New species of Trigonalyidae (Hymenoptera) from NW China

Abstract Four new species of Trigonalyidae are described and illustrated from Qinling Mts (Shaanxi, NW China): Bareogonalos xibeidai Tan & van Achterberg, sp. n., Jezonogonalos mandibularis sp. n., J. shaanxiensis sp. n., and Taeniogonalos paraclypeata sp. n. Orthogonalys hirasana Teranishi, 1929, is re-instated and reported from China. The female of Taeniogonalos alticola (Tsuneki, 1991) is described for the first time. In total, 18 species are known from Shaanxi province, 20 species for NW China, and eight described species are newly recorded for Shaanxi: Jezonogonalos luteata Chen et al., 2014, Orthogonalys hirasana Teranishi, 1929, O. elongata Teranishi, 1929, Pseudogonalos hahnii (Spinola, 1840), Taeniogonalos alticola (Tsuneki, 1991), T. formosana (Bischoff, 1913), T. taihorina (Bischoff, 1914), and Teranishia glabrata Chen et al., 2014. Poecilogonalos maga Teranishi, 1929, syn. n. and Taiwanogonalos claripennis Tsuneki, 1991, syn. n. are new synonyms of Taeniogonalos taihorina (Bischoff, 1914) and Taiwanogonalos alishana Tsuneki, 1991, syn. n. of Taeniogonalos alticola (Tsuneki, 1991). Revised keys to species of the genera Bareogonalos, Jezonogonalos, and Orthogonalys are included.


Introduction
Trigonalyidae (Hymenoptera) is a worldwide small family in its own superfamily Trigonalyiodea, with 115 recognized species Smith and Stocks 2005;Santos et al. 2012;Smith and Tripotin 2012;Smith et al. , 2015Yamane 2014). Most species of this family occur in tropical and subtropical regions and the family is absent in arctic and alpine habitats . Surprisingly, Trigonalyidae are fairly common at 1300-1500 m altitude in the Qinling Mts of Shaanxi (NW China). In total, 18 species from Shaanxi are recorded in this paper, which is 41% of the 44 spp. known from China.
Trigonalyidae are often misidentified; slender specimens (especially of Orthogonalys) with white banded antennae are 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 and the presence of unequal mandibles with 3-5 large teeth (Figs 10,36,57) and plantar lobes on the tarsal segments (Figs 12,29,91). In addition, the tarsal claws are cleft (bifurcate with the inner tooth larger than the outer one: Figs 24, 86) and as pointed out in Carmean and Kimsey (1998) the females have sparse white scales or specialized seta on the outside of the middle antennal segments.
Most Trigonalyidae develop as a hyperparasitoids on parasitoid wasp or fly larvae inside caterpillars and sawfly larvae. Primary endoparasitism of sawflies occurs, but the parasitoid still acts facultatively as a hyperparasitoid (Yamane and Terayama 1983;He and Chen 1986;Carmean and Kimsey 1998). Up to more than 2000 eggs may be laid 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 parasitoid wasp (Ichneumonidae or Braconidae) or fly (Tachinidae). Other species are brought into the nests of social Vespidae because they are inside the caterpillars used as prey by the wasps. Inside the nest they develop as primary endoparasitoids of the social wasp larvae.

Materials and methods
The specimens were mainly collected in Malaise traps, but a few by sweep net. The material was stored in 70% ethanol, prepared with the AXA method (van Achterberg 2009;van Achterberg et al. 2010) and glued on card points. Observations and descriptions were made with an Olympus SZX11 stereomicroscope and fluorescent lamps. Photographic images were made with the Keyence VHX-5000 digital microscope and processed with Adobe Photoshop CS5, mostly to adjust the size and background. Morphology. For other terminology used in this paper, see van Achterberg (1988van Achterberg ( , 1993, Hong et al. (2011) and Chen et al. (2014). Measurements were taken as indicated by van Achterberg (1988). Additional non-exclusive characters in the key are between brackets.
Material. Types and other examined specimens are deposited in the Northwest University, Xi'an, NW China (NWUX) and Naturalis Biodiversity Center, Leiden, Netherlands (RMNH).
