A revision of the Chinese Trigonalyidae (Hymenoptera, Trigonalyoidea)

Abstract The Chinese fauna of the family Trigonalyidae Cresson, 1887, is revised, keyed and fully illustrated for the first time. Eight genera of this family (Bakeronymus Rohwer, 1922, Bareogonalos Schulz, 1907, Jezonogonalos Tsuneki, 1991, re-instated, Lycogaster Shuckard, 1841, Orthogonalys Schulz, 1905, Pseudogonalos Schulz, 1906, Taeniogonalos Schulz, 1906 and Teranishia Tsuneki, 1991) are recorded from China. The genus Ischnogonalos Schulz, 1907, is synonymized with Taeniogonalos Schulz, 1906. In total 40 valid species are recognized. Twenty species are new for science: Jezonogonalos elliptifera sp. n., J. jiangliae sp. n., J. luteata sp. n., J. nigrata sp. n., Lycogaster angustula sp. n., L. flavonigrata sp. n., L. nigralva sp. n., Orthogonalys cheni sp. n., O. clypeata sp. n., O. robusta sp. n., Pseudogonalos angusta sp. n., Taeniogonalos bucarinata sp. n., T. cordata sp. n., T. geminata sp. n., T. sculpturata sp. n., T. triangulata sp. n., T. tricolorisoma sp. n., T. uncifera sp. n., Teranishia crenulata sp. n. and T. glabrata sp. n. Two species are reported new for China: Orthogonalys elongata Teranishi, 1929 and Nanogonalos flavocincta Teranishi, 1929 (renamed to Taeniogonalos subtruncata nom. n.). Seven new synonyms are proposed: Poecilogonalos yuasai Teranishi, 1938, and P. maga taiwana Tsuneki, 1991, of Taeniogonalos taihorina (Bischoff, 1914); Taiwanogonalos minima Tsuneki, 1991, and T. similis Tsuneki, 1991, of Taeniogonalos alticola (Tsuneki, 1991); P. intermedia Chen, 1949, and P. unifasciata Chen, 1949, of Taeniogonalos formosana (Bischoff, 1913). Six taxa are recognised as valid species: Bakeronymus seidakka Yamane & Terayama, 1983, Jezonogonalos laeviceps (Tsuneki, 1991), J. satoi (Tsuneki, 1991), Taeniogonalos alticola (Tsuneki, 1991), T. flavoscutellata (Chen, 1949) and T. gestroi (Schulz, 1908). Five new combinations are made: Jezonogonalos laeviceps (Tsuneki, 1991), comb. n., J. satoi (Tsuneki, 1991), comb. n., Taeniogonalos flavoscutellata (Chen, 1949), comb. n., T. gestroi (Schulz, 1908), comb. n. and T. lachrymosa (Westwood, 1874), comb. n. Lectotypes are designated for Lycogaster violaceipennis Chen, 1949, Poecilogonalos flavoscutellata Chen, 1949, P. rufofasciata Chen, 1949, and P. tricolor Chen, 1949.


Introduction
Trigonalyidae (Hymenoptera) is a worldwide small family in its own superfamily Trigonalyiodea, with 93 recognized species (Carmean and Kimsey 1998;Smith and Stocks 2005;Santos et al. 2012;Smith and Tripotin 2012;. Most species of this family occur in tropical and subtropical regions, but only one species occurs in Europe and the family is absent in arctic and alpine habitats (Carmean and Kimsey 1998). Carmean and Kimsey (1998) recognized two subfamilies: Orthogonalinae (corrected by Lelej (2003) to Orthogonalyinae) and Trigonalinae (correctly: Trigonalyinae) from their morphological phylogenetic analysis. The subfamily Trigonalyinae is subdivided in two tribes: Nomadinini and Trigonalini (correctly: Trigonalyini). According to their analysis the Orthogonalyinae are without derived characters and the recognition of the tribe Nomadinini makes the Trigonalyini paraphyletic. Therefore, we do not recognise these higher categories untill there is a better understanding of the evolution within this family. The distribution of the genera of the Trigonalyidae varies, but most genera known from China are wide spread. Bakeronymus occurs only in the Oriental Region and Pseudogonalos is only known from the Palaearctic Region. Jezonogonalos and Teranishia occur in the Oriental and East Palaearctic Regions, Bareogonalos and Lycogaster occur in the Oriental, East Palaearctic, Nearctic and Central American Regions, and Orthogonalys occurs in the Afrotropical, Oriental, East Palaearctic, Nearctic and Neotropical Regions. Taeniogonalos has a similar distribution but Taeniogonalos is also known from the Australian Region. The genus Trigonalys Westwood is most likely restricted to the Neotropical and Afrotropical Regions; the two Oriental species (Carmean and Kimsey 1998) are removed in this paper.
Trigonalyidae are often misidentified; slender specimens (especially of Orthogonalys) with white banded antennae may be mistaken for Ichneumonidae and robust specimens with black antennae for aculeate wasps (e.g. of the family Crabronidae). They can be identified by the combination of the well-developed costal cell of the fore wing (C1 in Fig. 1), the presence of mandibles with 3-5 large teeth (Figs 7,19,39,145,156,243,561,594) and the presence of plantar lobes on the tarsal segments (Fig. 5). In addition, the tarsal claws are cleft (bifurcate with the inner tooth larger than the outer one: Fig. 5) and as pointed out in Carmean and Kimsey (1998) the females have sparse white scales or specialized setae on the outside of the middle antennal segments.
Biologically, species of Trigonalyidae seem not to care about laying the eggs at or in the future food source as is normal for parasitoids. They deposit up to more than 2000 eggs on leaves, which are eaten eventually by caterpillars and sawfly larvae. In the digestive tract the eggs hatch, the mobile larva bores through the intestine wall to search for an eventually present larva of a parasitoid wasp (Ichneumonidae or Braconidae) or of a parasitoid fly (Tachinidae) to develop on it as a hyperparasitoid. The third larval instar has huge mandibles to kill possibly present other parasitoid larvae. Other species are brought into the nests of social Vespidae because they are inside the caterpillars used as prey by the wasps. In the wasp larvae they develop as primary endoparasitoids. Primary endoparasitism of sawflies occurs, but the parasitoid still acts facultatively as a hyperparasitoid (Yamane and Terayama 1983;He and Chen 1986;Weinstein and Austin 1991;Carmean and Kimsey 1998).  Table 1. List of the Chinese species of the family Trigonalyidae before this study with distribution in China.
Sichuan (Oriental) Lycogaster violaceipennis Chen, 1949 Zhejiang, Sichuan (Oriental) Orthogonalys cheni sp. n. Zhejiang, Hubei, Sichuan (Oriental) rently 16 species in 6 genera are known from China (Table 1), of which 15 species are considered valid; one is a synonym and two species are misidentified. In total, 40 valid species occurring in China and belonging to eight genera have been examined during this study. Twenty are new species and described in this paper and two species are new for China (one of which is renamed; Table 2). The actual number will be higher because several species from neighbouring countries are expected to occur also in China (e.g., in the northern provinces of China species known from Japan); in conclusion, the Chinese fauna is still clearly undersampled. All known species of Trigonalyidae from China are revised, illustrated and keyed. Unfortunately, several of the new and described species have males as holotype and the associated female is unknown. Collection at the type localities and the use of COI ("barcoding") will recover the conspecific female.

Material and methods
Descriptions of the species are a combination of comparative illustrations and a short diagnosis and description, followed by a summary of the distribution (only based on examined specimens unless otherwise indicated) and biology (as far as known, at least the phenology is given). Descriptions were made under either an Olympus SZ61 or SZ40 stereoscope, in combination with a 40 W LED lamp or a 27 W fluorescent lamp. Photographic images were made with an Olympus motorized stereomicroscope SZX12 and Stemi 2000-CS; the images were processed with both Image-Pro Plus and AnalySIS Extended Focal Imaging software, and plates were finished with ACDSee 10.0 and Photoshop CS, mostly to adjust the size and background.
Morphology. For other terminology used in this paper, see van Achterberg (1988van Achterberg ( , 1993 and Hong et al. (2011). Measurements are taken as indicated by van Achterberg (1988); width of the propodeal foramen is measured dorsally and its medial height in lateral view. Additional non-exclusive characters in the key are between brackets.
Material. Types and other specimens have been examined from the following institutions:

Species Distribution in China
Taeniogonalos triangulata sp. n.

Key to Oriental and Palaearctic genera of Trigonalyidae
Diagnosis. Body length 7.7-10.1 mm; antenna with 13-15 segments, of female third antennal segment 0.8 times as long as fourth segment and slender (Fig. 9); apical half of antenna of female and of male with hardly moniliform and symmetric segments (Fig. 9); male antenna without tyloids; antennal sockets closer to eyes than to each other; vertex with medio-longitudinal depression dorsally, widened anteriorly and posteriorly (Fig. 8); anterior ocellus slightly lower situated than posterior ocelli in depression (Fig. 8); clypeus not differentiated from face; mandibles narrow in anterior view, mandibular condyli far from eye and submedially attached to head (Fig. 10); maxillary and labial palpi with 4 and 2 segments, respectively and apical segment of labial palp slender and tapered; maxillary palp about 0.7 times as long as mandible and with distinct segments (Fig. 11); vein 1-SR of fore wing short (Fig. 17); hind basitarsus enlarged; hind trochanter without oblique groove; propodeal foramen rounded dorsally and with lamelliform rim (Fig. 12); first tergite subpetiolate and comparatively narrow basally (Fig. 14); protuberance of second sternite of ♀ small and forked, and of third sternite large and obtuse. Biology. Reared from the nest of Polistinae (Vespidae) (Yamane and Terayama 1983 Diagnosis. Eyes remain distinctly separated from mandibular condyli; fourth antennal segment slightly widened apically; middle lobe of mesoscutum with large yellow patch; truncate apical part of scutellum 0.7 times as wide as scutellum anteriorly; fore wing with three submarginal cells; apical half of hind basitarsus ivory; propodeum reticulate-rugose, with short rugae interconnecting transverse rugae. Description. Female paratype, length of body 10.1 mm (of fore wing 9.3 mm).
