown or epiphallus (ep), a pair of rather short lateral reddish brown distal third of hind femora, membranous processes (m) near epiphal- a few small marks of same colour on more lic apex, and ectoparameres (e) looking as proximal part of outer surface of hind fem- weakly sclerotised ventral plates narrow- ora, small brown or light brown dorsal spot ing to apex of genitalia and laterally fused on proximal and distal parts of both fore with epiphallus (Figs I: 8–10; V: 1–3). and middle tibiae (but in male from Vera- Female. General appearance as in male cruz, distal spot absent, and in one male including variability in colouration, but from Chiapas, proximal and distal spots of dorsal tegminal field with darkened pattern middle tibiae connected with each other by as in Fig. II: 2 (female from Chiapas lighter, brownish longitudinal line), several black- with darkened parts of antennae brown, ish or dark brown spots on dorsal surface three proximal segments of maxillary palpi of hind tibiae (long spot near their base, light brown, all basitarsi light, second seg- and small spot between each pair of tibial ment of fore and middle tarsi brownish, spines), two blackish or grayish brown ventral edge of pronotal lateral lobes com- (sometimes partly brownish / partly yel- pletely light, and other edges of pronotum lowish) proximal segments of fore and mid- with brown border), pronotum less dis- dle tarsi, and brown second segment and tinctly narrowing to head (its hind part sometimes also ventral part of basitarsus in only slightly wider than pronotal length), hind tarsi; tegmina with rather large black- dorsal tegminal field narrower and with 7 ish or grayish brown spots on dorsal field or 8 partly oblique longitudinal veins and of upper tegmen (Fig. II: 1) and somewhat sparse crossveins, venation of lateral tegmi- smaller and lighter similar spots on lower nal field with 12–14 oblique branches of Sc tegmen (latter spots in male from Veracruz (no S-shaped branches), and genital plate and in one male from Chiapas almost ab- with widely truncate apex. Ovipositor al- sent); abdomen with brown or light brown most completely dark brown (Fig. II: 3). tergites and proximal half of anal plate. Length in mm. Body: male 14–16, female Head rostrum almost twice as wide as scape, 13.5–17; body with wings: male 18–19.5, with very small (much smaller than lateral female 20–23; pronotum: male 2.9–3.1, fe- ocelli) and weakly distinct median ocellus. male 3.1–3.6; tegmina: male 14–15, female Pronotum clearly wider than its length, 14.2–16.5; hind femora: male 9–9.5, female distinctly narrowing to head, with weakly 9.4–9.8; ovipositor 5.8–6.2. angular (almost weakly arcuate) hind edge Notes. This cricket is characterised by of disc; metanotal gland with middle trans- its remarkably contrasting colouration verse process almost truncate at apex (Fig. (somewhat variable, but always with the III: 1). Fore tibiae with tympanal region rather large dark spots on the upper teg- hardly inflated (Fig. III: 2), outer tympa- men in male and both tegmina in female) num slightly longitudinal (almost round), as well as by the wings very slightly short- and inner tympanum moderately long (its ened, branches of longitudinal veins in the distal edge situated at middle of tibia) and female dorsal tegminal field not numerous, not very strongly slit-like (its membrane and male genitalia as in Figs V: 1–3. This visible). Tegmina with apical area weakly species is rather widely distributed in Cen- shortened, stridulatory apparatus as in tral America. It is synonymised with O. sci-

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 224 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI tulus from Honduras and indicated from “le of ventrolateral keel of these femora, and Mexique et l’Amérique centrale” by Sau- brown both spot around each spine of hind ssure (1874), recorded from a few southern tibiae (excepting spurs = apical spines) and states of Mexico by Saussure (1897), and small basal part of this spine; upper tegmen synonymised with O. eclectos from Costa with less spotted dorsal field (Fig. II: 4) and Rica here. Synonymy of O. scitulus and without darkened marks on lateral field; P. mexicanus is probably correct as their lower tegmen practically without spots; original descriptions do not contradict the abdominal sternites, genital plate and cerci material examined; however the larger O. light brown; head rostrum approximately signatus Walker, 1869 from “Orizaba” (Ve- 1.5 times as wide as scape; median ocellus racruz State of Mexico), also synonymised slightly larger, but distinctly smaller than by Saussure (1874) with P. mexicanus, lateral ones; fore tibiae with tympanal re- must be excluded from this synonymy as gion moderately inflated (as in Figs III: 3, it has a few large dark spots on the epicra- 4) and with inner tympanum longer (its nium not characteristic of P. mexicanus, but distal edge situated in distal half of tibia) presented in dark specimens of some larger and strongly slit-like (its membrane almost species of Paroecanthus (see below). Oro- invisible); fore and middle basitarsi slightly charis eclectos, judging by its photographs longer (for comparison see Figs III: 2 and (Otte, 2006), has the general appearance 3); tegmina with distinctly longer apical and male genitalia identical to those of the area (Fig. II: 4); hind wings also longer and specimens studied, but all tibiae of its ho- with distinctly longer hind part exposed lotype are with the long dark longitudinal behind tegmina; genitalia with somewhat line on the dorsal surface (it may be a most longer epiphallus (ep), distinctly larger and dark variant of colouration in this species; rather narrow membranous processes (m) in the specimens described here, this sur- on both sides of epiphallus, and ectoparam- face of tibiae is usually with the shorter eres (ectoparameral sclerites) isolated from dark spots, but one male from Chiapas has epiphallus (Figs IV: 1, 2; V: 4–6). the long darkish line on the dorsal surface Length in mm. Body 21; body with wings of middle tibiae). 29; pronotum 3.5; tegmina 20; hind femora 10.7. Paroecanthus aztecus aztecus Saussure, Notes. The male studied here is with 1874, stat. nov. the label inscription by Saussure’s hand – (Figs II: 4; IV: 1, 2; V: 4–6) “Paroec. aztecus”; it is one of specimens mentioned in the first redescription of this Material examined. One male; Mexico, Vera- species as a typical variant of P. aztecus cruz, “Atoyac, Veracruz, Schumann”; ZIN. (Saussure, 1897). It is impossible to exclude Redescription. Male. Colouration and that insufficiently described P.? signatus structure of body similar to those of P. mexi- (also from Veracruz State) is a synonym of canus, but distinguished by following char- this subspecies. acters: head without darkened stripe along clypeal suture, with completely light maxillary palpi and one brown dot under each Paroecanthus aztecus tuxtla subsp. nov. eye; pronotum with all edges having dark (Figs II: 5; III: 3; IV: 3, 4; V: 7–9) brown border (however this border in hind Holotype. Male; Mexico, Chiapas, environs and posteroventral parts partly grayish), of Tuxtla Gutierrez City, near reserve El Ocote, and with disc and lateral lobes lacking any 600–1000 m, primary forest, on leave, 19–24 large spots; legs yellowish with brown api- May 2006; coll. A. Gorochov & M. Berezin; ZIN. cal and light brown subapical parts of hind Paratype. Female; same data as for holotype; femora, dark brown line along distal third ZIN.

Figs IV (1–12). Paroecanthus; schematic. 1, 2, P. aztecus aztecus; 3, 4, P. aztecus tuxtla subsp. nov.; 5, 6, P. aztecus ocosingo subsp. nov.; 7, 8, P. oaxaca sp. nov.; 9, 10, P. simplex simplex subsp. nov.; 11, 12, P. simplex chamela subsp. nov. Distal half (1, 3, 5, 8–11) and distal part (2, 4, 6, 7, 12) of male genitalia from above (1, 3, 5, 7, 9, 11) and from below (2, 4, 6, 8, 10, 12). Abbreviations: a, anterolateral area of epiphallus; others, as in Figs I: 8–10.

Description. Male (holotype). Coloura- Paroecanthus aztecus ocosingo tion and structure of body very similar to subsp. nov. those of P. aztecus aztecus, but distinguished (Figs IV: 5, 6; V: 10–12) by following features: head without dark Holotype. Male; Mexico, Chiapas, Ocosingo dots under eyes as well as with brown both Distr., Selva Lacandona near border with Guate- stripe along clypeal suture and one dot on mala (between Biosphere reserve Montes Azules each gena near hind corner of subgena; all and Bonampak Natural Monument), environs of borders of pronotum very dark brown (al- Lacanja-Chansayab Vill., primary forest, at light, most blackish); legs with small dark spots at 20–27 May 2007; coll. M. Berezin & E. Tkatshe- apex of fore and middle femora and at base va; ZIN. of fore and middle tibiae, one brown dot on Paratype. Female; same data as for holotype; dorsal surface of fore tibiae, dark brown two ZIN. proximal segments of fore and middle tarsi, Description. Male (holotype). Coloura- tion and structure of body very similar to darkish apical part of distal segment of these those of P. aztecus tuxtla, but stripe along tarsi, dark brown both apex of hind basitar- clypeal suture lighter (light brown), dark sus and second segment of hind tarsus, and dots on genae absent, borders of pronotum somewhat darker (than in P. aztecus aztec- lighter (brown), two proximal segments us) marks on apical part of hind femora and of fore and middle tarsi only weakly dark- on hind tibiae; dorsal field of upper tegmen ened and with yellowish areas, colouration with more distinct spots than in P. aztecus of hind legs almost as in P. aztecus aztecus aztecus (Fig. II: 5); tegminal mirror slightly (however hind basitarsus with weakly shorter than in P. aztecus aztecus (Fig. II: 5); darkened apical part), colouration of tegmi- genitalia with epiphallic anterolateral areas na intermediate between those of P. aztecus (a) distinctly shorter than in P. aztecus az- aztecus and P. aztecus tuxtla, shape of tegmi- tecus and with membranous processes (m) nal mirror almost as in P. aztecus tuxtla, and on both sides of epiphallus (ep) rather wide genitalia with epiphallic anterolateral areas (Figs IV: 3, 4; V: 7–9). (a) much longer than in P. aztecus tuxtla Female. General appearance as in male, (Figs IV: 5, 6; V: 10–12). however with slightly darkened spots on Female. General appearance as in male, head behind eyes, lighter fore and middle however stripe along clypeal suture, dots legs (their tarsi with dorsal half of basitar- on genae, and borders of pronotum as in P. sus light), large darkish median spot on aztecus tuxtla, colouration and structure of disc, colouration and structure of tegmina tegmina as well as shape of genital plate and similar to those of P. mexicanus (but dark of ovipositor very similar to those of female marks smaller and less distinct, tegmina of P. aztecus tuxtla [but colouration of ovi- clearly longer, their dorsal field with 11 or positor light brown with brown ridges on 12 longitudinal veins and less sparse cross- apical (drilling) part of lower valves and veins, and their lateral field with 16 branch- brown lateral parts of rest of these valves es of Sc), and genital plate and ovipositor (see from below)]. almost identical to those of this species. Length in mm. Body: male 18, female 16; Length in mm. Body: male 22, female 17; body with wings: male 27, female 31.5; pro- body with wings: male 31, female 31; prono- notum: male 3.7, female 4; tegmina: male 21, tum: male 4, female 3.8; tegmina: male 23, female 23; hind femora: male 10.8, female female 22; hind femora: male 11.8, female 12.3; ovipositor 7.5. 11; ovipositor 7. Comparison. The new subspecies differs Comparison. Differences between the from P. aztecus aztecus mainly in the dis- new subspecies and P. aztecus aztecus are tinctly wider membranous processes (m) listed above. on both sides of epiphallus (ep), and from P.

Figs V (1–12). Paroecanthus. 1–3, P. mexicanus; 4–6, P. aztecus aztecus; 7–9, P. aztecus tuxtla subsp. nov.; 10–12, P. aztecus ocosingo subsp. nov. Male genitalia (1, 2), their distal half (3, 5, 8, 11) and their distal part (4, 6, 7, 9, 10, 12) from above (1, 5, 8, 11), from below (3, 6, 9, 12) and from side (2, 4, 7, 10).

Figs VI (1–10). Paroecanthus and Siccotrella. 1–3, P. oaxaca sp. nov.; 4–6, P. simplex simplex subsp. nov.; 7–9, P. simplex chamela subsp. nov.; 10, S. xadani sp. nov. Male genitalia (2, 3), their distal part (1, 6, 9) and their distal half (4, 5, 7, 8) from above (1, 4, 7), from below (2, 5, 8) and from side (3, 6, 9); head in front (10).

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 229 aztecus tuxtla, in the much longer epiphallic at base of oblique branches developed); fore anterolateral areas (a) (for comparison see basitarsi weakly (but distinctly) narrower Figs IV: 3 and 5). than in all previous subspecies (for comparison see Figs III: 3 and 4). Paroecanthus aztecus verapaz Male unknown. subsp. nov. Length in mm. Body 16.5; body with (Fig. III: 4) wings 25; pronotum 3.5; tegmina 19; hind femora 12; ovipositor 7.3. Holotype. Female; Guatemala, Vera Paz, Comparion. The new subspecies differs “S. Geronimo, Guatemala, Champion”; ZIN. from all the other subspecies of P. aztecus in Description. Female (holotype). Co- the more spotted fore and middle tibiae as louration and structure of body similar to well as narrower fore basitarsi. those of all previous subspecies of this spe- Notes. This specimen is with the label cies, but with following peculiarities: dorsal inscription by Saussure’s hand – “Paroec. part of head and of pronotum with rather aztecus var. a”; it was mentioned in the large brown areas [largest one (from me- first redescription of this species as “Var. a” dian ocellus to hind edge of vertex) almost (Saussure, 1897). reaching eyes and with light narrow stripes and dots in middle part; two much narrower Paroecanthus oaxaca sp. nov. longitudinal stripes behind each eye; medi- (Figs III: 5, 6; IV: 7, 8; VI: 1–3) an band on pronotal disc fused with somewhat wider spot near anterior edge of disc Holotype. Male; Mexico, Oaxaca, 35 km and with transverse band along hind edge NNE of Santa Cruz Huatulco Town (10 km N of disc (this median band with a few small of Xadani Vill.), 900–1000 m, partly primary and partly secondary forest, on leave, 7–11 May light spots along median line)]; epicranium 2006; coll. A. Gorochov & M. Berezin; ZIN. under rostrum, under eyes and under anten- Paratypes. One male, 2 females; same data as nae as well as upper part of clypeus with a for holotype; ZIN. few small brown spots and brownish dots; Description. Male (holotype). Coloura- antennae more or less yellowish with light tion and structure of body similar to those of brown proximal part (excepting three most P. aztecus. Body yellowish with brown spots basal segments which also yellowish) hav- on head dorsum between eyes, a few dots on ing a few moderately sparse brown rings; epicranium under antennae and under eyes each pronotal lateral lobe with brown both as well as on upper part of clypeus, proxi- rather large anterodorsal longitudinal spot mal part of antennae (excepting three most and a few small marks on rest of this lobe; proximal segments), and numerous marks femora with rather numerous brown dots on pronotum including wide stripe on disc and small spots as well as with dark brown along its hind edge, a few light brown spots apical part; fore and middle tibiae with on tegmina (almost as in P. a. aztecus), nu- brown areas on proximal half, dark brown merous darkish dots on legs (excepting tarsi distal part, and a few dark dots between and hind tibiae), dark brown spots at base these darkenings (Fig. III: 4); hind tibiae and at apex of fore and middle tibiae, dark with rather numerous darkish dots on dor- apical part of hind femora, large darkened sal surface and dark distal part of all other areas on pterothoracic and abdominal dor- surfaces; tarsi more or less light with small sum as well as on pterothoracic pleurites, darkened area at base of fore basitarsus and very numerous small darkened spots (middle tarsi missing) and darkish ventral and dots on lateral parts of abdominal ter- half of second segment of hind tarsus; teg- gites. Structure of head, thorax, legs and mina with almost uniformly yellowish dor- wings very similar to that of P. aztecus, but sal field (only a few small light brown spots metanotal gland with middle transverse

