Taeniogonalos Schulz, 1906

Kim, Jeong-Kyu & Tripotin, Pierre, 2024, Taxonomic review of the genus Taeniogonalos Schulz (Hymenoptera: Trigonalyoidea: Trigonalyidae) from Korea, with a description of the male of T. sauteri, Journal of Species Research 13 (3), pp. 269-287 : 270-272

Taeniogonalos Schulz, 1906

Taeniogonalos Schulz, 1906: 212 ; Weinstein and Austin 1991: 416; Carmean and Kimsey 1998: 65; Chen et al., 2014: 95 . Type species (by monotypy): Trigonalys maculata Smith, 1851 .

Poecilogonalos Schulz, 1906: 212 ; Marshakov, 1981: 105; Tsuneki, 1991: 46; Weinstein and Austin, 1991: 422; Lelej, 1995: 14. Type species (by monotypy): Trigonalys thwaitesii Westwood, 1874 . Synonymized by Carmean and Kimsey, 1998.

Nanogonalos Schulz, 1906: 211 ; Teranishi, 1929: 150; Marshakov, 1981: 107; Weinstein and Austin, 1991: 421. Type species (by monotypy): Nanogonalos enderleini De Santis, 1980 . Synonymized by Carmean and Kimsey, 1998.

Ischnogonalos Schulz, 1907: 11 ; Schulz, 1908: 33; Bischoff, 1933: 482; Bischoff, 1938: 11; Weinstein and Austin, 1991: 413; Carmean and Kimsey, 1998: 65. Type species (by monotypy): Trigonalys dubia Magretti, 1997 . Synonymized by Chen et al., 2014.

Lycogastroides Strand, 1912: 129 ; Weinstein and Austin, 1991: 413. Type species (by original designation): Lycogastroides gracilicornis Strand, 1912 . Synonymized by Carmean and Kimsey, 1998.

Lycogonalos Bischoff, 1913: 155 ; Weinstein and Austin, 1991: 415. Type species (by original designation): Lycogonalos flavicincta Bischoff, 1913 . Synonymized by Carmean and Kimsey, 1998.

Taiwanogonalos Tsuneki, 1991: 35 . Type species (by original designation): Taiwanogonalos alishana Tsuneki, 1991 . Synonymized by Carmean and Kimsey, 1998.

Diagnosis (characteristics applicable for both sexes unless sex specified). Body length 4.0 - 13.0 mm. Antenna 20 - 26 segmented (variable even in the same species), flagellomeres without whitish segments forming a band; male with a series of linear tyloids on mid flagellomeres. Clypeus semi-elliptically notched apicomedially. Lower frontal areas lateral to antennal sockets depressed, but those above antennal sockets flat. SAE, in dorsal view, low, usually dull triangular, and shelf between SAE weakly and roundly concave. Temple sparsely punctate (especially in posterior part) to punctate-reticulate, and more or less shiny. Occipital carina usually forming regular thin lamina complete to hypostomal carina at level of mandibular base, or sometimes dorsomedially forming a broadened lamina filling the excavated occiput. Vertex flattened without median depression dorsally. Apical segment of labial palp widened and obtuse, more or less triangular. Mandibles not distinctly separated from eye, malar space at most as long as length of antennal pedicel. Mesoscutum and scutellar disc distinctly sculptured (mostly punctate-reticulate and partially ridged) except on a smooth anteromedial declivity of the mesoscutum; scutellar trough with several distinct longitudinal ridges. Metanotal disc slightly convex laterally, and often sculptured. Anterior propodeal sulcus smooth laterally without bottom keel, and narrowed slit-like medially. Forewing fuscous in anterior apical part to anterior half. Hind trochanter vertically grooved subapically, forming an apical right-triangle compartment, and appearing to be made of two segments; fore trochanter not distinctly broadened apically in lateral view, and distinctly longer than hind trochanter (reaching twice as long as); hind tarsus slightly or not modified. Propodeal foramen triangular in shape and more or less arched dorsomedially, bordered by a dull but strong carina. T1 largely concave medially, with arcuate fine ridges in the anterior part of concavity. T2 basomedially depressed, more or less flattened, often shiny and less punctured, and sometimes depression extending posteriorly. In females, S2 distinctly convex (as in T2), sometimes with an apicomedian production (but always not forming paired small teeth); in males, S2 much less swollen than in females, and its larger apicomedian part flat to weakly concave with sparser punctures than remainder S2; S3 short, rarely with a vertical tubercle subapically, but always without a distinct ledge anteriorly. Last three metasomal segments bent backwards to S2 or downwards.

