Research Article |
Corresponding author: Zai-fu Xu ( xuzaifu@scau.edu.cn ) Academic editor: Andreas Köhler
© 2017 Valery M. Loktionov, Arkady S. Lelej, Zai-fu Xu.
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:
Loktionov VM, Lelej AS, Xu Z-f (2017) Discovery of the genus Nipponodipogon Ishikawa in the Oriental region, with description of two new species from China (Hymenoptera, Pompilidae). ZooKeys 692: 103-127. https://doi.org/10.3897/zookeys.692.12062
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The genus Nipponodipogon Ishikawa, 1965 is newly recorded from China (Guangdong, Hainan, and Yunnan) and the Oriental Region. Two new species, N. orientalis Loktionov, Lelej & Xu, sp. n. (Guangdong, Hainan, Yunnan) and N. shimizui Loktionov, Lelej & Xu, sp. n. (Guangdong, Yunnan), are described and illustrated. The updated key to the species based on
China, Deuterageniini , new species, Nipponodipogon , Oriental Region, Pepsinae
The family Pompilidae (spider wasps) is one of the largest families among the aculeate wasps in Hymenoptera. The family numbers around 5000 recent species in 125 genera and five subfamilies in the World (
One of such cleptoparasitic genera is Nipponodipogon Ishikawa, 1965, a representative of brood parasitic wasps.
Nipponodipogon, from the tribe Deuterageniini, subfamily Pepsinae, is distributed so far in the Eastern Palaearctic: in the Japanese Archipelago and the south of the Russian Far East.
In this paper we describe two new species of Nipponodipogon from China and enlarge the distribution of the genus to include China and the Oriental Region.
During the study of hymenopteran collection in South China Agricultural University, we examined more than 2300 specimens of Chinese spider wasps collected during last two decades from Jilin, Inner Mongolia, Ningxia, Gansu, Shaanxi, Henan, Zhejiang, Hebei, Fujian, Hunan, Guangdong, Hainan, Guangxi, Yunnan, Sichuan, and Guizhou. Of them only 14 specimens belonging to the genus Nipponodipogon were collected in 2006, 2010, and 2011 years in the Oriental part of China (Guangdong, Hainan and Yunnan) by yellow pan traps and sweeping nets. The following acronyms are used for the collections where type specimens are deposited:
IBSS Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far Eastern Branch of the Russian Academy of Sciences (former Institute of Biology and Soil Science), Vladivostok, Russia (curator Prof. Arkady Lelej).
To study male genitalic characters, genitalia were extracted after being previously softened. The muscles were removed in a sodium hydroxide solution (NaOH 10%). The genitalia were later placed in water to neutralize the NaOH and stored in micro vials filled with glycerin. Male genitalia were studied under a stereomicroscope in a depression slide.
Photographs of imagos and genitalia were taken with stereomicroscope SteREO Discovery.V12 and stacked using CombineZM software (
The terminology for morphology is mostly based on the glossary provided by the
F1, F2, F3 etc., the first, second, third flagellomeres, etc.;
MID the middle interocular distance;
OOD the distance between posterior ocellus and compound eye which is measured from above;
POD the postocellar distance which is measured from above;
S1, S2, S3 etc., the first, second, third metasomal sterna, etc.;
SMC2 the second submarginal cell of fore wing;
SMC3 the third submarginal cell of fore wing;
T1, T2, T3 etc., the first, second, third metasomal terga etc.;
UID the upper interocular distance.
Dipogon (Nipponodipogon)
Ishikawa, 1965: 89. Type species: Dipogon (Nipponodipogon) iwatai
Nipponodipogon
:
Female. Maxillary cardo with a few thin, pale bristles, the apex of these not extending beyond the maxillary lacinia. Antenna short, stout, and thickened toward middle of flagellum (fusiform); F1 less than 3× its width. Supra-antennal area of frons produced anteriorly into a frontal ledge overhanging the antennal radicle. Apical margin of labrum not or slightly emarginated medially. Metapleuron strongly convex above level of lateral face of pronotum and metapleuron (dorsal view). Metapostnotum narrow and practically linear, deeply sunken between the metanotum and propodeum. Crossvein cu-a of hind wing short and almost straight, forming obtuse angle with vein 1A. Male. Antenna slightly thickened medially, usually with F3–F11 triangularly produced beneath (except for N. orientalis Loktionov, Lelej & Xu, sp. n. and N. shimizui Loktionov, Lelej & Xu, sp. n.); F1 1.3–2.0× its width. Mandible with one subapical inner tooth. Body punctate. Exposed portion of hypopygium stick-like, compressed laterally; subbasal portion strongly widened (Figs
Nine species. Nipponodipogon hayachinensis (Ishikawa, 1968), ♀ (Japan: Honshu); N. iwatai (Ishikawa, 1965), ♀ & ♂ (Japan: Hokkaido and Honshu); N. kurilensis (Lelej, 1986), ♀ (Russia: Kuril Islands); N. mandibularis (Ishikawa, 1965), ♀ (Japan: Honshu); N. nagasei (Ishikawa, 1965), ♀ & ♂ (Japan: Hokkaido, Honshu and Kyushu); N. rossicus (Lelej, 1986), ♀ & ♂ (Russia: Primorskij Terr.); N. sudai Shimizu in Shimizu, Lelej & Loktionov, 2015, ♀ & ♂ (Japan: Hokkaido and Honshu) (
Palaearctic Region (Russia: Primorskij Terr., Kuril Islands; Japan: Hokkaido, Honshu, Kyushu) and Oriental Region (new record) (China: Guangdong, Hainan, Yunnan).
