Revision of the genus Woldstedtius Carlson, 1979 (Hymenoptera, Ichneumonidae, Diplazontinae) from Japan

Shunsuke Morishita; Kyohei Watanabe

doi:10.3897/dez.69.80492

Review Article

Revision of the genus Woldstedtius Carlson, 1979 (Hymenoptera, Ichneumonidae, Diplazontinae) from Japan

Shunsuke Morishita^‡, Kyohei Watanabe^§

‡ Hatchodori, Toyohashi, Japan

§ Kanagawa Prefectural Museum of Natural History, Odawara, Japan

Corresponding author: Shunsuke Morishita ( ezogengoroumodoki4.23@gmail.com )

Academic editor: Jose Fernandez-Triana

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Morishita S, Watanabe K (2022) Revision of the genus Woldstedtius Carlson, 1979 (Hymenoptera, Ichneumonidae, Diplazontinae) from Japan. Deutsche Entomologische Zeitschrift 69(1): 45-64. https://doi.org/10.3897/dez.69.80492

ZooBank: urn:lsid:zoobank.org:pub:C402CAE8-14FD-402A-AD05-54646B1493E8

Abstract

Japanese species of the genus Woldstedtius Carlson, 1979 are revised. Nine species are recorded from Japan, including two new species, W. alpicola sp. nov. and W. punctatus sp. nov. Woldstedtius biguttatus (Gravenhorst, 1829) is newly recorded from Japan. Taxonomic status of W. flavolineatus kuroashii (Uchida, 1957) is changed from the subspecies of W. flavolineatus (Gravenhorst, 1829) to a separated species. Woldstedtius holarcticus (Diller, 1969) is newly synonymized under W. kuroashii (Uchida, 1957). A key to Japanese species of this genus is provided.

Key Words

Asia, fauna, new species, parasitoid wasps, taxonomy

Introduction

The Ichneumonid subfamily Diplazontinae comprises 23 genera and more than 350 species worldwide (Yu et al. 2016). The host range of species in this subfamily appears to be notably narrow, with all confirmed hosts being species of hoverfly (Diptera, Syrphidae). The only exceptions in this regard are two species in the ichneumonid genus Bioblapsis Förster (Klopfstein 2014). The genus Woldstedtius Carlson, 1979 is a relatively large taxon comprising 44 species distributed in the Australasian, Holarctic, Neotropical, Oceanic, and Oriental regions (Balueva and Lee 2016; Vas 2016; Yu et al. 2016; Johansson 2020). Although the genus has been revised by Klopfstein (2014) for the Western Palearctic species, by Dasch (1964) for the Nearctic species and by Gauld et al. (1997) for the Costa Rican species, members in the Eastern Palearctic region have only been partially studied. Manukyan (2007), for example, provided a key to the six species of this genus known from Far East Russia, and Balueva and Lee (2016) have reviewed the South Korean species. In Japan, seven species have been formally recorded to date (Yu et al. 2016), however, we have found specimens of some unidentified species and identified a number of unresolved taxonomic problems. Nevertheless, given the large number of previous studies in other regions, this genus may be an adequate taxon for comparison of the fauna with that in other parts of the world.

The purpose of this study was to undertake a taxonomic review of the Japanese species of Woldstedtius. We have also produced a key to the Japanese species of this genus, which is presented herein.

Materials and methods

In this study, the dried specimens deposited in the following collections were examined:

AEIC American Entomological Institute, Logan, Utah, USA;

KPMNH Kanagawa Prefectural Museum of Natural History, Odawara, Kanagawa, Japan;

MU Meijo University, Nagoya, Aichi, Japan;

NSMT National Museum of Nature and Science, Tsukuba, Ibaraki, Japan;

SEHU Systematic Entomology, Hokkaido University, Sapporo, Japan;

TMNH Toyohashi Museum of Natural History, Toyohashi, Aichi, Japan;

ZSM Zoologische Staatssammlung München, Germany.

Stereomicroscopes (SMZ745 and SMZ800: Nikon, Tokyo) were used for observation. Photographs (Figs 1, 2, 4–7) were taken by digital camera (TG-4: Olympus, Tokyo) attached to the stereomicroscope (SMZ800). Photographs (Figs 3, 8) were taken by digital camera (CX6: RICOH, Tokyo) attached to the stereomicroscope (SMZ745). Digital images (Figs 1–9) were edited using Adobe Photoshop.

Morphological terminology mainly follows those established by Broad et al. (2018). The following abbreviations are used in description, diagnosis, and remarks: ocello-ocular line (OOL), postocellar line (POL), diameter of lateral ocellus (OD), segments of flagellomeres (FL), segments of maxillary palp (MP), metasomal tergites (T) and holotype (HT). The following abbreviations are used in material data: female (F), male (M), flight interception trap (FIT), light trap (LT), Malaise trap (MT) and yellow pan trap (YPT).

Results and discussion

Woldstedtius flavolineatus kuroashii (Uchida 1957) was originally described based on a single female, for which no morphological information, apart from the coloration of mesosoma and legs, is available. Therefore, we examined Japanese specimens of W. f. kuroashii, including the holotype, and compared these with the identified European specimens of W. f. flavolineatus deposited in ZSM (identified by Diller). On the basis of these comparison, we established that certain character states of W. f. kuroashii are clearly different from those of W. f. flavolineatus, and accordingly conclude that this subspecies should be treated as a separate species, W. kuroashii.

