Smicromyrme (Smicromyrme) lewisi Mickel, 1935,

Okayasu, Juriya, 2020, Velvet ants of the tribe Smicromyrmini Bischoff (Hymenoptera: Mutillidae) of Japan, Zootaxa 4723 (1), pp. 1-110: 9-24

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Smicromyrme (Smicromyrme) lewisi Mickel, 1935
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Smicromyrme (Smicromyrme) lewisi Mickel, 1935 

( Figs 1–8View FIGURE 1View FIGURE 2View FIGURE 3View FIGURE 4View FIGURE 5View FIGURE 6View FIGURE 7View FIGURE 8)

Mutilla japonica Cameron, 1900  (non Dalla Torre, 1897): 76, ♀, holotype ♀ (“Hakodate” [ĤM], Hokkaidô, Japan) [ BMNH]; André 1902b: 70, ♀  ; Schulz 1906: 158, ♀.

Mutilla cameroni Yano, 1932  (non Schulz, 1906): 326, ♂ ♀, syntypes ♂ ♀ (“Honshû, Kyûshû” [ĻḤ, ĘḤ], Japan) unknown; Kim et al., 1994: 255.

Smicromyrme lewisi lewisi Mickel, 1935: 288  , ♂ ♀, new name for Mutilla japonica Cameron, 1900  , holotype ♀ (“Hakodate” [Ĥ M], Hokkaidô, Japan) [BMNH], examined; Yasumatsu, 1938a: 85, ♀; Kim, 1963: 341; Kim, 1970: 793; Lelej 1985: 232, ♂ ♀; Lelej & Yamane 1992: 630, ♂ ♀; Takahashi & Matsumoto 1999: 23, ♀; Yamane 1999: 380, ♂ ♀; Haneda & Yamamoto 2000: 780, ♂ ♀, examined; Kubo 2000: 318, ♂; Nakamura & Haneda 2000: 119, ♂; Lelej 2002: 70; Tano et al. 2003: 16, ♂; Nagase 2004: 1280, ♂ ♀; Haneda et al. 2005: 57, ♂ ♀; Kurokawa 2005: 76, ♂ ♀; Kinota et al. 2005: 25, ♂; Negoro et al. 2005: 32, ♂ ♀; Terayama 2005: 8, 11, 16, ♂ ♀; Haneda et al. 2006a: 5, ♂ ♀; Haneda et al. 2006b: 40, ♂ ♀; Ohkusa 2006: 22, ♂; Ohkusa 2007: 30, ♂ ♀; Yamamoto & Haneda 2007: 15, ♂, examined; Ohkusa & Tajima 2008: 69, ♀; Fukasawa & Miyano 2010: 25; Ohkusa & Tajima 2010: 7, ♂ ♀; Terayama et al. 2011: 20, ♂ ♀; Ogawa et al. 2012: 38, ♀; Tsurusaki et al. 2012: 70; Hayashi et al. 2013: 25; Lelej et al. 2014: 393, ♂ ♀; Nagase et al. 2014: 56, ♂; Lo Cascio 2015: 558; Terayama et al. 2016: 168, ♂ ♀; Lelej 2017: 156; Okayasu et al. 2018: 304, ♀.

Smicromyrme lewisi yanoi Mickel, 1935: 289  , ♂ ♀, new name for Mutilla cameroni Yano, 1932  , holotype and allotype ♂ ♀ (“Honshû, Kyûshû” [ĻḤ, ĘḤ], Japan), unknown; Yasumatsu 1937: 68; Yasumatsu 1938a: 85, ♂; Yasumatsu 1938c: 351, ♀ non ♂ (Fig. 616-3); Yasumatsu 1951: 70, ♂; Kim, 1963: 341; Ishikawa 1965: 289, ♂ ♀; Kim, 1970: 793; Tsuneki 1972a: 16, ♀, synonymized under Smicromyrme rufipes lewisi Mickel  ; Lelej 1985: 233, ♂ ♀, synonymized under Smicromyrme lewisi lewisi Mickel  ; Hisamatsu 2004: 135.

Smicromyrme lewisi nigricula Mickel, 1935: 289  , ♂, holotype ♂ (“Jyôzankei” [aeƜữ], Hokkaidô, Japan) [University of Min- nesota, St. Paul, USA]; Lelej 1985: 233, ♂, synonymized under Smicromyrme lewisi lewisi Mickel. 

Smicromyrme rufipes rufipes: Chen 1957: 200  , ♂ ♀, examined, misidentification.

Smicromyrme momidziyama Tsuneki, 1972a: 17  , ♀, holotype ♀ (“Momidziyama” [ḢAƜ], Hokkaidô, Japan) [MNHAH], examined; Yoshida 1989: 651; Hashimoto & Nakanishi 1997: 31, ♀; Lelej 2002: 70, ♀, synonymized under Smicromyrme lewisi Mickel. 

Smicromyrme rufipes lewisi: Tsuneki 1972a: 16  , ♂ ♀, examined; Lelej 1976: 272, ♂ ♀; Lelej 1977b: 530, ♂ ♀; Lelej 1981: 78, ♂ ♀; Yoshida 1989: 651; Yamada 1990: 480; Kim et al. 1994: 255; Takahashi 1996: 28, ♂ ♀; Nakamura & Haneda 1997: 34, ♂ ♀; Matsuda & Nakamura 1999: 111, ♀; Nakamura et al. 2001: 83, ♂; Yamada 2001: 57, ♂ ♀; Kameyama et al. 2006: 74, ♀.

Smicromyrme rufipes lewisi  f. nigricula [invalid name according to Article 45.5 of ICZN (1999)]: Tsuneki 1972a: 16, ♂; Lelej 1977a: 91, ♂.

Smicromyrme rufipes lewisi var. nigricula  [invalid name according to Article 45.5 of ICZN (1999)]: Yoshida 1989: 651; Yamada 2001: 57.

Smicromyrme rufipes Mickel  (!): Nakamura et al. 2000: 61, ♀, lapsus calami.

Smicromyrme alberici: Shintani 2003: 17  , ♀, lapsus calami.

Diagnosis. MALE. Gena with large dense punctures; clypeus with median elevation and a pair of subapical teeth forming Y-shaped carina; antenna black. Mesopleuron beneath with precoxal longitudinal impunctate line; propodeum black. T1 longer than wide; T2 disc and T4–T5 with black setae; T7 without median longitudinal carina; genitalia in Fig. 5CView FIGURE 5. FE- MALE. Head black; frons and vertex with small confluent punctures; clypeus with median longitudinal tubercle apically slightly beyond the anterior margin of subapical transverse ridge. Mesosoma with transverse sulcus between propodeal spiracle and scutellar scale. Femora, tibiae and tarsi darkened apically. T2 with one narrow elongate basomedial spot of pale setae and apical narrow uniform band of pale setae; T2 disc with small sparse punctures; T3 with entire band of pale setae; T4 without pale setal band; striae of pygidial area parallel and reaching its apex.

Redescription. MALE (n = 179). Body length: 6.44 mm– 13.2 mm (average: 10.1 mm); fore wing length: 4.55 mm– 9.50 mm (average: 7.12 mm).

Color and setae. Head black, antennal tubercle sometimes brownish tinged; frons with dense short recumbent and sparse long erect golden to black setae; vertex with sparse short suberect and sparser long erect to golden to black setae except posterior margin with sparse long erect pale golden setae; gena with sparse short recumbent and sparser long erect pale golden setae; setae on malar space appressed; antennal scrobe glabrous; clypeus medially triangularly with sparse long erect golden to black setae, laterally with dense short appressed pale golden setae. Mandible black except median 1/6–1/4 reddish brown, with sparse long erect golden to black setae, basally also with dense long appressed pale golden setae. Prementum and stipes brownish black to black with sparse extremely short recumbent pale golden setae; palpi yellowish brown to brownish black with sparse extremely short erect golden setae. Antenna black; scape with dense long recumbent pale golden and sparse long erect black setae; pedicel with dense very short recumbent pale golden to black setae; flagellum with dense extremely short erect pale golden setae.

Mesosoma including tegula yellowish red except propleuron, ventral half of mesopleuron, metapleuron and propodeum black; anterior slope and dorsum of pronotum medially sometimes black; mesopleuron above mesopleural suture usually more or less yellowish red, sometimes entirely black; propodeum rarely partially reddish tinged; mesosoma entirely black in the nigricula form; pronotal collar with sparse short erect pale golden setae; dorsum of pronotum with sparse long recumbent pale golden and sparser long erect black setae; mesoscutum with sparse short suberect and sparser long erect black setae; tegula with sparse long recumbent black setae; scutellum and metascutellum with sparse long erect black setae; axilla and metanotal trough with dense short recumbent pale golden setae; dorsal propodeal face with sparse long erect pale golden setae, basomedially also with sparse short erect pale golden setae; propleuron, median part and ventral face of mesopleuron, ventral half of metapleuron and dorsal half of lateral propodeal face with sparse short recumbent and sparser long erect pale golden setae; lateral face of pronotum with sparse long recumbent pale golden setae; anterior part of mesopleuron, dorsal half of metapleuron and ventral half of lateral propodeal face with sparse short recumbent pale golden setae.