Notes. The first part of the name Trigonalyidae refers most likely to protuberance of the second sternite present in females of several spp. (but absent in the species treated in this paper, except Bareogonalos Schulz), which is triangular in lateral view.

1
Metanotum with large protuberance and long setae; left mandible with 2 subapical teeth (Fig. 10); maxillary palp with 6 segments; anterior half of scutellum flat or weakly convex and at same level as mesoscutum or slightly above it (Figs 5,16,24); fourth and fifth tergites largely smooth and shiny (Fig. 1) Surrounding membrane of veins 1-SR and 1-M of fore wing yellowish or infuscated as most of wing membrane (Figs 23,24); width of head in dorsal view 0.8-0.9 times maximum width of mesoscutum; third sternite of ♀ distinctly protruding medio-posteriorly in lateral view (Fig. 26) and apical part usually distinctly exposed in ventral view (Fig. 27); antenna with 21-23 segments (Fig. 28), rarely with 20 segments; mesoscutum with median groove posteriorly (Fig. 25); [very colour variable species  Diagnosis. Left mandible with two subapical teeth, ventral tooth shorter than dorsal tooth; maxillary palp with six segments; width of head in dorsal view 1.05-1.10 times maximum width of mesoscutum (Figs 4,15); anterior half of scutellum flat and at same level as mesoscutum; metanotum with large protuberance and long setae; wing membrane near veins 1-SR and 1-M of fore wing subhyaline as remainder of wing except darkened apical third of wing (Figs 3,14); hind basitarsus slender (Fig. 12); fourth and fifth tergites largely smooth and shiny; third sternite of ♀ in ventral view distinctly protruding medio-posteriorly (but obscured by second sternite). Differs from the only other known Palaearctic species, B. jezoensis (Uchida) mainly by the colour of the wing membrane (largely subhyaline vs largely yellowish in B. jezoensis; Fig. 23), the median groove of the mesoscutum (absent posteriorly vs present) and the wider head (1.05-1.10 times vs 0.80-0.90 times width of mesoscutum).
Metasoma. First tergite 0.3 times as long as apically wide (of paratype, obscured by propodeum in holotype), gradually narrowed basally, flat medially and straight apically; second tergite smooth and strongly shiny as following tergites, but with lateral patch of fine punctures; second sternite rather densely finely punctate and shiny, its medio-apical protuberance densely setose and widely truncate medially (Fig. 7), protuberance of third sternite similar but smaller and obscured by second sternite in ventral view (Fig. 7); down curved apex of metasoma nearly up to protuberance of second sternite (Fig. 8).
Legs. Hind coxa and femur smooth and shiny. Colour. Black; palpi, mandible apically, antenna (but scapus yellow ventrally and basal third of antenna brown), tegulae (but humeral plate brown), first tergite, veins (except yellowish veins 1-SR, 1-M and 1-SR+M of fore wing) and pterostigma dark brown; coxae basally black; femora (except yellow anterior face of fore femur, base and apex of middle and hind femora), and inner side of hind tibia (and partly ventrally) dark brown; 3 apical segments of hind tarsus and infuscate; pronotum dorso-apically, pair of wide patches on mesoscutum anteriorly, axilla, scutellum latero-posteriorly, dorso-apical patch of mesopleuron, pair of large antero-lateral patches on second-sixth tergites (touching each other on fourth and fifth tergites), pair of small subposterior patches on first sternite, large V-shaped patch on second sternite, and remainder of legs yellow; membrane of fore wing subhyaline, but apical 0.4 weakly infuscate (Fig. 3).
Variations. Length of body 8.0-9.3 mm, of fore wing 8.0-8.9 mm; antenna of ♀ with 20(2) or 21(2) segments; width of head in dorsal view 1.05-1.10 times maximum width of mesoscutum; mesoscutum sometimes entirely black, or with additional yellow spot medio-posteriorly; length of vein 1-M of fore wing 2.1-3.1 times as long as vein 1-SR; mandible entirely blackish brown or with some brownish patches subapically; antero-lateral yellow patches of metasoma either all separated from each other, or all touching medially, but sometimes only on second-fourth tergites being separated; yellow patch of second sternite may be separated into large patches; inner side of hind tibia largely dark brown or only apical third.