Metasoma. First tergite 0.9 times as long as apically wide, gradually narrowed basally, with large elliptical depression medially and concave apically; second tergite and sternite smooth; hypopygium incised apically.
Colour. Blackish brown or dark brown with rich yellow pattern (including mesopleuron; Figs 6,9,12,13,15); palpi and antenna (except largely yellow scapus) rather dark brown; patch on coxae, fore and middle trochanters partly, apex of fore and middle femora, fore tibia, base of middle and hind tibiae, all tarsi more or less pale yellow; pterostigma and marginal cell of fore wing and surroundings dark brown; remainder of wing membrane slightly infuscate.
Distribution. China (Taiwan). Notes. Only one species is known from China: B. seidakka Yamane & Terayama, 1983, re-instated. It was described as a subspecies of B. typicus Rohwer, 1922, from the Philippines, but comparison with specimens of this species with a paratype of B. seidakka shows considerable differences. For instance, the middle lobe of mesoscutum is entirely dark brown; the truncate apical part of the scutellum half as wide as the scutellum anteriorly; the fore wing has only two submarginal cells; the apical half of hind basitarsus is dark brown; the eyes are close to the mandibular condyli; the fourth antennal segment is distinctly widened apically and the propodeum has transverse rugae, without short interconnecting rugae. Therefore, we treat the taxon from Taiwan as an independent species, despite that some of the differences may be sex-related (e.g. the distance between the eye and the mandibular condylus). Schulz, 1907 http://species-id.net/wiki/Bareogonalos Figs 18-28 Bareogonalos Schulz, 1907: 18;Marshakov 1981: 104;Tsuneki 1991: 9;Weinstein and Austin 1991: 412;Carmean and Kimsey 1998: 60. Type species (by original designation): Trigonalys canadensis Harrington, 1896. Nippogonalos Uchida, 1929: 79;Tsuneki 1991: 4;Lelej 1995: 12;Weinstein and Austin 1991: 412. Type species (by original designation): Nippogonalos jezoensis Uchida, 1929. Synonymized by Bischoff 1938. Diagnosis. Body length 8-13 mm; antenna with 18-23 segments; antenna of ♂ without tyloids; area above supra-antennal elevations flat, superficially finely rugulose and elevations small or nearly absent (Fig. 19); apical segment of labial palp widened and obtuse, more or less triangular (Fig. 22); mandibular condyli remain far removed from eyes (Fig. 22); vertex normal, at most with slight median depression dorsally (Fig. 20); head with long setae, transverse and flattened dorsally (Fig. 20); metanotum weakly convex and spaced rugose medially (in B. huisuni: Fig. 23) or strongly lamelliform protuberant and densely sculptured; propodeum strongly areolate-rugose; hind trochanter simple, its dorsal triangular part not separated from basal part, but rarely partly differentiated (Tsuneki 1991); hind tarsus slightly or not modified; metasoma smooth and usually shiny or finely and rather densely punctate and rather dull; vein 1-SR of fore wing medium-sized to long (Fig. 28); protuberance of third sternite of ♀ similarly sized as protuberance of second sternite in lateral view or somewhat smaller (Fig. 26). Biology. Reared from Vespa, Vespula, Dolichovespula and Provespa spp. (Vespinae: Vespidae); the larva of at least one species has a final ectoparasitoid phase (Carmean 1991;Carmean and Kimsey 1998). Collected in August-October.

Bareogonalos
Notes. As pointed out by Tsuneki (1991) Nippogonalos differs by having the head much narrower than the mesoscutum (nearly as wide as mesoscutum in typical Bareogonalos). The name Nippogonalos could be used as subgeneric name in Bareogonalos for the species with an enlarged mesosoma resulting in a comparatively small head.

Key to Old World species of Bareogonalos Schulz, 1907
1 Metanotum with a thick and medially depressed protuberance and its setae long; head distinctly narrower than mesoscutum; anterior half of scutellum flat and at same level as mesoscutum; fourth and fifth tergites largely smooth and shiny; Japan Diagnosis. Head nearly as wide as mesoscutum (Fig. 23); anterior half of scutellum distinctly convex and above level of mesoscutum; metanotum weakly convex and without protuberance, its setae medium-sized (Fig. 23); protuberance of second sternite of ♀ densely setose and widely truncate medially (Fig. 26); metasoma largely finely and rather densely punctate and rather dull (Figs 25,27).
Description. Female paratype, length of body 11.0 mm (of fore wing 10.7 mm).
Head. Antenna with 20 segments, segments of apical half about as long as wide (Fig. 21); frons and vertex smooth and strongly shiny (Figs 19,20), with long yellowish setae; head gradually narrowed behind eyes and nearly as wide as mesoscutum (Fig. 23); dorsal length of eye 1.7 times length of temple (Fig. 20); temple smooth and shiny; occipital carina narrow lamelliform medio-dorsally and with a short crenula; supra-antennal elevations hardly developed as a thin rim and smooth; clypeus moderately emarginate and thick medio-ventrally.
Colour. Black or dark brown (including palpi) with yellow pattern (including mesopleuron; Figs 18,23,24,26); mandible largely chestnut brown; scapus and pedicellus yellow, third and following segments brown, but apical half of antenna dark brown; pronotum dorso-apically and dorsally, pair of wide patches on mesoscutum anteriorly, scutellum (except posteriorly), axilla partly, lateral patch of propodeum, basal 0.4 of second tergite (except small anterior patch and laterally), anterior quarter of 3 rd -5 th tergites (except laterally) and narrow median stripe of 4 th -7 th tergites, apex of fore and middle femora, base of fore and middle tibiae, yellow; hind coxa dorso-apically, inner side of middle trochanter and hind trochanter and trochantellus pale yellow; pterostigma and veins 1-SR, 1-M and more distal veins of fore wing light brown; membrane of anterior half of fore wing rather dark brown, remainder of wing membrane slightly infuscate.
Colour. Black; inner orbita narrowly orange-brown; apical half of supra-antennal elevations yellowish brown, clypeus with a narrow transverse orange-brown patch; minute patch of outer orbita, vertex posteriorly, occipital carina medially and mandible largely yellowish brown (Figs 39,40,43); mesosoma laterally black except for brown dorsal rim of pronotal side; middle lobe of mesoscutum with pair of narrow brown patches anteriorly; axilla with medium-sized brown patch; metanotum with pair of small lateral yellowish brown patches; metasoma dorsally black, with narrow orange-brown apical band at apex of all tergites and first two sternites (Figs 46,47,48); palpi dark brown; antenna brown, becoming darker apically; legs dark brown to black; pterostigma brown, remainder of wing membrane subhyaline.
Variation. Length of body 6.6-9.8 mm, of fore wing 5.8-8.7 mm; propodeum with pair of small brown spots; vertex coarsely reticulate-punctate; temple rugulosepunctate; second tergite with dense fine punctures posteriorly; length of vein 1-M of fore wing 1.5-1.6 times as long as vein 1-SR.
Male. Length of body 8.3-9.1 mm, of fore wing 6.8-7.8 mm; antenna with 23 segments, tyloids elliptical, 0.25 times as long as segment on 11 th -13 th segments and nearly circular on 14 th and 15 th segments; in general males are darker than females: vertex and outer orbita entirely black; mesosoma with or without brown patches; first and third tergites sometimes without narrow orange-brown apical band; genitalia extruded ( Fig. 52).
Head. Antenna incomplete, tyloids nearly circular, 0.1 times as long as segment on 10 th segment and 0.2 times as long as segments on 11 th -15 th segments (Fig. 56); frons rugose ( Fig. 54); vertex and temple largely smooth and shiny with sparse and fine punctures (Figs 55,58); head gradually narrowed behind eyes, eye in dorsal view 0.9 times as long as temple (Fig. 55); occipital carina strongly widened and lamelliform,  extensively crenulate dorsally; supra-antennal elevations strongly enlarged (about 0.7 times as long as scapus), outer side subvertical and largely smooth except for sparse punctures; clypeus slightly concave and thick medio-ventrally.
Metasoma. First tergite 1.1 times as long as apically wide, smooth and with distinct elliptical depression antero-medially, slightly convex posteriorly (Fig. 61); second-fifth tergites and all sternites largely smooth except for sparse superficial punctures ( Colour. Black; inner orbita narrowly, apex of supra-antennal elevation and malar space ivory (Fig. 54); mandibular teeth, palpi and tegulae dark brown; tibiae and tarsi rather brownish; pterostigma and apical half of first submarginal cell to anterior half of marginal cell of fore wing and area below that dark brown, remainder of wing membrane subhyaline.