Wings pale brown, with sparse short erect black, regularly distributed setae.

Legs dark brown to black; protibial spur dark brown; meso- and metatibial spurs yellowish white; procoxa, trochanters and femora with sparse short recumbent and sparser long erect pale golden setae; meso- and metacoxa with sparse long recumbent pale golden setae; tibiae with dense long recumbent and sparse long erect pale golden setae; tarsi with dense long appressed pale golden setae.

Metasoma black; T1, T2 around discal area, T3, basal half and lateral margin of T4, lateral slope of S1 and S2–S5 with sparse long erect pale golden setae; T2 disc, apical half of T4, T5–T7 and S6–S7 with sparse long erect black setae; posterior slope of S1 with dense short erect pale golden setae; S2 with sparse short recumbent and sparser long erect pale golden setae; S8 with sparse short recumbent and sparser long erect black setae; T1 and S2–S4 with apical fringe of sparse long pale golden setae; T2–T3 with apical fringe of dense long pale golden setae; T4–T6 and S5–S7 with apical fringe of sparse long black setae; segment 2 with lateral golden felt line on T and shorter on S.

Structure. Head 1.37–1.57 (average: 1.45) times wider than long; lateral margins strongly convergent posteriorly behind eye; vertex elevated posteriorly; POL/OOL = 0.50–0.78 (average: 0.64); OL:MOD:LOD (average in parentheses) = 1.00:0.92–1.40 (1.11):0.75–1.20 (0.94); GOI = 1.00–1.50 (average: 1.21); ocelli without posterior carina; lower frons without median carina or groove; antennal scrobe with distinct dorsal carina not reaching inner eye margin; frons with large confluent punctures; interocellar area and vertex posteriorly to it with minute sparse punctures; vertex laterally and gena with large dense punctures. Clypeus weakly convex, basomedially elevated, forming Y-shaped carina with subapical pair of teeth; anterior margin medially widely protruding, without median emargination; surface of clypeus with minute dense punctures, medially also with few large punctures, except apicomedial margin impunctate. Mandible narrowed to apex, without dorsal crook, apically pointed and bidentate; inner denticle distinctly smaller than apex; ventral margin of mandible excised into large tooth basally; surface of mandible with minute sparse punctures except basal outer face with small confluent punctures. Scape bending medially, with two parallel carinae beneath; surface of scape and pedicel with small dense punctures; F1 length 1.00–1.39 (average: 1.22) × its width, 0.55–0.74 (average: 0.64) × F2 length; F2–F11 almost same in width; length of F2–F10 each little longer than the following segment; F11 little longer than F10, conical, weakly compressed; surface of flagellum with minute dense punctures.

Mesosoma broadest at mesonotum including tegulae; ratio between head width:humeral width:pronotal width: mesonotal width (average in parentheses) = 64–83 (74):55–70 (63):75–91 (82):100; humeral angle weakly angulate; propodeum weakly convergent posteriorly; mesoscutal width 1.09–1.23 (average: 1.15) × its length, 0.60–0.74 (average: 0.65) × mesonotal width; tegula rounded posteriorly, at most reaching mesoscuto-scutellar articulation, 1.40–1.57 (average: 1.48) times longer than wide; mesoscutum distinctly convex; notauli long, reaching anterior margin of mesoscutum, weakly convergent posteriorly; parapsidal line short; parascutal carina well developed, forming distinct scale-like process on postero-lateral angle of mesoscutum; scutellum weakly convex without carina or impunctate line; mesopleuron with distinct transverse mesopleural suture, beneath weakly concave with short longitudinal precoxal impunctate line; dorsum of pronotum, mesoscutum and scutellum with large confluent punctures; tegula with small sparse, regularly distributed punctures; axilla and metanotal trough with minute dense punctures; metascutellum with small confluent punctures; dorsal propodeal face areolate, basally with median elongate tongue-like to triangular cell and large semicircular cell laterally to it, and with minute sparse punctures in basal cells; propleuron with small sparse punctures; lateral face of pronotum transversely puncto-striate; anterior part of mesopleuron, dorsal half of metapleuron and ventral half of lateral propodeal face with minute sparse punctures; median part of mesopleuron laterally, ventral half of metapleuron and dorsal half of lateral propodeal face with large rounded confluent cells; mesopleuron ventrally with small dense punctures.

Procoxa with minute dense and small sparse punctures; meso- and metacoxa with minute sparse and small sparse punctures; trochanters with minute sparse punctures; femora and tibiae with minute sparse punctures and small sparser punctures; tarsi with minute dense punctures; metacoxa with weak inner carina throughout; tibiae without outer spines; claws simple, without inner denticle.

T1 with distinct dorsum and with dorso-lateral subbasal rounded tubercle; T2 evenly convex, 1.33–1.61 (average: 1.44) times wider than long; S1 with straight median longitudinal carina with its outer margin subbasally weakly emarginate, posteriorly rounded without distinct angle or tooth; S2 without basomedial carina; T1 disc and lateral margin, T2 around discal area and S2 with large dense punctures; dorsum of T1, T3–T6 and S3–S8 with small dense punctures; T2 disc with large sparse punctures; T7 with small confluent punctures, medially with narrow elongate impunctate slight groove; S1 with large confluent punctures. Paramere down-curved with subapical inner protuberance; outer face of paramere with sparse short erect golden setae, basally longer; inner face of paramere dorsally with long appressed golden setae on basal half; parapenial lobe bifurcate, apically pointed; volsella without basal external lobe; cuspis short oval inner view, compressed and straight in ventral view; dorsal margin of cuspis with row of long, at most little shorter than cuspis length, erect golden setae throughout; apical half of ventral margin and apical half of inner face of cuspis with dense short, at most half of digitus length, erect golden setae; basal half of ventral margin and basal half of inner face of cuspis with dense long, as most half of cuspis length, erect golden setae; inner face of cuspis with developed golden setal bunch on deep depression; digitus long, stick-like, apically with 3–7 short erect golden setae little shorter than digitus length; penis valve with apical and subapical pointed lobes.

FEMALE (n = 183). Body length: 4.54 mm– 9.40 mm (average: 6.77 mm).

Color and setae. Frons, vertex and gena black except antennal tubercle reddish brown, shining among punctures; frons rarely medially reddish tinged; clypeus yellowish red to reddish brown; frons and vertex (except posterior margin) with sparse short recumbent and sparser long erect golden to black setae; posterior margin of vertex with sparse very short recumbent pale golden and sparser long erect pale golden to black setae; antennal scrobe with sparse short appressed pale golden setae; eye dorsally edged with long erect black setae; gena with sparse long recumbent and sparser long erect pale golden setae; setae on malar space appressed; clypeus with sparse long erect golden setae. Mandible yellowish red except apical 1/2–2/3 black, with sparse long erect pale golden setae. Prementum and stipes brownish black with sparse extremely short erect pale golden setae; palpi dark brown to brownish black with sparse extremely short erect pale golden setae. Scape dark brown with sparse long recumbent pale golden setae; pedicel yellowish red with sparse short recumbent pale golden setae; F1 yellowish red to brownish black; F2–F5 yellowish red to black, at least beneath brownish; F6–F10 dark brown to black; flagellum with dense extremely short recumbent pale golden setae.

Mesosoma yellowish red; pronotal collar black; dorsum of pronotum and propodeum dorsally variously black maculated ( Figs 8View FIGURE 8 B–E); pronotal collar with sparse long erect pale golden setae; thoracic dorsum with sparse short recumbent pale golden to black and sparser long erect golden to black setae; dorsal propodeal face with sparse long erect black setae on dorsal 1/3 and with sparse long erect pale golden setae on ventral 2/3; propleuron with sparse long erect pale golden setae; lateral face of mesosoma with sparse short appressed pale golden setae; postero-ventral angle of mesopleuron and metapleuron and lateral propodeal face also with sparse long erect pale golden setae.

Procoxa yellowish brown to black; meso- and metacoxa and trochanters yellowish red to yellowish brown; femora yellowish red except apical 1/4–3/4 dark brown to black; protibia dark brown to lack except basal 1/10–3/10 yellowish red; meso- and metatibiae yellowish red except apical 2/5–3/5 yellowish brown to dark brown; tarsi yellowish red to dark brown; protibial spur pale brown; meso- and metatibial spurs yellowish white; spines on tibiae and tarsi pale to dark brown; coxae with sparse short recumbent and sparser long erect pale golden setae; trochanters with sparse long erect pale golden setae; femora and tarsi with sparse long recumbent and sparser long erect pale golden setae; tarsi with sparse long appressed pale golden setae.