Biology.
Unknown. Collected in June-November. Distribution. China, Japan. Mesoscutum shiny and largely smooth; propodeum antero-laterally shiny and smooth; frons shiny and with wide smooth interspaces between punctures; mandible yellow except for orange brown teeth; third submarginal cell of fore wing narrower than second submarginal cell ..
Colour. Black; inner orbita vaguely partly orange brown; malar space and supraantennal elevations largely ivory; minute patch of outer orbita, vertex posteriorly, occipital carina medially dark brown; apex of scapus and pedicellus, and mandible largely pale brown but basally ivory and teeth dark brown (Figs 36,37); tegulae and pronotal lobe below it pale yellowish; first metasomal tergite apically broadly ivory and narrower laterally, first and second sternites with large ivory patch apico-laterally; with narrow ivory apical band at apex of sternites and of second and following tergites (Figs 29, 33); palpi rather dark brown; remainder of antenna blackish; fore leg (except black coxa and trochanter) yellowish brown; middle and hind legs mainly dark brown but coxae black (except ivory apex of hind coxa) and hind trochanter ivory with some faint infuscation; pterostigma brownish yellow, but anteriorly (except basally) and apically blackish; basal half of marginal cell and third submarginal cell dark brown and remainder of wing membrane subhyaline (Fig. 30).
Male. Unknown. Biology. Unknown. Distribution. China (Shaanxi). Collected at 1556 m. Etymology. Named after its conspicuously coloured mandible. Tan (Fig. 45); outer side of supra-antennal elevations subvertical, largely smooth, and elevations about 0.8 times as long as scapus (Fig. 45); frons moderately punctate (Fig. 46); supra-antennal elevations largely ivory dorsally (Fig. 45); mandible mainly dark brown, except ivory basal patch and pale brown base of teeth (Fig 46); metasoma dorsally largely smooth and largely black (Figs 39, 43); first tergite about 0.6 times as long as its apical width (Fig. 43); third sternite about 0.4 times as long as second sternite (Fig. 44). Close to J. satoi (Tsuneki) from Taiwan and J. mandibularis from Shaanxi, from which the new species differs because of the largely smooth supra-antennal elevations (with carinae or long grooves and some punctures in J. satoi and J. mandibularis), the densely rugose and rather matt scutellum (largely smooth and shiny), the scutellum somewhat above level of metanotum in lateral view and medially flattened (protruding far above level of metanotum and medially slightly longitudinally depressed), the mandible largely dark brown (largely pale brown and yellowish or ivory); the inner orbita of eye completely ivory (largely dark brown or with incomplete ivory stripe), and the middle lobe of mesoscutum with pair of ivory patches anteriorly (entirely black).
Wings. Fore wing: length of vein 1-M 1.5 times as long as vein 1-SR; third submarginal cell much wider anteriorly than petiolate second cell (Fig. 40).
Metasoma. First tergite 0.6 times as long as its apical width, smooth but basal depression anteriorly with some crenulae (Fig. 43); second and following tergites smooth except for punctulation and shiny; sternites rather densely finely punctate, with wide smooth interspaces; second sternite weakly curved in lateral view; third sternite about 0.4 times as long as second sternite (Fig. 44); hypopygium triangularly protruding in ventral view (Fig. 44).
Colour. Black; inner orbita narrowly ivory and connected to ivory malar space; pair of patches on clypeus, basal patch of mandible, large patch on supra-antennal elevations, pair of elongate patches on middle lobe of mesoscutum anteriorly, pair of narrow lines near tegulae, pair of small patches on metanotum medially, epipleura of first tergite, large patch apico-laterally on second tergite and narrow apical bands of sternites ivory; 12 th -22 nd antennal segments largely pale brown ventrally; mandible largely dark brown but teeth light brown basally (Figs 46); tegulae and trochanters mainly dark brown; palpi rather dark brown; legs blackish or dark brown, but fore femur apicoventrally and tibia ventrally yellowish; pterostigma basally yellow, and remainder dark brown; basal half of marginal cell and to a lesser degree area below it dark brown and remainder of wing membrane subhyaline (Fig. 40).