Key to species occurring in Korea

- S2 strongly swollen in profile, not flattened apicomedially. Antennae without tyloids ····························· ♀♀

- S2 less convex in profile and flattened or weakly concave. Antenna with linear tyloids···························· ♂♂

Females

1. S3 apicomedially with a strong vertical tubercle (arrows of Fig. 4B, D ), apical margin of tubercle weakly notched medially, forming a pair of dull teeth. Anterior half of forewing deeply fuscous ( Fig. 4E ): costal cell, radial cell, SMC1, SMC2, SMC3, SMC4 and upper two-third of DC1 clearly fuscous. T 4 - 6 with large hat-like to semicircular markings ( Fig. 4A ) ············································· T. subtruncata Chen et al.

- S3 simple apically, without tubercle. Only costal cell (often hyaline) and anterior apical part of forewing (usually including marginal cell and upper marginal part of SMC4) fuscous, or if more largely fuscous then at least DC1 and SMC1 hyaline ( Figs. 1C , 5B). T 4 - 6 without marking ( Figs. 1A , 5A), or with apical band ( Fig. 6A ) or paired markings ( Fig. 2A )······················· 2

2. S2 with a flat obtuse triangular or semi-circle apicomedian protuberance (arrow in Fig. 3B and in red circle of Fig. 3D ); upper frons with both transverse stripes (connecting to inner orbital markings) lateral to PO and a large cordate spot anterior to anterior ocellus ( Fig. 3C ) ······························ T. sauteri Bischoff

- Apical margin of S2 simple, without median tubercle. Upper frons without such stripes ( Figs. 1B , 2C, 4C, 5C), or if any (stripes often connected to spot anterior to median ocellus, and thus forming a bi-convex stripe across upper frons as in Fig. 6A, B ) then T 2 - 6 with bi-colored apical bands and paired markings that are reddish orange anteriorly and yellow posteriorly ( Fig. 6A, B )·········································································· 3

3. In a majority of specimens, almost entire mesoscutum (sometimes except median blackish part), axillae, scutellum, and very often upper half of mesopleuron and metapleuron concolorous pale to dark reddish ( Fig. 1A, C ); in a minority of specimens, mesosomal coloration much reduced, but still with traces of dark reddish coloration··························· T. fasciata (Strand)

- Corresponding parts of mesosoma not red to dark reddish colored, at most partially yellow to reddish orange ········································································· 4

4. SAE largely black, sometimes ferruginous apically, but a narrow apical rim opaque; SAEs more distantly spaced each other, DSAE ca. 0.44 × as long as distance between eyes at level of SAE ( Fig. 5C, D ). Notauli distinctly interrupted by transverse keels in its entire length. Head and mesosoma almost entirely black, with yellow marks reduced: at most mandible subapically, short stripes along inner and outer orbits, and spots on upper lateral margins of median lobe of mesoscutum yellow; only T 1 (sometimes absent) and T 2 with a distinct apical band ( Fig. 5A, D ) ······················· ·················································· T. taihorina (Bischoff)

- SAE yellow, with a broad translucent apical lamella; SAEs more closely spaced, DSAE ca. 0.33 × as long as distance between eyes at level of SAE ( Fig. 2C, D ). Notauli not distinctly interrupted by transverse keels, at most with indistinct bottom keel in its apical quarter. Head and mesosoma largely colored: at least clypeus, lower frons lateral to antennal sockets, lateral margins of median lobe of mesoscutum, axillae, scutellum, metanotum and propodeum with yellow to orange markings; metasomal terga largely colored, T 1 - 3 with apical bands, T 4 - 6 with paired large triangular spots (sometimes apical bands) ( Fig. 2A - D ; Fig. 6A, B ) ········································································· 5

5. Upper frons with transverse stripes lateral to PO, stripes often connecting to inner orbital markings and forming bi-convex stripe touching ocelli ( Fig. 6A, B ); scutellar disk colored at least laterally ( Fig. 6A ); apical markings of T 2 - 6 bi-colored, reddish orange anteriorly and yellow posteriorly ( Fig. 6A ) ························ ·························································· T. tricolor (Chen)

- Upper frons without such marking, at most with small spots lateral to PO ( Fig. 2C ); scutellar disk not colored ( Fig. 2A ); apical bands or markings of terga not bi-colored but concolorous deep yellow ( Fig. 2A, D ) ················································ T. formosana (Bischoff)

Males

1.F9 - 13 with tyloids, that on F9 variable in length (but usually developed in middle part of F), those on F10 - 12 developed in almost entire length of each F except basally and apically, and that on F13 maximally in basal half ( Fig. 1D ). Almost entire mesoscutum (sometimes except median blackish part) and scutellum concolorous, pale to dark reddish; in a minority of specimens, reddish mesonotal coloration much reduced, rarely completely black, but still antennae with tyloids on F9 - 13 (see also Diagnosis under this species) ················································ T. fasciata (Strand)

- More flagellomeres with tyloids, at least F8 with a well-developed tyloid. Mesoscutum and scutellum neither entirely colored nor reddish, their markings yellow to reddish orange ························································· 2

2. Anterior half of forewing (including costal cell, radial cell, SMC1, SMC2, SMC3, SMC4, and upper half of DC1) deeply fuscous ( Fig. 4E ). Each of T 4 - 6 with a large hat-like to semicircular markings. [F8 - 14 with tyloids, those on F8 - 12 developed in almost entire length (except basally and apically) of each F, that on F 13 in basal two-thirds, and that on F 14 in basal one-third ( Fig. 4F )]····················· T. subtruncata Chen et al.