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 230 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI process distinctly trilobate (Fig. III: 5), Description. Male (holotype). Coloura- fore and middle basitarsi slightly shorter tion and structure of body very similar to (for comparison see Figs III: 3, 4 and 6), those of P. oaxaca, but head dorsum with a and shape of tegminal mirror more similar pair of light brown additional longitudinal to that of P. a. aztecus. Abdominal apex as stripes near eyes and large dark brown area in P. mexicanus and P. aztecus, but genitalia along hind edge of epicranium, face with without distinct ectoparameres and mem- brown stripe along clypeal suture and with- branous processes on both sides of epiphal- out any darkened dots around this stripe, lus (ep) as well as with distinctly longer and borders of antennal cavities blackish, pro- rather narrow epiphallus (ventral surface of notum with darkened interrupted stripe genitalia with only a pair of not large mem- along edges of disc and a pair of brown dots branous convexities – possible traces of ec- on disc, darkish dots on legs almost absent toparameres) (Figs IV: 7, 8; VI: 1–3). or (on hind femora) weakly distinct, hind Variations. Paratype slightly lighter, i. tibiae with dark brown area at base and e. with smaller darkened marks including rather small (but distinct) spots around brown stripe on pronotal disc along its hind base of each spine and of both dorsal spurs, edge (this stripe narrow). metanotal gland with middle transverse Female. General appearance as in males, process weakly trilobate (almost rounded) but colouration and structure of tegmina as at apex (Fig. III: 7), fore and middle ba- well as shape of both genital plate and ovi- sitarsi hardly shorter (for comparison see positor very similar to those of P. aztecus Figs III: 6 and 8), genitalia with epiphallus (however colouration of ovipositor almost shorter and with sclerotised ectoparameres as in P. a. ocosingo). developed and widely fused with lateral Length in mm. Body: male 20–22, fe- epiphallic parts (Figs IV: 9, 10; VI: 4–6). male 17–18; body with wings: male 29–31, Variations. Sometimes dorsum of head female 30–31; pronotum: male 3.9–4, fe- lighter or almost uniformly yellowish, epi- male 3.8–4; tegmina: male 21–22, female cranium with darkish spots behind eyes, 22–22.5; hind femora: male 11.8–12.2, fe- and pronotum with larger or smaller dark- male 12–12.4; ovipositor 7–7.2. ened marks. Comparison. The new species differs Female. General appearance as in male, from P. aztecus in the male genitalia lacking but head dorsum in all specimens with dis- sclerotised ectoparameres and membranous tinct dark marks (as in holotype or with ad- processes on both sides of epiphallus; from ditional longitudinal stripes larger and dark P. mexicanus, in the same characters as well brown, or with narrow dark brown area as larger size, longer wings, and somewhat along hind edge of epicranium), stripe along different colouration; from P. aeschyntelos, clypeal suture sometimes weakly distinct, in the much narrower apex of male genitalia; pronotum sometimes with dark brown disc and from P. habros, in the larger size of body having a pair of large light brown areas, co- and darkened apical part of hind femora. louration and structure of tegmina as well as of genital plate and of ovipositor very Paroecanthus simplex sp. nov. similar to those of P. oaxaca. (Figs III: 7, 8; IV: 9, 10; VI: 4–6) Length in mm. Body: male 17–21, female 15–18; body with wings: male 28–31, Holotype. Male; Mexico, Oaxaca, 35 km female 29–31; pronotum: male 3.6–3.9, fe- NNE of Santa Cruz Huatulco Town (10 km N male 3.5–3.9; tegmina: male 19–21, female of Xadani Vill.), 900–1000 m, partly primary and partly secondary forest, on leave, 7–11 May 20–22; hind femora: male 10.8–12.2, female 2006; coll. A. Gorochov & M. Berezin; ZIN. 11.3–12; ovipositor 6.5–7. Paratypes. Five males, 3 females; same data as Comparison. The new species differs for holotype; ZIN. from P. aztecus and P. mexicanus in the male

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 231 genitalia lacking membranous processes on Length in mm. Body: male 18–20, fe- both sides of epiphallus; from P. oaxaca, in male 14–19; body with wings: male 27–29, the presence of rather large dark area at female 27–28; pronotum: male 3.6–3.8, base of hind tibiae, shorter epiphallus, and female 3.7–3.8; tegmina: male 19–20.5, fe- sclerotised ectoparameres developed; from male 19–20; hind femora: male 11–11.5, fe- P. aeschyntelos, in the distinctly shorter male 11–11.3; ovipositor 6.3–6.6. epiphallus; and from P. habros, in the dark- Comparison. The new subspecies is dis- ened apical part of hind femora. tinguished from P. simplex simplex mainly by the clearly trilobate middle transverse Paroecanthus simplex chamela process in male metanotal gland, less mem- subsp. nov. branous epiphallus, distinctly wider rami in (Figs III: 9; IV: 11, 12; VI: 7–9) male genitalia, and darker ovipositor.

Holotype. Male; Mexico, Jalisco, 19°33´N, Paroecanthus varius sp. nov. 105°5´W, biostation of Mexico University near Chamela Vill. (3–4 km from Pacific Ocean), dry (Fig. III: 10) forest on hills, on leave, 23–28 Nov. 2006; coll. A. Holotype. Female; Mexico, Chiapas, 130 km Gorochov & A. Ovtshinnikov; ZIN. WN of Tapachula City, environs of Ejido Las Paratypes. Three males, 2 females; same data Golondrinus Vill. near reserve El Triunfo, 800– as for holotype; ZIN. 1000 m, partly primary and partly secondary for- Description. Male (holotype). Coloura- est, at light, 13–17 May 2006; coll. A. Gorochov tion and structure of body very similar to & M. Berezin; ZIN. those of P. simplex simplex stat. nov., but Paratypes. Four females; same data as for ho- epicranium almost as in most light para- lotype; ZIN. types of P. simplex simplex [yellowish with Description. Female (holotype). Gen- small brown spots (one near median ocellus, eral appearance similar to that of P. aztecus, two on dorsum near hind edge of epicrani- P. oaxaca and P. simplex. Head yellowish um, and two under eyes) and light brown with dark brown triangular spot between stripe along lateral parts of clypeal suture], ocelli, 7 brownish wide longitudinal bands pronotum with only three light brown spots on dorsal half of epicranium behind previ- on its fore part and a pair of lateral brown ous spot and behind eyes (these bands with dots on middle part of disc, darkened areas a few oblique and transverse light lines and on middle legs and at base of hind tibiae narrow stripes), several small brown spots very small and lighter, metanotal gland with on face (under eyes, under antennae and middle transverse process clearly trilobate under rostrum) and on upper part of clyp- (almost as in P. oaxaca; Fig. III: 9), genitalia eus, dark brown proximal part of antennae with epiphallus less membranous in median (excepting three most proximal segments), part and with rami distinctly wider (Figs and light brown other parts of antennae; IV: 11, 12; VI: 7–9). pronotum yellowish with hardly darker Variations. Sometimes dorsum of head disc, brown median marks on disc along its with rather wide brown stripe along its hind anterior edge and along anterior half of its edge, and colouration of legs almost indis- median line, and brownish stripe on each tinguishable from that of P. simplex simplex. lateral lobe along its dorsal edge; legs yel- Female. General appearance as in males, lowish with several darkish dots on middle however colouration of tegmina and their femora and on hind femora, a few small structure as well as shape of both genital brown and brownish spots at apex of all plate and ovipositor very similar to those femora and at base and near base of fore and of P. oaxaca and P. simplex simplex, and co- middle tibiae, with one such spot only near louration of ovipositor almost uniformly base of hind tibiae, and with grayish brown brown (darker than in P. simplex simplex). spots around base of majority of spines in

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 232 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI hind tibiae (these spots partly fused with species in the clearly less inflated tympanal each other dorsally); tegmina yellowish part of fore tibiae (for comparison see Figs with semitransparent membranes of lat- III: 3, 4, 6, 8 and 10). From P. aeschynte- eral field and with small brown or brown- los and P. habros, it differs in the distinctly ish spots at base of each oblique branch more spotted legs. along lateral edge of dorsal field; all other parts of body yellowish (excepting more or Paroecanthus minor sp. nov. less uniformly brown ovipositor). Scape al- (Figs II: 6, 7; III: 11) most 1.5 times as wide as rostrum between antennal cavities; median ocellus much Holotype. Female; Mexico, Chiapas, environs smaller than lateral ocelli. Fore tibiae with of Palenque Town near Maya Archaeological Centre, ~200m, primary forest, on leave, 18–20 tympanal part hardly inflated (Fig. III: 10) Nov. 2006; coll. A. Gorochov & A. Ovtshinnikov; and tympana more or less intermediate be- ZIN. tween those of P. mexicanus and P. aztecus; Description. Female (holotype). Gen- fore and middle basitarsi almost as in P. sim- eral appearance more or less similar to that plex (see Figs III: 8 and 10); spines of hind of P. mexicanus, however colouration with tibiae comparatively long (as in all previous dark brown spot between ocelli, a pair of species). Tegmina with 11–12 longitudinal brown longitudinal bands from medial veins in dorsal field, 16–17 branches of Sc edges of eyes to hind edge of vertex, light in lateral field, and crossveins as in female brown median spot near hind edge of ver- of P. aztecus, P. oaxaca and P. simplex; hind tex, dark brown proximal part of antennae wings distinctly (but not strongly) exposed (excepting three yellowish most proxi- behind tegmina. Shape of genital plate and mal segments) interrupted by sparse light structure of ovipositor similar to those of all spots, brown short part of antennae near previous species. their dark part, light brown rest of anten- Variations. Colouration rather vari- nae interrupted by very sparse darkish able: two females with additional darkened spots, pattern of pronotum only slightly marks (wide brown stripe on disc along its lighter than in P. mexicanus (Fig. II: 6), hind edge; darkish dots on fore femora; more almost uniformly yellowish legs (but hind numerous such dots on other femora; almost femora with brown narrow subapical marks all veins of tegminal dorsal field brownish), and line along distal part of ventrolateral almost completely brown or blackish apical keel, and hind tibiae with darkened apical part of hind femora, and dark brown ovipos- part and small spot at base of spines and of itor (but one of these females almost with- dorsal spurs), brown pattern of tegmina as out dark spots on face and clypeus, except- in Fig. II: 6, and uniformly light brown ovi- ing brownish stripe along clypeal suture); positor (Fig. II: 7). Structure of bodyparts another female almost without brownish (including shape of head rostrum and of bands on head dorsum (excepting bands be- fore tibiae, structure of tympana, length of hind eyes) and with practically uniformly all wings, venation of tegmina, and shape of yellowish dorsal field of tegmina. genital plate and of ovipositor) very similar Male unknown. to that of female of P. mexicanus, but fore Length in mm. Body 14–16; body with and middle basitarsi extremely short, and wings 23–25; pronotum 3.4–3.7; tegmina dorsal field of tegmina with more numer- 17–18.5; hind femora 10.3–10.8; ovipositor ous crossveins (for comparison see Figs II: 6.2–6.4. 2 and 6; III: 2 and 11). Comparison. The new species differs Male unknown. from P. mexicanus in the wider rostrum of Length in mm. Body 11.5; body with head and distinctly shorter darkened part of wings 18; pronotum 3.1; tegmina 14; hind hind femora, and from all the other previous femora 8.5; ovipositor 4.7.

Comparison. The new species differs ish exposed part; other bodyparts (except- from P. mexicanus mainly in the less spotted ing brown ovipositor) uniformly yellowish. colouration of legs (especially in the yel- Rostrum of head narrow (scape almost twice lowish distal half of hind femora including wider than rostrum between antennal cavi- most part of their apical area) and shorter ties); median ocellus much smaller than lat- fore and middle basitarsi. From P. habros eral ocelli. Fore legs shorter than in all pre- and P. aeschyntelos, the new species differs vious species (pronotum 1.1 times as long in the pronotum with the darkened borders, as fore tibia, but in all previous species, this and from all the other congeners, in the ratio 0.8–0.9), with slightly inflated tym- same characters as from P. mexicanus as well panal part (Fig. III: 12) and with tympana as smaller size of body and (from majority of similar to those of P. aztecus, P. oaxaca, P. them) less inflated fore tibiae. simplex and P. varius (right fore leg damaged in nymphal stage – it shorter than left one Paroecanthus jalisco sp. nov. and with partly reduced tympana); fore and (Fig. III: 12) middle basitarsi almost as short as in P. mi- nor (see Figs III: 11 and 12); hind tibiae with Holotype. Female; Mexico, Jalisco, 19°33´N, rather short spines (length of longest spine 105°5´W, biostation of Mexico University near 0.7–0.8 mm, but in previous congeners, ex- Chamela Vill. (in 3–4 km from Pacific Ocean), cepting only smallest P. minor having similar dry forest on hills, on leave, 23–28 Nov. 2006; length of this spine, this length 1–1.2 mm). coll. A. Gorochov & A. Ovtshinnikov; ZIN. Tegmina with 13 longitudinal veins in dor- Paratype. Female; same data as for holotype; ZIN. sal field and 17–18 branches of Sc in lateral Description. Female (holotype). General field; hind wings strongly exposed behind appearance as in P. oaxaca and P. simplex. tegmina. Shape of genital plate and struc- Head yellowish with a few small brown spots ture of ovipositor similar to those of all pre- on epicranium under eyes and under anten- vious species. nae, light brown dot near hind corner of each Variations. Paratype with dorsal part subgena and stripe along clypeal suture, two of head hardly darkened (intermediate be- brownish longitudinal bands behind each tween yellowish and light brown) and that of eye, small darkish spots on dorsum along its pronotum almost light brown as well as with hind edge, and colouration of antennae as in distinct brown longitudinal band on upper P. oaxaca and P. simplex; colouration of pro- half of pronotal lateral lobes (this band al- notum similar to that of P. simplex chamela; most reaching hind edge of these lobes). legs yellowish with several darkish dots Male unknown. on all femora, small brown spots at apex of Length in mm. Body 14–15; body with fore and middle femora as well as at base of wings 25–27; pronotum 3.6–3.8; tegmina all tibiae (including somewhat larger sub- 17.5–19; hind femora 10.2–11.3; ovipositor basal spot on dorsal surface of fore tibiae), 5.3–5.7. dark brown apical part of hind femora, light Comparison. The new species differs brown wide transverse band behind middle from all the other congeners in the relative- of these femora, brownish apical part of all ly shorter fore legs and spines of hind tibiae, tibiae, and grayish brown tarsi (but with presence of wide light brown band behind light both proximal half of third segment of the middle of hind femora, and (from majority of them) narrow rostrum of head. all tarsi and area on dorsal and lateral surfaces of hind basitarsus); tegmina yellowish Paroecanthus selva sp. nov. with dorsal field having light brown vena- (Figs II: 8–10; III: 13) tion and brown short proximal parts of a few branches situated along lateral edge of Holotype. Male; Mexico, Chiapas, Ocosingo this field; hind wings transparent with gray- Distr., Selva Lacandona near border with Guate-