Metasoma black except S1 yellowish red to dark brown; T1 usually yellowish red to dark brown basally and black on apical margin, rarely on apical half or entirely black; T1, T2 around discal area, T3, S1 and S3–S6 with sparse long erect pale golden to golden setae; T2 disc with dense short recumbent and sparse short erect black setae; T2 below lateral felt line with sparse long recumbent and sparser long erect pale golden setae; T4–T5 with dense long appressed black (laterally pale golden) setae mixed with sparse long erect black golden setae; T6 with dense long erect golden setae except pygidial area glabrous; S2 with sparse short recumbent and sparser long erect golden setae; T1 with apical fringe of sparse long pale golden to golden setae; S2–S6 with apical fringe of sparse long golden setae; T2 with narrow elongate (1.23–2.25 times longer than wide, 1.74 on average) basomedial spot of dense appressed golden setae not reaching anterior margin of T2, and with apical narrow band of dense appressed golden setae slightly widened medially; minimum distance between spot and band on T2 0.68–1.54 (average: 0.99) × length of spot; T2 with lateral golden felt line; T3 with wide uniform band of dense appressed golden setae.

Structure. In dorsal view, outline of head pentagonal with anterior margin strongly protruding, 1.29–1.49 (average: 1.35) times wider than long; lateral margins strongly convergent behind eye; posterior margin truncate. In lateral view, outline of head rounded; gena wide with GOI = 0.73–1.03 (average: 0.87). Frons and vertex with small confluent punctures, intervals smooth; gena with small dense punctures; frons and vertex without median carina or groove; genal carina and hypostomal tooth absent; antennal scrobe without dorsal carina; eye oval, slightly protruding from head capsule; minimum distance between eyes 1.41–1.65 (average: 1.51) × eye height; eye height 1.45–1.97 (average: 1.70) × malar length. Clypeus with minute sparse punctures, medially elevated and forming longitudinal tubercle, with subapical transverse ridge; anterior margin of transverse ridge truncate, without notch or emargination; median tubercle slightly protruding beyond anterior margin of transverse ridge. Mandible slen- der, apically pointed, with 0–2 small subapical inner denticles; ventral margin not excised. Prementum without dome-like tubercle. Scape bending medially; scape and pedicel with minute sparse punctures; F1 length 1.15–1.45 (average: 1.32) × its width and 1.00–1.40 (average: 1.25) × F2 length; F2–F9 almost same in length and width; F10 slightly longer than F9 and slightly shorter than F1, conical and weakly compressed; surface of flagellum with minute dense punctures.

Mesosoma broadest at pronotal spiracles; lateral margins of mesosoma weakly crenulate, subparallel; ratio between head width:humeral width: pronotal width:mesonotal width:propodeal width:metasomal width (average in parentheses) = 100:78–95 (86):87–99 (93):77–90 (84):80–94 (88):115–133 (124); dorsal thoracic length 1.01–1.15 (average: 1.10) × pronotal width; anterior margin of dorsum of pronotum weakly arcuate; pronotum laterally weakly projecting, forming weak angle at pronotal spiracle; humeral angle weakly angulate, humeral carina weakly developed; scutellar scale small but distinct, nail-like, 1.13–3.93 (average: 2.07) times wider than long, 0.10–0.38 (average: 0.22) times wider than propodeum, 2.20–8.40 (average: 4.56) times wider than diameter of nearby punctures; transverse sulcus present between propodeal spiracle and scutellar scale; postscutellar carina absent; pronotal collar with minute dense punctures; thoracic dorsum with large confluent punctures; dorsal propodeal face puncto-reticulated by large confluent punctures; lateral face of mesosoma with minute dense punctures.

Procoxa with minute dense punctures; meso- and metacoxa with minute sparse punctures; trochanters, femora, tibiae and tarsi with minute sparse punctures; metacoxa with weak inner carina throughout; meso- and metatibia with a pair of longitudinal row of spines on outer face; protarsus with external comb of long spines; claws simple, without inner denticles.

T1 without distinct dorsum; T2 evenly convex, dorsal T2 length 0.75–0.95 (average: 0.86) × T2 width; S1 with low straight median longitudinal carina with its outer margin at most slightly emarginated medially; S2 without basomedial carina; T1 basally, T2 and S2 with small sparse punctures; T1 on apical 1/3, T3–T6 except pygidial area, S1 and S3–S6 with small dense punctures; pygidial area flattened, elongate; lateral margins of pygidial area distinctly carinate throughout, subparallel; surface of pygidial area with 16–24 longitudinal fine simple striae reaching its apex.

Type materials examined. Smicromyrme lewisi Mickel  : holotype, ♀ ( Figs 2View FIGURE 2 G–H, 6C–D, 7D–F, K), Hako- date, Hokkaidô, Japan, George Lewis leg. [ BMNH]. Smicromyrme momidziyama Tsuneki  : holotype, ♀ ( Figs 2View FIGURE 2 I–J, 6E–F, 7G–I, L), Mt. Momiji-yama, Sapporo, Hokkaidô, Japan, 3–4.VIII.1958, Katsuji Tsuneki leg. [ MNHAH (B1- 288588)].

Other materials examined. 180♂ 183♀. Japan  : 158♂ 173♀: [Hokkaidô]  1♂, Jyôzan [probably Jyôzankei], Sapporo , 31.VIII.1913, S. Matsumura leg. [ HUM]  ; 1♂, same place, 12.VII, collector unknown [ HUM]  ; 1♀, Fuji- nosawa, Chitose-shi , 22.VII.1928, T.Uchida leg. [ HUM]  ; 1♂ 1♀ (in couple; Fig.1View FIGURE 1), Zenibako, Otaru-shi , 26.VII.1938, Y. Sugihara leg. [ HUM]  ; 1♀ ( Fig. 8DView FIGURE 8), Sôunkyo , 17.VII.1953, T. Ishihara leg. [ EUM]  ; 2♂, Nukabira , 12.VII.1959, T. Nambu leg. [ NSMT]  ; 1♂, Jyôzankei , 16.VII.1959, K. Saito leg. [ NSMT]  ; 1♀, same place, 17.VII.1983, H. Makihara leg. [ NIAES]  ; 1♂, Hôheikyô, Jyôzankei , 6.VIII.1968, T. Nambu leg. [ NSMT]  ; 1♀, Sapporo , 10.VII.1973, T. Kumata leg. [ HUM]  ; 1♀, Shumarinai, Horokanai-chô , 22.VII.1980, H. Nishino leg. [ MTC]  ; 1♂, Horonuka , Ru- moe-shi, 10.VII.1990, Nishijima collection [ HUM]  ; 1♀, Chitose-ko, Chitose-shi , 12.VIII.1994, S. Hori leg. [ TUA]  ; 2♂, Setogawa-chô, Hakodate-shi , 1.VIII.2007, H. Suda leg. [ HSC]  ; 1♂, Hamanasu-no-oka-kôen, Hama-machi, Ishikari-shi , 43°15′29″N 141°21′45″E, 7.VII.2017, J. Okayasu leg. [ EUM]GoogleMaps  ; 1♂, Takasago , 24.VII, collector un- known [ HUM]  ; 1♂, Mt. Usuzan , 12.VIII, J. Yoshioka collection [ NSMT]. [Honshû,  Iwate] 1♂, Koiwai , 9.VIII.1975, T. Nambu leg. [ NSMT]. [Honshû,  Tochigi] 2♀, Nagaoka, Utsunomiya , 10.VIII.1980, K. Nakamura leg. [ TPM]  ; 1♀, same place, 21.IX.1980, K. Nakamura leg. [ TPM]  ; 1♂, Awano, Nakakasuo , 15.VII.1981, K. Kubota leg. [ TPM]  ; 1♀, Watarase, Nogi , 4.VII.1993, H. Ohkawa leg. [ TPM]  ; 1♀, Nakagawa River, Magino, Motegi-machi , 22.V.1994, K. Satoh leg. [ TPM]  ; 1♀, Nakagawa River, Nabekake, Kuroiso-shi , 2.VI.1994, K. Satoh leg. [ TPM]  ; 1♀, Yokokawa, Fujihara-machi , 21.VII.1997, K. Sato leg. [ TPM]  ; 1♂, Fubasami, Takanezawa-machi , 3.VIII.2006, K. Kagawa leg. [ TPM]. [Honshû,  Ibaraki] 1♂, Yatabe , 10–15.VII.1982, N. Fukuhara leg. [ NIAES]  ; 1♀, Ajigaura-kaigan , Hitachi- naka-shi, 23.VII.1997, K. Toyoda leg. [ TUA]  ; 1♀, same place, 15.VIII.1999, T. Toba leg. [ TUA]  ; 1♀, same place,