Female. Unknown. Biology. Unknown. Collected in June.
Head. Antenna unknown [male antenna with 21 segments (including scapus) glued on card with longitudinal tyloids on 10 th -15 th segments is most likely of holotype of T. alticola]; frons spaced punctulate; vertex largely smooth, sparsely punctulate and strongly shiny (Fig. 66), with rather short setae; temple largely smooth (Fig. 67); head gradually narrowed behind eyes, eye in dorsal view as long as temple (Fig. 66); occipital carina wide lamelliform medio-dorsally with one short carina and few short sublateral carinae; supra-antennal elevations strongly enlarged (about 0.6 times as long as scapus), outer side subvertical and largely smooth except for sparse punctures, latero-basally with pit-shaped depression (Fig. 66); clypeus slightly concave and thick medio-ventrally.
Metasoma. First tergite 0.7 times as long as apically wide, smooth and with indistinct depression basally (Fig. 71); second and following tergites smooth and strongly shiny (Fig. 71); sternites mainly smooth; second sternite weakly curved in lateral view; third sternite about half as long as second sternite (Fig. 72).
Colour. Black or dark brown; mandible (except apically), malar space largely, inner orbita narrowly, pronotum latero-dorsally, minute lateral patch of scutellum, posterior half of first segment largely (except medially), large patch laterally of second tergite, trochanters, trochantelli and base of femora ivory; palpi and tegulae yellowish brown; remainder of legs more or less brown or dark brown; pterostigma (but basally yellowish), veins and marginal cell (except apically) dark brown or brown; remainder of wing membrane subhyaline.
Head. Antenna with 25 segments, tyloids circular, 0.5 times as long as segment on 11 th -14 th segments and 0.2 times as long as segments on 10 th and 16 th segments (Fig.  77); frons densely punctate; vertex spaced and finely punctate (interspaces much wider than width of punctures) (Fig. 75), with short setae; temple largely smooth with sparse fine punctures (Fig. 79); head gradually narrowed behind eyes, eye in dorsal view 1.2 times as long as temple (Fig. 75); occipital carina moderately widened and lamelliform medio-dorsally, with many short carinae; supra-antennal elevations strongly enlarged  (about 0.7 times as long as scapus), outer side subvertical and largely smooth except for sparse punctures; clypeus slightly concave and thick medio-ventrally.
Colour. Black; mandible largely except brown teeth, inner orbita narrowly, apical half of supra-antennal elevations, pair of wide patches of clypeus, malar space pair of narrow patches of middle lobe of mesoscutum antero-laterally, small medial patch of metanotum, pair of small lateral patches on first and second tergites, apical margin of first and second sternites, base of fore trochanter and trochantellus, fore and hind trochanters and trochantelli ivory (Figs 74,75,80,83,84); metasoma ventrally black; palpi and antenna dark brown; ivory, fore tarsi and tibia yellowish brown, remainder of legs dark brown to black; pterostigma and anterior half of marginal cell dark brown, remainder of wing membrane subhyaline.
Metasoma. First tergite 0.6 times as long as apically wide, smooth and with shallow elliptical depression medially (Fig. 93); second to sixth tergites largely smooth with sparse and fine punctures; sternites densely and finely punctate; second sternite slightly convex in lateral view; third sternite about 0.3 times as long as second sternite (Fig.  95); hypopygium triangular in ventral view (Fig. 95).
Colour. Black; malar space, lateral margin of all tergites and posterior margin of second sternite ivory; teeth of mandible, antenna and tegulae dark brown; second tergite with pair of small pale yellow spots postero-laterally; legs dark brown to black; pterostigma and posterior half of first submarginal cell to anterior half of marginal cell and area below that dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 8.6-11.0 mm, of fore wing 6.8-8.7 mm; length of vein 1-M of fore wing 1.1-1.6 times as long as vein 1-SR; second to sixth tergites largely superficially coriaceous with sparse and fine punctures.
wing with more or less conspicuous dark brown patch below pterostigma (Fig. 101); metasoma of ♂ black or dark brown dorsally or nearly so (Fig. 105); first tergite 0.8 times as long as its apical width (Fig. 105); third sternite about 0.4 times as long as second sternite (Fig. 107).
Wings. Fore wing: length of vein 1-M 2.2 times as long as vein 1-SR and distinctly curved; veins r and 2-SR long (Fig. 101); second submarginal cell of fore wing medium-sized, parallel-sided and laterally about as high as wide anteriorly.
Metasoma. First tergite 0.8 times as long as apically wide, smooth and with wide depression basally (Fig. 105); second and following tergites largely smooth and strongly shiny (Fig. 105); sternites smooth, except for superficial punctures; second sternite weakly curved in lateral view; third sternite 0.4 times as long as second sternite (Fig. 107).
Male Diagnosis. Body length 5.5-15.0 mm; antenna with 22-24 segments, of ♀ widened medially ( Fig. 111, but hardly so in L. angustula sp. n.); antenna of ♂ without tyloids; supra-antennal elevations small, without depression dorsally (Figs 110,121,132,146); vertex convex and shiny; mandibular condyli close to level of eyes (Figs 113,122,135,149); apical segment of labial palp widened and obtuse, more or less triangular (Fig.  122); metanotum smooth, shiny and weakly convex (Figs 114,123,136,150); triangular dorso-apical part of hind trochanter separated by an oblique groove; fore trochanter subparallel-sided and distinctly longer than hind trochanter; hind tarsus slightly or not modified; basal half of third metasomal sternite with a posteriorly steep, smooth and complete transverse ledge (Figs 118,129,140; may be partly hidden under second sternite and rather low in L. violaceipennis); second sternite with pair of small triangular teeth on apical protuberance ( Fig. 118; but only with pair of lobe-shaped flaps in male of L. violaceipennis) and sometimes absent; epipleura of tergites laterally strongly pigmented; fifth sternite of ♀ distinctly emarginate medio-posteriorly (Fig. 118). Biology. In the New World reared as hyperparasitoid of Ichneumonidae in caterpillars of the families Saturniidae and Notodontidae (Carmean and Kimsey 1998). The record from Eumeninae nests probably concerns prey caterpillars that have been infested with both Ichneumonidae and Trigonalyidae.
Notes. It is clear from the original description (Magretti 1897) and from the redescription of Schulz (1908) that L. rufiventris (Magretti, 1897) from Myanmar was correctly assigned to the genus Lycogaster by Schulz (1908), Bischoff (1938) and Weinstein and Austin (1991). It has the medial third of the fore wing dark brown as L. flavonigrata and nigralva, but differs by the entire reddish metasoma, the absence of distinct separate teeth besides finely dentate lamelliform protuberance of second sternite and the distinctly convex vertex. The interpretation by Carmean and Kimsey (1998; based on specimens from central and southern India and retaining this species in the genus Trigonalys) seems to be incorrect considering the redescription of the holotype female by Schulz (1908)  Description. Holotype, female, length of body 6.7 mm (of fore wing 5.4 mm).
Colour. Black; metasoma somewhat brownish; mandibular teeth, palpi and tegulae dark brown; legs dark brown to black with tibiae and tarsi paler; pterostigma dark brown; wing membrane subhyaline.
Variation. Length of body 7.9 mm, of fore wing 6.1 mm; second tergite with pair of small ivory spots medio-laterally; second-fifth tergites rather densely and coarsely punctate.
Male. Length of body 8.7 mm, of fore wing 6.7 mm; antenna with 23 segments and without tyloids; vertex sparsely and rather coarsely punctate; sculpture of mesosoma coarser than that in female; second-fifth tergites and sternites densely and coarsely punctate; genitalia extruded.
Head. Antenna with 23 segments, segments after third segment distinctly widened and after 14 th segments becoming gradually slenderer (Fig. 124); frons coarsely and rather densely punctate and vertex spaced punctate (interspaces often much wider than width of punctures) and shiny (Figs 120,121), with short greyish setae; head subparallel-sided behind eyes, distinctly narrowed posteriorly and 1.3 times wider than mesoscutum (Fig. 121); dorsal length of eye 0.9 times length of temple (Fig. 121); temple smooth, shiny and with medium-sized whitish setae; occipital carina narrow lamelliform medio-dorsally and without crenula; supra-antennal elevations hardly developed as a thin rim and smooth; clypeus weakly emarginate and comparatively thin medio-ventrally.
Colour. Black (including mandible); palpi, fore tibia and tarsus, tegulae, pterostigma and veins dark brown; apex of middle trochantellus, hind trochanter and trochantellus, and narrowly base of hind femur ivory; pair of large lateral patches on second and third tergites and apically on second sternite yellow; fore wing largely dark brown but paler subbasally (Fig. 125).
Variation. Length of body 11.5-13.1 mm, of fore wing 8.9-10.0 mm; length of vein 1-M 1.9-2.1 times as long as vein 1-SR; outer side of fore tibia ivory; vein m-cu of fore wing subinterstitial or postfurcal by 0.2 times length of vein m-cu; third tergite with a pair of small to medium-sized yellow or ivory patches laterally.

Male. Unknown.