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 234 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI mala (between Biosphere reserve Montes Azules Length in mm. Body 17; body with and Bonampak Natural Monument), environs of wings 26.5; pronotum 4.2; tegmina 18.5; Lacanja-Chansayab Vill., primary forest, at light, hind femora 10; ovipositor 6. 20–27 May 2007; coll. M. Berezin & E. Tkatshe- Comparison. The new species is most va; ZIN. similar to P. jalisco, but distinctly differs Description. Male (holotype). Structure from it and from all the other congeners in of body similar to that of P. jalisco, but co- the rather dark (reddish brown) coloura- louration very different, reddish brown tion of body, much more dense crossveins in (rather dark) with following marks: head the dorsal field of tegmina, and presence of (Fig. II: 8) with dark brown genae and area numerous crossveins in their lateral field. behind each eye (latter area with narrow longitudinal light brown mark), yellowish antennae (scapes with a few brown spots) Genus Siccotrella gen. nov. and a pair of narrow longitudinal stripes Type species: Siccotrella mutabilis sp. nov. running along medial edge of eyes from up- Diagnosis. Crickets with general appear- per corner of antennal cavities to hind edge ance more or less similar to that of Paroe- of vertex, blackish maxillary palpi, and dark canthus, but more stocky. Head rather short, brown labial palpi; pronotum with dark with roundly angular (in profile) rostrum brown lateral lobes; legs with light brown approximately 1.5 times as wide as scape; third segment of all tarsi, yellowish hind ocelli situated in shape of triangle; median tibiae having darkish apical part, and dark- ocellus extremely small; lateral ocelli not ish two proximal segments of hind tarsi; large, but distinctly larger than median one. tegmina uniformly reddish brown with only Pronotum not very long (clearly wider than slightly lighter distal area; exposed part of its length) and distinctly (but not strongly) hind wings with colouration similar to that narrowing to head; male metanotal gland of distal tegminal area; ovipositor brown more or less similar to that of Paroecan- with two slightly darker transverse ridges thus, however with somewhat lower middle on ventral surface of ventral valves. Struc- process (Figs III: 14–17). Fore tibiae not ture of head rostrum and of ocelli similar inflated; outer tympanum not large, oval to that of P. jalisco. Pronotum 1.1 times (opened) and not immersed or hardly im- as long as fore tibia (as in P. jalisco). Fore mersed; inner tympanum absent (only short tibiae moderately inflated (Fig. III: 13) and longitudinal concavity on inner surface of with tympana as in P. aztecus, P. oaxaca, fore tibia presented) or strongly reduced (i. P. simplex, P. varius and P. jalisco; fore and e. very small, narrow, opened, and hardly or middle basitarsi similar to those of P. minor not immersed); hind legs somewhat shorter and P. jalisco (see Figs III: 11, 12 and 13); than in Paroecanthus. Tegmina with moder- hind tibiae with spines somewhat shorter ately shortened apical area (this area shorter than in P. jalisco (length of longest one 0.6 than in all known species of Paroecanthus; mm) (Fig. II: 9). Tegmina with dorsal field Figs VII: 1–6); hind wings also shortened – having 11–12 longitudinal veins and rather with apical part not exposed or very weakly dense crossveins (distinctly more numer- exposed behind tegmina. Abdominal apex ous than in all previous congeners), and similar to that of Paroecanthus (in both with lateral field having 15–16 branches of sexes, including ovipositor; Figs I: 1–3), Sc and numerous weak crossveins between but male genitalia lacking ectoparameres many of them; hind wings strongly exposed and with epiphallus more or less completely behind tegmina. Shape of genital plate and sclerotised and having a pair of dorsal finger- of ovipositor (Fig. II: 10) similar to that of like processes (Figs VIII: 1–15), and distal female of all previous congeners. part of ovipositor sometimes almost without Male unknown. large transverse ridges (S. xadani sp. nov.).

Figs VII (1–6). Siccotrella. 1, S. mutabilis sp. nov.; 2, S. sympatrica sp. nov.; 3, S. modesta modesta subsp. nov.; 4, S. modesta palenque subsp. nov.; 5, S. managua sp. nov.; 6, S. xadani sp. nov. Male upper tegmen (2–5) and its dorsal field (1); female tegmina from above (6).

Figs VIII (1–15). Siccotrella. 1–3, S. mutabilis sp. nov.; 4–6, S. sympatrica sp. nov.; 7–9, S. modesta modesta subsp. nov.; 10–12, S. modesta palenque subsp. nov.; 13–15, S. managua sp. nov. Male genitalia from above (1, 4, 7, 10, 13), from below (2, 5, 8, 11, 14) and from side (3, 6, 9, 12, 15).

Included species (named in original bi- rior half of genital plate dark brown; cerci, nomen). S. mutabilis sp. nov. (type species); paraprocts and distal half of genital plate Paroecanthus guatemalae Saussure, 1874 light brown. Median ocellus situated on from Guatemala; P. niger Saussure, 1874 hind edge of short and narrow median con- from Guatemala; P. toltecus Saussure, 1878 cavity of rostral dorsum. Hind edge of pro- from Mexico; P. sulcatus Saussure, 1897 notal disc weakly convex; metanotal gland from Guatemala; S. sympatrica sp. nov.; S. with comparatively wide middle process modesta sp. nov.; S. managua sp. nov.; S. truncate at anteromedian edge and having xadani sp. nov. a pair of rather distinct anterolateral pro- Comparison. The new genus is most re- jections (Fig. III: 14). Fore tibiae with very lated to the genus Paroecanthus, but well small and narrow inner tympanum which distinguished from it by the inner tympa- opened and hardly immersed; hind tibiae num absent or strongly reduced, hind legs with rather long spines (length of longest somewhat shorter, wings more shortened, one ~1 mm). Tegmina with dorsal field as and male genitalia lacking ectoparameres in photograph ( Fig. VII: 1) and with lat- and with the epiphallus more or less com- eral field having 20–21 branches of Sc more pletely sclerotised and having a pair of dor- or less similar to those of Paroecanthus in sal processes. shape; crossveins between these branches Etymology. The generic name is origi- absent; hind wings almost reaching apex of nated from siccus (Lat.) – stocky and the tegmina. Genitalia as in Figs VIII: 1–3. genus Sonotrella. Variations. Paratypes with rather variable colouration: dark parts in most dark Siccotrella mutabilis sp. nov. specimens almost black; but in most light (Figs I: 6; III: 14; VII: 1; VIII: 1–3) specimens, epicranium brown with almost light brown its lower parts, pronotum uni- Holotype. Male; Mexico, Chiapas, 130 km formly brown, legs almost completely light WN of Tapachula City, environs of Ejido Las brown, and thoracic sternites brown. Golondrinus Vill. near reserve El Triunfo, 800– Female. General appearance as in most 1000 m, partly primary and partly secondary forest, on leave, 13–17 May 2006; coll. A. Gorochov light males of this species, however wings & M. Berezin; ZIN. with only one tegminal vein (R) brown, area Paratypes. Six males, 1 female; same data as between tegminal Sc and R somewhat wider, for holotype; ZIN. dorsal tegminal field having 10 longitudinal Description. Male (holotype). Head dark veins and moderately sparse crossveins, lat- brown with light brown antennae (their eral tegminal field having 12–13 branches middle and distal parts almost yellowish) of Sc (more oblique and not S-shaped), and and mouthparts (upper part of clypeus and hind wings hardly exposed behind tegmina. small area at base of mandibles brown); Genital plate and ovipositor almost yellow- pronotum dark brown with brown disc; all ish with brownish drilling ridges on distal coxae and femora dark brown with almost part of ovipositor as well as with brown- light brown both apical part of hind femora ish apex and lateral border of distal part of and very small marks at apex of fore and ovipositor dorsal valves; structure of distal middle femora; all tibiae and tarsi uniformly part of ovipositor almost as in Fig. I: 1–3. light brown; tegmina uniformly yellowish, Length in mm. Body: male 14–15, female but with brown R, M and most part of Sc 13; body with wings: male 17–18, female 18; (excepting its branches); hind wings almost pronotum: male 3.1–3.4, female 3.5; tegmi- whitish; dorsum of pterothorax and of ab- na: male 12.5–13, female 12.8; hind femora: domen light brown with numerous brown male 10.5–11.2, female 11.3; ovipositor 5.8. and dark brown spots on abdomen; rest of Comparison. The new species is distin- abdominal tergites, all sternites and ante- guished from S. guatemalae comb. nov. by

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 238 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI the more uniform colouration of head, pro- tibiae; tegmina yellowish with dark brown notum and tegmina as well as by the longer M and brown other veins excepting light hind wings; from S. niger comb. nov., by the brown both middle part of Sc and majority uniformly light antennae and not dark tibi- of veins in stridulatory apparatus of lower ae as well as longer hind wings; from S. tolte- tegmen; hind wings whitish; other parts ca comb. nov., by the distinctly narrower of body dark brown with brown thoracic areas between proximal parts of oblique sternites, yellowish both metanotal gland branches in the middle part of dorsal tegmi- and numerous small spots on abdominal nal field in female and uniform colouration dorsum, more sparse light brown spots on of this field; and from S. sulcata comb. nov., abdominal sternites and genital plate, and by the more uniform colouration of male yellowish grey anal plate, paraprocts and tegmina and less transverse vein in mirror cerci (distal part of cerci yellowish). Ocelli of these tegmina. similar to those of S. mutabilis in shape, but median concavity on rostral dorsum longer Siccotrella sympatrica sp. nov. (reaching rostral apex). Structure of both (Figs III: 15; VII: 2; VIII: 4–6) pronotum and metanotal gland also similar to that of S. mutabilis, but middle process Holotype. Male; Mexico, Chiapas, 130 km of this gland distinctly narrower, clearly WN of Tapachula City, environs of Ejido Las rounded at anterior edge and with lateral Golondrinus Vill. near reserve El Triunfo, 800– (not anterolateral) projections (Fig. III: 1000 m, partly primary and partly secondary forest, on leave, 13–17 May 2006; coll. A. Gorochov 15). Inner tympanum absent (only short & M. Berezin; ZIN. longitudinal concavity presented); spines Paratype. Female; same data as for holotype; of hind tibiae somewhat shorter than in S. ZIN. mutabilis (length of longest one ~0.7 mm). Description. Male (holotype). Colou- Tegmina with dorsal field as in photograph ration of body light brown with following (Fig. VII: 2) and with lateral field having marks: dorsum of head hardly darker and 19–20 branches of Sc similar to those of S. with small darkish spot behind each ocellus, mutabilis in shape; hind wings distinctly a pair of yellowish longitudinal stripes along not reaching abdominal and tegminal api- medial edges of antennal cavities and of ces. Abdominal apex also similar to that of eyes (crossing lateral ocelli) as well as a pair this species, but genitalia distinguished by of similar stripes behind eyes, brown small epiphallus having wider distal part (Fig. marks near eyes around latter stripes, dark VIII: 4), more simple (not bulbous) apical brown proximal part of antennae (excepting part (Fig. VIII: 6) and less proximal posi- three most proximal segments), and yellow- tion of dorsal (finger-like) processes (Figs ish distal half of antennae; pronotum with VIII: 4, 6), and by distinctly shorter sclero- slightly darker anterior border and spots on tised longitudinal ribbons on both sides of disc, dark brown upper half of lateral lobes, guiding rod (Fig. VIII: 5). and brown ventral borders; legs with sparse Female unknown. darkish dots on all femora, dark brown Length in mm. Body 11.5; body with spots on apical part of hind femora, weakly wings 13.8; pronotum 2.6; tegmina 9.8; hind darkened fore and middle tibiae and basal femora 8.2. part of hind tibiae, brown second segment Comparison. The new species is most of fore and middle tarsi as well as most part similar in general appearance to S. guatema- of dorsal half of hind tibiae, and dark brown lae and S. sulcata, but it differs from them spots around bases of spines and of spurs in in the absence of dark spots on the tegminal hind tibiae (these spots partly fused with membranes. From all the other congeners, each other distally) as well as apical part of S. sympatrica differs in the more variegate hind basitarsi and rest of apical part in hind colouration and clearly smaller body.

Siccotrella modesta sp. nov. not darker than other pronotal parts, teg- (Figs I: 1–3; III: 16; VII: 3; VIII: 7–9) mina with all veins brown and with dorsal field having 9–10 longitudinal veins and Holotype. Male; Mexico, Chiapas, Ocosingo moderately sparse crossveins, tegminal Sc Distr., Selva Lacandona near border with Guate- mala (between Biosphere reserve Montes Azules with 11–12 oblique branches, width of area and Bonampak Natural Monument), environs of between tegminal Sc and R as well as shape Lacanja-Chansayab Vill., primary forest, 20–27 of genital plate and of ovipositor as in female May 2007; coll. M. Berezin & E. Tkatsheva; ZIN. of S. mutabilis (Figs I: 1–3), but coloura- Paratype. Female; same data as for holotype; tion of ovipositor darker (dark brown with ZIN. brown ventral median band between base of Description. Male (holotype). Coloura- ovipositor and base of its drilling part). tion of body almost uniformly light brown, Length in mm. Body: male 14.5, female however proximal part of antennae brown 14.5; body with wings: male 16, female 15; (excepting two most proximal segments pronotum: male 2.9, female 3.1; tegmina: which light brown), ventral edges of pro- male 12, female 10.8; hind femora: male 8.9, notum also brown, hind tibiae with gray- female 10; ovipositor 6. ish brown spots around bases of spines and Comparison. The new species is most spurs (these spots in distal part of tibia part- similar to S. sympatrica in the shape of male ly fused with each other dorsally), tegmina genitalia, but distinguished by the more almost yellowish with dark brown nar- uniform colouration and characters of male row stripe along R and brown other veins genitalia listed in the description. From S. (but in lower tegmen, majority of veins of mutabilis, the new species differs in the dif- stridulatory apparatus light brown), hind ferent shape of epiphallic apex (not bulbous wings whitish, and abdomen with small in profile); from S. niger, in the distinctly brown marks on tergites and weakly dark- lighter legs; and from S. tolteca, S. sulcata ened spots on sternites, on basal part of anal and S. guatemalae, in the absence of dark- plate and on apical part of genital plate. ened marks on tegminal membranes. Structure of ocelli, of head rostrum and of middle process of metanotal gland almost Siccotrella modesta palenque subsp. nov. as in S. sympatrica (however this process (Figs VII: 4; VIII: 10–12) somewhat wider and with less distinct lateral projections; Fig. III: 16). Legs as in S. Holotype. Male; Mexico, Chiapas, environs of sympatrica, but spines of hind tibiae hardly Palenque Town near Maya Archaeological Cen- longer (length of longest one ~0.8 mm). tre, ~200 m, primary forest, on leave, 18–20 Nov. Tegmina similar to those of both previous 2006; coll. A. Gorochov & A. Ovtshinnikov; ZIN. species, but their lateral field with 17–19 Description. Male (holotype). Coloura- branches of Sc which clearly more S-shaped tion and structure of body very similar to (especially in distal half of tegmen; Fig. VII: those of S. modesta modesta stat. nov., but 3); hind wings intermediate between those with following differences: in new sub- of S. mutabilis and S. sympatrica in length. species, abdomen almost uniformly light Abdominal apex (including genitalia) very brown, middle process of metanotal gland similar to that of S. sympatrica, however less rounded (roundly truncate) at ante- genitalia with: longitudinal sclerotised rib- rior edge, distal branches of tegminal Sc bons on both sides of guiding rod somewhat somewhat less S-shaped (Fig. VII: 4), and longer, proximal part of rami distinctly wid- genitalia with anterior (narrow) part of er (if to see from above), and distal part of rami distinctly shorter (this part slightly rami clearly bifurcated (Figs VIII: 7–9). longer than widened hind part of rami, Female. General appearance as in male, but in S. modesta modesta, narrow anterior however ventral edges of pronotum almost part of rami significantly longer than their