Woldstedtius holarcticus Diller, 1969 resembles W. kuroashii in terms of color and body structures. In Japan, Manukyan (2007) recorded this species from Kunashiri Island, although no additional specimens of the species have been recorded elsewhere in Japan. We compared the Japanese specimens of W. kuroashii, including holotype, with a paratype of W. holarcticus deposited in ZSM. As we detected no clear differences between the two taxa, we propose that W. holarcticus should be synonymized under the name W. kuroashii. Diller (1969) noted that females of W. holarcticus (= W. kuroashii) can be distinguished from those of W. flavolineatus with respect to coloration of the coxae, the proportions of the hind tibia and tarsus, and surface gloss of the body, and also established that the females of W. holarcticus (= W. kuroashii) can be distinguished from those of W. f. flavolineatus based on the medially flattened face (medially convex in W. f. flavolineatus). We confirmed this character state and established that the face is slightly convex medially in both W. kuroashii and W. f. flavolineatus, although this feature is weaker in the former than in the latter (Fig. 9A, B). However, given the overlap in the intraspecific variation of this character state, it cannot be reliably used to differentiate the two species. The leg coloration (with the exception of the coxae) of W. kuroashii collected from Honshu (including the holotype) is darker compared with that of some Japanese specimens collected from Hokkaido and female paratypes of W. holarcticus (blackish-brown to black) (orange to brown in some Japanese specimens collected from Hokkaido and the paratype of W. holarcticus). As a consequence of these comparisons, we detected no significant differences with respect to other character states of these specimens, and accordingly conclude that the observed variation is intraspecific variation within a single species.

In this study, we identified specimens of W. biguttatus from Japan for the first time. This species closely resembles W. flavolineatus, and indeed, some W. biguttatus specimens have been incorrectly recorded as W. flavolineatus e.g., Konishi et al. (2014) and Morishita et al. (2021). In contrast, we were unable to find any specimens of W. flavolineatus that have been collected in Japan. Given that some of the voucher specimens of the previous studies could not be reliably identified owing to inadequate labeling, we have certain reservations as to the Japanese distribution records of W. flavolineatus, and accordingly recommend a further re-examination of the distribution of this species.

On the basis of the aforementioned taxonomic treatments, we conclude that a total of nine species in the genus Woldstedtius have been found in Japan to date.

Taxonomy

Subfamily Diplazontinae Viereck, 1918

Genus Woldstedtius Carlson, 1979

Type

Bassus biguttatus Gravenhorst, 1829: 332. Original designation.

Diagnosis

According to Klopfstein (2014), this genus can be distinguished from other genera by the following combination of character states: antenna without tyloids in males; mesoscutum without notauli; fore wing areolet absent; hind tibia usually black with a white base, in males with light coloration often extending to half the length of the tibia, rarely hind tibia all dark or yellow or orange with a dark apex; ovipositor sheaths transversely truncate and open towards apex; very even, weakly coriaceous microsculpture.

Remarks

Japanese species can be identified by the following key.

Key to Japanese species of Woldstedtius (males of W. takagii and W. yokohamensis are unknown)

