Taeniogonalos formosana ( Bischoff, 1913 )

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 : 275-278

Taeniogonalos formosana ( Bischoff, 1913)

Mu-nui-gal-go-ri-beol (new Korean name) ( Fig. 2A - E )

Poecilogonalos formosana Bischoff, 1913: 151 , ♀ (holotype), Taiwan: Taihorn [Zoologisches Museum, Humboldt Universität, Berlin, Germany] .

Taeniogonalos formosana : Carmean and Kimsey, 1998: 67 (new combination).

Diagnosis. A black species with numerous yellow markings. It is distinct from the other Korean species of Taeniogonalos by the combination of a series of paired, well separated triangular spots on the T4 - 5, and a fully black scutellum.

Description. Female. Body length 4.4 - 13 mm, forewing length 3.7 - 11 mm.

Head. Occipital carina narrow in its entire length. Frons, vertex, temple, and gena anteriorly punctate-reticulate; mandible and clypeus densely punctate; gena posteriorly and occiput shiny, with moderate punctures; areas around posterior ocelli smooth.

Mesosoma. Mesoscutum punctate-reticulate, with longitudinal ridges posteriorly; scutellar disc longitudinally ridged anteriorly, irregularly reticulate posteriorly; propodeal dorsum obliquely ridged anteriorly, irregularly reticulate in remaining dorsum, propodeal side punctate-reticulate; propleuron densely punctate; pronotal side densely punctate anteriorly, longitudinally ridged posteriorly; mesopleuron punctate-reticulate, very often posterior declivity largely smooth and shiny; dorsal metapleuron moderately to densely punctate, ventral metapleuron sparsely to moderately punctate.

Metasoma. T1 0.56 × as broad as T2. T2 without median longitudinal depression. S1 1.5 × as broad as long. S2 more or less broadly and strongly beveled apicomedially, in profile, not evenly convex, highest posterior middle. T1 largely smooth, laterally scattered with punctures; T2 - 6 punctate-reticulate. S1 densely punctate laterally, moderately punctate medially; S2 densely punctate; S3 punctate-reticulate; S4 - 6 densely punctate.

Coloration. Body extensively pale yellow, yellow, deep yellow, or partially orange yellow in the following parts/ markings: posterior flat face of mandible, large lateral spots on clypeus, lower frons lateral to antennal sockets (linearly extending above along inner orbits), small spots anterior to anterior ocelli and anterolateral to posteri- or ocelli, paired transverse bands on vertex posteriorly (sometimes tinged with orange yellow, or almost entirely orange yellow, or much reduced) or a transverse band often with paired submesal and sublateral longitudinal stripes extending to ocellar region, stripes along outer orbits, pronotal dorsum, margins of lateral half of median lobe of mesoscutum, tegula, axillae, scutellum anterolateral to scutellar disc (sometimes lost), metanotal disc laterally, longitudinal thick sublateral stripes on propodeal dorsum, apical bands of T1 - 3 (that on T2 broadest; often those on T2 - 3 interrupted medially; in small specimens less than 4.0 mm in body length, apical band of T2 lost), paired large triangular spots on T4 - 5, T6 except medially, apical band of S1, apicolateral spots of S2. Inner face of all coxae, almost entire trochanters and trochantellus of all legs, basal and apical parts of all femora, inner faces of fore and mid tibia, and hind tibia basally pale yellow to yellow.

Antennae reddish brown ventrally, blackish brown dorsally. Legs except yellow coloration parts above mostly ferruginous.

Male. Much as in female except usual sexual dimorphic structures. Body length 6.5 - 9.2 mm, forewing length 5.5 - 8.0 mm. Antennae with tyloids as mentioned in the key. Larger apicomedian part of S2 very often weakly concave. Apical band of T2 often lost, or limited small apicolateral spots.

Biology. Taeniogonalos formosana is the second most commonly collected Trigonalid in Korea. It appears from mid-May to the end of September, and varies greatly in size, both clues pointing towards a wide range of potential hosts.

PT has studied the parasitoid complex of the Korean potter wasps ( Vespidae : Eumeninae) by collecting mud nests in winter, and has obtained this species on six occasions (seven specimens): four from Oreumenes decoratus nests, one from a small Eumenes sp. cell, the last from the nest of another, unidentified genus of Eumenid wasps.