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 240 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI widened hind part; for comparison see Figs previous congeners, but distinctly smaller VIII: 7–9, 10–12). than lateral ocelli; median concavity on Female unknown. rostral dorsum moderately long, running Length in mm. Body 13.5; body with from median ocellus to almost rostral apex. wings 15.5; pronotum 2.7; tegmina 11.3; Thorax similar to that of S. mutabilis, how- hind femora 8.8. ever middle process of metanotal gland ap- Comparison. Differences of the new sub- proximately as in S. modesta in width and species from nominotypical one are listed in with anterolateral projections and almost the description. angular anteromedian concavity (if to see from above) (Fig. III: 17). Legs similar to Siccotrella managua sp. nov. those of S. sympatrica and S. modesta, but (Figs III: 17; VII: 5; VIII: 13–15) length of longest spine of hind tibiae ~0.9 mm. Dorsal tegminal field as in Fig. VII: 5; Holotype. Male; Nicaragua, environs of Ma- lateral tegminal field with 19–20 branches nagua City, 27–28 Jan. 1985; coll. L. Medvedev; of Sc curved as in S. mutabilis and S. sympat- ZIN. rica; length of hind wings approximately as Description. Male (holotype). Coloura- in S. modesta. Abdominal apex also similar tion of body very light brown with follow- to that of previous congeners, however gen- ing pattern: dorsum of head with darkish italia with longer and almost S-shaped (in V-shaped spot between ocelli, a pair of yel- profile) apical part of epiphallus, dorsal fin- lowish stripes along medial parts of anten- ger-like epiphallic processes situated as in nal cavities (separated from these cavities S. mutabilis (but slightly longer), and proxi- by brown lines) and of eyes (these stripes mal part of rami widened approximately as running from rostral apex to hind edge of in S. modesta (Figs VIII: 13–15). vertex), and a pair of shorter longitudinal Female unknown. yellowish stripes behind eyes; rest of head Length in mm. Body 16; body with yellowish with brown proximal part of an- wings 17; pronotum 3; tegmina 12.5; hind tennae (excepting three most proximal seg- femora 10.3. ments which light brown); pronotum yel- Comparison. The new species differs lowish with light brown both median band from S. mutabilis, S. sympatrica and S. mod- on anterior half of disc and wide median esta in the longer and almost S-shaped (in area on hind half of disc, brown stripe on lat- profile) apical part of epiphallus; from S. eral lobes along their dorsal edge, and small niger, in the light legs; and from S. sulcata, darkish spot on anterior part of these lobes S. guatemalae and S. tolteca, in the absence under above-mentioned brown stripe; legs of darkened marks on tegminal membranes. with small brown marks on apical part of hind femora, two rather small darkish areas Siccotrella xadani sp. nov. on dorsoproximal part of hind tibiae, brown (Figs VI: 10; VII: 6; IX: 17) spots around spines and spurs of these tibiae Holotype. Female; Mexico, Oaxaca, 35 km (distal spots partly fused with each other NNE of Santa Cruz Huatulco Town (10 km N dorsally), and darkish ventral part of hind of Xadani Vill.), 900–1000 m, partly primary basitarsi; tegmina yellowish with brown R, and partly secondary forest, on leave, 7–11 May narrow stripe along M, two chords in dor- 2006; coll. A. Gorochov & M. Berezin; ZIN. sal field and some veins in basal area of this Description. Female (holotype). Epi- field, and light brown rest of veins; hind cranium dark brown with yellowish face, a wings whitish; abdomen yellowish with nu- pair of stripes from rostral apex to lateral merous darkish dots on abdominal tergites ocelli (separated from antennal cavities and brown median stripe on eighth sternite. only by brown lines), short stripes along Median ocellus slightly larger than in all medial edges of eyes, and lower parts of ge-

Figs IX (1–23). Different structures of body. 1–4, Angustitrella maculata sp. nov.; 5, 6, A. andensis sp. nov.; 7–9, A. trivialis sp. nov.; 10, A. columbia sp. nov.; 11, A.? borealis sp. nov.; 12, A. podagrosa; 13, 14, A. roosevelti; 15, 16, A. vicina; 17, Siccotrella xadani sp. nov.; 18–20, Ectotrypa veracruz sp. nov.; 21–23, Selvagryllus eltriunfo sp. nov. Male metanotal gland (1, 7, 10, 11); partly dorsal and partly inner view of fore tibia (2, 13); outer (3, 6, 16) and inner (5, 8, 12, 15, 19, 22) sides of fore tibia; outer tympanum (14, 20, 23); distal part of ovipositor from below (4, 9, 18, 21) and from above (17). [12, after Saussure (1897); 13, 14, after Rehn (1917); 15, 16, after Chopard (1912).]

Figs X (1–8). Angustitrella and Ectotrypa; male. 1, A. maculata sp. nov.; 2, A. trivialis sp. nov.; 3, A. columbia sp. nov.; 4, A.? borealis sp. nov.; 5, A. podagrosa; 6, A. vicina; 7, A. roosevelti; 8, E.? fallax (Sauss.). Tegmen (1, 2, 4) and its dorsal field (3, 5–8). [5, after Saussure (1897); 6, after Chopard (1912); 7, after Rehn (1917); 8, after Saussure (1874).]

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 243 nae, two light brown longitudinal stripes Comparison. The new species is most behind each eyes, and reddish brown area similar to S. tolteca, but distinguished from between ocelli, reversed V-shaped spot un- it by the dark brown pronotum and most der rostral apex (reaching median ocellus part of epicranium, and yellowish rest of and fused with previous area near this ocel- head (in S. tolteca, head and pronotum al- lus) and small marks under eyes and under most uniformly reddish brown, not dark, antennal cavities; antennae and mouthparts but with darkish marks on lateral lobes of almost completely yellowish; pronotum pronotum near its disc). It is impossible to almost uniformly dark brown; legs light exclude that S. xadani is only a subspecies brown with slightly darkened small areas of S. tolteca. at apex of hind femora, small darkish subbasal spot on dorsal surface of hind tibiae, Genus Angustitrella gen. nov. and hardly darkened rings around spines of these tibiae; tegmina light brown with Type species: Angustitrella maculata slightly lighter (yellowish) venation and sp. nov. narrow brown stripes along both sides of Diagnosis. Crickets with rather narrow some longitudinal veins in dorsal field (Fig. body (including head). Head slightly elon- VII: 6); hind wings whitish with very light gate and with roundly angular (in profile) brown apical part; rest of body more or less rostrum approximately 1.5 times as wide yellowish with dark brown ovipositor. Me- as scape; median ocellus indistinct; lateral dian ocellus extremely small, much smaller ocelli moderately large and weakly con- than lateral ocelli; rostral dorsum with very vex. Pronotum elongate (in male, it slightly short median concavity near median ocel- wider than its length; in female, it approxi- lus. Pronotum with more convex hind edge mately as wide as long), clearly narrowing of disc than in all previous congeners (Fig. to head in male, and with almost parallel VII: 6). Legs similar to those of S. sympat- lateral sides in female; metanotal gland with rica, S. modesta and S. managua, but length rather diverse middle process, but this gland of longest spine of hind tibiae as in S. mu- distinguished from that of Paroecanthus and tabilis. Tegmina with 9–10 longitudinal Siccotrella by distinctly longer (wider) areas veins in dorsal field (these veins in distal between two most hind fold of metanotum half of tegmen with more transverse proxi- (Figs IX: 1, 7, 10, 11). Fore tibiae moderate- mal parts, elongate cells between them, and ly or strongly inflated; outer tympanum not rather dense crossveins; Fig. VII: 6), 14–15 large, oval (opened) and hardly immersed; oblique branches of Sc in lateral field, and inner tympanum long, slit-like or almost area between Sc and R as in females of other slite-like (Figs IX: 2, 3, 5, 6, 8, 12–16); hind previous congeners; hind wings very weakly legs almost as in Paroecanthus. Tegmina exposed behind tegmina. Structure of ab- with rather long apical area and with mir- dominal apex also similar to that of these ror similar to that of some representatives of species, but genital plate with more round- Paroecanthus and Siccotrella (longitudinal, ed apex, and ovipositor distinguished from but not narrow; Figs X: 1–7); hind wings all congeners described above by somewhat not shortened, with hind part distinctly less distinct transverse ridges on ventral exposed behind tegmina. Abdominal apex surface of distal part and smaller denticles also similar to that of Paroecanthus and Sic- on lateral edge of distal part of dorsal valves cotrella, however male genital plate shorter (Fig. IX: 17). (moderately longer than anal plate), female Male unknown. genital plate with almost rounded or widely Length in mm. Body 20; body with truncate apex, and male genitalia with more wings 21; pronotum 3.8; tegmina 15; hind or less completely sclerotised epiphallus femora 11; ovipositor 6. lacking processes on dorsal part and with

Figs XII (1–8). Angustitrella; schematic. 1, 2, A. maculata sp. nov.; 3, 4, A. trivialis sp. nov.; 5, 6, A. columbia sp. nov.; 7, 8, A.? borealis sp. nov. Epiphallus with ectoparameres and guiding rod from above (1, 3, 5, 7) and from below (2, 4, 6, 8). Abbreviations: lo, posteromedial semimembranous lob- ule; others, as in Figs I: 8–10. ectoparameres developed (Figs XI: 1–18). X: 5 and 6) and are described from the dif- Included species (named in original bi- ferent regions, and their male genitalia are nomen). Angustitrella maculata sp. nov. not studied. So, synonymy of two these spe- (type species); Paroecanthus podagrosus cies is in need of examination. Angustitrella Saussure, 1897 from Guatemala and Pana- picipes, judging by Bruner (1916), has the ma; P. vicinus Chopard, 1912 from Guyana; fore tibiae similar to those of A. vicinus, but P. picipes Bruner, 1916 from Bolivia; P. roo- S. niger, judging by the original description sevelti Rehn, 1917 from Guyana; P. hesperus (Saussure, 1874), lacks inner tympanum. Hebard, 1924 from Ecuador; A. andensis This difference is of a generic level. sp. nov.; A. trivialis sp. nov.; A. columbia Two species from Trinidad mentioned sp. nov.; possibly P. versutus Otte, 2006 above were very briefly described in combi- from Costa Rica, “Bofana” mirifica Otte et nation with the new generic name “Bofana” Perez-Gelabert, 2009 and “B.” rufa Otte et (Otte & Perez-Gelabert, 2009), but this Perez-Gelabert, 2009 from Trinidad I., and name is unavailable because its authors did A.? borealis sp. nov. not fulfil the following articles of the Code: Angustitrella vicina comb. nov. and 13.1 – this article requests the presence of A. picipes comb. nov. were synonymised description, diagnosis or reference to one by Chopard (1968) with A. podagrosa of them for every new scientific name of comb. nov. and Siccotrella niger, respec- taxon after 1930, if it is not a new replace- tively. However A. vicina and A. podagrosa, ment name (however the authors did not judging by Saussure (1897) and Chopard give any description, diagnosis or refer- (1912), have some differences in shape of ence for their “new genus”, and they did tegminal mirror (for comparison see Figs not indicated its name as replacement one);

Figs XI (1–18). Angustitrella. 1–5, A. maculata sp. nov.; 6–10, A. trivialis sp. nov.; 11–15, A. columbia sp. nov.; 16–18, A.? borealis sp. nov. Male genitalia (1, 2, 5–7, 10–12, 15–18) and their distal part (3, 4, 8, 9, 13, 14) from above (1, 6, 11, 16), from below (2, 7, 12, 17), from side (5, 10, 15, 18), from above and slightly behind (3) and from opposite side (4), from behind (8) and from opposite side (9), and from below and slightly behind (13) and from opposite side (14).

Figs XIII (1–7). Ectotrypa; female. 1–3, E. veracruz sp. nov.; 4, 5, E. olmeca; 6, 7, E. brevis. Body from above (1, 4, 6); head in front (2); head with pronotum from side (3, 5, 7). [4, 5, after Saussure (1878); 6, 7, after Rehn (1905).]

13.4 – this article consider that united de- light brown rest of mouthparts (except- scription of genus and species is possible, if ing brown labium and maxillary palpi, but this genus is monotypical, and if a concrete these palpi with almost light brown spots) indication about such kind of description and sparse rings on antennal flagellum; pro- is given (however the authors included in notum brown with a pair of reddish brown their “new genus” three species simultane- spots on disc and dark brown lateral lobes; ously, and they did not give any indication pterothorax with light brown dorsum, about such kind of description). Also these brown lateral sides and intermediate (be- authors indicated Paroecanthus olmecus as tween brown and light brown) colouration the type species of “Bofana” as well as of of sternites; legs yellowish with dark brown the genus Paroecanthus. Later Otte (2009) distal third of femora, brown long subbasal corrected the latter mistake and indicated spot on dorsal part of tibiae (in fore tibiae, “B.” mirifica as a type species of “Bofana”, this spot occupying areas around tympana but he gave neither generic description nor also; Figs IX: 2, 3), darkish apical part of fore diagnosis again. However this action does and middle tibiae as well as two proximal not make the latter name available because segments of fore and middle tarsi, brown the above-mentioned articles of the Code spots around spines and spurs of hind tibiae are not fulfilled as in Otte & Perez-Gelabert as well as apical part of hind basitarsi and (2009). So, at present “Bofana” is still an most part of second segment of hind tarsi, unavailable name. and darkish distal part of third segment Comparison. The new genus is related to of all tarsi; tegmina almost transparent, the genera Paroecanthus and Siccotrella, but but with partly whitish and partly grayish clearly distinguished from them by narrow- venation as well as with very light brown er both head and pronotum (latter one espe- membranes in dorsal field of upper tegmen; cially narrow in female), fore tibiae strongly hind wings whitish; abdomen brown with inflated, inner tympanum large and slit-like light brown majority of sternites and base or almost slit-like (from only Siccotrella), of cerci as well as with dark brown genital wings not shortened (also from Siccotrella), plate and paraprocts. Head and pronotum male genitalia with the epiphallus more or weakly pubescent (not shining) and not less completely sclerotised (from Paroecan- rugous; rostral dorsum with slight narrow thus) and with the ectoparameres developed and rather short median concavity; prono- (from Siccotrella). tal disc without distinct concavities; lat- Etymology. The generic name is originat- eral lobes of pronotum slightly crumpled ed from angustus (Lat.) – narrow and the (not flat); metanotal gland as in Fig. IX: 1. genus Sonotrella. Fore tibiae moderately inflated (Figs IX: 2, 3); inner tympanum almost slit-like (with Angustitrella maculata sp. nov. well visible membrane), and distal edge (Figs IX: 1–4; X: 1; XI: 1–5; XII: 1, 2) of this tympanum situated near middle of tibia; spines of hind tibiae moderately short Holotype. Male; Ecuador, western slopes of (length of longest one ~0.6 mm). Tegmina Andes Mts, 10 km E of Agua Blanca Vill. (vil- as in Fig. X: 1; hind wings reaching base of lage in environs of Puerto Lopez Town), San distal third of hind tibiae. Genitalia with Sebastian Forest, 700 m, cloud primary forest, on leave, 26–29 Oct. 2005; coll. A. Gorochov & rather short epiphallus having widened an- A. Ovtshinnikov; ZIN. terior part, rounded hind part and distinct Paratypes. Three females; same data as for median notch at anterior edge; ectoparam- holotype; ZIN. eres in shape of rather narrow arcuate plate Description. Male (holotype). Head almost fused with epiphallus laterally and brown with dark brown genae, face of epi- having long medial part (Figs XI: 1–5; XII: cranium, clypeus and mandibles, and with 1, 2).