1	Hind coxa white with a brown dorsal stripe in females (Fig. 4A). Face blackish-brown to black with a white median spot, this spot connected with white clypeus in females (Fig. 4B), entirely white in males (Fig. 4F). Scutellum with large white marking (Fig. 4C). Latero-median carina of T I short, along less than basal 0.3 of T I (Fig. 9H)	W. karafutensis (Uchida, 1957) (female and male)
–	Above combination of character states lacking. Hind coxa entirely black or entirely orange in females (Figs 1A, 2A, 3A, 5A, 6A, 7A, 8A). Coloration of face various in females. Scutellum entirely black or black with a whitish-yellow to yellow apical spot (Figs 1C, 2C, 3C, 5C, 6C, 7C, 8C). Latero-median carina of T I long, along more than basal 0.4 of T I (Fig. 9E–G, I–K)	2
2	Hind coxa entirely black in both sexes (Figs 1A, 5A, 6A, E, 7A, 8A)	3
–	Hind coxa entirely orange in females (Figs 2A, 3A), entirely or largely yellow to reddish-yellow and usually with a blackish-brown dorsal stripe in males (Figs 1E, 2E, 3 E, 5E)	7
3	Inner orbits divergent downward (Figs 7B, 9C). Face 2.6–2.9 × as broad as high. Basal 0.2 of propodeum weakly protruded in lateral view (Fig. 9D). Yellow shoulder marks of mesoscutum absent (Fig. 7A)	W. takagii (Uchida, 1957) (female)
–	Inner orbits almost parallel (Figs 1B, 5B, 6B, 8B). Face less than 2.5 × as broad as high. Basal 0.2 of propodeum not protruded in lateral view. Yellow shoulder marks of mesoscutum present (Figs 1A, 5A, 6A, 8A)	4
4	Scutellum coarsely and densely punctate (separated by ca. 0.8–1.3 × their diameter) and entirely black (Fig. 6C). Propodeum rugulose (Fig. 6D)	W. punctatus sp. nov. (female and male)
–	Scutellum finely and sparsely punctate (separated by ca. 1.5–2.5 × their diameter) and black with a whitish-yellow to yellow apical spot (Figs 1C, 5C, 8C). Propodeum coriaceous or at least finely rugulose	5
5	Face black with a pair of yellow spots along inner orbits (Fig. 8B). Scutellum finely and densely punctate (separated by ca. 1.0 × their diameter)	W. yokohamensis (Uchida, 1957) (female)
–	Face black with a whitish-yellow to yellow median spot (Figs 1B, 5B). Scutellum finely and sparsely punctate (separated by ca. 1.5–2.5 × their diameter) (Figs 1C, 5C)	6
6	T I 1.4–1.5 × as long as maximum width. Propodeum finely rugulose (Fig. 1D). Hind trochanter black (Fig. 1A). Pleural carina of propodeum entirely present	W. alpicola sp. nov. (female)
–	T I 1.1–1.25 × as long as maximum width. Propodeum coriaceous (Fig. 5D). Hind trochanter white (Fig. 5A). Pleural carina of propodeum absent posteriorly	W. kuroashii (Uchida, 1957) (female)
7	Inner orbits divergent downward. Antenna with 19–21 (rarely 22) flagellomeres. Metasoma entirely black or black with some orange markings in females	W. citropectoralis (Schmiedeknecht, 1926) (female and male)
–	Inner orbits almost parallel in females (Figs 2B, 3B), weakly divergent downward in males (Figs 1F, 2F, 3F, 5F). Antenna with 22–25 flagellomeres. Metasoma entirely black in females	8
8	Mesoscutum without yellow shoulder marks in females (Fig. 2A). Face entirely black or black with a small median yellow or brown spot in females (Fig. 2B). Mesopleuron with a yellow ventral marking (Fig. 2E) in males. Mesosternum black in males (Fig. 2E)	W. biguttatus (Gravenhorst, 1829) (female and male)
–	Mesoscutum with yellow shoulder marks in females (Fig. 3A). Face black with a large yellow median spot in females (Fig. 3B). Mesopleuron with a large whitish-yellow to yellow marking; it is enlarged anteriorly in males (Figs 1E, 3E, 5E). Mesosternum yellow in males (Figs 1E, 3E, 5E)	9
9	Propodeum finely rugulose (e.g., Fig. 1D), with a complete pleural carina. Bases of T IV to T VII each with a transverse yellow band	W. alpicola sp. nov. (male)
–	Propodeum coriaceous (e.g., Figs 3D, 5D), with a pleural carina absent posteriorly in both sexes. Coloration of T IV to T VII various in males	10
10	Bases of T III and T IV each with a transverse yellow basal band in males. T I entirely coriaceous	W. flavolineatus (Gravenhorst, 1829) (female and male)
–	T III with a pair of whitish-yellow apical spots (sometimes these spots connected each other). T IV and T V each with a transverse whitish-yellow apical band. T I coriaceous, with irregular rugae laterally	W. kuroashii (Uchida, 1957) (male)

Woldstedtius alpicola sp. nov.

http://zoobank.org/E030CAC0-73B1-4BDC-BD63-F5E6BD3BAC4B

Figs 1A–F, 9E

Type series

Holotype : F, Japan, Honshu, Mie Pref., Inabe City, Mt. Fujiwaradake, 1 Jun 2021, S. Morishita leg. (KPMNH). Paratypes: Japan: [Honshu] 1 F, Yamanashi Pref., Koshu City, Mt. Daibosatsu, Kaminikkawa-toge, 16 Jun 2007, K. Watanabe leg. (TMNH); 1 M, Kanagawa Pref., Yamakita Town, Mt. Oomuroyama–Mt. Kanyuudousan, 15 Jun 2008, H. Kawai leg. (KPMNH); 1 F, Nagano Pref., Otaki Vil., Mt. Ontakesan, 13–25 Jul 2015, S. Shimizu leg. (MT) (KPMNH).

Figure 1.

Woldstedtius alpicola sp. nov. (A–D. female, holotype; E, F. male, paratype) ― A, E. Habitus; B, F. Head, frontal view; C. Mesonotum, dorsal view; D. Propodeum, dorsal view.

Description

Female (n = 3). Body length 8.5–9.2 (HT: 8.5) mm, polished, coriaceous and covered with silver setae.

Head 0.51–0.54 (HT: 0.53) × as long as wide. Clypeus 2.0–2.35 (HT: 2.0) × as broad as high, slightly convex basally in lateral view. Face 2.1–2.4 (HT: 2.4) × as broad as high, densely punctate, convex medially in lateral view and separated from clypeus by shallow clypeal sulcus. Inner orbits almost parallel (Fig. 1B). Length of malar space 0.9–1.15 (HT: 0.9) × as long as basal mandibular width. POL 1.9–2.1 (HT: 1.9) × as long as OD. OOL 0.9–1.18 (HT: 0.9) × as long as OD. POL 1.86–1.95 (HT: 1.86) × as long as OOL. Antenna with 24–25 (HT: 24) flagellomeres. FL I 1.29–1.35 (HT: 1.35) × as long as FL II. MP IV1.42–1.53 (HT: 1.53) × as long as MP V.

