Statilia maculata ( Thunberg, 1784 )

Shim, Jaeil & Song, Jeong-Hun, 2024, A taxonomic review of the order Mantodea in Korea based on morphology and DNA barcodes, ZooKeys 1206, pp. 1-43 : 1-43

Statilia maculata ( Thunberg, 1784)

Figs 52–62 , 63–70

Mantis maculata Thunberg, 1784: 61.

Pseudomantis haanii Saussure, 1871 a: 37 , 1871 b: 400.

Statilia maculata var. hyalina Giglio-Tos, 1927: 410.

Statilia haani var. major Werner, 1922: 154.

Statilia maculata continentalis Werner, 1935: 495.

Statilia maculata ( Thunberg, 1784): ESK and KSAE 1994: 44; Kim 2010: 31; Kim 2021: 65. Korean record.

Statilia nemoralis ( Saussure, 1870): Jeon et al. 1999: 227 (misidentification); Kim 2010: 31 (misidentification). Korean record.

Specimens examined.

[KsNU] South Korea: CN: 1 ♂, Mt. Bongsoo , Daechung-myeon, Yesan-gun, 18 V 2017 Hongjoon Choi ; JB: 1 ♀, Eunpa Lake , Jigok-dong, Gunsan-si, 3 IX 2016; Donghwan Na ; 1 ♀, Miryong-dong , Gunsan-si, 27 VI 2017, Eunhye Jeon ; [ NASIC] South Korea: GW: 1 ♂, Ssangyong , Yeongwol-gun, 30 IX 1999, Miae Kim ; GG: 1 ♀, Mt. Umyeon , Seocho-gu, Seoul, 1 IX 1991, Jiyung Oh ; 1 ♀, Jamsil 6 - dong , Songpa-gu, Seoul, 20 X 1997, Soyeon Kim ; 1 ♀, Mt. Nam , Hada-ri, Heungcheon-myeon, Yeoju-si, 3 IX 2000, Yeongbo Lee ; 1 ♀, Seodun-dong , Suwon-si, 4 IX 2000, Taewoo Kim ; 1 ♂, Seodun-dong , Suwon-si, 20 IX 2000, Taewoo Kim ; 1 ♂, Seodun-dong , Suwon-si, 20 IX 2001, Jeonghun Hwang ; ♂ 1, Haguidong , Uiwang-si, 24 IX 2003, Jeongsun Lee ; 1 ♂, Mt. Gwanggyo , Jangan-gu, Suwon-si, 10 X 2003, Mikyeong Ahn ; CB: 1 ♀, Mt. Nam , Cheongju-si, 8 VIII 2000, Hyea Lee ; 1 ♂, Geumseok-ri , Geumwang-eup, Eumseong-gun, 14 IX 2019, Seong-Gyu Lee ; CN: 1 ♂ 1 ♀, Coastal Dune , Sindu-ri, Wonbuk-myeon, Taean-gun, 1 IX 2005, Yeongbo Lee ; GB: 1 ♀, Mt. Sobaek , Punggi-gun, 5 IX 2000, Miae Kim ; 1 ♂ 3 ♀, Mt. Angi , Songhyeon-dong, Andong-si, VIII 2022, Jaeil Shim ; 2 ♂ 6 ♀, Hotel Interburgo , Manchon-dong, Suseong-gu, Daegu-si 29 X 2023, Jaeil Shim ; GN: 1 ♂, Sinhyeon-ri , Geoje island, Geoje-si, 11 IX 2002, Miae Kim ; 2 ♂ 1 ♀, Mundong Waterfall , Geoje-si, 19 IX 2006, Miae Kim ; ♀ 1 (green), Recreational Forest , Dongbu-myeon, Geojedo, Geoje-si, 29 IX 2020, Wonjun Sung ; JB: 2 ♂ 4 ♀, Gueok , Yongjin-myeon, Wanju, 13 VIII 2013, Hanjun Bae ; 1 ♂, Iseo-myeon , Wanju-gun, 26 VII 2016, Taeman Han ; 1 ♀, Iseo-myeon , Wanju-gun, 29 VII 2014, Kyusuk Lee ; 1 ♂ 1 ♀, Apartment , Iseo-myeon, Wanju-gun, 6 VIII 2015, Kyusuk Lee ; 1 ♀ (green), National Institute of Agricultural Sciences , Iseo-myeon, Wanju-gun, 24 VIII 2016, Hyeop Lee ; 1 ♀, Miryong-dong , Gunsan-si, 27 VI 2017, Eunhye Jeon ; 2 ♂, Iseo-myeon , Wanju-gun, 23 VIII 2019, Jaeil Shim ; 1 ♂ 2 ♀, Mt. Moak , Gui-myeon, Wanju-gun, 7 IX 2019, Jaeil Shim ; 7 ♂ 5 ♀, Jangsu-eup , Jangsu-gun, 5 IX 2019, Jaeil Shim ; JN: 3 ♀, Geumgok-dong , Buk-gu, Gwangju-si, 20 III 2021, Jaeil Shim ; 4 ♀, Near Yeosu Airport , Sinpung-ri, Yulchon-myeon, Yeosu-si, IX 2021, Byeongmin Jeong ; 2 ♂ 1 ♀, Is. Bogil-do , Bogil-myoen, Wando-gun, 2 III 2023, Jaeil Shim (reared from ootheca) ; 8 ♂ 9 ♀, Island Yeoseodo , Yeoseo-ri, Cheongsan-myeon, Wando-gun, VI 2023, Jaeil shim (reared from Ootheca) ; JJ: 1 ♂, Seonheul-ri , Jeju-si, 28 IX 2000, Taewoo Kim ; 1 ♂, Sinae 1 - ri , Jeju-si, 18 X 2001, Mikyeong Ahn ; 1 ♂, Bijarim Forest , Jeju-si, 22 IX 2006, Miae Kim ; Japan: 1 ♂, Asakura , Kyushu, 24 VIII 2013, Sangwook Park ; [ NIBR] South Korea: GG: 1 ♀, Namhansanseong , Seongnam, 28 IX 1997, Jeong Yun Chang .