The representatives of the genus Nipponodipogon are brood parasitic wasps. Nipponodipogon nagasei and N. iwatai brood-parasitize species of Deuteragenia Šustera, 1912 (tribe Deuterageniini), and N. iwatai brood-parasitizes species of Auplopus Spinola, 1841 (tribe Auplopodini). Female of N. nagasei routinely lays up to five eggs on a single host spider, all of which develop into adult wasps without larval cannibalism, instead all spider wasps previously studied lay only one egg on a host spider (
Holotype. CHINA: ♀, Guangdong, Nankunshan, 4–6.VI.2011, Zai-fu Xu, No. 2016001247 (
Female. Mesosoma completely yellow orange (Figs
Female. Body length 6.1–8.0 mm; fore wing length 4.7–6.2 mm. Head and metasoma black; sometimes clypeus along anterior margin brownish; antenna black, except flagellomeres 3–10 muddy yellow ventrally and sometimes scape and pedicel yellowish-brown ventrally; mandible brownish subapically. Mesosoma completely yellow orange (Figs
Head and mesosoma matt. Frons, vertex, and mesosoma, except propodeum, finely and densely punctate. Pronotum anteriorly, laterally and collar finely striate and punctate. Mesopleuron with denser punctures. Upper mesopleuron and metapleuron finely and densely striate. Lateral side of metanotum with several regular oblique striae. Propodeum strongly and densely punctate with fine transverse rugae posteriorly and much stronger rugae posterolaterally (Figs
Body with gray pubescence mostly short, but longer on clypeus, mesopleuron, propodeum posterolaterally and coxae. Body without setae except the following: upper frons sometimes with one long erect setae and a few shorter ones; clypeus with a few long suberect setae anteriorly; coxae and T1 basally with scattered short erect setae; S2–S5 with scattered longer erect setae posteriorly; T6 and S6 with denser long erect pale setae.
Width of head in frontal view 1.1–1.2× its height. Vertex weakly convex between eye tops (Fig.
Pronotum with anterior declivity flattened, not distinctly differentiated from dorsum; dorsum in dorsal view slightly narrowing anteriorly; shoulder gently rounded; juncture between dorsal and lateral faces narrowly and roundly raised; posterior margin weakly and arcuately emarginate medially. Mesoscutum slightly sloped anteriorly; posterolateral margin not reflexed; parapsidal sulcus finely impressed. Discs of mesoscutellum and metanotum barely raised above level of mesoscutum and propodeum (Fig.
Fore wing (Fig.
Male. Body length 3.7–4.6 mm; fore wing length 3.5–4.1 mm. Body black; antenna black with scape brown or black ventrally and flagellum weakly brown ventrally; mandible brown subapically; pro- tibia and tarsi brown; spurs of pro- and mesotibia brown, spurs of metatibia dark brown (Fig.
Body mostly punctate and somewhat polished. Frons, discs of pronotum, mesoscutum, mesoscutellum, and metanotum finely and densely punctate. Pronotum laterally polished and indistinctly punctate. Mesopleuron with coarser punctures than frons. Upper mesopleuron striate. Lateral side of metanotum with several regular oblique striae. Metapleuron finely punctate. Propodeum more or less matt, finely and densely punctate with weak dense transverse striae posterolaterally. Metasomal segments finely punctate. S1 with several longitudinal rugae basally. Transverse groove on S2 weak, gently arcuate, not connected medially. S6 with scattered setiferous pores (Fig.
Width of head in frontal view 1.1× its height. Vertex moderately convex between eye tops (Fig.
Pronotum with anterior declivity weakly concave, more differentiated from dorsum than in female; dorsum in dorsal view narrowing anteriorly; shoulder gently rounded; juncture between dorsal and lateral faces roundly raised; posterior margin weakly and arcuately emarginate. Mesoscutum slightly sloped anteriorly; parapsidal sulcus finely impressed. Discs of mesoscutellum and metanotum somewhat more strongly raised above level of mesoscutum and propodeum than in female. Metapostnotum (Fig.