Mesosoma. Lateral aspect of pronotum strigose anteriorly. Mesoscutum finely and densely punctate (separated by ca.1.0 × their diameter) (Fig. 1C). Scutellum finely and sparsely punctate (separated by ca.1.5–2.5 × their diameter) (Fig. 1C). Anterior part and lower half of mesopleuron coarsely and densely punctate. Speculum smooth. Sternaulus weakly impressed. Propodeum finely rugulose (Fig. 1D), rounded in lateral view and pleural carina entirely present. Fore wing length 7.4–8.5 (HT: 7.4) mm. Nervellus intercepted below middle. Hind femur 4.7–5.3 (HT: 4.7) × as long as maximum depth in lateral view. Hind tibia 7.5–8.0 (HT: 8.0) × as long as maximum depth in lateral view. Ratio of length of hind first to fifth tarsomeres 1.0: 0.6: 0.4: 0.2: 0.2–0.3 (HT: 0.2).

Metasoma. T I rectangular in dorsal view (Fig. 9E), 1.4–1.5 (HT: 1.4) × as long as maximum width, irregular rugulose laterally, sometimes striate between latero-median carinae. Latero-median carina present basal ca. 0.5 of T I (Fig. 9E). T II 0.6–0.73 (HT: 0.73) × as long as maximum width, striate anteriorly and strigose laterally.

Coloration (Fig. 1A–D). Body (excluding wings and legs) black. Face with a large yellow spot medially. Palpi, subtegular ridge and mesepisternum yellow. Mandible yellow, except for apex and base. Lateral aspect of pronotum with a yellow spot posteriorly. Mesoscutum with yellow shoulder marks. Tegula tinged with yellow anteriorly. Scutellum with a yellow spot apically. Wings hyaline. Veins and pterostigma blackish-brown except for yellowish-brown wing base. Legs blackish-brown to black. Fore trochanter, trochantellus, apex of hind femur and base of hind tibia tinged with white. Fore femur, tibia and tarsomeres orange to brown. Apex of hind trochantellus and base of hind femur tinged with reddish-brown.

Male (n = 1). Similar to female. Body length (excluding antennae) 8.1 mm. Face 2.5 × as broad as high. Length of malar space 0.8 × as long as basal mandibular width. POL 2.2 × as long as OD. OOL 1.2 × as long as OD. POL 2.1 × as long as OOL. Inner orbits weakly divergent downward (Fig. 1F). Fore wing length 6.3 mm.

Coloration (Fig. 1E, F). Body (excluding wings and legs) black. Clypeus, palpi, face, ventral surface of antenna, malar space, propleuron, epicnemium, mesosternum, tegula, subtegular ridge and mesepisternum yellow. Gena tinged with yellow ventrally. Mandible yellow, except for apex. Lateral aspect of pronotum tinged with yellow ventrally and posteriorly. Mesopleuron with a large yellow marking, it enlarged anteriorly. Scutellum with a yellow spot apically. T IV to T VII with a transverse yellow band posteriorly. Wings hyaline. Veins and pterostigma brown to blackish-brown except for yellow wing base. Legs yellow to yellowish-brown. Hind coxa and trochanter each with a blackish-brown stripe dorsally. Hind trochantellus and tibia tinged with blackish-brown.

Distribution

Japan (Honshu).

Bionomics

Host unknown. Adults were collected in broad-leaved forest at altitudes of ca. 1,000–2,000 meters.

Etymology

The species name refers that this species inhabits alpine region.

Remarks

This species resembles W. kuroashii but can be distinguished from the latter by the following combination of character states: pleural carina of propodeum entirely present in both sexes (absent posteriorly in both sexes of W. kuroashii); propodeum finely rugulose in both sexes (coriaceous in both sexes of W. kuroashii); T I 1.4–1.5 × as long as maximum width in both sexes (1.1–1.25 in females, 1.14–1.26 in males of W. kuroashii). This species also resembles a Korean species, W. pallidus Balueva & Lee, 2016 (male is unknown), but can be distinguished by the following combination of character states in females: a large yellow median spot of face present (absent in W. pallidus); antenna with 24–25 flagellomeres (22–23 in W. pallidus); scutellum finely and sparsely punctate on coriaceous background (entirely coriaceous in W. pallidus); propodeum finely rugulose (coriaceous in W. pallidus).

Woldstedtius biguttatus (Gravenhorst, 1829)

Figs 2A–F, 9F

Bassus biguttatus Gravenhorst, 1829: 332.

Bassus rufipes Gravenhorst, 1829: 337. Name preoccupied.

Bassus confusus Woldstedt, 1874: 63. Synonymized by Morley (1906).

Syrphoctonus flavolineatus: Konishi et al. 2014: 491. Misident (at least in part).

Woldstedtius flavolineatus: Morishita et al. 2021: 53. Misident.