Female. General appearance as in male, only from A. hespera having the distance but tegmina completely very light brown between base of hind basitarsus and base of with partly whitish and partly grayish lon- this spur ~2 times as long as this spur; in the gitudinal venation as well as with brown new species, this ratio is ~1.5). crossveins (part of these crossveins almost light brown, but sometimes some of them Angustitrella andensis sp. nov. dark brown in dorsal field), abdominal ter- (Figs IX: 5, 6) gites somewhat lighter, genital plate com- Holotype. Female; Ecuador, western pletely yellowish, ovipositor brown with slopes of Andes Mts, 10 km E of Agua Blan- dark brown lateral parts, dorsal tegminal ca Vill. (village in environs of Puerto Lopez field with 10–11 longitudinal veins and Town), San Sebastian Forest, 700 m, cloud rather numerous crossveins, and lateral teg- primary forest, on leave, 26–29 Oct. 2005; minal field with somewhat wider Sc-R area coll. A. Gorochov & A. Ovtshinnikov; ZIN. and 13–15 oblique branches of Sc as well as Description. Female (holotype). General with crossveins slightly less numerous than appearance similar to that of female of A. in previous field. Genital plate short and maculata, but head uniformly brown with with apex widely truncate as well as with only proximal part of antennae dark brown posterolateral corners rounded; oviposi- excepting three brown most proximal an- tor with distal part not wider than rest of tennal segments (middle and distal parts of ovipositor and with only small denticles on antennae missing), pronotal disc uniformly lateral edges of this part (transverse ridges brown, fore and middle femora with yel- on dorsal and ventral surfaces of this part lowish proximal half and dark brown dis- absent; Fig. IX: 4). tal half, hind femora dark brown with yel- Length in mm. Body: male 15, female lowish proximal third, all tibiae with dark 14–17; body with wings: male 21, female brown proximal part and gradual transition 22–23.5; pronotum: male 2.9, female 3.1– to brown other parts (Figs IX: 5, 6), all 3.3; tegmina: male 15.5, female 16–16.5; tarsi brown, dorsal field of tegmina practi- hind femora: male 9, female 9.4–9.8; ovi- cally uniformly light brown, lateral field positor 7.4–7.6. of tegmina more or less uniformly brown, Comparison. The new species differs ventral part of abdomen (including genital from A. podagrosa, A. vicina, A. picipes, A.? plate and cerci) also brown, ovipositor light versuta comb. nov., A.? mirifica comb. nov. brown, and distal edge of inner (slit-like) and A.? rufa comb. nov. in the legs con- tympanum situated slightly distad than trastingly spotted, and additionally from middle of fore tibia (Fig. IX: 5). two the latter species, in the apical part Male unknown. of epiphallus lacking distinct bifurcation. Length in mm. Body 13.5; body with From A. roosevelti comb. nov. and A. hes- wings 23; pronotum 2.9; tegmina 16.5; hind pera comb. nov. having the legs more or less femora 8.8; ovipositor 6.3. spotted, the new species differs in the infla- Comparison. The new species is similar tion of fore tibiae shorter and less strong, to A. maculata, but distinguished by the inner tympanum shorter (situated almost above-listed characters of colouration and only in proximal half of tibia) [in A. roos- slightly more distal position of distal edge evelti, distal edge of this tympanum situated of inner tympanum. From A. roosevelti and distinctly distad than the middle of tibia A. hespera also having spotted legs, the new (for comparison see Figs IX: 2 and 13), and species differs in the same characters as A. in A. hespera, inflation of fore tibiae and in- maculata (see Comparison after the de- ner tympanum longer than even in A. roos- scription of A. maculata), and from all the evelti], and outer spur of hind basitarsi long other congeners, in the femora contrast- (the latter character separates A. maculata ingly bicolourous.

Angustitrella trivialis sp. nov. (slit-like) tympanum situated distinctly dis- (Figs I: 5; IX: 7–9; X: 2; XI: 6–10; XII: 3, 4) tad than middle of tibia (approximately as in A. podagrosa, A. roosevelti and A. vicina; Holotype. Male; Peru, Loreto Department, see Figs IX: 8, 12, 13, 15), spines of hind bank of Morona River near its mouth (not far tibiae slightly shorter (length of longest one from Puerto Morona Town), ~200 m, primary forest, at light, 20–23 Jan. 2010; coll. A. Goro- ~0.4 mm), venation of tegmina as in photo- chov; ZIN. graph (Fig. X: 2), hind wings longer (reach- Paratypes. One female; Peru, Ucayali De- ing apical part of hind tibiae), and genitalia partment, ~35 km NWW of Atalaya Town on with distinctly narrower anterior part of Ucayali River, environs of Sapani Vill., ~300 epiphallus and clearly shorter ectoparam- m, primary forest, at light, 26–31 Oct. 2008; eres fused with epiphallus laterally (Figs XI: coll. A. Gorochov, M. Berezin, L. Anisyutkin, E. 6–10; XII: 3, 4). Tkatsheva & V. Izersky; ZIN. One male; Eastern Variations. Apical segment of all palpi Ecuador, 70 km SE of Lago Agrio Town, envi- almost light brown; hind femora with more rons of S. Pablo de Kantesiya Vill. on Aguarico River, lowlying primary forest, on leave, 10–17 (than in holotype) gradual transition from Nov. 2005; coll. A. Gorochov & A. Ovtshinnikov; light brown proximal part to dark brown ZIN. One female; Southern Ecuador, bank of distal part; upper tegmen with darkened Morona River near border with Peru, environs spot near plectrum. of Puerto Morona Vill., ~300 m, primary forest, Female. General appearance as in male, at light, 5–15 Jan. 2010; coll. A. Gorochov; ZIN. but genae of head and pronotal lateral lobes Description. Male (holotype). Head and in one female completely dark brown, teg- pronotum reddish brown with dark brown mina completely light brown or with dorsal antennae (excepting three most proximal field light brown and lateral field brown, segments: scape almost light brown, two light part of hind femora varied from light other segments brown) and lower and hind brown to brown, hind tibiae and all tarsi parts of pronotal lateral lobes, grayish brown sometimes partly brown, most part of ovi- palpi, and light brown rest of mouthparts positor dark brown (as in A. maculata) or (but upper part of clypeus reddish brown); light brown (almost as in A. andensis), ve- legs dark brown with light brown coxae, nation of tegmina similar to that of A. mac- most part of hind femora (however their ulata and A. andensis (however crossveins distal part dark) and spines and spurs of between branches of Sc somewhat more hind tibiae; tegmina light brown with some- sparse), structure of genital plate also simi- what lighter dorsal field of upper tegmen lar to that of these species, and shape of ovi- and almost transparent dorsal field of lower positor distinguished from that of A. macu- tegmen; hind wings whitish; pterothorax lata only by somewhat wider base of distal and abdomen brown (their dorsal part and part (Fig. IX: 9). pterothoracic sternites lighter, almost light Length in mm. Body: male 12–13, fe- brown). Structure of body similar to that male 11–14; body with wings: male 19–20, of A. maculata, but pronotum and dorsum female 19.5–22; pronotum: male 2.5–2.6, of head hardly pubescent (almost shining) female 2.5–2.8; tegmina: male 12.5–13, fe- and weakly (but distinctly) rugous, prono- male 13.5–15; hind femora: male 7.5, female tal disc with a few weakly distinct concavi- 7.5–8; ovipositor 4.9–5.3. ties, metanotal gland with somewhat deeper Comparison. The new species is simi- concavity behind middle process and me- lar to A. podagrosa, A. vicina, A. roosevelti dian fusion of two distal folds of metanotum and A. hespera in the strongly inflated fore (in A. maculata, median parts of these folds tibiae, but distinguished from the first spe- well separated from each other; for compari- cies by the shorter mirror in male tegmina son see Figs IX: 1 and 7), fore tibiae strongly and somewhat more proximal position of inflated (Fig. IX: 8), distal edge of inner inner tympanum (for comparison see Figs

IX: 8 and 12; X: 2 and 5); from A. vicina, in graph (Fig. X: 3), and genitalia with much the more oblique both mirror and diagonal wider anterior part of epiphallus and small vein in male tegmina as well as less oblique (but distinct) median notch at epiphallic (more longitudinal) branches in apical area apex as well as with distinctly longer ecto- of these tegmina (see Figs X: 2 and 6); and parameres (Figs XI: 11–15; XII: 5, 6). from A. roosevelti and A. hespera, in the uni- Female unknown. form colouration of fore and middle legs. Length in mm. Body 14; body with The latter character and more inflated fore wings 21; pronotum 2.4; tegmina 14.5; hind tibiae of A. trivialis distinctly separate this femora 8. species from A. andensis and A. maculata Comparison. The new species is most (from the latter species, A. trivialis differs similar to A. trivialis, but distinguished by also in the ectoparameres clearly fused with the lighter fore and middle legs and above- the epiphallus). From A. picipes, the new mentioned characters of male metanotal species differs in the smaller size of body; gland and of male genitalia. From all the from A.? mirifica and A.? rufa, in the shape other congeners, the new species differs in of epiphallus lacking characteristic elongate the same characters as A. trivialis (see Com- posterolateral lobes; and from A.? versuta, parison of A. trivialis with all the other spe- in the rounded (not angular) lateral edges cies) in combination with the not dark fore of base of ovipositor distal part. and middle femora which are lighter than Notes. The above-mentioned females head and pronotum. (paratypes) collected in different localities and having some differences in colouration Angustitrella? borealis sp. nov. (including colouration of ovipositor) may (Figs IX: 11; X: 4; XI: 16–18; XII: 7, 8) belong to different subspecies. Holotype. Male; Mexico, Chiapas, environs of Tuxtla Gutierrez City, near reserve El Ocote, Angustitrella columbia sp. nov. 600–1000 m, primary forest, at light, 19–24 May 2006; coll. A. Gorochov & M. Berezin; ZIN. (Figs IX: 10; X: 3; XI: 11–15; XII: 5, 6) Paratypes. Two females; Mexico, Chiapas, Holotype. Male; Colombia, “Mosquitero, Rio Ocosingo Distr., Selva Lacandona near border Magdalena, Woronov”, 9 May 1926; ZIN. with Guatemala (between Biosphere reserve Montes Azules and Bonampak Natural Monu- Description. Male (holotype). General ment), environs of Lacanja-Chansayab Vill., pri- appearance similar to that of holotype of A. mary forest, at light, 20–27 May 2007; coll. M. trivialis, but genae and face of epicranium Berezin & E. Tkatsheva; ZIN. as well as pronotal lateral lobes complete- Description. Male (holotype). General ly dark brown, antennae almost uniformly appearance more or less similar to that of all brown, palpi and femora light brown (how- previous species of Angustitrella, but char- ever hind femora with brown apical part), acterised by following characters: head and tibiae and tarsi brown (with light brown pronotum reddish brown with dark brown apical tarsal segment and spurs of hind ba- proximal part of antennae (excepting three sitarsus), pterothorax and abdomen more or most proximal segments which brown), less uniformly light brown, metanotal gland pronotal lateral lobes and a pair of spots with clearly smaller and distinctly less deep on pronotal disc, and brown mouthparts concavity behind middle process (this con- (excepting reddish brown middle part of cavity also clearly smaller and less deep maxillary palpi and almost light brown la- than in A. maculata) as well as with some- brum) and middle part of antennae (their what narrower areas between lateral parts distal part missing); fore and middle legs of two distal folds of metanotum (Fig. IX: dark brown with almost blackish apical part 10), length of longest spine of hind tibiae of femora and base of tibiae, brown trochan- ~0.5 mm, dorsal field of tegmina as in photo- ters and most part of third tarsal segments,

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 251 and light brown claws; hind legs with brown brown lateral field and uniformly light basal part, reddish most part of femora and brown dorsal field, tegminal venation simi- small dorsal subbasal spot on tibiae, dark lar to that of female of A. maculata, and brown apical part of femora and most part abdominal apex almost as in female of A. of tibiae and tarsi, reddish brown lower part trivialis. of proximal half of tibiae, and light brown Length in mm. Body: male 15, female distal part of third tarsal segments; tegmina 13–15; body with wings: male 23.5, female almost uniformly light brown with slightly 21–23; pronotum: male 2.7, female 2.7–2.9; darker majority of veins (excepting light tegmina: male 16, female 14–15.5; hind brown oblique veins in stridulatory ap- femora: male 7.8, female 7–7.7; ovipositor paratus) and semitransparent stridulatory 5–5.3. areas of lower tegmen; hind wings whitish Comparison. The new species is most with light brown venation and membranes similar to A.? mirifica and A.? rufa, but it of distal part; pterothorax and abdomen differs from them in the slightly narrower brown with hardly lighter areas on metano- mirror of male tegmina, somewhat shorter tum and two proximal sternites of abdomen, posterolateral lobes of epiphallus, and dis- and slightly darker apical part of anal and tinctly wider notch between these lobes. genital plates; external structure of body From A. podagrosa, A. vicina, A. picipes, A. similar to that of A. trivialis, but pronotum roosevelti, A. hespera, A. maculata, A. an- and dorsum of head practically without densis and A.? versuta, the new species dif- pubescence (clearly shining) and more dis- fers in the shining and very rugous dorsum tinctly rugous, pronotum also with more of both head and pronotum as well as more noticeable concavities on disc and some- crumpled pronotal lateral lobes, and from what more crumpled lateral lobes, meta- all the other congeners with known male notal gland with concavity behind middle genitalia, in the genital characters listed in process approximately as in A. mutabilis and Description. Differences in male genitalia with two distal folds of metanotum almost between the new species (+ two the above- as in A. trivialis, however areas between lat- mentioned most similar species) and all the eral parts of these folds wider and median other congeners here studied are rather parts of these folds (fused with each other) significant; these differences indicates that narrower (Fig. IX: 11), spines of hind tibiae this genus may be divided into two subgen- hardly longer (length of longest one ~0.5 era or even two related genera. mm), tegmina with slightly longer mirror (Fig. X: 4), and hind wings hardly longer Genus Ectotrypa Saussure, 1874 (reaching basal part of hind tarsi). Genita- Chremon Rehn, 1930, syn. nov. lia (Figs XI: 16–18; XII: 7, 8) distinguished from those of all previous species of Angusti- Type species: Ectotrypa olmeca Saussure, trella by presence of elongate posterolateral 1874. epiphallic lobes directed aside and back- Note. This genus is characterised by wards and of a pair of posteromedial semi- the elongate and rather narrow body (Figs membranous lobules (lo), straight ectopara- XIII: 1, 4, 6), head with the elongate hind meres clearly separated from epiphallus, part (behind eyes), and long and narrow larger guiding rod, and absence of curvature pronotum (Figs XIII: 1, 3–7) which is dis- of distal part of genitalia in profile (this cur- tinctly longer than its width. The latter vature characteristic of all previous species character is presented in female, but in pre- of Angustitrella with known genitalia; for viously unknown male, pronotum might be comparison see Figs XI: 5, 10, 15, 18). of somewhat different shape. In all the pre- Female. Colouration and structure of vious genera of the subtribe Paroecanthina, body as in male, however tegmina with pronotum is shorter than its width or (only