Redescription.

Measurements (mm): Total length (vertex to tip of abdomen) ♂ 35.2–43.1, ♀ 43.7–57.7; head width ♂ 4.5–5.0, ♀ 5.4–5.8; head length ♂ 2.9–3.2, ♀ 4.1–4.3; pronotum width ♂ 2.9–3.2, ♀ 4.4–4.6; pronotum length ♂ 12.1–14.0, ♀ 14.0–15.3; forewing (tegmina) length ♂ 26.4–31.2, ♀ 33.1–36.1. Male (Figs 52 , 55 , 57 , 63–68 ) Medium sized, body slender. Coloration (Figs 52 , 53 ): Beige to dark brown. Head (Fig. 55 ): Triangular. Head width 1.5–1.6 × length. Vertex flat; dark transverse stripe on dorsal apex. Vertex and juxtaocular bulge with sparse, pale, transversely striped pattern. Compound eye large, globular, protruding laterally; in live specimens, dorsal and lateral surface with two brightly colored lines. Ocelli large, oblong. Antenna filiform, slightly longer than pronotum. Lower frons posterior apex protruding very slightly. Epistomal sulcus slightly concave. Prothorax (Fig. 57 ): Pronotum slender, narrow; length 4.1–4.3 × as long as maximum width; dorsal surface smooth. Pronotum lateral margin with numerous denticles. Medial keel protruding. Metazone 3 × as long as prozone. Membranous area between basis of forecoxa attachment, shiny and pinkish pearl in color (in live specimens). Furcasternite (Fig. 59 ) anterior area with rectangular dark spot, remaining posterior area reddish brown to magenta. Forelegs (Prothoracic legs) (Figs 62 – 65 ): Coxa very slender, triangular; dorsal margin (Figs 59 , 62 ) with 6–8 white spines; proximal area of ventral surface (Fig. 59 ) with large and shiny rectangular black spot. Coxal lobes divided from each other. Femur ventral surface shiny, with two black spots (Figs 62 , 63 ), large rectangular spot preceding the tibial spur groove, linear spot (transverse line) distad to tibial spur grooves; tibial spur groove (Fig. 63 ) with white or whitish yellow pattern; distal area of ventral surface (Fig. 62 ) pale pink to magenta (in live specimens). Spination formula: Avts = 11; Pvts = 6–7; Avfs = 14; Pvfs = 4; Ds = 4. Tarsomere ventral surface (euplantula) dark brown. Basal rim of Pvts with dark spot. In 14 Avfs (Figs 63 , 64 ): spines 2, 4, 6, 8, 10, 12, and 14 larger in size than remaining Avfs; spines 1, 2, 4, 6, 8, 10, 12, and 14 black, with small dark spot at the base. Meso- and metathoracic legs: Tarsi 5 - segmented. Wings (Fig. 66 ): Forewing completely surpassing the end of abdomen. Stigma elongate, slightly protruding. Hindwing costal area reddish brown; discoidal area with darkish smoky mottled pattern but cross veins clearly transparent. Abdomen: Cerci setose, not flattened, brown, with 14 segments. Male subgenital plate (coxosternite IX) (Fig. 67 ) irregular rhomboidal, inter-stylar margin deeply notched in V-shape. Styli rather long. Male genitalia (Fig. 68 ): Right phallomere forming a V-shape pva; pia sclerotized and weakly wrinkled; fda triangular. Left phallomere with elongate and curved paa, surface smooth, distal end blunt and slightly swollen; afa sclerotized, small and irregularly rough, surface weakly granulated; L 4 B C-shaped. Ventral phallomere (Fig. 68 ) irregular rhomboidal; sdpm sclerotized, wide triangular; sdpl anteriorly curved hook-shaped with blunt apex, margin between sdpm and sdpl flat or slightly concave. Female (Figs 53 , 56 , 58 , 59 ). Similar to male, with following differences. Coloration (Fig. 53 ): Body and forewing color beige to dark brown. Head (Fig. 56 ): width 1.3 × as long as length. Antenna 2 × longer than prozone length. Prothorax (Figs 58 ): Pronotum length 3.1–3.3 × as long as maximum width. Green morphotype female (Figs 54 , 60 , 61 ). Prothorax: Rectangular black spot (Fig. 60 ) of furcasternite absent or very weakly developed, remaining posterior area greenish. Forlegs (Prothoracic legs) (Figs 60 , 61 ): Tibial spur grooves (Fig. 61 ) with rectangular pale spotted pattern. Distal area of femur ventral surface pale pink or green. Wings (Fig. 54 ): Hindwing transparent or anal area with few mottled dark spots. Ootheca (Figs 69 , 70 ). Measurements (mm): Length 20.4–28.9; maximum width 6.1–10.2; maximum height 6.4–8.3; length of emergence area 15.3–24.2; width of emergence area 2.2–3.5. Identification: Fusiform in shape, nearly hemispherical in cross-section. Proximal end with medial elevation of emergence area. Ootheca attached by its ventral surface. External wall bright brown. External coating weakly covering lateral zone of emergence area; beige in color. Exhibiting ~ 25– 50 egg chambers clearly delimited by visible prominently oblique lips (Fig. 70 ). Distal end of ootheca narrowed into residual process, greatly elongated, and attached to substrate. Nymph. First instar nymph: Body dark brown, leg with few brightly colored stripes. Mid to last instar nymph: Forefemur distal area (pale pink to magenta) covered by mottled black or brown spots.