Materials examined

JAPAN: [Hokkaido] 1 F, Hokkaido, Nemuro, Shibetsu, Rubesu, 25–28 Aug 1971, K. Yamagishi leg. (MU); 1 F, Hokkaido, Sapporo, 20 May 1967, K. Kusigemati leg. (SEHU). [Honshu] 1 F, Gunma Pref., Katashina Vil., Marunuma, Yuzawa, 2 Jul 2008, K. Watanabe leg. (KPMNH); 1 F, Saitama Pref., Honjo City, Okubo-yama, 1 Apr 2001, N. Shimizu leg. (KPMNH); 1 F, Saitama Pref., Satte City, Makinoji, 6 Apr 2009, S. Yoshizawa leg. (KPMNH); 2 F, Tokyo, Chiyoda, Imperial Palace, Fukiagegyoen 20 May–19 Jun 1996, K. Konishi leg. (MT) (NSMT); 1 F, Tokyo, Chiyoda, Imperial Palace, Fukiagegyoen, Kajuen, 17–24 Sep 2009 (MT) (NSMT); 1 F & 1 M, ditto, 14–21 Oct 2009 (MT) (NSMT); 1 M, ditto, 13–20 Apr 2010 (MT) (NSMT); 1 F, ditto, 11–18 May 2010 (MT) (NSMT); 1 M, Tokyo, Chiyoda, Imperial Palace, Fukiagegyoen, Otakinagare, 29 Mar–6 Apr 2009 (MT) (NSMT); 1 F, ditto, 12–17 May 2011 (MT) (NSMT); 1 F, ditto, 17–24 May 2011 (MT) (NSMT); 1 F, Tokyo, Chiyoda, Imperial Palace, Biol. Inst., nr. Paddy field, 12–17 May 2011 (MT) (NSMT); 1 F, Tokyo, Chiyoda, Imperial Palace, Dokan-shinmichi, 7–14 Oct 2009 (MT) (NSMT); 1F & 1 M, ditto, 21–28 Oct 2009 (MT) (NSMT); 4 F & 1 M, ditto, 28 Oct–4 Nov 2009 (MT) (NSMT); 2 F & 1 M, ditto, 11–18 Nov 2009 (MT) (NSMT); 1 M, ditto, 18–24 Nov 2009 (MT) (NSMT); 1 F & 4 M, ditto, 24 Nov–7 Dec 2009 (MT) (NSMT); 1 M, ditto, 3–17 Mar 2010 (MT) (NSMT); 3 M, ditto, 29 Mar–6 Apr 2010 (MT) (NSMT); 2 F, ditto, 6–13 Apr 2010 (MT) (NSMT); 1 F, ditto, 27 Apr–4 May 2010 (MT) (NSMT); 3 F, ditto, 4–11 May 2010 (MT) (NSMT); 6 F & 2 M, ditto, 25 May–1 Jun 2010 (MT) (NSMT); 1 M, Tokyo, Chiyoda, Imperial Palace, Dokan-bori, 23 Mar 2010, K. Watanabe leg. (MT) (NSMT); 2 M, Tokyo, Ome City, Mt. Otsukayama, 1 Jun 2008, M. Irie leg. (KPMNH); 3 M, Tokyo, Ome City, Mt. Mitakesan, 1 Jun 2008, M. Gunji leg. (KPMNH); 1 F, Kanagawa Pref., Fujisawa City, Ishikawa, 11 May 2001, I. Waki leg. (NSMT); 1 F, Kanagawa Pref., Ebina City, Sagamigawa, 13 Apr 2006, M. Ooishi & R. Watanabe leg. (YPT) (KPMNH); 1 F, Kanagawa Pref., Chigasaki City, Serizawa, 10 May 2013, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Atsugi City, Funako, Tokyo University of Agriculture, 6 May–7 Jun 2016, Y. Kato & S. Koizumi leg. (MT) (KPMNH); 1 M, ditto, 22 Apr–16 May 2016, Y. Kato & S. Koizumi leg. (MT) (KPMNH); 1 M, Kanagawa Pref., Atsugi City, Tokyo University of Agriculture, 2 Apr 2009, H. Katahira leg. (KPMNH); 1 M, Kanagawa Pref., Oiso Town, Koma, Komayama, 16 Apr 2016, K. Watanabe leg. (KPMNH); 4 M, Kanagawa Pref., Nakai Town, Zoushiki, 16 Apr 2019, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Kiyokawa Vil., 4 Oct 2008, Y. Oogane leg. (KPMNH); 2 F, Kanagawa Pref., Hadano City, Koubouyama, 1 May 2016, K. Watanabe & H. Utsugi leg. (KPMNH); 2 M, ditto, 18 Apr 2010, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Hadano City, Chimura, Mt. Zukkoyama, 16 Apr 2017, K. Watanabe leg. (KPMNH); 1 M, ditto, 7 May 2017, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Tennoujione, 29 Jun 2013, (FIT) (KPMNH); 3 M, Kanagawa Pref., Kaisei Town, Kanaishima, 22 Mar 2016, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Odawara City, Kuno, 24 Mar 2014, K. Watanabe leg. (KPMNH); 1 M, ditto, 24 Mar 2014, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Odawara City, Kamisoga, 29 Apr 2017, K. Watanabe leg. (KPMNH); 1 F, Kanagawa Pref., Yamakita Town, Hinokiboramaru, 6 Aug 2014, T. Taniwaki leg. (KPMNH); 1 F, Kanagawa Pref., Yamakita Town, Mt. Oomuroyama–Mt. Kanyuudousan, 15 Jun 2008, H. Kawai leg. (KPMNH). 1 F, Yamanashi Pref., Koshu City, Enzanushioku, Sagashio, 12 Jun 2010, K. Watanabe leg. (KPMNH); 1 F, Yamanashi Pref., Hokuto City, Masutomi, Biwakubo-sawa, 28 Jul 2007, K. Watanabe leg. (KPMNH); 1 F, Yamanashi Pref., Yamanakako Vil., Hirano, Mikuni-toge, 8 Aug 2020, S. Morishita leg. (TMNH); 3 F, Shizuoka Pref., Shizuoka City, Umegashima, 3 Jun 2001, T. Sugiyama leg. (YPT) (MU); 2 F, ditto, 18 Jun 2001, T. Sugiyama leg. (YPT) (MU); 2 F, ditto, 18 Jun–2 Jul 2001, T. Sugiyama leg. (MT) (MU); 1 F, ditto, 17 Jul–5 Aug 2001, T. Sugiyama leg. (MT) (MU); 1 F, Aichi Pref., Shitara Town, Tsuguhonsawa, 7 May–26 Jun 2020, S. Morishita leg. (MT) (TMNH); 1 