On three occasions, one imago of T. formosana was found enclosed and dead in the cell, unable to get through the mud wall that apparently represented a deadly barrier. Due to poor collecting conditions, no remains, or evidence on the nature of the secondary host where collect- ed, and it could not be determined with certainty. We have not secured clear evidence of the Trigonalid wasps having consumed any of the Eumenid larvae. It may also have emerged from a parasitoid (wasp or fly) present internally in one of the preys.

On the last three occasions, the Trigonalid wasp emerg- ed from one of the fly pupae present in the parasitized cell, all in O. decoratus nests. On each case the batch of fly larvae that occupied the cell had devoured all the contents before pupation. On one occasion the fly larvae had invaded the adjacent cell by boring a hole in the partition wall.

Surprisingly, these flies are not Tachinidae , as expected, but are belongs to the closely related family Sarcophagidae , subfamily Miltogramminae ( Fig. 7 ), a group of kleptoparasitic flies known to deposit their eggs or young larvae in the open cells of solitary wasps and bees, or on the prey they carry in. This fly family has never been mentioned in literature as a potential host of Trigonalids; all Diptera yet recorded as secondary hosts belong to the family Tachinidae , and therefore entered the nest as an internal parasitoid of one of the stored prey.

Despite a relatively large number of Miltogramminae pupae present in each contaminated cell (20, 12 and 4), usually only one specimen of T. formosana emerged from the cell (2 in one case). The flies are of medium size (body length 6.5 - 7 mm, quite large for Miltogramminae ) and the T. formosana imagos were almost of the same size (6 - 7 mm long), showing that they had absorbed the full content of the fly larvae.

Three Trigonalid wasps emerged in spring, in synchrony with the flies, by digging an emergence hole in the pupa. On another occasion, the Trigonalid imago was found already fully formed but dead in the pupa in late fall, at the moment of collecting the nest. Three other similar pupae were found in the cell, from which the Miltogramminae flies emerged normally in the next spring.

The occurrence of this species in mud nests seems to be accidental, and may not be beneficial to it, as suggested by the numerous occasions where the adult wasp was found dead in the cell. This species seems poorly adapted to escape the mud nests. As a general rule, the accidental trapping of parasitoids is common in potter wasp nests ( Johnson et al., 2023).

Breeding records. South Korea ·[DJ] ♀? (minute), Wadong, found dead on 20 iii 1995 in Oreumenes decoratus nest; ♀, Wadong (vegetable garden in forested area), found dead on 27 iv 1995 in the closed cell of a small Eumenes sp. [CN] ♀, Kapsa, Gongju-si, found dead on 25 ii 1996 in the mud nest of an unidentified Eumenid (not Eumenes or Oreumenes ) [CB] ♂, Pyeongsan-ri, Dongi-myeon, Okcheon, emerged 17 iv 2022, ♀ emerged 27 iv 2022 from a single cell of O. decoratus containing 12 Miltogramminae pupae.; Saesan-ri, Dongi-myeon, Okcheon, ♂ emerged 27 iv 2022 from an O. decoratus nest of 4 connected cells, containing about 20 Miltogramminae pupae; Hangok-ri, Yongsan-myeon, Yeongdong, ♀ dead on 5 iii 22 in a Miltogramminae pupa among a batch of 4 pupae found in a cell of O. decoratus in forest.