Biological notes.

Statilia maculata is found throughout the Korean Peninsula. This species has shown a remarkable adaptability to urban and suburban environments, and is often observed on building walls and streetlights, exhibiting positive phototaxis. They are known to deposit their oothecae under stones and in cracks of tree bark. Notably, S. maculata is capable of producing a hissing sound by rubbing its hindwings and abdomen together. This species typically hatches in early June, with adults emerging in the middle of August.

Distribution.

China, Japan, Nepal, South Korea, Taiwan. Invasive in Eastern USA and Russia.

Remarks.

Statilia maculata is predominantly found in the eastern Palearctic regions ( S. maculata has also been introduced to Eastern USA and Russia), whereas S. nemoralis is described from the Philippines and, erroneously, from various Southeast Asian countries ( Ehrmann 2002; Patel and Singh 2016; Schwarz et al. 2018). These two species, with S. nobilis ( Brunner de Wattenwyl, 1893) , are frequently confused in the taxonomic literature, leading to numerous misidentifications (see details in Schwarz et al. 2018); for example, both S. nobilis and S. maculata have been erroneously reported as S. nemoralis ( Wang 1993; Jeon et al. 1999; Oshima 2017; Schwarz et al. 2018; Oshima 2020; Shcherbakov and Govorov 2020). Notably, the green morph is relatively uncommon in female S. maculata (Fig. 54 ) but prevalent in both sexes of S. nobilis and S. nemoralis ( Zhu et al. 2012; Ehrmann and Borer 2015; Schwarz et al. 2018).

Jeon et al. (1999) initially reported S. nemoralis in Korea based on two female specimens lacking black spots on the furcasternite. Differences in the male genitalia, specifically the margins of the sdpl and sdpm, are reliable for distinguishing between S. maculata and S. nobilis ( Schwarz et al. 2018) . On the other hand, the lack of a black spot on the furcasternite is a common feature of the green morph of S. maculata . The occurrence of the Philippine Statilia nemoralis is continental SE Asia needs confirmation. Subsequent examinations revealed that the specimens initially identified by Jeon et al. (1999) as S. nemoralis were, in fact, misidentifications of S. maculata .

Subfamily Tenoderinae Brunner de Wattenwyl, 1893

Figures

Figures 52–62. Habitus, head, pronotum and foreleg of Statilia maculata 52 male dorsal aspect 53 female dorsal aspect 54 female dorsal aspect (green morpho-type) 55 male face 56 female face 57 male pronotum 58 female pronotum 59 furcasternite (brown morpho-type) 60 furcasternite (green morpho-type) 61 femur ventral aspect (green morpho-type) 62 foreleg ventral aspect (live specimens). Abbreviation: Tsg = tibial spur groove. Red arrows = dark spot of furcasternite.

Figures 63–70. Foreleg, wings, male genitalia and ootheca of Statilia maculata 63 foreleg ventral aspect 64 foreleg interior aspect 65 foreleg dorsal aspect 66 wings 67 subgenital plate 68 male genitalia 69 ootheca (dorsal aspect) 70 ootheca (lateral asepct). Abbreviation: Ts = tibial spur Tsg = tibial spur groove. Red arrow = stigma.