F, ditto, 17 Jun 2019, S. Morishita leg. (TMNH); 1 F, ditto, 5 Jul 2019, S. Morishita leg. (TMNH); 1 F, Aichi Pref., Shinshiro City, Tsukudeiwanami, 17 May 2019, S. Morishita leg. (TMNH); 1 M, ditto, 2 May 2019, S. Morishita leg. (TMNH); 3 F & 4 M, Aichi Pref., Toyohashi City, Hosoya, Kitahosoya, 6 Apr 2019, S. Morishita leg. (TMNH); 1 F, Aichi Pref., Toyohashi City, Unoya, Nabeyamashita, 10 Apr 2019, S. Morishita leg. (TMNH); 1 F & 1 M, ditto, 30 Apr 2019, S. Morishita leg. (TMNH); 2 M, Aichi Pref., Toyohashi City, Oiwa, Taimatsu-toge, 6 Apr 2021, S. Morishita leg. (TMNH); 1 F, Aichi Pref., Toyohashi City, Imure, Takayama, 2 Jun–6 Jun 2019, S. Morishita leg. (MT) (TMNH); 1 F, Aichi Pref., Toyohashi City, Nishiiwata, 12 Jul 2018, S. Morishita leg. (TMNH); 1 M, ditto, 3 May 2019, S. Morishita leg. (TMNH); 3 M, Aichi Pref., Toyokawa City, Mikami-cho, 15 Sep 2019, S. Morishita leg. (TMNH); 1 M, Aichi Pref., Kasugai City, Takagi, 19 May 2017, M. Sugiura leg. (MU); 2 F, Aichi Pref., Toyota City, Sanage, 30 Apr–6 May 2002, M. Kiyota leg. (MT) (MU); 1 F, Aichi Pref., Asuke Town, Tanoshiri, 15–24 May 2005, Y. Nishimura leg. (MT) (MU); 1 F, ditto, 15–21 Jun 2005, Y. Nishimura leg. (MT) (MU); 1 F, ditto, 15–21 Jun 2005, J. Yamagiwa leg. (MT) (MU); 1 F, ditto, 13–19 Jul 2005, Y. Nishimura leg. (MT) (MU); 1 F, Aichi Pref., Nisshin City, Komenogi, 28 May–3 Jun 2011, H. Seo leg. (MT) (MU); 1 F, Aichi Pref., Nisshin City, Nokata, 28 May–3 Jun 2011, H. Seo leg. (MT) (MU); 1 F, Aichi Pref., Ichinomiya City, 14–20 Oct 2006, C. Ueshima leg. (MT) (MU); 1 M, Gifu Pref., Kani City, Katabira 3–9 Apr 2004, K. Ito leg. (MT) (MU); 1 M, ditto, 10–16 Apr 2004, K. Yamagishi leg. (MT) (MU); 2 F, ditto, 17–23 Apr 2004, K. Ito leg. (MT) (MU); 1 M, ditto, 1–7 May 2004, K. Ito leg. (MT) (MU); 1 M, ditto, 15–21 May 2004, K. Ito leg. (MT) (MU); 1 F, Nagano Pref., Kawakami Vil., Azusayama, 14 Jun 2015, K. Watanabe leg. (KPMNH); 1 F, Nagano Pref., Nagawa Town, Daimon, 27 Aug 2011, S. Fujie leg. (KPMNH); 1 F, Nagano Pref., Otaki Vil., Mt. Ontakesan, Hakkaisan, 8 Aug 2010, K. Watanabe leg. (KPMNH); 1 F, ditto, 25 Jun–15 Jul 2015, S. Shimizu leg. (MT) (KPMNH); 1 F, Toyama Pref., Mt. Jodosan, 7 Jun 1972, M. Watanabe leg. (KPMNH); 1 M, Toyama Pref., Toyama City, Inonedani, 7–14 Jul 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, ditto, 8–15 Sep 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, ditto, 15–22 Sep 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, Toyama Pref., Toyama City, Kamegai, 1–8 Sep 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, ditto, 8–15 Sep 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, Toyama Pref., Toyama City, Jurodani, 7–14 Jul 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, Toyama Pref., Nanto City, Kamimomose, 11–18 Aug 2009, M. Watanabe leg. (MT) (KPMNH); 1 M, ditto, 1–8 Sep 2009, M. Watanabe leg. (MT) (KPMNH); 4 M, ditto, 15–29 Sep 2009, M. Watanabe leg. (MT) (KPMNH); 2 F, Ishikawa Pref., Nomi City, Mitsukuchi, 30 Apr–13 May 2011, R. Ishiguro leg. (MT) (MU); 1 F, ditto, 6–21 Oct 2011, H. Fukutomi leg. (MT) (MU); 1 F, Ishikawa Pref., Hakusan City, Sannomiya, 17–23 May 2009, H. Fukutomi leg. (MT) (MU); 1 F, ditto, 24–30 May 2009, H. Fukutomi leg. (MT) (MU); 1 F & 1M, Mie Pref., Taiki Town, Takihara, 20–31 May 2019, T. Nishimura leg. (MT) (MU); 1 M, Kyoto Pref., Yawata City, Yawatahayashinomoto, 15 May 2015, K. Watanabe leg. (KPMNH); 2 F, Osaka Pref., Chihayaakasaka Vil., Mt. Kongosan, 7–15 Jul 2012, S. Fujie leg. (MT) (KPMNH); 1 F, Hyogo Pref., Kobe City, Motoyama Town, Hokura-jinjya, 5 May 2012, K. Watanabe leg. (KPMNH); 1 F, Tottori Pref., Wakasa Town, Mt. Hyonosen, 17 Jul 2011, K. Watanabe leg. (KPMNH). [Shikoku] 1 F & 1 M, Tokushima Pref., Zennyuji-toh, 13–22 May 2003, H. Otsuka leg. (MT) (MU). [Kyushu] 1 F, Kagoshima Pref., Sakurajima, 13 May 1973, K. Kusigemati leg. (SEHU); 1 F, Kagoshima Pref., Terayama, 19 Apr 1973, K. Kusigemati leg. (SEHU); 1 F, Kagoshima Pref., Uearata, 18 Oct 1973, K. Kusigemati leg. (SEHU). BULGARIA: 1 M, Batak, 20 Jul 1966 (ZSM). GERMANY: 1 M, Stolzenau, 8 Aug 1945 (ZSM). RUSSIA: 1 F, Altay, Lake Telezkoe, Chulishman, 6 Aug 1989, A. Tereshkin leg. (ZSM).