Material examined (around 250 specimens from South Korea, including the following). South Korea ·[SL] Mt. Daemosan, Gangnam-gu, 23 vii 1996 (SH Kim) [ GG] ♀, Mt. Chukryeongsan , Sudong-myeon, Namyangju-si, 12 vii 1980 ( JI Kim) ; ♀, same locality, 28 ix 1980 ( HG Park) ; ♀, Gunpo-si , 24 vi 1986 ( JJ An) ; ♀, Gwangleung , Pocheon-si, 22 ix 1990 ( EJ Ryu) ; ♀, Gangssibong , Pocheon-si, 28 vi 1998 ( JD Yeo) ; ♀, Gwonseon-gu , Suwon-si, 2 ix 2000 (JN Gang) ; ♀, Palya-ri , Jinjeop-eup, Namyangju-si, 12 ix 2007 (SB Ha) ; ♀, Mt. Maguksan , Anseong-si, 23 vi 2009 ( JK Kim) ; ♀, Osammi-dong , Osan-si, 6 ix 2011 ( JK Kim) ; ♀, Bangchuk-ri , Hyeondeok-myeon, Pyeongtaek-si, 21 vi 2014 (OC Kwon) ; ♀, Geumchon-ri , Sinpyeong-myeon, Dangjin-si, 24 viii 2014 (OC Kwon) [GW] ♀, Gangchon , 4 ix 1982 ( HG Kim) ; 2♀♀, Yonsei Univ. Campus , Maeji-ri, Wonju-si, 11 vii 1996 ( HW Byun) ; ♀, same locality, 19 vii 1996 ( HW Byun) ; ♀, same locality, 15 vi 2007 (HW Byun & HY Han) ; 2♀♀, Mt. Balgyosan , Eoron-ri, Hongcheon-gun, 4 ix 1998 ( YG Park) ; ♀, Yongmunsa , Yangpyeong-gun 5 ix 1998 ( YG Park) ; Duwibong , Dangok , Chodong-ri , Sindong-eup, Jeongseon-gun, 22 vii 2000 ( ES Kim) ; ♀, Durobong , Yeongok-myeon, Gangleung-si, 19 viii 2001 (WM Kim) ; ♀, Seo-myeon , Chuncheon-si, 20 v 2004 ( JM Go & JY Yang) ; ♀, Muleunggyegok , Samhwa-dong, Donghae-si (37°28 ʹ 02 ʺ N 129°01 ʹ 53 ʺ E), 1 ix 2009 (SW Suk & YB Lee) [ CB] ♀, Mt. Minjujisan , Yeongdong-gun, 3 vi 1989 ( GC Jeong) ; ♀, Uipung-ri , Danyang-gun, 3 viii 1995 ( JY Cha) ; ♀, Mt. Namsan , Chungju-si, 28 viii 2000 ( JD Yeo) ; ♀, Gacheon-ri , Eomjeong-myeon, Chungju-si, 28 ix 2005 ( JS Lee) [ DJ] ♀, Yongeun-dong , 9 vi 1988 (GH Choi) [ CN] ♀, Hwahak-ri , Annam-myeon, Okcheon-gun, 17 ix 2007 (SB Ha) ; ♀, Daegok-ri , Haemi-myeon, Seosan-si, 22 vi 2013 ( CH Jang) ; 2♀♀, Dongmun-ri , Taean-gun, 21 viii 2014 ( JK Kim) ; ♀, Yonghyeon-ri , Unsan-myeon, Serosan-si, 24 viii 2014 ( JK Kim) ; 2♀♀, ♂, National Institute of Ecology , Maseo-myeon, Secheon-gun (38°01 ʹ 47.21 ʺ N 126°43 ʹ 36.02˝E), 22 v - 5 vi 2017 (Malaise trap) (OC Kwon) ; ♀, 17♂♂, same locality, 16 vi - 5 vii 2017 (Malaise trap) (OC Kwon) ; ♀, 3♂♂, same locality, 5 - 15 vii 2017 (Malaise trap) (OC Kwon) ; ♂, same locality, 15 vii - 1 viii 2017 (Malaise trap) (OC Kwon) ; 2♀♀, same locality, 12 - 27 viii 2017 (Malaise trap) (OC Kwon) ; ♀, same locality, 27 viii - 9 ix 2017 (Malaise trap) (OC Kwon) ; ♀, 30 v - 14 vi 2018 (Malaise trap) (OC Kwon); 3♀♀, 2♂♂, same locality, 14 - 27 vi 2018 (Malaise trap) (OC Kwon) ; 2♂♂, same locality, 27 vi - 16 vii 2018 (Malaise trap) (OC Kwon) ; 5♀♀, ♂, same locality, 16 - 24 vii 2018 (Malaise trap) (OC Kwon) ; ♀, 24 vii - 8 viii 2018 (Malaise trap) (OC Kwon); ♂, 8 - 22 viii 