Biology. Unknown. Collected in April, July, August and October at 650-850 m. Distribution. China (Fujian, Jiangxi, Yunnan). Etymology. Name derived from "flavus" (Latin for "yellow) and "niger" (Latin for "black") because of the yellow-black pattern of the second sternite.
Head. Antenna with 23 segments, segments after third segment distinctly widened and after 14 th segments becoming gradually slenderer (Fig. 133); frons coarsely and rather densely punctate and vertex spaced punctate (interspaces often much wider than width of punctures) and shiny (Figs 131,132), with short greyish setae; head subparallel-sided behind eyes, distinctly narrowed posteriorly and 1.1 times wider than mesoscutum (Fig. 132); dorsal length of eye 0.7 times length of temple (Fig. 132); temple smooth, shiny and with medium-sized whitish setae; occipital carina narrow lamelliform medio-dorsally and without crenula; supra-antennal elevations hardly developed as a thin rim and smooth; clypeus weakly emarginate and comparatively thin medio-ventrally.
Colour. Black (including mandible); apex of all trochanters ivory, all tarsus, tegulae, pterostigma and veins dark; pair of large lateral patches on second tergite and small patches on basal third tergite and apically on second sternite yellow; fore wing largely brown but paler subbasally (Fig. 134
Colour. Black; mandible largely ivory except dark brown teeth; palpi, pterostigma and veins dark brown; mesosoma largely reddish brown except for ventral half of mesopleuron and metapleuron and posterior half of propodeum dark brown to black; outer side of fore femur, large part of middle and hind femur, all coxae and tarsus dark brown, remainder of legs ivory; pair of large lateral patches on second tergite yellow; fore wing largely dark brown, but paler subbasally (Fig. 148). Variation. Length of body 12.3 mm, of fore wing 9.7 mm; legs darker; length of vein 1-M of fore wing 2.0 times as long as vein 1-SR.
Colour. Black; mandible largely (but teeth dark brown), entire inner orbita narrowly, narrow stripes behind eyes posteriorly on vertex, large posterior spot on middle lobe of mesoscutum, broad median longitudinal stripe on scutellum, large median spot on metanotum, stripe on upper pronotum and lower pronotum, broad patch on lower mesopleuron, small spot on metapleuron, large lateral spots on propodeum, coxae dorsally, fore and mid trochanters and trochantelli, postero-lateral margin of second-fifth tergites and metasoma ventrally largely, ivory; clypeus black; metasoma dorsally largely dark brown, apical 0.7 of first and second tergite brownish; anterior margin of sternites with narrow transverse dark brown band; palpi yellowish brown; antenna black with a small subapical ivory band; coxae dark brown ventrally; remainder of legs yellowish brown; pterostigma dark brown; wing membrane subhyaline.
Variation. Length of body 5.8-7.0 mm, of fore wing 5.1-6.2 mm; ivory stripe on posterior vertex reduced into small spot or absent; lateral ivory spots on propodeum large or reduced; ivory patch on lower mesopleuron small; metasoma largely dark brown to black with first and second tergites and sternites largely yellowish brown; clypeus with pair of ivory spots laterally; legs darker; length of vein 1-M of fore wing 2.0-2.3 times as long as vein 1-SR.
Male. Length of body 5.6-8.1 mm, of fore wing 4.5-6.3 mm; antenna with 24-25 segments; ivory stripe along inner orbita narrow or broad; ivory stripe along outer orbita reduced or extending to malar space; clypeus largely dark brown with pair of ivory spots, largely ivory or entirely ivory; upper mesopleuron with ivory spot or absent; propodeum with large lateral spots or absent; legs and metasoma similar to that of female; genitalia extruded ( Fig. 167).
Colour. Head black with inner orbita ivory up to malar space; mesosoma black; metasoma largely dark brown to nearly black, posterior 0.3 of first tergite and sternite, posterior 0.8 of second tergite, entirely third tergite and second sternite, anterior half of third tergite reddish-yellow; mandible largely ivory with teeth reddish-yellow to dark brown; palpi yellowish brown; antenna black to dark brown; trochanters and trochantelli ivory, coxae and hind femur black, remainder of legs yellowish brown with tarsi paler; pterostigma brown; wing membrane subhyaline.
Variation. Length of body 7.8-9.6 mm, of fore wing 6.9-8.4 mm; frons sparsely and superficially punctate; vertex with small ivory spots posteriorly; clypeus with small yellowish brown spot medially or ivory ventrally; length of vein 1-M of fore wing 2.1-2.3 times as long as vein 1-SR.
Description. Redescribed after a female from Sichuan, length of body 7.4 mm (of fore wing 6.2 mm).
Colour. Black; mandible largely (but teeth dark brown), broad stripes of inner and outer orbita, clypeus largely (except for median dark brown spot), large posterior spot on middle lobe of mesoscutum, broad median longitudinal stripe on scutellum, large median spot on metanotum, stripe on upper pronotum and lower pronotum, oblique stripe on upper mesopleuron and broad patch on lower mesopleuron, small spot on metapleuron, large lateral spots on propodeum, trochanters and trochantelli and outer side of coxae, posterior and lateral margin of all metasomal tergites and sternites, ivory or pale yellow; remainder of metasoma largely dark brown to nearly black; palpi yellowish brown; antenna black with a large ivory band subapically; fore and mid tibiae and all tarsi yellowish brown, remainder of legs dark brown to nearly black; pterostigma dark brown; wing membrane subhyaline.
Variation. Length of body 5.8-8.3 mm, of fore wing 4.7-7.4 mm; upper mesopleuron with large interrupted ivory patch or an oblique stripe or mesosoma laterally entirely (Japan) or largely black; ivory patch of mesoscutum present or absent; ivory patches on posterior and lateral margin of all tergites and sternites comparatively small or large; clypeus entirely ivory or with dark median spot; inner and outer orbita widely ivory but largely black in holotype of O. debilis; hind coxa entirely dark brown or largely ivory dorsally; length of vein 1-M of fore wing 1.9-2.5 times as long as vein 1-SR.  Male. Length of body 7.4 mm, of fore wing 6.0 mm; antenna with 24 segments; ivory stripe along outer orbita reduced or wide (as in holotype of O. hirasana); lateral mesosoma and propodeum entirely black; metasoma largely black, only with posterolateral margin of first and second tergites and sternites dark brown (but absent in holotype of O. hirasana); ventral half of clypeus ivory; genitalia extruded (Fig. 219).
Head. Antenna incomplete, with 26 segments according to original description; frons and vertex coarsely punctate and partly with smooth interspaces equal to width  of punctures or wider; temple smooth; head subparallel-sided behind eyes, eye in dorsal view 1.1 times as long as temple (Fig. 222); occipital carina narrow, non-lamelliform and smooth dorsally (Fig. 222); supra-antennal elevations medium-sized (about 0.5 times as long as scapus), outer side oblique and punctate, apically smooth and nonlamelliform (Fig. 222); clypeus nearly straight and thick medio-ventrally.
Wings. Fore wing: length of vein 1-M 2.7 times as long as vein 1-SR (Fig. 224); second submarginal cell 1.6 times longer than third cell.
Colour. Black; mandible, clypeus, face, curved patch behind eye dorsally, triangular patch behind stemmaticum, ventral half of outer orbita and inner orbita largely and widely, large posterior patch on middle lobe of mesoscutum and on scutellum, metanotum medially, large patches on propodeum medio-posteriorly and laterally, pronotal side dorsally and ventrally, two large patches on mesopleuron and large patch on metapleuron, first metasomal tergite and sternite largely, second tergite laterally, second sternite largely, second-fourth tergites latero-posteriorly, fifth and sixth tergites posteriorly, seventh tergite, third-fifth sternites laterally, pale yellow or ivory; remainder of metasoma dark brown; antenna black with a wide ivory band (apex of 10 th and 12 th -19 th segments) subapically and scapus and pedicellus largely dark brown; fore leg missing; middle and hind legs brownish-yellow, but coxae narrowly basally, hind tibia (except basally) and hind tarsus dark brown; pterostigma and veins dark brown; wing membrane subhyaline.
Colour. Black; mandible largely (but teeth orange-brown or black), two patches at inner orbita, clypeus faintly medially, large posterior spot on middle lobe of mesoscutum, broad median longitudinal stripe on scutellum, large median spot on metanotum, small posterior spot and large lateral spots on propodeum, narrow stripe on posterior  margin of first metasomal tergite and anterior margin of second tergite, almost entirely fifth and sixth tergites and first and second sternites and antero-latero margin of third sternite, ivory; palpi yellowish brown; basal half of antenna black, medially ivory and dark brown apically (Fig. 234); femur, tibia and tarsus of fore leg and tarsus of hind leg yellowish brown, remainder of legs dark brown to nearly black; pterostigma of fore wing dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 9.0-12.6 mm, of fore wing 7.7-10.8 mm; antenna of ♀ with 27-29 segments; frons without ivory stripes or stripes continuous, clypeus with pair of ivory spots, entirely ivory or black, vertex and outer orbita with ivory spots or stripes; pronotal side with ivory stripe antero-dorsally; propodeum entirely black; antenna medially orange-yellow; legs darker or paler; length of vein 1-M of fore wing 1.2-3.1 times as long as vein 1-SR.