Figure 2.

Woldstedtius biguttatus (Gravenhorst, 1829) (A–D. female; E, F. male) ― A, E. Habitus; B, F. Head, frontal view; C. Mesonotum, dorsal view; D. Propodeum, dorsal view.

Description

Female (n = 100). Body length 4.5–7.2 mm, weakly polished, coriaceous and covered with silver setae.

Head 0.5–0.54 × as long as wide. Clypeus 2.0–2.2 × as broad as high, slightly convex basally in lateral view. Face 2.0–2.35 × as broad as high, densely punctate, convex medially in lateral view, separated from clypeus by shallow clypeal sulcus. Inner orbits almost parallel (Fig. 2B). Length of malar space 1.0–1.1 × as long as basal mandibular width. POL 1.8–2.3 × as long as OD. OOL 1.0–1.3 × as long as OD. POL 1.7–2.1 × as long as OOL. Antenna with 22–25 flagellomeres. FL I 1.25–1.33 × as long as FL II. MP IV 1.25–1.33 × as long as MP V.

Mesosoma. Lateral aspect of pronotum rugulose anteriorly. Mesoscutum finely and sparsely punctate (separated by ca. 1.5–3.0 × their diameter) (Fig. 2C). Scutellum finely and sparsely punctate (separated by ca. 1.5–2.5 × their diameter) (Fig. 2C). Anterior and lower parts of mesopleuron coarsely and sparsely punctate. Sternaulus weakly impressed. Propodeum rounded in lateral view, without rugae (Fig. 2D), without carinae except for anterior part of pleural carina. Fore wing length 3.8–6.5 mm. Nervellus intercepted below middle. Hind femur 4.22–4.5 × as long as maximum depth in lateral view. Hind femur 8.0–8.5 × as long as maximum depth in lateral view. Ratio of length of hind first to fifth tarsomeres 1.0: 0.6: 0.4–0.5: 0.2–0.3: 0.3.

Metasoma. T I rectangular in dorsal view (Fig. 9F), 1.0–1.2 × as long as maximum width, rugulose laterally. Latero-median carina present basal ca. 0.5 of T I. T II 0.66–0.88 × as long as maximum width, striate anteriorly and strigose laterally.