Fore wing (Fig.
The female of new species is similar to those of Nipponodipogon kurilensis, N. sudai, and N. shimizui sp. n. by having outer apicoventral corner of metafemur produced triangularly (Fig.
Male of new species is closely related to that of N. shimizui sp. n. by some morphological characters including shape of hypopygium and genitalia, but easily distinguishes in propodeum with fine transverse striae posterolaterally (Fig.
In spite of the fact that females and males were collected in different locations (two males from Yunnan and five females from Guangdong and Hainan) and have different coloration (mesosoma completely yellow orange in female vs completely black in male), we consider that they are opposite sexes of same species. Male of new species has propodeum with fine transverse striae posterolaterally that correlates with strong transverse rugae on propodeum posteriorly, especially in posterolateral portion in female (vs male without any striae, female with fine transverse striae in Nipponodipogon shimizui sp. n.). Such coloration differences in female and male of new species are not exception and occur in widely distributed Palaearctic species Arachnotheutes rufithorax (Costa, 1881) (
The name of the new species refers to the first record of the genus in the Oriental Region.
China (Guangdong, Hainan, Yunnan).
Holotype. CHINA: ♀, Guangdong, Nanling, 8–17.VIII.2010, Hua-yan Chen, yellow pan traps, No. 2016001839 (
Female. Outer apicoventral corner of metafemur produced triangularly (Fig.
Female. Body length 5.2–6.4 mm; fore wing length 4.3–5.1 mm. Head, mesosoma and metasoma black; sometimes clypeus along anterior margin dark brown; antenna black, except F3–F10 muddy yellow ventrally and scape yellowish-brown ventrally; mandible brownish subapically. Legs yellowish-brown or brown with procoxa laterally, profemur externally, meso- and metafemur, tibiae apically and tarsi somewhat darker (Fig.
Head and mesosoma matt. Frons, vertex and mesosoma, except propodeum, finely and densely punctate. Pronotum laterally and finely striate and punctate. Mesopleuron with denser and coarser punctures then on disc of pronotum. Upper mesopleuron rugose. Metapleuron finely and densely striate. Lateral side of metanotum with several regular oblique striae. Propodeum strongly and densely punctate with fine transverse rugae posteriorly. Metasoma somewhat polished, except T6 and S6 matt. T1–T5 with fine punctures; T6 finely shagreened, without distinct setiferous pores; S6 less shagreened, than T6, with scattered setiferous pores located posteriorly and postero-laterally; S1–S5 with somewhat larger punctures than on T1–T5. S1 with several longitudinal rugae medially. Transverse groove on S2 gently arcuate.
Body with gray pubescence mostly short, but longer on propodeum posterolaterally. Body without setae except the following: upper frons sometimes with one long erect setae; clypeus with a few long suberect setae anteriorly; S2–S5 with scattered long or short erect setae posteriorly; T6 and S6 with denser long erect pale setae.
Width of head in frontal view 1.1–1.2× its height. Vertex weakly convex between eye tops (Fig.
Pronotum with anterior declivity flattened, not distinctly differentiated from dorsum; dorsum in dorsal view slightly narrowing anteriorly; shoulder gently rounded; juncture between dorsal and lateral faces narrowly and roundly raised; posterior margin weakly and arcuately emarginate medially (Fig.
Fore wing (Fig.
Male. Body length 3.8 mm; fore wing length 3.4 mm. Body black; antenna black with scape brown ventro-apically and flagellum indistinctly brownish ventrally; mandible brown subapically; protibia and protarsi brown; spurs of tibia brown (Fig.
Width of head in frontal view 1.1× its height. Vertex moderately convex between eye tops (Fig.
Pronotum with anterior declivity weakly concave, more differentiated from dorsum than in female; dorsum in dorsal view narrowing anteriorly; shoulder gently rounded; juncture between dorsal and lateral faces roundly raised; posterior margin arcuately emarginate. Parapsidal sulcus finely impressed. Discs of mesoscutellum and metanotum somewhat stronger raised above level of mesoscutum and propodeum than in female. Metapostnotum longer and not deeply sunken between metanotum and propodeum, as in female; somewhat narrowing in middle; metapostnotum length 0.25× metanotum length medially. Propodeum evenly convex with posterior declivity not differentiated from dorsum; posterior surface evenly convex.
Fore wing (Fig.
The female of new species is similar to those of Nipponodipogon kurilensis, N. sudai and N. orientalis sp. n. by having outer apicoventral corner of metafemur produced triangularly (Fig.