Male. Unknown. Biology. Unknown. Collected in June and August at 1500-1760 m. Distribution. China (Shaanxi, Guangxi). Etymology. Named after the comparatively robust body of this species: from "robustus" (Latin for "hard and strong like an oak").  (1845) and later authors, and this action could made it available according to Article 11.6.1 of the ICZN Code (1999: "However, if such a name published as a junior synonym had been treated before 1961 as an available name and either adopted as the name of a taxon or treated as a senior homonym, it is made available thereby but dates from its first publication as a synonym"). However, these authors refer to  Schulz, 1906: 209;Weinstein and Austin 1991: 424;Tsuneki 1991: 3;Lelej 1995: 12;Carmean and Kimsey 1998: 72. Type species (by monotypy): Trigonalis (!) hahnii Spinola, 1840.
Colour. Black; malar space nearly black, somewhat brownish; mandibular teeth, palpi, tegulae, fore and mid tibiae and tarsi, pterostigma and anterior half of marginal cell of fore wing dark brown, remainder of wing membrane pale brown.
Male  Diagnosis. Notauli comparatively wide and coarsely crenulate; malar space ivory; middle mesoscutal lobe less convex than of P. angusta and rather matt, largely densely punctate.
Description. Redescribed after a female from Liaoning, length of body 10.9 mm (of fore wing 8.2 mm).
Colour. Black; malar space ivory; mandibular teeth, palpi, tegulae, fore and mid tibiae and tarsi dark brown; pterostigma and posterior half of first submarginal cell to anterior third of marginal cell of fore wing and area below it dark brown, remainder of wing membrane pale brown.
Biology. Hyperparasitoid of Ichneumonidae in caterpillars and reared from Diprionid sawflies (Carmean and Kimsey 1998). Collected in China in April, June and July.
Notes. The enigmatic genus Ischnogonalos Schulz, 1907, is synonymized; it fits well in the genus Taeniogonalos as redefined in this paper. Unfortunately, the holotype of the type species could not be found. It is aberrant because of the comparatively slender first metasomal tergite, the posteriorly protruding propodeum, the swollen male antennal segments and the pair of minute teeth medio-posteriorly on the second metasomal sternite.
Description. Holotype of T. alticola, male, length of body 6.0 mm (of fore wing 5.4 mm).
Female. Unknown. Variation. Holotypes of T. minima and of T. similis are very similar to the holotype of T. alticola. T. minima has the antenna with 22 segments and longitudinal tyloids on 10 th -13 th segments (Fig. 283); mesoscutal lobes largely smooth and shiny or rugose; mesopleuron smooth (except antero-dorsally) to obliquely finely rugulose; seventh tergite brown or ivory; length of body 4.8-6.0 mm and of fore wing 4.4-5.4 mm.
Description. Holotype, female, length of body 5.9 mm (of fore wing 5.2 mm). Head. Antenna with 22 segments; frons spaced punctate (interspaces about equal to width of punctures or less); vertex largely smooth except some superficial punctures (Fig. 305), with medium-sized setae; temple largely smooth except some sparse punctation (Fig. 308); head gradually narrowed behind eyes, eye in dorsal view 1.5 times as long as temple (Fig. 305); occipital carina strongly widened (medio-dorsally about 1.5 times as wide as posterior ocellus) and lamelliform medio-dorsally, with one strong median carina and few weak carinae laterally; supra-antennal elevations medium-sized (about 0.4 times as long as scapus) and their outer side oblique (Fig. 305); clypeus moderately concave and thick medio-ventrally.
Colour. Black; apex of scapus, inner and outer orbita narrowly, middle mesoscutal lobe antero-laterally, small anterior spot of lateral lobe, metanotum with small lateral spot, first tergite with faint patch postero-laterally and second tergite with large postero-lateral patch yellow (Fig. 311); apical 0.4 of antenna, tegulae, remainder of scapus and basally palpi dark brown, remainder of antenna and palpi apically brown; mandible dark brown but subbasally and near base of teeth brown; apex of trochanters narrowly and trochantelli largely pale yellow and remainder dark brown; fore femur apically and most of tibia yellowish brown, remainder of fore leg dark brown; apex of metasoma narrowly brown; apical patch of fore wing, pterostigma and veins dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 4.9-10.4 mm, of fore wing 4.7-9.0 mm; antenna of ♀ with 21 (1), 22 (10), 23 (6) or 24 (2) segments; yellow patches of inner orbita wide triangular or linear, mandible brown to largely yellow; lateral brownish patch of vertex present or absent; patch lateral mesoscutal lobes postero-laterally and axilla black or yellow; metanotum black or yellow medially and its lateral patch narrow or wide; propodeum black or with pair of large yellow patches; tegulae dark brown, brown or yellow; first and second sternites latero-posteriorly black or orange-brown; only first and second tergites or all tergites yellow or orange-brown latero-posteriorly, patches sometimes connected except for a small brown triangle; second tergite largely smooth to largely moderately densely punctate; vertex largely smooth except some superficial punctures to largely coarsely punctate; length of vein 1-M of fore wing 1.3-1.6 times as long as vein 1-SR; width of depression in front of occipital carina medio-dorsally 1.0-1.5 times width of posterior ocellus and crenulation varies from strong to weak. Male. Length of body 5.0-8.4 mm, of fore wing 4.2-7.7 mm; antenna with 21 (1), 22 (16), 23 (10) or 24 (3) segments, tyloids linear, 0.7 times as long as segment on 11 th -14 th segments and about half as long on 10 th and 15 th segments with a remnant on 16 th segment; genitalia internal, sometimes extruded (Fig. 316); in general males are darker than females: head entirely black or orbita narrowly yellow and mesosoma black or pronotum dorsally slightly yellow; tegulae brown or yellow; metasoma coloured as holotype or darker; sculpture varies as in females, but in general less strongly developed.
Colour. Black; head anteriorly with moderate orange-brown pattern (Fig. 318); remainder of head black except for yellowish brown outer orbita (Fig. 322); mesosoma laterally black except for ventral and dorsal brown small patches of pronotal side and middle mesoscutal lobe strongly contrasting with both lateral lobes, yellow laterally and orange-brown medially (Fig. 323); scutellum largely orange-brown, only medio-posteriorly and narrowly black laterally (Fig. 323); axilla with small brown patch; metasoma dorsally black, first three tergites with posterior orange-brown stripes, other tergites with large orange-brown patch medially; metasoma ventrally black (Fig. 323), first sternite with orange posterior brown stripes black basally, second and third sternites with pair of small orange-brown patches latero-posteriorly; palpi and antenna (but ventrally brown) dark brown; patch on middle and hind coxae, all trochanters partly, apex of all femora, fore and middle tibia tarsi dark brown, remainder of legs black; pterostigma and anterior half of fore wing largely dark brown (Fig. 321), remainder of wing membrane hyaline.

Male. Unknown.
Biology. Unknown. Collected in June. Distribution. China (Yunnan). Etymology. From "cordatus" (Latin for "heart-shaped") after the cordate shape of the yellow and orange-brown middle lobe of the mesoscutum.
Wings. Fore wing: length of vein 1-M 1.5 times as long as vein 1-SR (Fig. 338); second submarginal cell slightly longer than third cell.
Colour. Blackish brown; mandible, triangular patch along inner orbita, supra-antennal elevations, nearly entire clypeus, medium-sized stripe along outer orbita and interrupted transverse stripe on posterior vertex, anterior margin of pronotum broadly and dorsally with a narrow stripe, metanotum partly, fore coxa ventrally, middle coxa dorsally largely, trochanters largely, trochantelli, fore tibia anteriorly, base and apex of middle femur, base of hind femur, base of middle and hind tibia, pale yellow or ivory; small medial patch of vertex brown; mesosoma largely orange-brown dorsally and orange-brown laterally, but darkened ventrally; mesosternum dark brown, meta-  Figures 353-360. Taeniogonalos fasciata f. interrupta (Chen, 1949), holotype, female. 353 Antenna 354 fore and hind wings 355 head lateral 356 mesosoma dorsal 357 mesosoma lateral 358 metasoma dorsal 359 metasoma lateral 360 metasoma ventral. soma blackish brown but broad band on posterior margin of first tergite and of second tergite and of first sternite, broadly interrupted band on posterior margin of second sternite and sixth tergite yellowish brown; palpi yellowish brown, but labial palp partly dark brown; antenna dark brown, but scapus with yellowish ventro-apically (Fig. 331); remainder of legs dark brown; pterostigma brown, but veins, marginal cell and area below it dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 5.2-13.4 mm, of fore wing 2.2-12.2 mm; antenna of ♀ with 22 (5), 23 (1), 24 (2), 25 (4) or 26 (2) segments; vertex largely smooth with sparse fine punctures behind stemmaticum to densely punctate; lateral lobe of mesoscutum longitudinally rugulose to coarsely rugose; propodeum mainly rugulose to coarsely rugose; yellow patches of inner orbita wide triangular or linear, supraantennal elevations largely black only apically brownish to entirely yellow; transverse stripe on posterior vertex orange-yellow or reduced; pair of yellow spots on clypeus large, small or absent or clypeus entirely yellow; mesoscutum and its scutellum more or less with extensive dark brown pattern, propodeum largely orange-brown, largely dark brown except for long pale yellow stripes laterally (Fig. 356) or entirely black; mesosoma largely black laterally or dorsal half orange-brown; yellow stripe on posterior margin of second tergite narrow or interrupted medially, yellow band on posterior margin of first sternite narrow to wide, yellow posterior band of second sternite reduced to wide; legs sometimes largely dark brown; length of vein 1-M of fore wing 1.4-1.8 times as long as vein 1-SR; second metasomal tergite more or less with a shallow depression medio-basally; second sternite usually densely punctate, but sometimes sparsely so.