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 252 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI in some females) as long as wide. The other mon repentinus Rehn, 1930 (type species characteristic features of Ectotrypa are fol- of Chremon), Ch. apithanon Otte et Per- lowing: legs long and thin (especially fore ez-Gelabert, 2009, Ch. anembatos Otte et and middle ones; Figs IX: 19; XIII: 1, 4, 6), Perez-Gelabert, 2009, Ch. jaiqui Otte et with the tibiae distinctly longer than fem- Perez-Gelabert, 2009, Ch. eribombos Otte ora; fore tibiae hardly inflated in the tym- et Perez-Gelabert, 2009, Ch. procax Otte panal region and with the outer tympanum et Perez-Gelabert, 2009 from Hispaniola oval (opened) and not immersed, and inner I.; E. veracruz sp. nov.; possibly Carsidava tympanum almost slit-like, rather short cinerascens Walker, 1869 (type species of (not longer than outer one) and weakly im- Carsidava) from Brazil, Paroecanthus fal- mersed (Fig. IX: 19, 20); distal part of ovi- lax Saussure, 1874 from Cuba, P. sporadi- positor rather narrow (somewhat narrower cos Otte et Perez-Gelabert, 2009, P. scinax than in the previous genera of Paroecanthi- Otte et Perez-Gelabert, 2009, P. valens Otte na) and having several distinct oblique den- et Perez-Gelabert, 2009 from Hispaniola I., ticles on the lateral edges (these denticles and P. strongylops Otte et Perez-Gelabert, are somewhat larger and more oblique than 2009, P. otaros Otte et Perez-Gelabert, in the previous genera; Fig. IX: 18). 2009, P. hwinanus Otte et Perez-Gelabert, No one male included in Ectotrypa by 2009 from Jamaica. previous authors, but there are two similar genera (with known males) having the pro- Ectotrypa veracruz sp. nov. notum longer than its width. One of these (Figs IX: 18–20; XIII: 1–3) genera is Carsidava Walker, 1869, an enig- matic genus synonymised with Paroecan- Holotype. Female; Mexico, Veracruz, 15–20 thus by Saussure (1874). However, judging km NE of Catemaco Town, Los Tuxtlas (biostation of Mexico University) in 2 km from coast by its pronotal shape, Carsidava may be of Mexican Gulf, partly primary and partly sec- a synonym of Ectotrypa or a distinct ge- ondary forest on hills, on leaves, 6–17 Nov. 2006; nus (Carsidava differs from Ectotrypa by coll. A. Gorochov & A. Ovtshinnikov; ZIN. the legs “rather short and stout”; Walker, Paratype. Female; same data as for holotype; 1869). Another genus is Chremon Rehn, ZIN. 1930 including a few species with the long Description. Female (holotype). Head pronotum (in their males, pronotum may be black with dark brown most part of clyp- from hardly longer than its width to slightly eus as well as genae under hind half of eyes, wider than its length). Legs of Chremon are reddish brown maxillary palpi and genae similar to those of Ectotrypa in shape (its under anterior half of eyes (but with black fore tibiae may be from hardly to almost spot near each eye), brown other mouth- moderately inflated), and its ovipositor parts (excepting light brown lateral parts of is also more or less similar to that of Ecto- clypeus), yellowish rostral area (Figs XIII: trypa. Chremon is a more probable synonym 1–3) and scapes, and light brown rest of an- of Ectotrypa. Its males are with the tegmi- tennae; pronotum black with a pair of dark nal stridulatory vein curved as in the other brown spots on disc and brown borders; legs representatives of Paroecanthina and with yellowish with blackish stripes along each the tegminal mirror possibly varying from ventral keels of femora (in hind femora, slightly elongate (oval) to possibly very these stripes from dark brown to brown), long (narrow; Fig. X: 8), but their genitalia rose tibiae (excepting yellowish small basal are insufficiently studied. part, subbasal spot, distal part, and spines So, Ectotrypa may include the follow- and spurs), brownish small marks on api- ing species (named in original binomen): cal part of hind femora and on outer side of E. olmeca (type species) from Mexico; E. their distal third, and brown two longitudi- brevis Rehn, 1905 from Costa Rica; Chre- nal parallel stripes on inner side of distal half

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 253 of hind femora; tegmina with light brown anembatos comb. nov. and E. jaiqui comb. dorsal field having hardly lighter venation, nov. in the presence of characteristic dark reddish lateral part of this field and Sc-R stripes along the ventral keels of femora, but area, yellowish Sc (including its branches), it differs from them by the most part of head greyish brown (rather dark) membranes dark and pronotum almost uniformly dark. between Sc branches, and greyish (lighter) From all the other true and possible con- stripe along ventral edge of lateral field geners, the new species differs in the more (Figs XIII: 1, 3); venter of body yellowish contrast colouration (with the dark head with brown anterior part and small darkish and pronotum as well as two dark stripes marks on abdominal pleurites; rest of abdo- along the ventral keels of femora). men yellowish with reddish brown areas on hind tergites as well as rose anal plate, para- Genus Selvagryllus Otte, 2006 procts and cerci, and dark brown ovipositor having light (almost yellowish) median part Type species: Selvagryllus spizon Otte, from base of ovipositor to base of its distal 2006. part (Fig. IX: 18). Shape of head typical of Note. This genus is practically not de- this genus (Figs XIII: 1–3); rostrum be- scribed, but its author (Otte, 2006) indi- tween antennal cavities slightly narrower cated that for the recognition of Selvagryl- than scape; lateral ocelli very small; median lus, it is necessary to see the recognition of ocellus indistinct. Pronotum approximately type species, and he included in this genus 1.2 times as long as wide; its lateral lobes low only one species. Such “description” may be (Figs XIII: 1, 3). Fore tibiae hardly inflated considered an united description of a new in tympanal region (Fig. IX: 19). Tegmina genus and a new species (Comission on Zo- insignificantly not reaching apex of hind ological Nomenclature, 1999: article 13.4). femora; dorsal field with 9–10 somewhat In the species description, Otte mentioned oblique longitudinal veins and numerous only a single character suitable for the ge- crossveins; lateral field with 15–16 oblique neric recognition: “large inner and large branches of Sc and a few crossveins between outer tympana”. Possibly he wished to write some of these branches; hind wings reach- that the both tympana (inner and outer) are ing tegminal apex (not exposed behind opened (not slite-like). His photographs tegmina) (Fig. XIII: 1). Genital plate mod- (Otte, 2006: Figure 116) show that the erately short, with roundly truncate apex; stridulatory vein in male tegmen of S. spi- ovipositor as in Fig. IX: 18. zon is strongly S-shaped (as in all the other Variations. Paratype slightly smaller, genera of Paroecanthina), its male genitalia with black spot near anteroventral edge are weakly similar to those of Angustitrella? of each eye larger (reaching subgena), and borealis, and its general appearance is some- without any rose areas (fore and middle tib- what similar to a female described below as iae as well as epiproct, paraprocts and cerci S.? eltriunfo sp. nov. yellowish). So, this genus includes S. spizon (type Male unknown. species) from Costa Rica and possibly a few Length in mm. Body 16–17; body with other species with the both tympana opened wings 18–19.5; pronotum 3.8–4; tegmina (they are given in original binomen): Paroe- 12–13; hind femora 11.5. canthus foraminatus Saussure, 1878 from Comparison. The new species is similar Cuba; P. olmecus Saussure, 1897 from Mex- to E. olmeca in the black pronotum, but dis- ico; P. tibialis Saussure, 1897 from Panama; tinguished by the darker head (in E. olmeca, S.? eltriunfo sp. nov. Eneoptera annulata head light with only the face darkened). Scudder, 1869, described from Central E. veracruz is also similar to E. repentinus America without any data on its tympana, comb. nov., E. apithanon comb. nov., E. may also belong to this genus because head

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 254 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI colouration of S.? eltriunfo sp. nov. is rather (Fig. IX: 23); inner tympanum oval (not similar to that of a specimen determined by long), slightly immersed (Fig. IX: 22); hind Nickle (1992: Fig. 11.42) as Scudder’s spe- tibiae and tarsi as in Fig. XIV: 4. Tegmina cies; however this determination is prob- reaching proximal part of hind femora, with lematic because Nickle’s picture is not in ac- 9–10 weakly oblique longitudinal veins and cordance to Scudder’s original description. numerous crossveins in dorsal field, and with 12–13 distinctly oblique branches of Selvagryllus? eltriunfo sp. nov. Sc and only a few crossveins between them; (Figs IX: 21–23; XIV: 1–4) hind wings weakly (but distinct) exposed behind tegmina. Genital plate and oviposi- Holotype. Female; Mexico, Chiapas, 130 km tor similar to those of Ectotrypa veracruz WN of Tapachula City, environs of Ejido Las (for comparison see Figs IX: 18 and 21). Golondrinus Vill. near reserve El Triunfo, 800– 1000 m, partly primary and partly secondary for- Male unknown. est, at light, 13–17 May 2006; coll. A. Gorochov Length in mm. Body 14; body with & M. Berezin; ZIN. wings 18; pronotum 2.7; tegmina 12; hind Description. Female (holotype). Co- femora 8.5; ovipositor 5.8. louration of body yellowish with follow- Comparison. From S. spizon, the new ing marks: head with characteristic brown species differs in the almost completely pattern on dorsum of head and behind eyes light lateral lobes of pronotum; from S.? (Figs XIV: 1–3) as well as with a few brown- foraminatus comb. nov., in the absence of ish dots under eyes and small light brown dark marks on tegmina, on legs and on ab- spot near medial corner of each subgena; domen; from S.? olmecus comb. nov., in the pronotum with four dark brown longitu- distinctly shorter inner tympanum; from S.? dinal stripes interrupted in middle part (a tibialis comb. nov., in the not spotted an- pair near median line of disc and a pair along tennae; and from S.? annulatus comb. nov., lateral edges of disc) and with a few brown in the absence of dark marks on face, on ros- and light brown spots between these stripes trum of head, and on tegmina. (Figs XIV: 2, 3); legs with two brown longitudinal lines in middle part of outer surface Subtribe Tafaliscina Desutter, 1988, of hind femora, with a few small brownish stat. nov. marks on ventral and dorsal surfaces of hind femora as well as on inner and outer This subtribal name originates from Ta- surfaces of distal part of these femora, with faliscinae Desutter, 1988. At the first time dark brown spot near base of each spine Tafaliscinae was mentioned by Desutter and of each spur in hind tibiae, and with (1987) as a new subfamily of her family En- brown spot between spurs of hind basitarsi eopteridae, but without any description or (Fig. XIV: 4); ovipositor dark brown with diagnosis (unavailable name). The diagno- almost light brown median stripe (from sis of Tafaliscinae was published in another basal part of ovipositor to base of distal part paper (Desutter, 1988), and thus the name of ovipositor). Head not elongate; rostrum Tafaliscinae becomes available since 1988. between antennal cavities approximately This subtribe is distinguished from the twice narrower than scape; median ocellus subtribe Paroecanthina by the lateral part absent; lateral ocelli distinct, medium-sized. of male stridulatory vein (lacking stridula- Pronotum clearly wider than its length, tory teeth) short and less strongly S-shaped with roundly convex hind edge. Legs rather (if tegminal stridulatory apparatus is pre- short; tibiae slightly longer than femora in served; Figs I: 7; XV: 3, 9, 12), sclerites of fore and middle legs, and slightly shorter, male genitalia are not reduced (ectoparam- in hind legs; fore tibiae not inflated; outer eres are well sclerotised and clearly separat- tympanum almost round, hardly immersed ed from the epiphallus; endoparameres are

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 255 well developed, rather large and with dis- (these processes with membranous medial tinct apodemes; mold of spermatophore at- part) and a pair of short medial projections tachment plate is also developed) excepting on hind edge (Figs XIV: 10–12). guiding rod which is reduced and/or fused Included species. Only type species. with the above-mentioned mold (Figs XIV: However it is impossible to exclude that 10–12; XV: 4–6; XVI: 3–5, 7–9, 15–17; some species (with the both tympana de- XVII: 5–7; XVIII: 3–5), and distal part of veloped and male genitalia unstudied) in- ovipositor usually with the more obtuse or cluded in the genus Amblyrhethus (A. manni truncate apex (Figs XV: 8, 10, 11, 13, 14, 17, Rehn, 1917 from Brazil; A. natalensis Rehn, 18, 21–24; XVI: 10, 11; XIX: 7, 8). 1917 from Brazil; A. ponderosus Hebard, Tafaliscina includes the following gen- 1928 from Panama; Amblyopus capitatus era: Tafalisca Walker, 1869; Amblyrhethus Saussure, 1878 from unknown locality) Kirby, 1906; Pseudogryllus Chopard, 1912, may belong to this genus. gen. resurr.; Perutrella gen. nov.; Mexitrypa Comparison. The new genus is similar gen. nov.; Brazitrypa gen. nov. to Amblyrhethus in the presence of stridulatory apparatus in male tegmina, but dis- Genus Perutrella gen. nov. tinguished by the inner and outer tympana well developed and epiphallus with the long Type species: Perutrella originalis sp. nov. and narrow posterolateral processes. From Diagnosis. Crickets with general appear- all the other genera of Tafaliscina, the new ance partly similar to that of Angustitrella genus differs in the presence of tympana and partly, to that of Amblyrhethus. Head and of stridulatory apparatus as well as in not elongate, with small lateral ocelli and the same character of male genitalia. indistinct median ocellus; rostrum slightly Etymology. The generic name is origi- wider than scape and roundly angular in nated from Peru and the genus Sonotrella. profile. Pronotum transverse, with weakly convex hind edge, and in male, distinctly Perutrella originalis sp. nov. narrowing to head (Fig. XIV: 5); male (Figs I: 7; XIV: 5–12) metanotal gland reduced, with only a group of short hairs on central convexity of meta- Holotype. Male; Peru, Junin Department, Satipo Prov., ~25 km SE of Satipo Town, envi- notum (Fig. XIV: 6). Legs short and stout; rons of Rio Venado Vill., ~1200 m, partly pri- femora of fore and middle legs hardly short- mary and partly secondary forest, at light, 20–23 er than their tibiae; fore tibiae (Fig. XIV: 7) Oct. 2008; coll. A. Gorochov, M. Berezin, L. An- somewhat widened, but compressed (flat- isyutkin, E. Tkatsheva & V. Izersky; ZIN. tened) laterally; both tympana oval, opened Description. Male (holotype). Coloura- (outer tympanum almost not immersed, but tion grayish, rather light, however with fol- inner one hardly immersed); middle tibiae lowing marks: head with a few hardly dark- less widened, but similarly flattened; hind ened longitudinal bands on hind two thirds femora much longer than hind tibiae; hind of vertex and rather small dark brown spots basitarsi also long (Fig. XIV: 8). Male teg- on genae (near eyes) and on antennal flagel- mina completely developed, with stridula- lum; pronotum with a few dark brown marks tory apparatus having all structures similar on disc (Fig. XIV: 5) and brown stripe on to those of Paroecanthina excepting stridu- lateral lobes along their dorsal edge; legs latory vein which typical of Tafaliscina (see with numerous dark dots on femora and Figs I: 7; XIV: 9). Male genital plate rather tibiae, but fore tibiae having a pair of addi- short, approximately 1.5 times as long as tional somewhat larger dark spots on dorsal anal plate (Fig. XIV: 8); male genitalia half near tympana, middle and hind tibiae with short epiphallus having a pair of long having additional subbasal dark band, hind and rather narrow posterolateral processes femora having dark dots partly fused with

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 256 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI each other (in form of short oblique stripes) Genus Amblyrhethus Kirby, 1906 and apical part distinctly darkened, and tarsi having distal part of third segment Amblyopus Saussure, 1878; nec Amblyopus De- jean, 1835 (Coleoptera). darkened; tegmina almost semitransparent, but upper tegmen with majority of veins Type species: Amblyopus brevipes Sau- brown, rest of veins light brown and many ssure, 1878. membranes having different spots or short Note. The genus, judging by its type spe- stripes from dark to weakly darkened (Fig. cies, differs from Perutrella in the absence XIV: 9), and lower tegmen with more uni- of inner tympanum (outer tympanum pres- formly light (almost completely semitrans- ent), S-shaped part of male stridulatory parent) basal and stridulatory areas. Meta- vein shorter and less curved (Fig. XV: 3), notal gland as in Fig. XIV: 6. Both tympana and male genitalia without posterolateral rather large (outer tympanum almost equal processes and with the distinctly larger me- to inner one in size); hind tibiae almost dial projections of its hind edge (Figs XV: twice shorter than hind femora and approx- 4–6). It includes type species and prob- imately twice longer than hind basitarsi, ably Amblyopus depressus Saussure, 1878 with short and thick spines and spurs, and (both from Colombia). Some other species with only a few small denticles; hind ba- were included in this genus by their authors sitarsi with large denticles and long spurs (Amblyopus capitatus Saussure, 1878; Am- (these spurs distinctly longer than spurs of blyrhetus manni Rehn, 1917; A. natalensis hind tibiae) (Figs XIV: 7, 8). Tegmina long, Rehn, 1917; A. ponderosus Hebard, 1928), with rather narrow mirror and other charac- however systematic position of these spe- ters as in Fig. XIV: 9; hind wings also long, cies is not clear; they may belong to Amblyr- extending to apex of hind legs. Anal plate hethus as well as to another genus (see the with rounded apex and a pair of S-shaped paragraph about composition of Perutrella) dorsal keels separating median concavity or even to different genera. Amblyrhetus from rather narrow lateral parts; genitalia nodifer Chopard, 1956 from Peru is also as in Figs XIV: 10–12. included in Amblyrhethus by its author; it Female unknown. has the tympana as in Perutrella and male Length in mm. Body 16.5; body with genitalia more or less similar to those of A. wings 24.5; pronotum 2.8; tegmina 18; hind brevipes (Figs XV: 15, 16), and thus its sys- femora 9.3. tematic position is not very clear also. Comparison. The new species differs from all the other similar or related species in the Amblyrhethus brevipes (Saussure, 1878) clearly longer and narrower mirror in male (Figs XV: 1–8) tegmina (from Amblyrhetus? manni and A.? natalensis; compare Figs XIV: 9; XV: 9, 12) as Lectotype. Male; Colombia, “Columb., Mori- well as distinctly longer hind basitarsi (from tz”, “Amblyopus brevipes Sss., male”, “997”, “Am- A.? ponderosus and A.? capitatus; Figs XIV: 8; blyopus Sauss.”, “male, co-typus”, “Amblyopus brevipes Sauss.*, male / Mem. Soc. Phys. Nat. XV: 19, 20). Its male genitalia are similar to Hist. Genève 25, 1878, p. 599”; MNH. those pictured by Desutter (1987: Figs 12– Other material examined. Three females; 14) as belonging to Tafalisca lurida Walker, Colombia, “Columbien, Moritz”, “Cat. № 997”, 1869 (this determination is erroneous, and determined by unknown person as “Amblyopus these genitalia may belong to an unknown brevipes Sauss.”; MNH. species of Perutrella), but distinguished by Note. The species was described after the epiphallus distinctly more curved in pro- syntypes from Colombia (male from MNH file and hind edge of ectoparameres (if to see and female from “Musée de Leipzig”; Sau- from below) not rounded, but with the short ssure, 1878). This male is here designated as and almost angular projection. lectotype of this species.