13.X.2002, H. Ono leg. [ TUA]; 10♂ 2♀, same place  , 18.VII.2003, H. Ono leg. [ TUA]; 1♂, Sadanami, Kitaibarakishi   , 18–30.VII.2002, H. Goto leg., MST [ FFPRI]; 7♂, same place  , 30.VII–13.VIII.2002, H. Goto leg., MST [FF- PRI]; 1♂, same place  , 13–27.VIII.2002, H. Goto leg., MST [ FFPRI]; 1♂, same place  , 27.VIII–10.IX.2002, H. Goto leg., MST [ FFPRI]; 1♂, Okami, Satomi-mura   , 12–26.VIII.2003, S. Makino et al. leg., MST [ FFPRI]; 2♂, same place  , 29.VII–12.VIII.2003, S. Makino et al. leg., MST [ FFPRI]. [Honshû, Saitama] 1♀, Togasaki   , 21.VI.1945, collector unknown [ NIAES]; 1♀, Kita-akikawa, Okutama   , 27.VIII.1965, S. Katsuya leg. [ NIAES]; 1♂, Hodosan   , 8.VIII.1966, T. Nambu leg. [ NSMT]; 1♂, Nagatoro   , 21.VII.1968, T. Nambu leg. [ NSMT]; 2♂, same place  , 22.VII.1969, T. Nambu leg., K. Tsuneki det. 1972 as Smicromyrme rufipes lewisi Mickel  [ NSMT]; 1♂, Nakatsug- awa  , 17.VIII.1969, T. Nambu leg. [ NSMT]; 1♂, Hikawa-jinjya, Konosu   , 28.VI.1970, T. Nambu leg. [ NSMT]; 1♂, Shinpukuji, Chichibu   , 14.VII.1973, T. Nambu leg. [ NSMT]; 1♂, Minano   , 3.VIII.1973, T. Nambu leg. [ NSMT]; 1♂, Hikigun   , 14.VII.1974, M. Terayama leg. [ MTC]; 1♂, Mizuba–Kawamata, Ôtaki   , 4.VIII.1974, T. Nambu leg. [ NSMT]; 5♂, Mumeisenshi, Ogose   , 23.VII.1981, T. Nambu leg. [one in MTC and four in NSMT]; 3♂, Kasayama, Ogawa   , 13.VII.1994, M. Uchida leg. [ NSMT]; 1♀, Sugaya, Ranzan-machi   , 17.VII.1996, K. Toyoda leg. [ TUA]. [Honshû, Tôkyô] 1♀, Meguro   , IX.1914, M. Yamanaka leg. [ NSMT]; 1♂, Meguro   , 21.VII.1916, M. Yamanaka leg. [ NSMT]; 1♀, Meguro   , 17.IX.1916, M. Yamanaka leg. [ NSMT]; 1♂, Tachikawa   , 1.VII.1928, K. Sato leg. [ NSMT]; 2♂, Meguro   , VII.1929, M. Yamanaka leg. [ NSMT]; 1♀, Totuka   , 23.VII.1937, S. Asahina collection. [ NSMT]; 1♀, same place  , 22.VIII.1938, S. Asahina collection. [ NSMT]; 1♀, Nishiogikubo   , 5.VIII.1938, M. Kawai leg. [ NIAES]; 1♀, same place  , 15.VII.1939, M. Kawai leg. [ NIAES]; 1♀, same place  , 3.VII.1940, M. Kawai leg. [ NIAES]; 1♂ 1♀, same place  , 4.VII.1940, M. Kawai leg. [ NIAES]; 1♂, same place  , 13.VII.1941, M. Kawai leg. [ NIAES]; 1♂ 1♀, Fuchû   , 11.VIII.1939, Nagasawa leg. [ NIAES]; 1♂, same place  , 15.VIII.1939, Nagasawa leg. [ NIAES]; 1♂, same place  , 18.VIII.1939, Nagasawa leg. [ NIAES]; 1♂, same place  , 5.IX.1939, Nagasawa leg. [ NIAES]; 1♂, Yoyogi   , 19.VIII.1952, A. Katô leg. [ TUA]; 1♀, same place  , 25.IX.1955, collector unknown [ TUA]; 1♂, Haramachi near Tamagawa, Ôta-ku   , 26.VII.1953, S. Katsuya leg. [ NIAES]; 1♀, Kanamachi   , 30.VII.1953, S. Yoshioka leg. [ NSMT]; 4♂ 8♀, Mt. Takao-san   , 2.VIII.1953, J. Minamikawa leg. [ NIAES]; 1♀  , 6.IX.1953, S. Katsuya leg. [ NIAES]; 1♂, Ômori, Magome, Ôta-ku   , 24.VII.1955, S. Katsuya leg. [ NIAES]; 1♀, Mt. Takao   , 27.VII.1955, A. Habu leg. [NI- AES]; 1♀, Setagaya   , 18.VIII.1957, S. Katsuya leg. [ NIAES]; 1♂, Takao-machi, Hachiôji-shi   , 16.VII.1965, H. Suda leg. [ HSC]; 1♀, Katakura, Hachiôji   , 2.VIII.1989, M. Tomokuni leg. [ NSMT]; 1♀, Kanamachi   , 20.VI, J. Yoshioka collection [ NSMT]; 1♀, same place  , 26.VI, J. Yoshioka collection [ NSMT]; 1♀, same place  , 2.VII, J. Yoshioka col- lection [ NSMT]; 1♀, same place  , 7.VII, J. Yoshioka collection [ NSMT]; 1♀, same place  , 21.VII, J. Yoshioka col- lection [ NSMT]; 1♀, same place  , 22.VII, J. Yoshioka collection [ NSMT]; 7♂ 5♀, same place  , 25.VII, J. Yoshioka collection [ NSMT]; 5♂, same place  , 26.VII, J. Yoshioka collection [ NSMT]; 1♂ 2♀, same place  , 30.VII, J. Yoshioka collection [ NSMT]; 1♀, same place  , 11.VIII, J. Yoshioka collection [ NSMT]; 1♀, same place, 12, J. Yoshioka col- lection [ NSMT]; 1♀, Hachiôji-shi   , 3.VIII.1998, M. Terayama leg. [ MTC]; 1♂, Suginami   , 25.VII, J. Yoshioka col- lection [ NSMT]; 1♂, Meguro, M. Yamanaka leg. [ NSMT]. [Honshû, Chiba] 1♀, Narashino-shi   , 18.VII.1948, A. Katô leg. [ TUA]; 1♂, Amaha-machi, Futtsu-shi   , 10.VIII.1958, H. Suda leg. [ HSC]; 1♂ 1♀, Shônan-machi, Hi- gashikatsushika, Kashiwa-shi   , 25.VII.1966, H. & T. Suda leg. [ HSC]; 1♂ 1♀, Seiwa-mura, Kimitsu-shi   , 9.VIII.1966, H. & T. Suda leg. [ HSC]; 2♂, Usui, Sakura-shi   , 6.VIII.1968, H. Suda leg. [ HSC]; 1♀, Chônan-machi, Chôsei-gun   , 16.VII.1970, H. Suda leg. [ HSC]; 1♂ 1♀, Ôja-machi, Sakura-shi   , 14.VIII.1970, H. Suda leg. [ HSC]; 1♀, Iida, Sakura-shi   , 15.X.1970, H. Suda leg. [ HSC]; 1♂, Shibayama-machi, Sanbu-gun   , 6.VIII.1971, H. Suda leg. [ HSC]; 2♂, Mabashi   , 20.VIII.1973, S. Katsuya leg. [ NIAES]; 1♀, same place  , 21.VII.1974, S. Katsuya leg. [ NIAES]; 5♀, same place  , 28.VII.1974, S. Katsuya leg. [ NIAES]; 4♂ 5♀, same place  , 26.VII.1975, S. Katsuya leg. [ NIAES]; 1♂, Kawamata–Kusagawara, Kimitsu-shi   , 28.VI.1980, H. Suda leg. [ HSC]; 1♂, Narita, Narita-shi   , 25.VII.1980, H. Suda leg. [ HSC]; 1♂, Takada-chô, Chiba-shi   , 5.VIII.1995, H. Suda leg. [ HSC]; 1♂, Yagi, Sakura-shi   , 31.V.1998, H. Suda leg. [ HSC]; 1♂, Kagobeta, Youkaichiba-shi   , 21.VII.1999, T. Ito leg. [ EUM]; 1♂ 1♀, Horinouchi, Ichikawa-shi   , 6.VIII.2002, H. Suda leg. [ HSC]; 1♀, same place  , 3.IX.2003, H. Suda leg. [ HSC]; 1♂, Kitakokubun, Ichikawa-shi   , 6.VIII.2002, H. Suda leg. [ HSC]; 1♀, same place  , 3.IX.2003, H. Suda leg. [ HSC]; 1♀, Kônodai, Ichikawa-shi   , 4.IX.2002, H. Suda leg. [ HSC]; 1♂, Tsuchiuki, Sakura-shi   , 4.VIII.2004, H. Suda leg. [ HSC]; 1♀, same place  , 2. IX.2004, H. Suda leg. [ HSC]; 2♂ 1♀, Ueno, Nagara, Chôsei-gun   , 10.VIII.2004, H. Suda leg. [ HSC]; 1♂, Ki, Shiroishi   , 26.VII.2006, H. Suda leg. [ HSC]; 1♀, Shishiba, Shiroi-shi   , 10.IX.2006, H. Suda leg. [ HSC]; 1♀, same place  , 28.IX.2006, H. Suda leg. [ HSC]; 1♀, Rokuizô, Nagara, Chôsei-gun   , 17.VIII.2009, H. Suda leg. [ HSC]. [Honshû, Kanagawa] 1♂ 1♀ (in couple), Hinokiboramaru, Mt. Tanzawa-san   , 5.VIII.1985, N. Ohbayashi leg. [ EUM]. [Honshû, Yamanashi] 1♂ 4♀, Ikari-chô, Kôfu-shi   , 24.VIII.1971, H. Suda leg. [ HSC]; 1♂, same place  , 10.VIII.1977, H., T. & Y. Suda leg. [ HSC]; 1♂, Ôtsuki-chô, Ôtsuki-shi   , 19.VII.1997, H. Suda leg. [ HSC]. [Honshû, Nagano] 1♂, Karuizawa-machi, Kitasaku   , 23.VIII.1967, T. & H. Suda leg. [ HSC]; 1♀, Ôdaira-kôgen, Kiso   , 19.VII.1975, Y. Hori leg. [ TUA]; 2♂, Taira, Ômachi-shi   , 12.VIII.1976, H., T. & Y. Suda leg. [ HSC]; 1♂, Komagatake-kôgen, Komaganeshi   , 6.VIII.1980, H., T. & Y. Suda leg. [ HSC]; 1♂, Torii-touge, Sanada-machi   , 23.VII.2005, K. Nakamura leg. [ TPM]; 1♂ 1♀, Kisofukushima, Mitsuhashi leg. [ NSMT]. [Honshû, Gifu] 1♂, Tochibora, Kamioka   , 5.VIII.1971, T. Nambu leg. [ NSMT]; 3♀, Hirugano   , 20.VII.1975, H. Yamada leg. [ TUA]. [Honshû, Shizuoka] 3♂, Kanaya   , 3.VIII.1947, J. Minamikawa leg. [ NIAES]; 9♂, same place  , 10.VIII.1947, J. Minamikawa leg. [ NIAES]; 2♂, same place  , 13.VIII.1947, J. Minamikawa leg. [ NIAES]; 3♂, same place  , 6.VIII.1948, J. Sonan leg. [ NIAES]; 1♂, same place  , 24.VII.1950, J. Minamikawa leg. [ NIAES]; 1♂, same place  , 25.VII.1950, J. Minamikawa leg. [ NIAES]; 1♂, same place  , 21.IX.1952, J. Minamikawa leg. [ NIAES]; 2♂, same place  , 19.VIII.1954, J. Minamikawa leg. [NI- AES]; 2♂, Mt. Hakkô-san   , 24.VII.1948, J. Minamikawa leg. [ NIAES]; 1♂, Kasuisai   , 8.VIII.1948, J. Minamikawa leg. [ NIAES]; 1♂, Ogasayama   , 20.VIII.1948, J. Minamikawa leg. [ NIAES]; 2♂, Mochimune   , 15.VII.1950, J. Mi- namikawa leg. [ NIAES]; 2♂, Shimoda, Izu   , 21.VIII.1957, T. Nambu leg. [ NSMT]; 1♀, same place  , 21.VIII.1957, T. Nambu leg., K. Tsuneki det. 1972 as Smicromyrme rufipes lewisi Mickel  [ NSMT]; 2♂, Shirahama, Izu   , 21.VIII.1957, T. Nambu leg. [ NSMT]; 1♂, Dainititoge, Abe-gun   , 5.VIII.1959, J. Minamikawa leg. [ NIAES]; 1♂, Abe-tôge   , 8.VIII.1960, J. Minamikawa leg. [ NIAES]; 1♀, Fumoto, Fujinomiya-shi   , 11.IX.2003, K. Iijima leg. [ TUA]; 1♂, Ita, Numazu-shi   , 10–30.V.2013, H. Handa leg., MST [ TUA]; 2♂, same place  , 30.V–29.VI.2013, H. Handa leg., MST [ TUA]; 8♂, same place  , 29.VI–23.VIII.2013, H. Handa leg., MST [ TUA]. [Honshû, Aichi] 1♀, Jôkôji, Seto   . 