- Forewing with an anterior apical darkened patch (anterior one-third of forewing weakly fuscous in sole specimen of T. tricolor , but DC1 hyaline). T 4 - 6 without such markings: completely black or with discontinuous paired markings·············································· 3

3. SAE largely black, sometimes ferruginous apically, but narrow apical rim opaque; SAEs more distantly spaced each other, DSAE ca. 0.44 × as long as distance between eyes at level of SAE. Head and mesosoma almost entirely black, with yellow marks reduced: at most mandible subapically, and short stripes along inner and outer orbits yellow; only T 1 - 2 with apical markings: a continuous band on T 1 (often absent), and a pair of oval spots distantly spaced on T 2. [F8 - 13 with tyloids, those on F8 - 11 developed in almost entire length of each F, that on F 12 in basal two-thirds, and that on F 13 in basal one-fifths ( Fig. 5E )]······················· ·················································· T. taihorina (Bischoff)

- SAE yellow, with a broad translucent apical lamella; SAEs more closely spaced each other, DSAE at most 0.37 × as long as distance between eyes at level of SAE. Head and mesosoma largely colored: at least clypeus, lower frons lateral to antennal sockets, lateral margins of median lobe of mesoscutum, axillae, scutellum, metanotum and propodeum with yellow to orange markings; all terga colored apically ····························· 4

4.F8 - 16 with tyloids, those on F8 - 14 developed in almost entire length of each F, that on F 15 in basal two-thirds, and that on F 16 in basal half ( Fig. 2E ). Upper frons lateral to PO without transverse stripe, at most with small spots anterolateral to PO; scutellar disk not colored···································· T. formosana (Bischoff)

- At most F8 - 14 with tyloids ( Figs. 5E , 6D ). Upper frons lateral to PO with transverse stripes connecting to inner orbital markings; scutellar disk yellow to reddish yellow at least laterally ····················································· 5

5. Apical markings on metasomal terga bi-colored, reddish orange anteriorly and yellow posteriorly. Larger species, more than 8 mm in body length ······················ ·························································· T. tricolor (Chen)

- Apical markings on terga not bi-colored but concolor yellow, at most apical band of T 2 anteriorly evanescently tinged with reddish brown. Smaller species, less than 6 mm in body length················· T. sauteri Bischoff

Figures

Fig. 4. Taeniogonalos subtruncata Chen et al. A, General habitus in dorsal view, ♀. B, General habitus in lateral view, ♀. C, Head in frontal view, ♀. D, S2-4, with a subapical tubercle on S3, ♀. E, Forewing,♀. F, F6-15, ♂. Scale bars: 1 mm.

Fig. 1. Taeniogonalos fasciata(Strand). A, General habitus in dorsal view, ♀. B, Head in frontal view, ♀. C, General habitus in lateral view, ♀. D, F8-14, ♂. Scale bars: 1 mm.

Fig. 6. Taeniogonalos tricolor (Chen).A, General habitus in dorsal view, ♀. B, Head in frontal view, ♀. C, General habitus in dorsal view, ♂. D, F7-14, ♂. Scale bars: 1 mm.

Fig. 2. Taeniogonalos formosana (Bischoff).A, General habitus in dorsal view, ♀. B, General habitus in lateral view, ♀. C, Head in frontal view,♀. D, Head in dorsal view, ♀. E, F7-21, ♂. Scale bars: 1 mm.

Fig. 3. Taeniogonalos sauteri Bischoff. A, General habitus in dorsal view, ♀. B, General habitus in lateral view, ♀. C, Head, in frontal view, ♀. D, S2-4 in ventral view, with semicircular flat production on apical margin of S2. E, General habitus in dorsal view, ♂. F, Head in front-dorsal view, ♂. G, F5-20, ♂. Scale bars: 1 mm.

Fig. 5. Taeniogonalos taihorina (Bischoff). A, General habitus in dorsal view, ♀. B, General habitus in lateral view, ♀. C, Head in frontal view, ♀. D, Head in dorsal view, ♀. E, F7-14, ♂. Scale bars: 1 mm.