Variations. Length of body 8.7-10.2 mm, of fore wing 7.1-8.1 mm; antenna of ♀ with 25(2) or 27(1) segments; metanotum black or with pair of ivory spots medially; ivory patches of clypeus and mesoscutum rather small to large; mesoscutum sometimes with minute ivory spot medio-posteriorly; second submarginal cell of fore wing petiolate or sessile anteriorly; length of vein 1-M of fore wing 1.5-1.7 times as long as vein 1-SR; pterostigma largely yellowish brown or largely dark brown; mandible dark brown or ivory subbasally; whitish setae of mesosoma long or medium-sized.
Male. Unknown.   Fig. 51, rarely with small patches in males); first metasomal tergite of ♀ with brownish or ivory apical transverse band, at most with narrow brown median line; vein 1m-cu of fore wing connected to first submarginal cell (Fig. 50) (Fig. 62) or interstitial, rarely connected to second submarginal cell; scutellum moderately to coarsely punctate (Figs 64, 71)  Temple with large to medium-sized ivory patch (Figs 49, 58), if mediumsized or small then frons laterally widely ivory or yellowish (Fig. 56); thirdfifth metasomal tergites of ♀ with pair of well-differentiated large triangular ivory patches; posterior half of third metasomal sternite of ♂ ivory or largely so (Fig. 54); pronotum and mesopleuron often with rich pattern of ivory or pale yellowish patches (Figs 49, 52)  Length of eye 0.9-1.1 times temple in dorsal view; mesoscutum densely (rugulose-)punctate, hardly shiny and robust in dorsal view; temple often entirely black; basally second sternite of ♀ ivory; posterior half of propodeum with some transverse rugae; fifth and sixth metasomal tergites more or less ivory medio-apically; slightly more robust species (♀); medially third sternite 0.4-0.7 times as long as second sternite (Fig. 73)  Notes. Orthogonalys centrimaculata Bischoff, 1951, from N. Vietnam (Sa Pa, Lao Cai) will run in the key to O. robusta, but its vertex has a medio-posterior pale patch (absent in O. robusta), the mesoscutum is very finely and densely transversely rugulose or coriaceous, matt (mainly finely remotely (rugulose-)punctate and with satin sheen), and hind tibia and tarsus are yellowish brown (dark brown). It runs to O. formosana if only the colouration of the vertex is considered, but it has the mesoscutum is very finely and densely transversely rugulose or coriaceous, matt (coarsely punctate and shiny in O. formosana) and hind tibia yellowish brown (apical two-thirds dark brown).

Orthogonalys clypeata Chen, van Achterberg, He & Xu, 2014
Orthogonalys clypeata Chen et al., 2014: 67-71 (diagnosis, description, distribution). Notes. The series shows a considerable variation in the shape of the second and third submarginal cells, but the third cell remains wider anteriorly than the second cell. The size difference is also considerable, e.g., the length of the fore wing (3 ♀) is 4.6-8.2 mm.
The new species runs in the key to Orthogonalys by Chen et al. (2014) to O. clypeata Chen, van Achterberg, He & Xu, 2014, and differs as follows: mesoscutum coriaceousrugulose and anterior pair of smooth stripes inconspicuous (densely to remotely punctate and anterior pair of smooth stripes rather conspicuous in O. clypeata), area behind stemmaticum with some fine rugae and posteriorly aciculate (smooth), frons coarsely punctate (at most moderately punctate and often largely smooth), and pterostigma of ♀ dark brown (largely brownish yellow).
Wings. Fore wing: length of vein 1-M 2.2 times as long as vein 1-SR (Fig. 62); second submarginal cell twice as long as third cell.