Figs XIV (1–12). Different structures of body. 1–4, Selvagryllus? eltriunfo sp. nov., female; 5–12, Perutrella originalis sp. nov., male. Head in front (1); anterior part of body from above (2, 5) and from side (3); hind tibia, dorsal view (4); metanotum from above (6); fore tibia, outer view (7); middle and hind legs together with abdominal apex from side (8); tegmen (9); genitalia from above (10), from below (11) and from side (12).

Figs XV (1–24). Amblyrhethus and Tafalisca. 1–8, A. brevipes; 9–11, A.? manni; 12–14, A.? natalensis; 15–18, A.? nodifer; 19, A.? ponderosus; 20–22, A.? capitatus; 23, 24, T.? crypsiphonus. Head with pronotum from above (1); hind tibia, inner view (2, 19) and inner-dorsal view (20); dorsal field of male tegmen (3, 9, 12); male genitalia from above (4, 15), from below (5) and from side (6); same without both anterior part and endoparameres from below (16); distal part of ovipositor from side (7) and from below (8), and its left half from above (10, 13, 17, 21, 23) and from below (11, 14, 18, 22, 24). [2, 20–24, after Saussure (1878); 9–14, after Rehn (1917); 15–18, after Chopard (1956); 19, after Hebard (1928).]

Redescription. Male (lectotype). Co- from the same island; this work shows that louration yellowish grey with following Tafalisca is characterised by the male geni- marks: numerous darkish dots on majority talia more or less similar to those of Amblyr- of bodyparts; brown pattern on dorsum of hethus, but with the posteromedial projec- head and of pronotum (as in Fig. XV: 1); tions of epiphallus very large (lobe-like), yellowish antennae having sparse segments strongly inflated and semimembranous of flagellum darkened; brownish dorsal part (Figs XVI: 3–5, 7–9, 15–17). All the oth- of fore and middle tibiae; a few small dark er species with different or unknown male spots on hind tibiae; light brown tegmina genitalia, included in this genus by different having small brown spots on dorsal field; authors (Eades et al., 2011) and similar to yellowish cerci; almost brown abdomen. Tafalisca in the absence of stridulatory ap- Structure of body more or less similar to paratus in male tegmina and of tympana, that of Perutrella originalis, but outer tym- belong to other genera or are in need of a panum somewhat shorter, inner tympa- restudy. The preservation of traces of male num absent, hind tibiae clearly longer (but stridulatory apparatus in T. lurida and T. shorter than hind femora) and with more rico (see Otte & Perez-Gelabert, 2009: Fig- numerous denticles, hind basitarsi distinct- ures 413, B and 419, B) supposes that these ly shorter (Fig. XV: 2), tegmina shorter apparatus and tympana might be reduced (reaching only abdominal apex) and with independently in Tafalisca and in some somewhat different venation of dorsal field other genera probably related to Tafalisca (Fig. XV: 3), hind wings not exposed be- in connection with the similar structure of hind tegmina, anal plate without any keels hind tibiae (their denticles are rather long and without concavity between them, and and more or less similar to their spines in genitalia as in Figs XV: 4–6. shape; Figs XVI: 1). Female. General appearance similar to The genus Metrypa was described by male, but tegminal dorsal field with 9–10 Brunner-Wattenwyl (1873) without inclu- oblique longitudinal veins and numerous sion of any species. Saussure (1874) syn- and somewhat irregular crossveins (form- onymised Metrypa and Tafalisca, however ing slightly cellular venation). Ovipositor with usage of the junior name as a valid one; with distal part as in Figs XV: 7, 8. he also included three species in this genus. Length in mm. Body: male 15, female One of these species is T. lurida, and two 16–17; pronotum: male 2.6, female 2.8–3.2; other species are with unclear generic posi- tegmina: male 11; female 11–12; hind femo- tion as some their important characters (for ra: male 8, female 9–10; ovipositor 5.5–5.9. example, characters of male genitalia) are unstudied up to now. Since this publication, Genus Tafalisca Walker, 1869 Metrypa is always considered a synonym of Tafalisca, but its type species is not desig- Type species: Tafalisca lurida Walker, nated. For exact fixation of this synonymy, 1869. I designate here T. lurida as a type species Metrypa Brunner-Wattenwyl, 1873 of Metrypa. So, this species is now the type (synonymised by Saussure, 1874). species of both Tafalisca and Metrypa. Metrypus Saussure, 1878 (unjustified Tafalisca includes the following species: emendation). T. lurida (type species); T. gnophos Otte et Note. The genus originally was described Perez-Gelabert, 2009 from Cayman Islands; after a single female from Hispaniola I. T. periplanes Otte et Perez-Gelabert, 2009 (Walker, 1869: “St. Domingo”). Recently and T. rico Otte et Perez-Gelabert, 2009 from this species was very briefly redescribed Puerto Rico; T. eleuthera Otte et Perez-Gela- by Otte & Perez-Gelabert (2009) on base bert, 2009 from Bagamas; T. ecuador sp. nov.; of holotype, studied by them, and of a male T. mexico sp. nov.; T. proxima sp. nov.

Tafalisca ecuador sp. nov. slightly oblique veins as well as numerous (Figs XVI: 1–5) and irregular crossveins; tegminal lateral field with very narrow distal part and 5–6 Holotype. Male; Western Ecuador, envi- almost parallel veins as well as without dis- rons of Puerto Lopes Town, low hills near Agua tinct crossveins; hind wings reaching only Blanca Vill. (not far from Pacific coast), dry forest, at night, in cavity inside of dry bush branch, middle of 4th abdominal tergite. Anal plate 24–26 Oct. 2005; coll. A. Gorochov & A. Ovt- simple (short and with rounded apex); geni- shinnikov; ZIN. tal plate almost twice longer than anal plate Description. Male (holotype). Colou- and with roundly angular apex; genitalia as ration light brown with following marks: in Figs XVI: 3–5. head with brown large spot on dorsum, Female unknown. somewhat smaller transverse spot on hind Length in mm. Body 17; pronotum 2.7; part of vertex, a few small spots between tegmina 7.2; hind femora 8.8. antennal cavities and along clypeal suture, Comparison. The new species differs and yellowish genae, palpi, labrum and an- from all the above-mentioned congeners tennae (but scape with small brownish me- (excepting two new species) in the distinct- dial spots); pronotum with yellowish lateral ly shorter distal part of ectoparameres, and lobes and lateral parts of disc, brown stripe from the other species included in Tafalisca along dorsal edge of lateral lobes, and gray- (Eades et al., 2011), in the wings shortened ish brown transverse spot along hind edge and/or male genitalia different. of disc; legs with numerous almost indistinct darkish dots, brownish grey small Tafalisca mexico sp. nov. subbasal spot on dorsal surface of fore and (Figs XVI: 6–11) middle tibiae as well as a few areas on apical part of hind femora, and grayish brown most Holotype. Male; Mexico, Jalisco, 19°33´N, part of dorsal surface of hind tibiae; tegmina 105°5´W, biostation of Mexico University near Chamela Vill. (3–4 km from Pacific Ocean), with yellowish stripe on dorsal field along dry forest on hills, on dry bush branch, at night, its lateral edge, brownish stripe on lateral 23–28 Nov. 2006; coll. A. Gorochov & A. Ovt- field along its dorsal edge, and light (almost shinnikov; ZIN. transparent) rest of lateral field; hind wings Paratypes. Three females; same data as for whitish; sternites of metathorax and of first holotype; ZIN. abdominal segment brown; most part of Description. Male (holotype). General abdominal tergites and of anal plate dark appearance similar to that of T. ecuador, but brown. Rostrum of head roundly angular distinguished by following features: dor- in profile, slightly narrower than scape; sum of head more uniformly brown; genae ocelli indistinct. Pronotum transverse, with with a few brown marks; dark median spot almost straight hind edge (Fig. XVI: 1); along clypeal suture distinctly larger; an- metanotal gland with very large and rather tennal flagellum light brown; pronotal disc deep concavity having smaller median con- brown with light brown lateral parts; pro- vexity in its hind two thirds (almost as in notal lateral lobes with brown line along Fig. XVI: 13). Legs rather thick and stout; anterior, ventral and posterior edges; dorsal tibiae hardly longer than femora in fore and tegminal field brown with distinct whitish middle legs, but somewhat shorter, in hind stripe along its lateral and apical edges; lat- legs; tympana absent; hind tibiae much lon- eral tegminal field with brown majority of ger than hind basitarsi and with longest veins and band along dorsal edge; middle spines almost twice longer than longest tibiae slightly darkened; apical part of hind denticles (Fig. XVI: 1). Tegmina shortened, femora dark brown; hind tibiae and tarsi reaching middle of 5th abdominal tergite; almost uniformly greyish brown (however their dorsal field with 8–9 longitudinal and spines and some denticles slightly lighter);

Figs XVI (1–17). Tafalisca. 1–5, T. ecuador sp. nov.; 6–11, T. mexico sp. nov.; 12–17, T. proxima sp. nov. Male body from above (1); male tegmen (2, 6, 12); male genitalia (3–5) and their distal half (7–9, 15–17) from above (3, 7, 15), from below (4, 8, 16) and from side (5, 9, 17); distal part of ovipositor from above (10) and from below (11); male metanotal gland from above (13); spermatophore from side (14).

Figs XVII (1–7). Mexitrypa ovtshinnikovi sp. nov.; male. 1, left tegmen; 2, metanotal gland from above; 3, hind tibia, outer view; 4, abdominal apex from above; 5–7, genitalia from above (5), from below (6) and from side (7).

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 263 anal plate yellowish with darkish apex; Description. Male (holotype). Head yel- sternites of hind half of abdomen with dark lowish with dark brown dorsum (having a brown hind part; genital plate dark brown; pair of light stripes along medial edges of cerci yellowish grey; tegmina and hind antennal cavities and of eyes) and rostrum, a wings longer, reaching hind part of 7th few small dark marks on genae and on frons and 6th abdominal tergites respectively; (under rostrum, eyes and antennal cavities) genitalia with sclerotised part of epiphalus as well as on subgenae and on upper part somewhat shorter, a pair of sclerotised rib- of clypeus, and distinct brown stripe along bons at apex of genitalia somewhat longer, clypeal suture; pronotum dark brown with ventromedian sclerotization near this apex yellowish lower half of lateral lobes (hav- slightly narrower and (in profile) with an- ing narrow dark stripe along anterior, ven- gular (not widely rounded) ventral projec- tral and posterior edges); legs light brown tion, ectoparameres without spine-like pro- with a few brownish dots on fore and middle jection at proximal corner (see in profile), femora, small brown marks on fore coxae and endoparameres with distinctly shorter and on subbasal part of dorsal surface of fore apodemes (for comparison see Figs XVI: tibiae, greyish brown middle and hind tibiae 3–5, 7–9). as well as most part of hind tarsi and reticu- Female. Colouration and structure of lar ornament on outer surface of hind fem- body (excepting genital structures) as in ora, almost dark brown distal part of these male, but tegmina with somewhat longer femora, and dark all second tarsal segments; widened part of lateral field having more tegmina brown with yellowish white lon- numerous parallel veins (7–8). Genital gitudinal stripe and light grey membranes plate rather short, with widely rounded in lower half of lateral field (Fig. XVI: 12); (almost truncate) apex, light lateral parts hind wings very light brown; pterothorax and dark median area; ovipositor almost brown with slightly lighter metanotal gland uniformly brown or dark brown, with distal (Fig. XVI: 13) and light sternites (but meta- part as in Fig. XVI: 10, 11. thoracic sternite with large brown median Length in mm. Body: male 18, female spot); abdomen with dark brown tergites, 18–22; pronotum: male 3, female 3.5–4; sternites, and anal and genital plates, light tegmina: male 10, female 10.5–12; hind pleural area and cercal base (but rest of cerci femora: male 9.8, female 12–13.5; ovipositor greyish brown). External structure of body 8.7–9.3. similar to that of T. ecuador and T. mexico, Comparison. The new species differs however tegmina reaching base of 8th ab- from T. ecuador in the above-mentioned dominal tergite, tegminal dorsal field with features (including light male anal plate 9–10 longitudinal veins, lateral tegminal and dark male genital plate; in T. ecuador, field with 7–8 longitudinal veins and several male anal plate dark, and male genital plate distinct crossveins, and hind wings reaching light). From all the other true and possible apex of 6th abdominal tergite. Genitalia with congeners, the new species differs in the sclerotised part of epiphallus as in T. mexico, same characters as T. ecuador. ectoparameres and endoparameres as well as sclerotised ribbons at apex of genitalia as in Tafalisca proxima sp. nov. T. ecuador, ventromedian sclerotization near (Figs XVI: 12–17) genital apex angular in profile (almost as in T. mexico) and rather wide (almost as in T. Holotype. Male; Mexico, Oaxaca, 35 km ecuador) (Figs XVI: 15–17); spermatophore NNE of Santa Cruz Huatulco Town (10 km N of Xadani Vill.), 900–1000 m, partly primary and as in Fig. XVI: 14. partly secondary forest, in cavity inside of dry li- Female unknown. ana, at night, 7–11 May 2006; coll. A. Gorochov Length in mm. Body 21.5; pronotum 3.7; & M. Berezin; ZIN. tegmina 12.5; hind femora 11.8.