Coloration (Fig. 2A–D). Body (excluding wings and legs) black. Face sometimes with a small yellow or brown median spot. Palpi, tegula and upper part of mesepisternum yellow. Mandible yellow, except for apex and base. Lateral aspect of pronotum with a yellow spot posteriorly. Subtegular ridge tinged with yellow anteriorly. Scutellum with a yellow spot apically. Wings hyaline. Veins and pterostigma brown to blackish-brown except for yellowish-brown wing base. Legs orange. Apex of fore coxa and apex of hind femur tinged with black. Trochanters and trochantelli yellow. Hind tibia and tarsomeres black. Base of hind tibia tinged with white.

Male (n = 71). Similar to female. Inner orbits weakly divergent downward (Fig. 2F). Length of malar space 0.83–1.0 × as long as basal mandibular width.

Coloration (Fig. 2E, F). Body (excluding wings and legs) black. Clypeus, palpi, face, ventral surface of antenna, malar space, propleuron, epicnemium, tegula, subtegular ridge and mesepisternum yellow. Gena tinged with yellow ventrally. Mandible yellow, except for apex. Lateral aspect of pronotum with a yellow spot ventrally and posteriorly. Mesoscutum with yellow shoulder marks. Mesopleuron with a yellow marking ventrally. Scutellum with a yellow spot apically. T III and T IV each with a pair of yellow spots anteriorly (sometimes these spots united into a single spot). Wings hyaline. Veins and pterostigma brown to blackish-brown except for yellowish-brown wing base. Legs yellow. Fore and mid femora, tibiae and tarsi orange. Hind coxa with a blackish-brown stripe dorsally. Hind femur brown to blackish-brown. Base of hind trochantellus tinged with blackish-brown. Hind tibia and tarsomeres black. Base of hind tibia tinged with whitish-yellow.

Distribution

Japan (Hokkaido, Honshu, Shikoku, and Kyushu). Outside Japan, this species is widely distributed in Palearctic region (Yu et al. 2016).

Bionomics

Host unknown in Japan. Outside of Japan, the following three hoverfly species have been recorded as hosts: Eupeodes lapponicus (Zetterstedt, 1838); Neocnemodon fulvimanus (Zetterstedt, 1843); Sphaerophoria scripta (Linnaeus, 1758) (Yu et al. 2016). Anthonomus pomorum (Linnaeus, 1758) and Loxostege sticticalis (Linnaeus, 1761) have also been recorded as hosts (Yu et al. 2016), but are considered doubtful records. Most adults were collected from various open habitats (e.g., grasslands, meadows and paddy fields).

Remarks

This is the first record of this species from Japan. This species resembles W. flavolineatus, but can be distinguished from the latter by the following combination of character states: face entirely black or black with a small yellow or brown median spot in females (black with a large yellow median spot in females of W. flavolineatus); pronotum black with yellow ventral corner in males (black with yellow ventral and hind parts in males of W. flavolineatus); yellow shoulder marks absent in females (present in females of W. flavolineatus); mesopleuron with a yellow marking ventrally in males (with a large yellow marking; it is enlarged anteriorly in males of W. flavolineatus).

Woldstedtius citropectoralis (Schmiedeknecht, 1926)

Bassus abdominator Bridgman, 1886: 336. Name preoccupied.

Homocidus citropectoralis Schmiedeknecht, 1926: 3412.

Material examined

No specimens available.

Distribution

Japan (Kunashiri Is.). Outside Japan, this species is widely distributed in Holarctic region (Yu et al. 2016).

Remarks

This species was recorded from Japan by Manukyan (2007), but no additional specimens were found in this study.

Woldstedtius flavolineatus (Gravenhorst, 1829)

Remarks

This species is divided into two subspecies, W. flavolineatus flavolineatus and W. f. nigroscutellatus (Habermehl, 1925). The latter subspecies has been recorded from Germany and Netherlands (Habermehl 1925; Teunissen 1948). Klopfstein (2014) noted this subspecies could not be examined by her because type specimens were not available. Judging from the original description of the latter (Habermehl 1925), this subspecies could readily be synonymized under the former.

Woldstedtius flavolineatus flavolineatus (Gravenhorst, 1829)

Figs 3A–F, 9G

Bassus flavolineatus Gravenhorst, 1829: 337.

Homocidus flavolineatus Uchida, 1957: 251.

Bassus interruptus Holmgren, 1858: 359. Synonymized by Thomson (1890).

Bassus bimaculatus Holmgren, 1858: 360. Synonymized by Thomson (1890).

Bassus agilis Cresson, 1868: 111. Synonymized by Dasch (1964).

Bassus frontalis Cresson, 1868: 111. Synonymized by Dasch (1964).

Mesoleius junctus Provancher, 1883: 10. Synonymized by Dasch (1964).

Materials examined

Italy: 1 F Sud-tirol, Coltina d’Ampezzo, 29 Jul 1933, E. Bauer leg. (ZSM). Bulgaria: 1 M Vitoscha, 24 Jul 1966 (ZSM).

Figure 3.

Woldstedtius flavolineatus flavolineatus (Gravenhorst, 1829) (A–D. female; E, F. male) ― A, E. Habitus; B, F. Head, frontal view; C. Mesonotum, dorsal view; D. Propodeum, dorsal view.