Male. Length of body 5.8-11.1 mm, of fore wing 5.1-10.8 mm; antenna with 22 (3), 23 (1), 24 (3) segments, tyloids linear, in specimen with 22 or 23 segments: 0.8 times as long as segment on 11 th -14 th segments and 0.2 times as long as segment on 15 th segment; in specimens with 24 segments: 0.8 times as long as segment on 12 th -14 th segments and 0.2 times as long as segment on 15 th segment; second sternite of ♂ partly flattened medio-posteriorly; genitalia internal; head and apex of metasoma of male darker than of female; otherwise variation as in female.
Metasoma. First tergite 0.7 times as long as apically wide, smooth and with distinct elliptical depression medially (Fig. 369); second-sixth tergites densely reticulatepunctate; sternites densely punctate; second sternite distinctly convex in lateral view, medio-apical protuberance triangularly protruding behind apical margin of sternite and posteriorly truncate (Fig. 370); third sternite about 0.2 times as long as second sternite (Fig. 371); hypopygium triangular in ventral view. Colour. Yellow with black markings; frons largely yellow, area above and between supra-antennal elevations largely black but with a heart-shaped yellow spot below anterior ocellus, black marks medio-ventrally extending to base of clypeus; clypeus entirely yellow; mandible yellow with teeth dark brown; vertex largely black with a V-shaped yellow patch posteriorly and postero-lateral area above occipital carina yellow; temple entirely yellow; mesosoma laterally black with following yellow markings: broad oblique stripe on lower propleuron, stripe on upper pronotum and lower pronotum, broad oblique but medially interrupted stripe on mesopleuron and metapleuron; mesosoma dorsally yellow with following black markings: broad median stripes on middle lobe of mesoscutum, lateral lobes of mesoscutum, broad median stripe on propodeum; metasoma yellow with following black markings: anterior half of first tergite and sternite, anterior 0.7 of second tergite except for broad median patch, anterior 0.8 of third tergite, small median patch on fourth and fifth tergites, broad antero-lateral patches on second sternite, anterior half of third sternite, entirely fourth and fifth sternites; palpi yellowish brown; antenna dark brown; dorsal coxae, trochanters and trochantelli, base and apex of femora yellow, remainder of legs dark brown; pterostigma dark brown, marginal cell more or less dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 7.0-8.4 mm, of fore wing 6.7-7.5 mm; antenna of ♀ with 25 segments; length of vein 1-M of fore wing 1.4-1.6 times as long as vein 1-SR; palpi yellow; antenna yellowish brown; legs paler; dark brown patch of marginal cell of fore wing weakly to distinctly pigmented.
Male. Length of body 7.6-8.5 mm, of fore wing 6.0-6.7 mm; antenna with 25-26 segments, tyloids linear, 0.7 times as long as antennal segment on 11 th -14 th segments and about half as long on 10 th and 15 th segments with a remnant on 16 th segment; genitalia internal.
Wings. Fore wing: vein 1-M as long as vein 1-SR (Fig. 376), distinctly curved and vein 1-SR widened anteriorly; second submarginal cell about as long as third cell.
Colour. Black; orbita yellow, narrow but widened near antennal sockets, supra-antennal elevations largely yellow and connected to yellow patch (Fig. 374), clypeus with pair of large yellow patches; head posteriorly brownish-yellow, including two pairs of oblique patches; mandibles largely dark brown with some yellow near base of teeth; mesosoma laterally black except for dorsal and ventral yellow patches of pronotal side; middle lobe of mesoscutum with pair of triangular yellow patches anteriorly; tegulae yellow; scutellum laterally and axilla with pair of large yellow patches; metanotum with a large medial and a large lateral patch yellow; propodeum pair of large yellow patches (Fig. 378); first tergite ivory medio-posteriorly, second tergite widely yellow posteriorly, third tergite with pair of small yellow patches postero-laterally; fourth-sixth tergites with pair of large yellow patches; metasoma ventrally black but apical margin of first sternites yellowish and apex of hypopygium brownish (Fig. 382); palpi and apical half of antenna brown; remainder of antenna mainly light brown; apex of trochanters, trochantelli largely, anterior stripe on fore tibia, base of all femora and apex of fore and middle femora, base of middle and hind tibiae pale yellow; remainder of tibiae and tarsi more or less dark brown; pterostigma and marginal cell of fore wing largely and area below it dark brown; remainder of wing membrane slightly infuscate.
Male. Similar to the pale and dark (= "unifasciata") forms of the female, but vertex and frons largely black; second sternite distinctly flattened medially; tyloids linear.
Variation. The pale and typical form has the supra-antennal sockets largely yellow (Fig. 374), but these are largely or entirely black or dark brown in the dark form (Fig.  385). The vertex is partly yellowish brown or orange posteriorly (Fig. 374), sometimes including posterior part of frons, but rather obscurely so in the dark form (Fig.  385); lobes of mesoscutum sometimes largely orange laterally; yellow medial patch of metanotum is sometimes divided into two patches; metasoma finely to coarsely punctate dorsally; second-fifth sternites laterally black or partly orange; second submarginal cell of fore wing 1.0-1.5 times as long as third cell.
Notes. P. intermedia Chen, 1949 (♀), and P. unifasciata Chen, 1949 (♂), are synonymized here with T. formosana, because they share the less steeply protruding supra-antennal protruberances (angle about 35°; Figs 374, 385, 396), the more or less brownish or yellowish vertex posteriorly, head anteriorly and mesosoma dorsally with a yellowish pattern and differ only in colour from T. formosana (Bischoff) and intermediates are present. If the melanistic form should be named, then f. intermedia (Chen) is available; T. lachrymosa (Westwood, 1874), comb. n. from Philippines has a similar colour pattern but a different venation according to the original description. Taeniogonalos taihorina (Bischoff, 1914) is very similar to the dark form of T. formosana, but differs by having the area between the supra-antennal protuberances distinctly concave and the protuberances more steeply protruding laterally (angle about 45°), the pronotum laterally, the head anteriorly and the mesosoma dorsally entirely black. Diagnosis. Outer side of supra-antennal elevations oblique and elevations about 0.3 times as long as scapus (Fig. 408); occipital carina strongly widened medio-dorsally and distinctly lamelliform (Fig. 408); head posteriorly entirely black and dorsally punctate with distinct smooth interspaces (Fig. 408); middle mesoscutal lobe similarly coloured to lateral lobes, yellow latero-posteriorly, black or yellow laterally and black medially (Fig. 412); scutellum with pair of medium-sized yellow patches medially (Fig.  412); anterior half of fore wing only below pterostigma or subapically dark brown and remainder subhyaline (Fig. 410); mesosoma with moderately limited yellowish pattern dorsally (Fig. 412); second sternite of female distinctly convex ( Fig. 415; probably less of ♂); third sternite of female without protuberance (Fig. 416); metasoma with limited pale pattern (Figs 414, 415, 416).
Head. Antenna with 26 segments; frons densely punctate and no smooth interspaces; vertex punctate with large smooth interspaces but patch behind stemmaticum and near eyes with narrow interspaces and distinctly shiny (Fig. 408), with rather long setae; temple largely smooth with sparse fine punctures (Fig. 411); head gradually narrowed behind eyes, eye in dorsal view 1.4 times as long as temple (Fig. 408); occipital carina strongly widened medio-dorsally and distinctly lamelliform, smooth (Fig. 408); supra-antennal elevations medium-sized (about 0.3 times as long as scapus), outer side oblique and largely smooth except for sparse punctures; clypeus slightly concave and thick medio-ventrally.
Colour. Black; frons with narrow yellow stripes along inner orbita extend to malar space, supra-antennal elevations yellow, clypeus with pair of yellow patches, remainder of head black; mandibles largely yellow with dark brown teeth; mesosoma laterally black except for dorsal yellow patches of pronotal side; middle lobe of mesoscutum with pair of elongate yellow patches anteriorly, lateral lobes of mesoscutum with narrow yellow patch along outer margin; scutellum with pair of wide yellow patches anteriorly; metanotum with one large medial yellow patch; propodeum pair of small yellow patches (Fig. 412); metasoma dorsally black, lateral margin of first tergite ivory and remainder black, second tergite with pair of large yellow patches postero-laterally, third tergite with pair of small yellow patches postero-laterally; metasoma ventrally black (Fig. 416), apical margin of all sternites ivory; palpi and antenna dark brown; patch on middle and hind coxae, fore and middle trochanters largely, anterior stripe on fore femur, base of all femora and apex of fore and middle femora, fore tibia anteriorly, base of middle and hind tibiae pale yellow; remainder of tibiae and tarsi more or less dark brown; pterostigma and apical half of marginal cell of fore wing and area below it dark brown; remainder of wing membrane slightly infuscate.