Male of new species is closely related to that of N. orientalis sp. n. by having some morphological characters including shape of hypopygium and genitalia, but can be easily distinguished in having propodeum without any striae posterolaterally (Fig.
In spite of females and males were collected in different locations (one male in Yunnan and six females in Guangdong), we consider that they are opposite sexes of the same species. Male S6 of new species lacks setiferous pores posteromedially (Fig.
It is a pleasure to name this species after the well-known taxonomist Dr. Akira Shimizu (Tokyo Metropolitan University, Japan).
China (Guangdong, Yunnan).
(based on
Females
1 | Outer apicoventral corner of metafemur produced triangularly (Figs |
2 |
– | Outer apicoventral corner of metafemur rounded ( |
5 |
2 | Posterolateral portion of propodeum with strong transverse rugae (Figs |
N. orientalis Loktionov, Lelej & Xu, sp. n. |
– | Posterolateral portion of propodeum with fine transverse striae or punctures (Figs |
3 |
3 | Vertex between eye tops strongly convex ( |
N. kurilensis (Lelej) |
– | Vertex between eye tops slightly convex (Fig. |
4 |
4 | Mesoscutum not raised along midline ( |
N. sudai Shimizu |
– | Mesoscutum raised along midline (Fig. |
N. shimizui Loktionov, Lelej & Xu, sp. n. |
5 | Transverse groove on S2 nearly straight ( |
N. iwatai (Ishikawa) |
– | Transverse groove on S2 subangulate ( |
6 |
6 | Mandible short, its apex and two additional teeth rounded, basal tooth vestigial ( |
N. mandibularis (Ishikawa) |
– | Mandible normal-sized, its apex and two additional teeth pointed, basal tooth distinct ( |
7 |
7 | Vertex strongly convex between eye tops; hence head in frontal view nearly circular in outline ( |
N. hayachinensis (Ishikawa) |
– | Vertex not very strongly convex between eye tops; hence head in frontal view not circular in outline ( |
8 |
8 | Ocelli forming right- or obtuse-angled triangle and gena strongly receding posteriorly ( |
N. nagasei (Ishikawa) |
– | Ocelli usually forming acute-angled triangle and gena roundly receding posteriorly ( |
N. rossicus (Lelej) |
Males (unknown for N. kurilensis, N. mandibularis, and N. hayachinensis)
1 |
T1 distinctly petiolate basally (Figs |
2 |
– | T1 not petiolate basally. Lateral hook on S6 not compressed laterally and not thin, but claw-like, curved and pointed to apex | 4 |
2 |
F3–F11 produced triangularly beneath, forming serrated profile. Lateral hook on S6 large, strongly compressed laterally and thin, subtriangular in profile (Fig. |
N. sudai Shimizu |
– |
F3–F11 not produced triangularly beneath, not forming serrated profile. Lateral hook on S6 not compressed laterally nor thin, but claw-like, curved and pointed to apex (Figs |
3 |
3 | Propodeum with fine transverse striae posterolaterally (Fig. |
N. orientalis Loktionov, Lelej & Xu, sp. n. |
– | Propodeum without any striae (Fig. |
N. shimizui Loktionov, Lelej & Xu, sp. n. |
4 | Ocellar triangle acute- to right-angled. Meso- and metatibial spurs dark brown. Exposed portion of hypopygium compressed laterally with ventral face flattened and polished, broad basally, tapering apically ( |
N. iwatai (Ishikawa) |
– | Ocellar triangle obtuse-angled; or if right-angled, meso- and metatibial spurs stramineous. Exposed portion of hypopygium completely compressed laterally and very thin, its ventral face linear ( |
5 |
5 | Genitalia with long setae on anterior margin of paramere ( |
N. nagasei (Ishikawa) |
– | Genitalia with short setae on anterior margin of paramere ( |
N. rossicus (Lelej) |
Nipponodipogon sudai Shimizu, paratype. 44 Mesoscutellum, metanotum, metapostnotum, propodeum and base of T1, dorsal view 45T1, dorsal view 46S6, ventral view 47 Genitalia, ventral view 48 Hypopygium and S7, ventral view 44 Female 45–48 Male. Scale bars 0.1 mm for 44–46; 0.25 mm for 47, 48.
We would like to thank Dr. Akira Shimizu (Tokyo Metropolitan University, Japan) for the gift of valuable comparative material. We are grateful to Andreas Köhler, Eduardo dos Santos and anonymous reviewers for appraising the manuscript and useful suggestions that have improved it. This study supported by the National Basic Research Program of China (No. 2013CB127600) and the Russian Found of Basic Research (No. 15-29-02466, 16-54-0041, 17-04-00259).
July 18, 2017 Prof. Xu Zai-fu died suddenly after a serious illness. He was only 52 years old. We indebted him for his kindness and support of Hymenoptera research in China.