2018 (Malaise trap) (OC Kwon); 3♀♀, same locality, 4 - 9 ix 2018 (Malaise trap) (OC Kwon) ; 3♀♀, same locality, 27 v - 2 vi 2019 (Malaise trap) (OC Kwon) ; ♂, same locality, 2 - 9 vi 2019 (Malaise trap) (OC Kwon) ; 3♀♀, ♂, same locality, 9 - 17 vi 2019 (Malaise trap) (OC Kwon) ; ♀, ♂, 17 - 23 vi 2019 (Malaise trap) (OC Kwon); 3♀♀, 3♂♂, same locality, 23 vi - 1 vii 2019 (Malaise trap) (OC Kwon) ; ♀, 5 - 12 viii 2019 (Malaise trap) (OC Kwon); 3♂♂, same locality, 12 - 19 viii 2019 (Malaise trap) (OC Kwon) ; ♂, same locality, 19 - 26 viii 2019 (Malaise trap) (OC Kwon) ; ♀, same locality, 5 - 9 ix 2019 (Malaise trap) (OC Kwon) ; ♀, same locality, 14 - 21 ix 2019 (Malaise trap) (OC Kwon) ; 2♂♂, same locality, 1 - 8 vi 2020 (Malaise trap) (OC Kwon) ; ♀, ♂, same locality, 8 - 15 vi 2020 (Malaise trap) (OC Kwon) ; ♀, ♂, same locality, 15 - 22 vi 2020 (Malaise trap) (OC Kwon) ; ♂, same locality, 22 - 29 vi 2020 (Malaise trap) (OC Kwon) ; ♀, same locality, 29 vi - 6 vii 2020 (Malaise trap) (OC Kwon) [GB] Mt. Juwangsan , 30 vii 1983 ( HG Park) ; ♀, Cheongpyeong , Cheongsong-gun, 16 vii 1988 ( HJ Oh) ; ♀, Hyeonnae-ri , Gigye-myeon , Buk-gu, Pohang-si (36°11 ʹ 06 ʺ N 129°04 ʹ 03 ʺ E), 30 viii 2011 (SW Suk et al.) ; ♀, ♂, Dodeok-ri , Angye-myeon, Uiseong-gun (36°25 ʹ 49.02 ʺ N 128°27 ʹ 35.70 ʺ E), 21 vi - 5 vii 2017 (Malaise trap) ( JK Kim) ; ♂, same locality, 2 - 16 viii 2017 (Malaise trap) (OC Kwon) [ GN] Jangdangol , Mt. Jirisan, Hadong-gun, 1 viii - 8 ix 2001 (Malaise trap) ( JW Lee) ; ♂, Baetaegogae , Yeongpo-ri , Wondong-myeon, Yangsan-si (36°26 ʹ 23.2 ʺ N 128°57 ʹ 28.2 ʺ E), 11 - 25 vi 2019 (Malaise trap) (JA Jeon) [JN] ♀, Sokhyeon-dong , Suncheon-si, (34°58 ʹ 47.01 ʺ N 127°27 ʹ 40.563 ʺ E), 4 - 18 vii 2017 (Malaise trap) (OC Kwon) ; ♀, same locality, 1 - 15 viii 2017 (Malaise trap) (OC Kwon) ; ♂, Ondang-ri , Gwanggui-myeon, Gurye-gun (35°17 ʹ 17.77 ʺ N 127°26 ʹ 40.03 ʺ E), 15 - 29 v 2018 (Malaise trap) (OC Kwon) .

Distribution. Korea (SL, GG, GW, CB, DJ, CN, GB, GN, JN; new record), China (Jilin, Shaanxi, Ningxia, Henan, Fujian, Zhejiang, Guangdong, Sichuan, Yunnan, Guizhou, Tibet), Russian Far East (Amurskaya Oblast, Primorskii Krai, South Sakhalin, Kuril Islands: Kunashir, Shikotan), Japan (Hokkaido, Honshu), Taiwan.

Remarks. Chen et al. (2014) synonymized T. intermedia and T. unifasciata as color forms of T. formosana . Body markings of all the Korean materials herein are primarily yellow, as shown in intermedia - and unifasciata -form. No typical formosana -form with primarily reddish-orange coloration is found in Korea.

Figures

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. 7. T. formosana freshly emerged from the pupa of his Miltogramminae host, near a dead specimen of the fly.