6.VI.1975, H. Yamada leg. [ TUA]; 1♀, Tokoku, Moriyama, Nagoya-shi   , 5.VII.1975, H. Yamada leg. [ TUA]; 1♀, Mt. Sanage-yama, Seto   , 17.VII.1975, H. Yamada leg. [ TUA]; 2♀, Sobue   , 19.IX.1975, H. Yamada leg. [ TUA]; 1♂, Asahiyawata , 650 m alt  ., 18–28.VII.1998, M. Ozawa leg., MST [ TUA]. [Honshû, Niigata] 1♂, Himekawa   , 17.VIII.2001, T. Tsuru leg. [ TUA]; 1♀, same place  , 18.VIII.2001, T. Tsuru leg. [ TUA]. [Honshû, Fukui] 1♂, Mt. Monjusan   , 12.VIII.1962, K. Tsuneki leg. [ NMNH]. [Honshû, Kyôto] 1♀, Shûgakuin   , 8.VI.1959, T. Shyôji leg. [ NSMT]; 1♀, Mt. Aobasan   , 28.VII.1970, K. Tsuneki leg. [ NMNH]. [Honshû, Nara] 1♀, Tenri-city   , 24.VII.1959, I. Miyagi leg. [ EUM]; 1♀, Miyataki, Yoshino   , 9.VIII.1974, S. Ueda leg. [ ZMUC]; 2♀, Mt. Kasuga-yama, Zoushi-chô, Nara-shi   , 21.VIII.2003, T. Tsuru leg. [ TUA]. [Honshû, Wakayama] 1♀, Seto   , 5.VII.1929, H. Katayama leg., C. E. Mickel det. 1934 as Smicromyrme lewisi yanoi Mickel  [ ELKU]. [Honshû, Hyôgo] 1♂, Harima   , 20.VI.1912, S. Matsumura leg. [ HUM]; 1♂, Higakyôkoku, Higami   , 17.VII.1972, S. Ueda leg. [ ZMUC]; l ♂, Mt. Senzan, Awajishima Island   , 8.VIII.1974, S. Ohkusa leg. [ ZMUC]; 1♂, Hiromineyama, Himeji-shi   , 22.VII.1991, H. Suda leg. [ HSC]; 1♀, Narutodai, Awaji-shima Island   , 1.VIII.1998, collector unknown [ EUM]; 1♂ 1♀, Kuzaki, Harima, M. Yamanaka collection [ NSMT]. [Honshû, Okayama] 1♂, Hiruzen-kôgen   , 21.VII.1996, Y. Kawakami leg. [ EUM]; 6♂ 1♀, same place  , 22.VII.1996, Y. Kawakami leg. [ EUM]. [Kasaoka Islands] 1♀, Ôura, Kitagi-jima Island   , 30.VI.2003, T. Ishikawa leg. [ TUA]. [Honshû, Shimane] 1♂, Sashiumi, Izumo-shi , 35°20.234′N 132°39.909′EGoogleMaps  , 16–17.VII.2017, H. Yoshitomi leg. [ EUM]. [Honshû, Hiroshima] 1♂, Sandankyô   , 26.VII.1979, T. Nambu leg. [ NSMT]; 1♀, Akashi, Miyauchi, Hatsukaichi-shi   , 13–15.VIII.2015, S. Imada leg. [ EUM]. [Honshû, Yamaguchi] 1♀, Iwakuni-shi   , 5.IX.2004, M. Terayama leg. [ MTC]. [Oki Islands] 1♂, Saigô, Dôgo   , 16.VIII.1963, H. Kadowaki leg. [ NIAES]. [Shikoku, Tokushima] 1♀, Koyadaira, Mima-shi   , 13.VII.1925, E. Gallois leg. [ HUM]; 3♂, Yoshida, Kaifu   , 11.VII.1999, Y. Kawakami leg. [ EUM]. [Shikoku, Ehime] 1♂, Tarumi, Matsuyama   , 6.VIII.1948, M. Miya- take leg. [ EUM]; 1♂, Matsuyama   , 3.VIII.1951, M. Miyatake leg. [ EUM]; 1♀, same place  , 4.VII.1953, F. Takechi leg. [ EUM]; 1♀, Dôgo, Matsuyama   , 5.IX.1955, M. Miyatake leg. [ EUM]; 1♀, Oda-chô   , 16.VI.1984, E. Yamamoto leg. [ EUM]; 1♀, same place  , 13.VII.1996, E. Yamamoto leg. [ EUM]; 1♀, same place  , 10.VIII.1996, E. Yamamoto leg. [ EUM]; 2♀ ( Fig. 8CView FIGURE 8), same place  , 7.VI.1997, E. Yamamoto leg. [ EUM]; 1♀, same place  , 21.VI.1997, E. Yamamoto leg. [ EUM]; 1♂, same place  , 15.VII.1999, E. Yamamoto leg. [ EUM]; 1♂, Koyayama, Odamiyama   , 22.VIII.1991, E. Yamamoto leg. [ EUM]; 1♀, Mt. Takanawa-san, Matsuyama-shi   , 2.X.2004, T. Kitano leg. [ EUM]; 1♂, Yurano, Nimyô, Kumakôgen-chô   , 28.VI.2006, E. Yamamoto leg. [ EUM]; 1♂ 1♀ (in couple; Figs 2View FIGURE 2 A–B, E–F, 3A–B, 4A–C, G, 5A, C, E, 6A–B, 7A–C, J), Sugezawa-machi, Matsuyama-shi   , 21.V.2016, K. Kuroda leg. [ EUM]. [Shikoku, Kôchi] 1♂, Kuroson, Shimanto-shi   , 29.VII.1954, A. Katô leg. [ TUA]; 4♂, Sakihama, Muroto-shi   , 3.IX.1995, Y. Kawakami leg. [ EUM]; 2♂, same place  , 14.IX.1995, Y. Kawakami leg. [ EUM]; 2♂, same place  , 15.IX.1995, Y. Kawakami leg. [ EUM]; 1♂, same place  , 2.VI.1996, Y. Kawakami leg. [ EUM]; 5♂, Mt. Kajigamori   , 14.VII.1996, Y. Kawakami leg. [ EUM]; 1♀, Egawasaki, Shimanto-shi   , 3.VIII.1997, Y. Kawakami leg. [ EUM]; 1♂, Hitsuzan, Kôchi-shi   , 6.VI.1998, Y. Kawakami leg. [ EUM]; 1♀, same place  , 5.IX.2002, Y. Kawakami leg. [ EUM]; 2♀, same place  , 4.VI.2006, Y. Kawakami leg. [ EUM]; 1♀, same place  , 6.VI.2006, Y. Kawakami leg. [ EUM]; 1♀, Inabu, Nangoku-shi   , 4.IX.2001, T. Beppu leg. [ EUM]; 2♀, Mishima, Shimanto-chô   , 28.VII.2002, Y. Kawakami leg. [ EUM]; 1♀, same place  , 4.VIII.2002, Y. Kawakami leg. [ EUM]; 5♀, same place  , 14.IX.2004, Y. Kawakami leg. [ EUM]; 1♂, same place  , 22.VII.2007, Y. Kawakami leg. [ EUM]; 5♀, Irugi, Sakihama-chô, Muroto-shi   , 19.IX.2005, Y. Kawakami leg. [ EUM]; 12♂, Hageyama, Tosa-shi   , 16.VII.2006, Y. Kawakami leg. [ EUM]; 1♀, same place  , 24.VII.2006, Y. Kawakami leg. [ EUM]; 10♂, same place  , 21.VII.2007, Y. Kawakami leg. [ EUM]; 1♂, Shimonokae, Tosashimizu-shi   , 5.VII.2007, Y. Kawakami leg. [ EUM]; 1♀, Ioki, Aki-shi   , 8.IX.2008, Y. Kawakami leg. [ EUM]. [Shikoku] 1♂, Koguchi   , 26.VII.1958, I. Miyagi  leg. [ EUM]. [Kyûshû, Fukuoka] 1♂, Mt. Hikosan–Morizane ( Buzen )  , 18.VII.1929, Esaki & Fujino leg., B. Petersen det. 1981 as Smicromyrme rufipes lewisi Mickel  [ ZMUC]; 1♀, Chikuzen   , 3.IX.1930, K. Yasumatsu leg. [ TARI]; 1♀, same place  , 3.IX.1930, K. Yasumatsu leg., B. Petersen det. 1984 as Smicromyrme rufipes lewisi Mickel  [ ZMUC]; 1♀  , 14.VI.1931, K. Yasumatsu leg. [ ELKU]; 1♀, Kurate, Sengoku   , 23.V.1948, M. Yano leg. [ ELKU]; 1♂, Mt. Hikosan ( Bungo )  , 5.VIII.1951, K. Yasumatsu leg. [ ELKU]; 1♀, Kurate, Ohorima   , 11.VII.1953, M. Yano leg. [ ELKU]; 1♀, Inunaki   , 2.IX.1955, Y. Hirashima leg. [ ELKU]; 1♂, Mt. Hiko   , 7.VII.1978, K. Takeno leg. [ ELKU]; 1♀, same place  , 2.VII.1981, K. Taeno leg. [ ELKU]; 4♂, Inugatake ( Buzen )  , 14.VII.1988, Hori, Kawahara & Yasumatsu leg. [ ELKU]; 2♂, Mt. Hiko, Soeda, Tagawa   , 21.VII.2013, R. Ito leg. [ ELKU]; 1♀, Nokono-shima Island, Fukuoka-shi   , 3.VII.2015, S. Imada leg. [ EUM]; 1♂, Nata-kaigan Be- ach, Higashi-ku   , 9.VII.2017, T. Kawano leg. [ ELKU]. [Kyûshû, Ôita] 1♀, Sobosan ( Bungo )  , 4.VIII.1931, K. Ya- sumatsu leg. [ ELKU]; 1♀, Furuno, Hasama-machi, Ôita-gun   , 14.IX.2000, T. Miyake leg. [ EUM]. [Kyûshû, Miyazaki] 1♂, Hyûga   , 4.VIII.1938, S. Asahina collection [ NSMT]. [ Kyûshû , Kumamoto] 1♀  , 23.VII.1907, H. Kawamura leg. [ HUM]; 2♀, Tomioka-shiki, Amakusa   , 16.VI.1931, Esaki & Hori leg. [ ELKU]; 3♀, same place  , 19.VI.1931, Esaki & Hori leg. [ ELKU]; 1♂, Koga, Ueki Town, Kita Ward , Kumamoto  City, 26.VI.2016, K. Narita leg. [ EUM]. [Kyûshû, Kagoshima] 1♂, Chiran   , 12.VII.1963, Y. Kurosawa leg. [ NSMT (NSMT-Hym No. 4871)]; 1♀ ( Fig. 8BView FIGURE 8), Sata-misaki, Tajiri, Sata-magome , Minami-ôsumi-chô  , 16.VII.2011, H. Yoshitake leg. [ EUM]. [Tsushima] 1♂ 1♀, Tsushima   , 31.VII.1937, Yoshida leg. [ NSMT]. [Gotô Islands] 1♀, Nozaki-jima Island   , 30.VII.1971, S. Okadome leg. [ ZMUC]. [Izu Islands] 1♂, Kamitatsu, Motomachi, Izu-ôshima Island   , 17.VIII.2012, K. Tsuji leg., YPT [ TUA]; 1♀, Motomachi, Izu-ôshima Island   , 17.VIII.2012, K. Tsuji leg. [ TUA]. [ Japan] 1♀, Ôsato   , 18.VII.1903, M. Yamanaka collection [ NSMT]; 1♂, Iyosano-san   , 21.VII.1915, S. Matsumura leg. [ HUM]; 1♂, Sumiyoshi   , 21.VII.1934, H. K. leg. [ NSMT]; 1♀, Sumiyoshi   , 30.VII.1934, H. K. leg. [ NSMT]; 1♂, Ryoosui-ji   , VIII.1939, T. Sato leg. [ NIAES]. [Probably Japan] 1♂ [ ELKU]; 2♂ 1♀, M. Yamanaka leg. [ NSMT]. Russia: 1♂ 1♀: 1♂ 1♀, Semenovka , Amur Oblast  , 6.VII.1975, A. S. Lelej leg. [ ZMUC]. China: 5♂ 1♀: 1♂, Beijing  , 30.V.1938, K. Tsuneki leg. [ MNHAH]; 1♂, Apaka , Inner Mongolia  , 10.VI.1939, K. Tsuneki leg. [ MNHAH]; 1♂, same place  , 28.VI.1939, K. Tsuneki leg. [ MNHAH]; 1♂, Harbin , Heilongjiang  , 7.VII.1954, V.Alin leg., vend. Eing. Nr. 22, 1954, B. Petersen det. 1987 as Smicromyrme strandi lewisi Mickel  [ ZMUC]; 1♂ 1♀, Longtong, Jinan , Shan- dong  , 700 m alt., collector unknown, C. E. Mickel det. as Smicromyrme rufipes (Fabricius)  [ TARI]. South Korea: 16♂ 7♀: 1♀, Genzan   , 2.VIII.1919, E. Gallois leg., A. S. Lelej det. 2001 as Smicromyrme lewisi Mickel  [ HUM]; 2♀, Suwon   , 10.VII.1924, K. Sato leg. [ NSMT]; 2♂, same place  , 10.VI.1925, K. Sato leg. [ NSMT]; 1♂, same place  , 1.VIII.1927, K. Sato leg. [ NSMT]; 1♀, same place  , IX.1928, K. Sato leg. [ NSMT]; 1♂ 1♀, same place  , VI.1930, K. Sato leg. [ NSMT]; 1♂, same place  , VI.1930, K. Sato leg., B. Petersen det. 1981 as Smicromyrme rufipes lewisi Mickel  [ ZMUC]; 1♂, same place  , VI.1931, K. Sato leg. [ NSMT]; 3♂, same place, K. Sato leg. [ NSMT]; 1♀, Taiden, Tiunan   , 29.VIII.1933, K. Okazaki leg. [ ZMUC]; 1♂, Mt. Soyosan, Dongducheon , Gyeonggi  , 15.VI.1941, collector unknown [ MNHAH]; 1♂, 1♂ 1♀ (in couple), same place  , 18.VI.1943, collector unknown [ MNHAH]; 1♂, Seoul  , 20.VIII.1942, collector unknown [ MNHAH]; 2♂, Pyongyang Hotel garden, South Pyongyang   , 14.VIII.1971, S. Horvatovich & J. Papp leg. [ MNHAH]; 1♂, Unkano   , 19.VII, T. Uchida leg. [ HUM]. D.P.R. Korea: 1♀ ( Fig. 8EView FIGURE 8), Mt. Myohyang, Around Habiro , 200–550 m alt  ., 30.VI.2009, C. Han leg. [ EUM]  .