Colour. Yellow with black pattern; frons largely yellow, area above and between supra-antennal elevations largely black but with a heart-shaped yellow spot below anterior ocellus, black marks medio-ventrally extending to base of clypeus; clypeus yellow with ventral margin dark brown; mandible yellow with teeth and latero-ventral margin black; vertex largely black with a V-shaped yellow patch posteriorly and posterolateral area above occipital carina yellow; upper temple yellow and lower margin black; mesosoma laterally black with following yellow markings: broad oblique stripe on propleuron, stripe on upper pronotum and lower pronotum, broad oblique but medially interrupted stripe on mesopleuron and metapleuron; mesosoma dorsally black with following yellow markings: antero-lateral stripes on middle lobe of mesoscutum, axilla, antero-lateral half, metanotum, broad longitudinal lateral stripes on propodeum; metasoma black with following yellow markings: broad patch on posterior margin of first tergite and anterior margin of second tergite, narrow continuous stripe on posterior an lateral margin of second tergite, postero-lateral broad patches on fourth to sixth tergites, posterior half of first sternite, rather broad stripe on posterior margin of second sternite; palpi dark brown; antenna dark brown with first five segments darker; coxae dorsally, trochanters and trochantelli, base and apex of femora and base of tarsi yellow, remainder of legs dark brown to black; pterostigma dark brown, marginal cell more or less dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 5.5-9.3 mm, of fore wing 4.2-8.1 mm; length of vein 1-M of fore wing 1.2-1.4 times as long as vein 1-SR.
Male. Length of body 7.4-9.1 mm, of fore wing 5.8-6.9 mm; antenna with 24 segments, tyloids linear, 0.8 times as long as segment on 10 th -16 th segments and 0.5 times as long as segment on 17 th segment; genitalia internal.
Notes. As pointed out by Schulz (1908) specimens from Indonesia and Myanmar differ from both examined female syntypes of T. thwaitesii (Westwood, 1874) from Sri Lanka. He created a new name for the specimens from Indonesia and Myanmar: Poecilogonalos pulchella var. gestroi. After examination of a type of T. thwaitesii, Dr D. Smith (in litt.) kindly informed one of us (CvA) that T. thwaitesii (auctt.) from North India up to China is indeed a species different from the typical T. thwaitesii from Sri Lanka and possibly South India. Therefore, we use the first available name for this species, T. gestroi (Schulz, 1908) which is a new combination.
Colour. Black; mandible medially, apices of fore and middle trochanters, hind trochanter and trochantellus and bases of femora largely ivory; second tergite with indistinct pair of small brownish patches posteriorly (Fig. 436); antenna (but darkened basally), palpi and tegulae mainly brown; fore tibia and tarsus, middle tibia basally brownish-yellow and remainder of legs dark brown; subapical patch of fore wing infuscate; pterostigma and veins brown, remainder of wing membrane subhyaline.

Additional material.
No material available for this study; reported from China by Lelej (2003).
Diagnosis. Supra-antennal elevations 0.1-0.4 times as long as scapus and outer side of elevations oblique (Fig. 451); occipital carina lamelliform, wide; head black, but supra-antennal elevation anteriorly, two patches of clypeus, inner and (narrowly) outer orbita yellow; metanotum with lateral yellow patch; entire fore wing dark brown (Fig. 449); second metasomal sternite of female simple, only slightly convex (Fig. 449); second and third sternites of male with flattened and shiny posterior patch; second tergite and sternite with wide yellow band apically, and remainder of metasoma largely black (Fig. 450); length of body 14.5 mm.
Description (translation of short redescription in Russian by Marshakov (1981); only males known). Male. Antenna with 23-25 segments, 10 th -16 th segments with tyloids; two spots on clypeus, a narrow band on inner and outer orbita of eyes, two spots on mesoscutum, [most of] postscutellum, band on first [mainly ventrally] and second [metasomal] segments, anterior and middle tarsi and tibiae yellow; [remainder of] legs light brown, antenna rusty yellow to light brown. Specimens from Amurskoj oblast have the scutellum marked with yellow, the wings are very slightly darkened and the background colour of the first and second [metasomal] segments is black (with brown-red hue in specimens from Mongolia); the head, the mesoscutum and the propodeum dorsally have large spots; the mesopleuron is finely densely punctate, with almost no interspaces between the punctures, matt; the frontal keels are like a "7". The length of the body is 9-11 mm.
Biology. Unknown. Distribution. China (Inner Mongolia); Russia; Mongolia; Korea (Lelej 2003). Notes. Taeniogonalos flavocincta (Teranishi, 1929) was synonymized with T. mongolica by Marshakov (1981), but the holotype of T. flavocincta is not synonym with T. mongolica. The male holotype of T. flavocincta differs as follows: the frons, vertex and mesoscutum coarsely reticulate-punctate (moderately spaced punctate with distinct smooth interspaces in T. mongolica; Fig. 451), the clypeus black (with pair of ivory patches; Fig. 451), the metanotum with only a medial ivory patch (with medial and pair of lateral patches; Fig. 450), the mesoscutum black anteriorly (with pair of ivory patches; Fig. 450), the second and third tergites densely and coarsely punctate, rather matt (largely smooth and strongly shiny; Fig. 450) and the posterior half of the fore wing subhyaline (infuscate; Fig. 449). From the translated redescription of Marshakov above it is clear that his series is mixed and consists of several species; the presence of T. mongolica in Inner Mongolia needs reconfirmation.
Description. Redescribed after the female from Fujian, length of body 12.7 mm (of fore wing 10.3 mm).
Colour. Head largely dark brown with following orange-brown markings: stripes along inner and outer orbita (including malar space), lateral area of clypeus and area above clypeus, V-shaped patch behind stemmaticum; mesosoma largely dark brown to black with following yellowish brown makings: antero-lateral stripes on middle  largely black with following orange-brown or yellowish brown markings: narrow stripe on posterior margin of first tergite and sternite, broad stripe on posterior margin of second and third tergites, posterior 0.7 of fourth and almost entirely fifth and sixth tergites, postero-lateral spots on second sternite; mandible dark brown; palpi dark brown; basal half of antenna (including scapus) yellowish brown, otherwise dark brown; legs dark brown to nearly black; pterostigma and anterior half of fore wing dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 7.8-12.7 mm, of fore wing 6.4-10.3 mm; antenna of ♀ with 24-27 segments; orange-brown markings on body paler or darker; length of vein 1-M of fore wing 1.0-1.2 times as long as vein 1-SR.
Wings. Fore wing: length of vein 1-M 1.6 times as long as vein 1-SR (Fig. 467); second submarginal cell 1.7 times as long as third cell.
Metasoma. First tergite 0.4 times as long as apically wide, smooth and with distinct elliptical depression medially (Fig. 471); second-sixth tergites densely and rather finely reticulate-punctate but exposed anterior rim of tergites superficially coriaceous; second sternite rather densely and coarsely reticulate-punctate with distinct smooth interspaces, other sternites largely smooth; second sternite strongly convex, medio-apical protuberance triangularly protruding behind apical margin of sternite in lateral view, apically triangular in posterior view and truncate in ventral view (Figs 472,473); third sternite about 0.1 times as long as second sternite (Fig. 473); hypopygium triangular in ventral view (Fig. 473).
Colour. Head tricoloured, frons below supra-antennal elevations largely yellow except for interantennal area and middle of clypeus dark brown, above elevations blackish brown; with orange-brown patch in front of anterior ocellus and transverse stripes beside lateral ocelli and pair of oblique stripes branching off mainly orange-brown posterior part of head (Fig. 465); orbita largely yellow, but inner orbita with wide triangularly yellow patch ventrally; clypeus yellow but medially and ventral margin dark brown; mandible yellow with teeth and largely basally dark brown; mesosoma largely black but pronotal side dorsally and ventrally, middle lobe of mesoscutum laterally, axilla largely, large lateral spots on metanotum and propodeum yellow, antero-lateral 0.6 of scutellum and metanotum largely, pair of large patches on propodeum and tegulae yellow; metasoma blackish brown but first tergite medio-posteriorly, posterior band of first sternite and of second and third tergites, pair of postero-lateral patches on fourth-seventh tergites yellow; posterior margin of second sternite pale yellowish; palpi and antenna brown; coxae dorsally partly, trochanters dorsally and apically, trochantelli, base of femora narrowly and tibiae subbasally ivory, remainder of legs dark brown; pterostigma and marginal cell of fore wing and area below cell dark brown, remainder of wing membrane largely subhyaline.
Variation. Length of body 5.5-7.6 mm, of fore wing 4.7-6.5 mm; length of vein 1-M of fore wing 1.2-1.6 times as long as vein 1-SR; upper frons largely black with small triangular yellow patch below anterior ocellus; clypeus weakly concave medioapically, often nearly straight; area besides stemmaticum yellow or orange or dark brown; propodeum with reversed comma-shaped lateral patch or oval patch; scutellum entirely orange-brown or with brown or black patch or all black; metasoma very variable in colour (see figures in Tsuneki 1991), may be darker than holotype, bands and patches on tergites of variable size, second-sixth sternites entirely black, with small yellowish brown postero-lateral patch on third sternite or fourth-sixth tergites of ♀ yellow, with medial dark brown patch or only anteriorly dark brown; basal half of second tergite sometimes medially yellow or blackish; mesopleuron black or with two fine yellow stripes. The holotype of T. pictipennis has the scutellum entirely yellow, the mesopleuron and the metapleuron with an elongate yellow patch postero-ventrally, the scutellar sulcus smooth and narrow, the vertex behind the posterior ocelli densely sculptured (without smooth interspaces between punctures) and the second sternite densely punctate. The differences seem to be intraspecific and, therefore, we accept the synonymy of this species with T. sauteri.