Comparison. The new species differs Comparison. The new genus is related from T. ecuador and T. mexico in the pe- to Tafalisca, but distinguished by the fea- culiarities of colouration [anal plate dark tures listed above. Separation of these gen- (not as in T. mexico), and genital plate also era from each other is connected with the dark (not as in T. ecuador)] and above-men- possibility of independent lose of tegminal tioned characters of male genitalia. From all stridulatory apparatus and tympana in Ta - the other true and possible congeners, the falisca (see Note on this genus above) and new species differs in the same features as T. in the new genus. If this preliminary hy- ecuador and T. mexico. pothesis is erroneous, the new genus may be considered a subgenus of Tafalisca. Genus Mexitrypa gen. nov. Etymology. The generic name is origi- Type species: Mexitrypa ovtshinnikovi nated from Mexico and the genus Metrypa. sp. nov. Diagnosis. Structure of body similar to Mexitrypa ovtshinnikovi sp. nov. that of Tafalisca: head not elongate; prono- (Figs XVII: 1–7) tum wider than its length and with almost Holotype. Male; Mexico, Veracruz, 15– straight hind edge; male metanotal gland 20 km NE of Catemaco Town, Los Tuxtlas (bio- developed (Fig.XVII: 2); legs short and station of Mexico University) in 2 km from coast stout, without any tympana, with tibiae of Mexican Gulf, partly primary and partly sec- hardly longer than femora in fore and mid- ondary forest on hills, on bark of dry tree branch, dle legs, and with hind tibiae having den- at night, 6–17 Nov. 2006; coll. A. Gorochov & ticles rather long and spine-like (Fig. XVII: A. Ovtshinnikov; ZIN. 3); male tegmina without stridulatory ap- Description. Male (holotype). Coloura- paratus; male genitalia similar to those of tion dark brown with following marks: head Amblyrhethus, but with apex of epiphal- with small lightish stripes behind eyes and lus distinctly bifurcate (i. e. with a pair of near lateral ocelli, yellowish areas on mem- lobe-like projections). However this genus branes of antennal cavities, small spot on ge- distinguished from Tafalisca by following nae under eyes, areas on distal half of labrum characters: pronotum distinctly narrowing and on apical segment of maxillary palpi, to head; hind tibiae horter (hind femora grayish brown frons under rostral apex, an- approximately 1.5 times as long as these tennal flagellum and most part of maxillary tibiae); hind basitarsi longer (their length palpi, and light brown rest of mouthparts; intermediate between those of Tafalisca legs with light brown coxae, yellowish dor- and Perutrella in relation to length of hind sal half of fore and middle tibiae (covered tibiae; for comparison see Figs XIV: 8; XVI: by dark dots and diverse somewhat larger 1; XVII: 3); dorsal tegminal field with wider marks), grayish brown hind femora (having areas between longitudinal veins and dense darker apical part, small light subapical dor- cellular venations between them (Fig. sal spot, and larger dark dorsal spot situated XVII: 1); male genitalia with lobe-like hind near it, but slightly proximad), and whitish projections not very large and not inflated very small subbasal mark on hind tibiae; (Fig. XVII: 5–7). tegmina grayish brown with grayish and Included species. Only type species. But whitish marks as in Fig. XVII: 1; hind wings it is impossible to exclude that Tafalisca light grey; rest of thorax grayish brown; ab- lineatipes Bruner, 1916 from Jamaica, T. domen black with light brown anal plate evimon Otte et Perez-Gelabert, 2009 from (having several small dark marks) and small Trinidad I., and some of the other species, lightish spots on cerci. Scape almost 1.5 included now in Tafalisca (Eades et al., times as wide as rostrum between antennal 2011) without study of their male genitalia, cavities; lateral ocelli small; median ocellus may belong to this genus. indistinct. Metanotal gland with central

Figs XVIII (1–9). Brazitrypa longiapex sp. nov. 1, male metanotum from above; 2, male abdominal apex from above; 3–5, male genitalia from above (3), from below (4) and from side (5); 6, female genital plate from below; 7–9, distal part of ovipositor from side (9), and its left half from above (7) and from below (8). Abbreviations: a, possible apodeme of mold of spermatophore attachment plate; ae, apodeme of endoparamere; en, endoparamere; gp, genital plate; s, mold of spermatophore attachment plate; others, as in Figs I: 8–10.

Figs XIX (1–8). Different structures of body. 1, 2, Brazitrypa? haani, male; 3–8, Pseudogryllus elongatus gigas subsp. nov., female. Body from above (1); hind tibia, outer view (2); head in front (3); pronotum from above (4); genital plate from below (5); most part of hind leg, outer view (6); distal part of ovipositor from above (7) and from below (8). [1, 2, after Saussure (1878).]

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 267 concavity distinctly smaller than in all spe- (Fig. XVIII: 1); genitalia with epiphallic an- cies of Tafalisca described here, and com- terolateral parts very long and very strong- pletely crossed by low median keel (Fig. ly curved (Figs XVIII: 3, 5), with epiphallic XVII: 2). Hind tibiae as in Fig. XVII: 3. posterolateral projections moderately long, All wings almost equal in length, extending flat and almost acute (Figs XVIII: 3–5), behind abdominal apex and reaching proxi- and with mold of spermatophore attach- mal part of hind tibiae; tegminal venation as ment plate long or having a long apodeme in Fig. XVII: 1. Abdominal apex with anal (Figs XVIII: 3, 4); ovipositor apex narrowly plate as in photograph (Fig. XVII: 4), geni- rounded or acute (Fig. XVIII: 7–9) (trun- tal plate approximately twice longer than cate or obtusely rounded in Tafalisca). previous plate and narrowing to roundly Included species (named in original bi- angular apex, and genitalia having lateral nomen). B. longiapex sp. nov. (type spe- and ventral parts of hind half of epiphallus cies); Tafalisca paulista Rehn, 1918 and T. membranous as well as ectoparameres more paranaensis Dias et Mello, 2010 from Brazil or less similar to those of T. mexico in shape (Dias & Mello, 2010; Figure 5); possibly (Figs XVII: 5–7). Metrypa haani Saussure, 1874 also from Female unknown. Brazil and some of the other species with Length in mm. Body 19; body with unknown male genitalia from the list of wings 23; pronotum 3.7; tegmina 16; hind Tafalisca species presented now in the elec- femora 12.2. tronic catalogue of Orthoptera (Eades et Comparison. The new species differs al., 2011). from T.? lineatipes and T.? evimon (having Note. If B.? haani comb. nov. belongs to the more or less similar male genitalia) in this genus, the hypothesis about indepen- the lateral parts of epiphallus membranous dent lose of tegminal stridulatory apparatus (from T.? lineatipes) and apical part of in some related genera of Tafaliscina will epiphallus less deeply bifurcate (from T.? be supported by two evidences: presence of evimon) [it is interesting to note that male traces of such apparatus in some species of genitalia of T.? evimon is somewhat similar Tafalisca and in one species of Brazitrypa. to those pictured by Desutter (1988: Fig. Etymology. The generic name is originat- 52–54) as belonging to Amblyrhethus man- ed from Brazil and the genus Metrypa. ni]. From all the other similar species, the new species differs in the characteristic Brazitrypa longiapex sp. nov. (spotted) colouration of tegmina. (Figs XVIII: 1–9) Etymology. The species is named in hon- Holotype. Male; Brazil, “Espirito-Santo Bra- or of A.B. Ovtshinnikov, photographer of in- sil, ex coll. Fruhstorfer”, “haani Sauss.”; MIZ. sects and one of collectors of this specimen. Paratype. Female; same data as for holotype, but without species determination; MIZ. Genus Brazitrypa gen. nov. Description. Male (holotype). Size of body approximately as in Acheta domesticus Type species: Brazitrypa longiapex sp. L. Colouration light brown with head dor- nov. sum and pronotum brown (but each prono- Diagnosis. External body structure simi- tal lateral lobe having light brown longitu- lar to that of Tafalisca and Mexitrypa (in- dinal spot along ventral edge), longitudinal cluding the absence of stridulatory appa- median line on outer surface of hind femora ratus in male and of tympana, and charac- dark, area in apical part of these femora teristic armament of hind tibiae), but male darkened, all veins and area between Sc metanotal gland, male genitalia and apex of and R in tegmina intensively brown, hind ovipositor different: gland with the charac- wings whitish, abdomen from brown to teristic median convexity instead concavity dark brown (excepting light brown cerci).

Head rather high (general shape of body Genus Pseudogryllus Chopard, 1912, more or less cylindrical); rostral apex weak- gen. resurr. ly roundly angular (very obtuse) in profile. Pronotum rather long (approximately as Type species: Pseudogryllus elongatus long as wide) and with almost parallel lat- Chopard, 1912. eral sides; metanotal gland lacking special- Note. This genus was described for a sin- ised hairs (its structure as in Fig. XVIII: gle female from French Guyana (Chopard, 1). Legs stout (but not very short); hind 1912). Later it was synonymised with Ta - tibiae somewhat shorter than hind femora; falisca by the same author (Chopard, 1968). armament of these tibiae approximately as But necessity of this synonymy is problem- in Tafalisca and Mexitrypa; hind basitarsi atic. Pseudogryllus is similar to Tafalisca, much shorter than hind tibiae and with Mexitrypa and Brazitrypa in the character- large denticles (three outer and two in- istic spine-like denticles on the hind tibiae ner) and very large spurs (however these and in the absence of stridulatory apparatus spurs distinctly not reaching apex of third in male tegmina and of tympana; however it segment of hind tarsi, approximately as in differs from them in the clearly larger body as well as significantly longer denticles Figs XIX: 1, 2). Tegmina reaching abdomi- of hind tibiae (which are almost indistin- nal apex and without traces of stridulatory guishable from the true spines) and spurs apparatus (dorsal tegminal field with longi- of hind basitarsi (these spurs almost reach tudinal veins almost parallel to its median the apex of third segment of hind tarsi) (for line and with crossveins moderately dense); comparison see Figs XVI: 1; XVII: 3; XIX: hind wings hardly exposed behind tegmina. 1, 2 and 6). Male genitalia of Pseudogryllus Abdomen with anal plate simple and almost are unstudied, and thus its relationship to triangular, and with genital plate long (Fig. the above-mentioned genera is unclear. So, XVIII: 2); genitalia as in Figs XVIII: 3–5. at present it is most reasonable to consider Female. General appearance as in male, Pseudogryllus a separate genus including but abdomen light brown with dorsum of only type species. hind tergites, anal plate, part of genital plate (Fig. XVIII: 6), and paraprocts darkened. Genital plate as in Fig. XVIII: 6; oviposi- Pseudogryllus elongatus gigas subsp. nov. tor slightly shorter than hind femora, with (Figs XIX: 3–8) acute apex and partly smooth dorsal and Holotype. Female; Colombia, “Rio Coquetá, ventral surfaces of distal part (Figs XVIII: A.S. Woronov”, VII.1926; ZIN. 7–9); colouration of ovipositor brown Description. Female (holotype). General (rather dark), but with yellowish median appearance very similar to that of P. e. elon- stripe running from base of ovipositor to gatus stat. nov. Colouration yellowish with base of its distal part. following marks: arcuate stripe on vertex Comparison. The male described was (Fig. XIX: 3), apical part of all femora, mid- determined by an unknown person as B.? dle and hind tibiae, all tarsi, longitudinal haani, but it differs from this species in the veins of tegmina, and median part of geni- longer wings and absence of traces of strid- tal plate brown; antennal flagellum, fore ulatory apparatus in male tegmina (in B.? tibia, dorsolateral spot near middle of hind haani, wings are significantly not reaching femur (Fig. XIX: 6), and sternites of hind the abdominal apex, and male tegmina are half of body light brown; spot on rostrum with the traces of stridulatory apparatus; (Fig. XIX: 3), a pair of spots on pronotal Fig. XIX: 1). From B. paulista comb. nov. disc (Fig. XIX: 4), all borders of pronotum, and B. paranaensis comb. nov., B. longia- rather large subapical spot on ventrolateral pex is distinguished by the distinctly longer part of hind femur (fused with brown apical male genital plate and epiphallus. part and three narrow longitudinal stripes:

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 216–270 A.V. GOROCHOV. TAXONOMY OF PODOSCIRTINAE: AMERICAN PAROECANTHINI 269 near median line of outer surface and along (project No 10-04-00682), Presidium of the Rus- distal half of each ventral keel; Fig. XIX: 6), sian Academy of Sciences (Program “Biosphere narrow stripes along tegminal R and M, lat- origin and evolution of geo-biological systems”), eral parts of genital plate, and most part of and the Ministry of Education and Science of the Russian Federation. ovipositor (excepting slightly lighter narrow median stripe running from ovipositor base to its middle part) dark brown; rest of REFERENCES tegmina very light, yellowish grey (almost Bruner L. 1916. XV. South American crickets, whitish); exposed part of hind wings grey. Gryllotalpoidea, and Achetoidea. Annals of Shape of head and pronotum as in Fig. XIX: the Carnegie Museum, 10: 344–428. 3, 4; all ocelli developed, medium-sized (al- Brunner-Wattenwyl C. 1873. Disposito most equal to each other in size), situated in generum Gryllodeorum secundum. In: H. shape of triangle. Tibiae of fore and middle Saussure. Système des Gryllides. Par Mr. legs slightly longer than their femora; hind Ch. Brunner de Wattenwyl. Mittheilungen femur approximately 1.6 times as long as der Schweizerischen Entomologischen Gesell- hind tibia; this tibia with very long spines schaft, 4(4):163–170. Chopard L. 1912. Contribution a la faune des and denticles (especially on inner keel); Orthoptères de la Guyane Française (2e mé- hind basitarsus much shorter than hind tib- moire. Gryllidae). Annales de la Société ento- ia, with three outer and one inner denticles mologique de France, 81: 401–432. (Fig. XIX: 6). Tegmina somewhat extending Chopard L. 1956. Some crickets from South behind apex of hind femora, with 7 parallel America (Grylloidea and Tridactyloidea). longitudinal veins in dorsal field and dense Proceedings of the United States National cellular venation between all longitudinal Museum, 106(3366): 241–293. veins; hind wings distinctly exposed behind Chopard L. 1968. Gryllides. Orthopterorum Cat- tegmina. Genital plate and ovipositor as in alogus, 12: 213–500. Figs XIX: 5, 7, 8. Desutter L. 1987. Structure et évolution du complexe phallique des Gryllidea (Orthop- Male unknown. tères) et classification des genres Néotropi- Length in mm. Body 32; body with caux de Grylloidea. Première partie. Annales wings 41; pronotum 6; tegmina 29; hind de la Société entomologique de France (nou- femora 19; ovipositor 19. velle série), 23(3): 213–239. Comparison. The new subspecies dif- Desutter L. 1988. Structure et evolution du fers from P. e. elongatus in the longer spines complexe phallique des Gryllidea (Orthop- and especially denticles on the inner keel of teres) et classification des genres Neotropi- hind tibia (in the new subspecies, they lon- caux de Grylloidea. Deuxieme partie. An- ger than those on the outer keel of this tibia, nales de la Société entomologique de France but in the nominotypical subspecies, arma- (nouvelle série), 24(3): 343–373. Dias P.G.B.S. & Mello F. de A.G. de. 2010. A ments on the both keels of hind tibia are al- new species of Tafalisca Walker, 1869 from most equal in size; see Chopard, 1912: Figs the Iguaçu National Park, Brazil (Grylloi- in p. 413) and presence of three (but not dea, Eneopteridae, Tafaliscinae). Zootaxa, two) outer denticles on the hind basitarsus. 2453: 48–54. Eades D.C., Otte D., Cigliano M.M. & Braun, ACKNOWLEDGEMENTS H. 2011. Orthoptera Species File Online. http://osf2.orthoptera.org/HomePage.aspx I thank all my colleagues and friends help- Gorochov A.V. 1986. On system and morpho- ing me during my work in America and collect- logical evolution of the cricket family Gryl- ing many specimens used for this study. Also I lidae (Orthoptera) with description of new am grateful to K.K. Günther and I. Dorandt taxa. Communication 1. Zoologicheskiy Zhur- (MNH) as well as A. Liana (MIZ) for supplying nal, 65(4): 516–527 (In Russian). the additional material. This work is supported Gorochov A.V. 2002. Taxonomy of Podoscir- by the Russian Foundation for Basic Research tinae (Orthoptera: Gryllidae). Part 1: the

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