The nigricula form. 4♂: Japan  : 3♂ [Hokkaidô]  1♂ ( Figs 2View FIGURE 2 C–D, 3C–D, 4D–F, H, 5B, D, F), Jyôzankei , 16.VII.1959, K. Sato leg. [ NSMT]  ; 1♂, Momijiyama, Ishikari , 5.VIII.1968, T. Nambu leg. [ NSMT]  ; 1♂, Setogawachô, Hakodate-shi , 1.VII.2007, H. Suda leg. [ HSC]. South Korea  : 1♂, Andong , Gyeongsangbuk-do, 4.VI.1970, H. Yamada leg. [ TUA]  .

Confirmed distribution. Japan: Hokkaidô, Honshû (Iwate, Tochigi, Ibaraki, Saitama, Tôkyô, Chiba, Kana- gawa, Yamanashi, Nagano, Gifu, Shizuoka, Aichi, Niigata, Fukui, Kyôto, Nara, Wakayama, Hyôgo, Okayama, Shimane, Hiroshima, Yamaguchi), Shikoku (Tokushima, Ehime, Kôchi)  , Kyûshû (Fukuoka, Ôita, Miyazaki, Kuma- moto, Kagoshima), Oki Islands (Dôgo), Awaji-shima Island, Tsushima Island , Gotô Islands ( Nozaki-jima Island ), Izu Islands ( Izu-ôshima Island )  . Russia. South Korea. D.P.R. Korea. China.