Colour. Head black with pale yellow stripes along inner orbita and apex of supraantennal elevations and dorsal-medial occipital carina brown; mesosoma orange-brown dorsally (Fig. 491); mesosoma largely orange-brown laterally except for anterior and dorsal yellow patches of pronotal side and dark brown propleuron; metasoma with moderately wide orange-yellow apical band at apex of first tergite and sternite, second tergite with broad orange-yellow band at posterior margin, remainder of metasoma black; palpi dark brown; mandible black with teeth dark brown; basal half of antenna (except scapus) yellowish brown, scapus and apical half of antenna dark brown (Fig.  488); trochanters and trochantelli and base of femora of all legs pale yellow, tibiae and tarsi of fore and mid leg yellowish brown, mid and hind coxae orange-brown, remainder of legs dark brown; pterostigma dark brown, remainder of wing membrane subhyaline.
Male. Unknown. Biology. Unknown. Collected in July.
Description. Female from Shaanxi, length of body 12.4 mm (of fore wing 9.8 mm).
Colour. Black; palpi dark brown; mandible black with teeth dark brown; antenna dark brown; frons with small yellowish brown spots along inter orbita, remainder of head black except for posterior vertex with small yellowish brown spots and narrow yellowish brown stripes along outer orbita; apex of supra-antennal elevations yellowish brown; tegulae dark brown; mesosoma laterally black except for antero-dorsal yellowish brown patch of pronotal side; metanotum with one medium-sized medial yellowish brown patch; hind trochanter, trochantellus and base of hind femur ivory, remainder of legs dark brown to black with tarsi paler; anterior half of fore wing dark brown, posterior half subhyaline; metasoma with moderate wide orange-brown apical band at apex of second tergite and sternite, third tergite with narrow orange-brown band at posterior margin, fourth to sixth tergites with large orange-brown patches posteriorly.  Male. Holotype male is very similar to the redescribed female except shape of second sternite. It has length of body 10.7 mm (of fore wing 8.1 mm); brown antenna with 23 segments with longitudinal tyloids on 10 th -16 th segments; outer orbita largely and pronotum dorso-posterioly ivory, head posteriorly and third tergite apically black; second sternite of male distinctly shallowly depressed medio-posteriorly; third sternite of male flat medially; Biology. Unknown. Collected in May and July. Distribution. China (Shaanxi), Korea. Etymology. Renamed after the blunt (subtruncate) protuberance of the second sternite of the female: from "sub" (Latin for "under, less") and "truncus" (Latin for "cut off").
Wings. Fore wing: vein 1-M 1.3 times as long as vein 1-SR (Fig. 522), distinctly curved and vein 1-SR widened anteriorly; second submarginal cell about twice as long as third cell.
Colour. Black; metasoma largely black with wide orange-brown band at apex of second tergite and sternite; mandible nearly black, somewhat brownish; antenna dark brown to black, becoming darker apically; tibiae and tarsi yellowish brown, remainder of legs dark brown to nearly black; pterostigma and anterior half of fore wing dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 8.8-10.1 mm, of fore wing 7.1-8.2 mm; length of vein 1-M of fore wing 1.4-1.6 times as long as vein 1-SR.
Male. Length of body 8.2-9.7 mm, of fore wing 6.8-7.9 mm; antenna with 23 segments, tyloids linear, 0.8 times as long as segment on 10 th -14 th segments and 0.5 times as long as segment on 15 th segment; genitalia extruded.
Colour. Head largely dark brown with following yellow or orange-brown markings: stripes along inner and outer orbita (including malar space), lateral corner of clypeus, V-shaped patch behind stemmaticum and oblique patches on postero-lateral vertex; mesosoma largely black with following yellow or yellowish brown makings: lateral stripes on middle lobe of mesoscutum, tegulae, lateral stripes on scutellum, metanotum, small spots on upper and lower pronotum, dorsal spot on mesopleuron, small lateral spots on propodeum; metasoma largely black with following dark brown or yellow markings: narrow stripe on posterior margin of first-fourth tergites and first sternite, large median patch on fifth and sixth tergites yellow, narrow stripe in front of yellow stripe on first-fourth tergites dark brown; mandible largely yellow with anteroventral corner and teeth dark brown; palpi dark brown; basal half of antenna yellowish brown, apical half dark brown; legs dark brown to black; pterostigma and anterior half of fore wing dark brown, remainder of wing membrane subhyaline.
Variation. Length of body 8.6-12.1 mm, of fore wing 8.4-11.7 mm; antenna of ♀ with 25-28 segments; head largely orange-brown or reddish brown with similar but reduced black markings as in lectotype; clypeus largely yellow with middle dark brown; lower pronotum with or without yellow spot; mesopleuron with or without orange-brown spot; lateral spot on propodeum larger or absent; metasoma extensively varies, yellow stripe on posterior margin of second tergites broader or absent, thirdsixth tergites with reddish brown or dark brown stripe on posterior margin, second sternite with broad reddish brown stripes on posterior margin; antenna entirely dark brown; outer side of hind coxa, trochanter, trochantellus and base of femur ivory; length of vein 1-M of fore wing 1.1-1.4 times as long as vein 1-SR; sixth tergite with whitish or yellowish pubescence; head besides stemmaticum with orange transverse patch; frons also posteriorly and partly anteriorly orange.
Male. Length of body 7.8-12.4 mm, of fore wing 7.2-11.1 mm; antenna with 25-28 segments, tyloids linear, 0.8 times as long as segment on 10 th -12 th segments and as long as segment on 13 th -15 th segments with a remnant on 16 th segment; genitalia internal, sometimes extruded; second sternite with shallow concave and shiny medioposterior area; males more or less coloured as T. sauteri, but sometimes with pale head as females, colour varies as in females.
Description. Holotype, male, length of body 10.2 mm (of fore wing 8.9 mm). Head. Antenna with 25 segments, tyloids nearly as long as segment present on 10 th -13 th segments, on 14 th and 15 th segments shorter tyloids; frons densely punctate and no smooth interspaces; vertex punctate with large smooth interspaces but patch behind stemmaticum and near eyes with narrow interspaces and distinctly shiny (Fig.  573), with rather long setae; temple largely smooth with sparse fine punctures (Fig.  577); head gradually narrowed behind eyes, eye in dorsal view 1.9 times as long as temple (Fig. 573); occipital carina narrow medio-dorsally; supra-antennal elevations medium-sized (about 0.3 times as long as scapus), outer side oblique and largely smooth except for sparse punctures; clypeus slightly concave and thick medio-ventrally.
Colour. Black; head anteriorly with extensive pale yellow pattern (Fig. 572); remainder of head black except pale yellow outer orbita and pair of posterior patches on the vertex pale yellow (Fig. 573); mesosoma laterally black except for anterior and dorsal yellow patches of pronotal side; middle lobe of mesoscutum with pair of wide pale yellow patches anteriorly; axilla with large pale yellow patch; metanotum with one large medial and pair of large lateral pale yellow patches; propodeum with pair of large elongate pale yellow patches (Fig. 578); metasoma dorsally orange-brown, first tergite posteriorly largely pale yellow and remainder black, second tergite postero-laterally, fifth tergite laterally and most of sixth tergite (except medial brown patch) pale yellow; metasoma ventrally tricoloured (Fig. 581): first sternite black basally, narrowly orangebrown and yellow apically, second sternite black, medio-posteriorly and laterally orange-brown except for narrow yellow apical margin, remainder of metasoma ventrally mainly orange-brown; palpi and antenna (but ventrally brown) dark brown; patch on middle and hind coxae, fore and middle trochanters largely, anterior stripe on fore femur, base of all femora and apex of fore and middle femora, fore tibia anteriorly, base of middle and hind tibiae pale yellow; remainder of tibiae and tarsi more or less dark brown; pterostigma and apical half of marginal cell of fore wing and area below it dark brown; remainder of wing membrane slightly infuscate.
Female. Unknown. Variation. Paratype from Guizhou: length of body 8.2 mm and of fore wing 7.0 mm, antenna with 25 segments and tyloids similar to holotype; pale patches of vertex and of mesoscutum slightly smaller than of holotype and orange-brown part of second sternite is smaller. The paratype from Yunnan has the pale patches of vertex and of mesoscutum absent and median patch of metanotum reduced to 2 small pale patches.
Colour. Black; palpi dark brown; mandible dark brown; antenna dark brown; frons with narrow yellowish brown stripes along inter orbita, remainder of head black; tegulae dark brown; legs dark brown to black with tibia and tarsi paler; metasoma with moderate wide orange-brown apical band at apex of second tergite and sternite; pterostigma and anterior half of fore wing dark brown, ventral half subhyaline.
Colour. Black; malar space brownish; postero-lateral margin of first tergite and sternite, second and third tergites and sternites largely reddish-yellow; palpi dark brown; mandible with large basal ivory spot, otherwise orange-brown to dark brown; antenna black with a small ivory band subapically; apex of fore femur, tibia and tarsus, ventral side of mid and hind tibiae yellowish brown, remainder of legs dark brown to black; pterostigma dark brown; wing membrane subhyaline.