Colour. Black; palpi and tegulae pale yellow; inner orbita (except dorsally) ivory and connected to broadly ivory malar space; basal half of mandible ivory, apical half pale brown, but teeth dark brown; apical quarter of antenna brownish ventrally; first tergite latero-posteriorly, apical half of second tergite (and medio-anteriorly protruding into black area), pair of large triangular spots on third tergite latero-posteriorly, first sternite laterally, second sternite and fourth sternite laterally and medially yellow (Fig. 69); hind trochanter and trochantellus white; coxae and hind femur black; fore and middle trochanters, base of femora, hind tibia and tarsus dark brown; remainder of legs yellowish brown; pterostigma dark brown; wing membrane subhyaline.
Biology. Reared as hyperparasitoid of Ichneumonidae in caterpillars and from Diprionidae . Collected in April-August. Distribution. Palaearctic region, but two species (P. harmandi Schulz, 1907 and P. hahnii (Spinola)) occur in the North Oriental region.
Biology. Reared as hyperparasitoid of parasitoid wasps (Ichneumonidae and Braconidae) and parasitoid flies (Tachinidae) in caterpillars, but some species are primary parasitoids of Pergid sawflies in Australia (Raff 1934;Carne 1969;He and Chen 1986;Weinstein and Austin 1995;Carmean and Kimsey 1998). Collected mainly in April-October, rarely in November or January.
Distribution. This genus occurs in all major regions, but is unknown from Europe and western Nearctic region. Most of the species occur in the East Palaearctic, Northeast Oriental, and Neotropical regions .
Revised part of key to Chinese species of the genus Taeniogonalos Schulz (for first part, see Chen et al. 2014) 15 Occipital carina strongly widened medio-dorsally (in ♀ up to 1.5 times as wide as diameter of posterior ocellus) and with pair of circular carinae; third antennal segment of ♀ dark brown ventrally; second sternite of ♀ strongly convex and distinctly punctate medially; anteriorly vertex of ♀ finely punctate and with distinct smooth interspaces; mesosoma of ♀ with limited yellowish pattern or entirely black (as in ♂ Second metasomal sternite of ♀ less convex medially and in lateral view its ventral border gradually sloping posteriorly (Fig. 81), strongly shiny medially and third sternite smooth anteriorly or nearly so; second sternite of ♂ evenly slightly convex medio-posteriorly; propodeum comparatively narrow, almost triangular in dorsal view (especially ♂, less so in ♀; Fig. 82) and with nearly straight lateral margins; [vertex largely smooth and often strongly shiny, black, without brownish patch; outer half of lateral mesoscutal lobe usually partly smooth and rather shiny; small species with hind trochanter of ♀ often yellow (Fig. 90) Head. Antenna with 20 segments; frons coarsely punctate, with smooth interspaces narrower than punctures and with some rugae anteriorly (Fig. 88); vertex largely smooth and strongly shiny posteriorly, but spaced moderately punctate anteriorly (Fig.  87); temple smooth except some punctures near eye and mandible (Fig. 89); head gradually narrowed behind eyes, eye in dorsal view 1.1 times as long as temple (Fig.  87); occipital carina narrow, non-lamelliform and smooth dorsally (Fig. 87); supraantennal elevations medium-sized (about half as long as scapus), outer side subvertical anteriorly and with few rather small punctures and apically with few striae (Fig. 87); clypeus slightly concave medio-ventrally and with blunt tubercle above it.
Wings. Fore wing: length of vein 1-M 1.7 times as long as vein 1-SR (Fig. 82); second submarginal cell 1.3 times as long as third cell.
Colour. Black; palpi dark brown basally and pale brown apically; antenna pale brown, but scapus and apical 0.4 of antenna darkened; inner orbita slightly brownish near level of antennal sockets; mandible mainly dark brown, but medially pale brown; sixth tergite yellowish ivory; robust hind trochanter and trochantellus mainly white except some slightly brownish small patches; fore and middle tibiae and tarsi brownish yellow, but middle tibia dark brown posteriorly; remainder of legs, tegulae and pterostigma dark brown; wing membrane subhyaline except for infuscated patch near apex of fore wing (Fig. 82).
Notes. The holotype of T. alishana fits better with T. alticola than with T. taihorina considering its sculpture. The male holotype has been collected at the same day and locality as the male holotype of T. alticola.