Other localities in the literature. Japan: Honshû (Aomori, Miyagi, Toyama, Ishikawa, Shiga, Tottori) ( Tsuneki 1972a; Gôukon 1993; Takahashi 1996; Yamada 2001; Negoro et al. 2005; Haneda et al. 2006a; Ohkusa & Tajima 2010; Ogawa et al. 2012; Tsurusaki et al. 2012; Nagase et al. 2014), Okushiri-tô Island ( Munakata 1987), Ôsumi Islands (Tanega-shima Island, Iô-jima Island) ( Terayama et al. 2011). Kuril Islands (Kunashiri-tô Island) ( Lelej 1985). Mongolia. Terayama (2005) and Terayama et al. (2011, 2016) listed North America in the Distribution section of Sm. lewisi  , however, this record is doubtful because the distribution of this species is limited in Eurasia and its surrounding islands of east than Khakassia.

Occurrence in Japan. March–October.

Biology. This species is known as a parasitoid of ground-nesting apoid wasps, including Cerceris hortivaga Kohl, 1880  , Tachysphex nigricolor nigricolor ( Dalla Torre, 1897)  and Gorytes tricinctus ( Pérez, 1905)  ( Hymenoptera  : Crabronidae  ) ( Tsuneki 1965; Gôukon 1993). Lelej (1985) and Lelej et al. (2014) collected Sm. lewisi  from colonies of Oxybelus bipunctatus Olivier, 1812  ( Crabronidae  ) and bees of the genera Colletes Latreille, 1802  and Hylaeus Fabricius, 1793  ( Hymenoptera  : Colletidae  ) in Primorsky Krai, Russia and Tuva. Lelej et al. (2001) mentioned Cerceris  sp. ( Crabronidae  ) and a solitary bee ( Hymenoptera  : Apoidea) as possible hosts. Yasumatsu (1938c) noted that “ Smicromyrme lewisi yanoi Mickel, 1935  seems to prey on Bembix niponica Smith, 1873  ” ( Crabronidae  ) (translated from Japanese). In addition, Yamada (2001) collected 25 males and six females in a habitat of B. niponica  and T. nigricolor  . Bembix niponica  is also a possible host of Sm. lewisi  , but there is no direct evidence. Gyotoku (1941) collected four males of Sm. lewisi  at a light in Fukuoka, Kyûshû.

Comparison. The male of this species is similar to that of Sm. rufipes ( Fabricius, 1787)  in having the clypeus with median elevation and a pair of subapical teeth forming Y-shaped carina, the antenna black, the mesopleuron beneath with precoxal longitudinal impunctate line, the propodeum black, T1 longer than wide and T7 without median longitudinal carinae, but differs in having the T2 disc, T4 and T5 with black setae (pale golden in Sm. rufipes  ) and the shape of the parapenial lobe (see Fig. 5CView FIGURE 5 and Fig. 64.6 in Lelej 1985).

The female of this species is similar to that of Sm. chuchianus (Tsuneki, 1993)  , Sm. rufipes  and Sm. strandi 

( Zavattari, 1913) in having the head black, the clypeus with a median longitudinal tubercle apically slightly beyond the anterior margin of transverse ridge, the mesosoma with a transverse sulcus, T2 with one basomedial spot of pale setae, T3 with an entire pale setal band, T4 with black setae and the striae of the pygidial area parallel and reaching its apex, but differs in having the frons and vertex with small confluent punctures (with dense but separated punctures in Sm. chuchianus  and Sm. rufipes  ), the basomedial pale spot on T2 narrow elongate (ovate in others), the apical pale band on T2 narrow uniform (abruptly widened medially in Sm. chuchianus  ) and the T2 disc with small sparse punctures (small confluent in Sm. strandi  ). The following specimens were also examined for comparison: Sm. chuchianus  : holotype, ♀, Chuchi, Chiayi, Taiwan, 29–30.VII.1976, Tadao Murota leg. [MNHAH]; Sm. strandi  : 1♀, Chenkou, Chungan, Fujian, 5.VIII.1943, T. C. Maa leg., B. Petersen det. 1987 as Smicromyrme strandi strandi (Zavattari)  [TARI].

Synonymic notes. Yano (1932) illustrated a female of Sm. lewisi  with a short Japanese description in “ Iconographia Insectorum Japonicorum Editio Prima ” under the name Mutilla cameroni Yano. Unfortunately  , this name is valid according to ICZN (1999). The original description was:

“Hitohoshi-Aribachi [Japanese name] (new name). Mutillidae  . Female. Head black. Antenna, clypeus and mandible dark ferruginous. Mesosoma and legs ferruginous, sometimes partially more or less blackish. Metasoma black, basal half of first tergum frequently ferruginous. Head and mesosoma with large coarse punctures and metasoma shiny, with minute punctures. Body covered with long setae, dorsum with blackish setae, side and venter with pale setae. Second tergum of metasoma with median elongate oval spot near anterior margin; narrow band on posterior margin of second tergum and wide band on posterior margin of third tergum consisting of dense pale setae. Body length 4–9 mm. Male winged, black. Dorsum of prothorax and mesothorax ferruginous. Body covered with long black setae, both second and third tergum of metasoma with wide band consisting of dense pale setae on posterior margin. Wing tinged with grayish back and veins black. Body length around 12 mm. Distributed in Honshû and Kyûshû. (Yano)” (translated from Japanese; Fig. 8AView FIGURE 8).

Yano put “ Ḿü ” (= new name) just after the Japanese name of Mutilla cameroni  . This Japanese word is used to denote both new Japanese name and new replacement name. However, Yano did not refer to Camron’s species Mutilla japonica  , so this “new name” should be regarded as indicating a new Japanese name for a newly erected taxon but not a new name for Mu. japonica  . Although the syntypes of Mu. cameroni  were not designated, the original description and the illustration fit the characteristics of Sm. lewisi  . The name Mutilla cameroni  was preoccupied by Schulz (1906), so Mickel (1935) proposed a new name Sm. lewisi yanoi  . At that time, Mickel in his key separated Sm. l. lewisi  and Sm. l. yanoi by the propodeum (with well-defined black maculation dorsally in Sm. l. lewisi  ; entirely yellowish red in Sm. l. yanoi). Later Tsuneki (1972a) and Lelej (1985) synonymized Sm. l. yanoi under the nominotypical form. This was reasonable because the black maculation on the propodeum varies greatly and both the lewisi  form and yanoi form have been found in the same locality.

Chen (1957) placed Sm. l. yanoi as a junior synonym of Sm. strandi  . His materials were composed of males only. The male of Sm. strandi  can be easily separated from that of Sm. lewisi  by having the gena with small sparse punctures, the median elevation and subapical teeth on clypeus separated, the mesosoma yellowish red except the propleuron and mesopleuron ventrally black and the mesopleuron beneath without a precoxal longitudinal impunctate line, although these two species are identical in genitalic morphology. The true Sm. lewisi  was not included in Chen’s specimens of Sm. strandi  , while those of Sm. rufipes  were actually Sm. lewisi  . Full species status of Sm. lewisi  is supported in that Sm. lewisi  and Sm. strandi  share an overlapping distribution in southern China, suggesting that there may be premating barriers between these two species. The following specimens of Sm. strandi  were examined: 1♂, Guling, Lushan, Jiangxi, China, 3.VIII.1935, O. Piel leg., B. Petersen det. 1987 as Smicromyrme strandi strandi (Zavattari)  [TARI]; 1♂, Shui Pei Kai, Shaowu, Fujian, China, 24.V.1944, T. C. Maa leg., B. Petersen det. 1987 as Smicromyrme strandi strandi (Zavattari)  [TARI].

The body of Sm. momidziyama Tsuneki, 1972  is covered with golden setae instead of black setae ( Fig. 16View FIGURE 16). The color of setae in mutillids sometimes varies greatly. In females of Sm. lewisi  , Nemka limi ( Chen, 1957)  , N. wotani  , N. yasumatsui (Mikel, 1936)  , stat. rev. et comb. nov. and Mickelomyrme hageni  , the setae on frons, vertex and thoracic dorsum continuously changes from black to golden. Potentially, black setae can be easily altered to golden ones, so Sm. momidziyama  is regarded as a mutant of Sm. lewisi  . Tsuneki (1972) also referred to the smaller posterior punctate area on the dorsal propodeal face and the narrower and longer pygidial area as diagnostic. These characters also vary and Sm. momidziyama  is not distinguishable from Sm. lewisi  . Therefore, Lelej’s (2002) synonymization of Sm. momidziyama  under Sm. lewisi  was reasonable.

Remarks. The holotype of Sm. lewisi  has a large depression on the frons below the right eye.

Terayama et al. (2011) reported that 2/3 of specimens from Hokkaidô fit the nigricula form. In the present study, only three males of this form collected in Japan were examined. Although males of the typical form are widely distributed throughout Palaearctic Japan, the nigricula form can be found in Hokkaidô and northern Honshû only. In Hokkaidô, males of both forms are sometimes collected in the same place. There are no morphological differences between these two forms, including the male genitalia, as Tsuneki (1972a) had already mentioned. One additional specimen of the nigricula form from South Korea was also examined. This record implies that this form is an environmentally affected phenotype rather than a genetically endemic population.

HUM

Humboldt University Zoologisches Museum

NSMT

National Science Museum (Natural History)

NIAES

National Institute for Agro-Environmental Sciences

PRI

College of Eastern Utah

AES

University of Alaska

NMNH

Smithsonian Institution, National Museum of Natural History

ZMUC

Zoological Museum, University of Copenhagen

TARI

Taiwan Agricultural Research Institute

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hymenoptera

Family

Mutillidae

Genus

Smicromyrme

Loc

Smicromyrme (Smicromyrme) lewisi Mickel, 1935

Okayasu, Juriya 2020
2020
Loc

Smicromyrme alberici:

Shintani, T. 2003: 17
2003
Loc

Mutilla cameroni

Kim, C. W. & Lee, J. W. & Park, J. S. & Kim, B. J. & Park, J. C. 1994: 255
1994
Loc

Smicromyrme rufipes lewisi var. nigricula

Yamada, M. 2001: 57
Yoshida, M. 1989: 651
1989
Loc

Smicromyrme momidziyama

Lelej, A. S. 2002: 70
Hashimoto, Y. & Nakanishi, A. 1997: 31
Yoshida, M. 1989: 651
Tsuneki, K. 1972: 17
1972
Loc

Smicromyrme rufipes lewisi: Tsuneki 1972a: 16

Kameyama, T. & Nishi, M. & Nakamura, S. 2006: 74
Nakamura, S. & Kameyama, T. & Katayama, S. 2001: 83
Yamada, M. 2001: 57
Matsuda, S. & Nakamura, S. 1999: 111
Nakamura, S. & Haneda, Y. 1997: 34
Takahashi, H. 1996: 28
Kim, C. W. & Lee, J. W. & Park, J. S. & Kim, B. J. & Park, J. C. 1994: 255
Yamada, H. 1990: 480
Yoshida, M. 1989: 651
Lelej, A. S. 1981: 78
Lelej, A. S. 1977: 530
Lelej, A. S. 1976: 272
Tsuneki, K. 1972: 16
1972
Loc

Smicromyrme rufipes lewisi

Lelej, A. S. 1977: 91
Tsuneki, K. 1972: 16
1972
Loc

Smicromyrme rufipes rufipes:

Chen, C. - W. 1957: 200
1957
Loc

Smicromyrme lewisi lewisi

Okayasu, J. & Williams, K. A. & Lelej, A. S. 2018: 304
Lelej, A. S. 2017: 156
Terayama, M. & Suda, H. & Murota, T. & Tano, T. 2016: 168
Lo Cascio, P. 2015: 558
Nagase, H. & Harada, K. & Ito, H. & Yago, M. 2014: 56
Hayashi, M. & Kadowaki, H. & Matsuda, T. & Fukaya, H. & Chikami, Y. 2013: 25
Ogawa, H. & Nosaka, M. & Hashii, N. & Yokoyama, M. & Tsurusaki, N. 2012: 38
Tsurusaki, N. & Hayashi, M. & Miyanaga, R. & Ichisawa, K. & Kawakami, Y. 2012: 70
Terayama, M. & Suda, H. & Murota, T. & Haneda, Y. & Tano, T. 2011: 20
Fukasawa, S. & Miyano, S. 2010: 25
Ohkusa, S. & Tajima, T. 2010: 7
Ohkusa, S. & Tajima, T. 2008: 69
Ohkusa, S. 2007: 30
Yamamoto, E. & Haneda, Y. 2007: 15
Haneda, Y. & Nozaka, C. & Tano, T. & Kurokawa, H. & Murota, T. 2006: 5
Haneda, Y. & Nozaka, C. & Tano, T. & Kurokawa, H. & Murota, T. 2006: 40
Ohkusa, S. 2006: 22
Haneda, Y. & Inoue, S. & Nozaka, C. & Tano, T. & Kurokawa, H. & Murota, T. 2005: 57
Kurokawa, H. 2005: 76
Kinota, K. & Haneda, Y. & Tano, T. 2005: 25
Negoro, H. & Haneda, Y. & Tano, T. 2005: 32
Terayama, M. 2005: 8
Nagase, H. 2004: 1280
Tano, T. & Kurokawa, H. & Murota, T. & Nozaka, C. 2003: 16
Lelej, A. S. 2002: 70
Haneda, Y. & Yamamoto, E. 2000: 780
Kubo, K. 2000: 318
Nakamura, S. & Haneda, Y. 2000: 119
Takahashi, H. & Matsumoto, Y. 1999: 23
Yamane, Sk. 1999: 380
Lelej, A. S. & Yamane, Sk. 1992: 630
Lelej, A. S. 1985: 232
Kim, C. W. 1970: 793
Kim, C. W. 1963: 341
Yasumatsu, K. 1938: 85
Mickel, C. E. 1935: 288
1935
Loc

Smicromyrme lewisi yanoi

Hisamatsu, M. 2004: 135
Lelej, A. S. 1985: 233
Tsuneki, K. 1972: 16
Kim, C. W. 1970: 793
Ishikawa, R. 1965: 289
Kim, C. W. 1963: 341
Yasumatsu, K. 1951: 70
Yasumatsu, K. 1938: 85
Yasumatsu, K. 1938: 351
Yasumatsu, K. 1937: 68
Mickel, C. E. 1935: 289
1935
Loc

Smicromyrme lewisi nigricula

Lelej, A. S. 1985: 233
Mickel, C. E. 1935: 289
1935