Tenodera sinensis Saussure, 1871

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

Tenodera sinensis Saussure, 1871

Figs 88–97 , 98–102

Mantis mandarinea Saussure, 1871 a: 289 .

Tenodera aridifolia var. sinensis Saussure, 1871 b: 417 .

Tenodera aridifolia (Stoll, 1813) : ESK and KSAE 1994: 44 (misidentification). Korean record.

Tenodera sinensis Saussure, 1871: Kim 2010: 31; Kim 2021: 65. Korean record.

Specimens examined.

[KsNU] South Korea: JB: 1 ♂, Eunpa Lake , Gunsan-si, 8 XI 2016, Donghwan Na ; [ NASIC] South Korea: GW: 2 Nymphs, Balsan 2 - ri , Chuncheon-si, 12 VI 1998, Sungsoon Jang ; 1 ♀, Jinburyeong , Jinbu-myeon, Pyeongchang-gun, 29 IX 2000, Taehwa Kang ; 1 Nymph, Ingye-ri , Okgye-myeon, Gangneung-si, 15 VIII 2002, Jingoo Yeo ; 3 ♂, Mt. Odae , Hongcheong-gun, 29 VIII 2019, Jaeil Shim ; GG: 1 ♂, Mijang-ri , Samjuk-myeon, Anseong-si, 17 IX 2000, Yeongbo Lee ; 1 ♂, Mt. Cheolma , Incheon, 26 VIII 2001, Taewoo Kim ; 1 Nymph, Shihwa , Siheung-si, 30 VII 2003, Jaecheon Son ; 1 Nymph, Temple Jeondeungsa , Onsu-ri, Gilsang-myeon, Ganghwa-gun, Incheon, 4 IX 2009, Yeongbo Lee ; 1 ♀, Jikdong-ri , Sohol-eup, Pocheon-si, 7 IX 2011, Yeongbo Lee ; 2 Nymphs, Temple Bogwangsa , Gwangtan-myeon, Paju-si, 10 VII 2013, Yeongbo Lee ; 3 Nymphs, World Cup Park , Seongsan-dong, Mapo-gu, Seoul, 21 VII 2013, Yeongbo Lee ; 2 Nymphs, Noel Park , Sangam-dong, Mapo-gu, Seoul, 30 VII 2013, Yeongbo Lee;) ; 1 ♂, Island Gureopdo , Gureop-ri, Deokjeok-myeon, Incheon, 28 VI 2023, Jaeil Shim, Wonjun Sung (reared from nymph) ; CB: 1 Nymph, Magok-ri , Bongyang-eup, Jecheon-si, 13 VII 2005, Taehwa Kang ; 1 ♀, Gamgok-myeon , Eumseong-gun, 31 VIII 2019, Byeongmin Jeong ; 3 ♂, Geumseok-ri , Geumwang-eup, Eumseong-gun, 14 IX 2019, Seong-Gyu Lee ; 1 ♂, Jeongbang-ri , Annae-myeon, Okcheon-gun, 12 X 2000, Yeongbo Lee ; CN: 1 ♀, Mt. Sikjang , Daejeon, 14 VIII 2019, Geonheyok Kim (reared from nymph) ; 4 ♂, Coastal Dune, Sindu-ri , Wonbuk-myeon, Taean-gun, 12 VIII 2023, Jaeil Shim, near the grassland ; GB: 1 Nymph, Geumgok-ri , Byeonggok- myeon, Yeongdeok-gun, 1 VII 2009, Yeongbo Lee, Hoyeon Jeong ; GN: 4 ♀, Mt. Noja , Dongbu- myeon, Island Geojedo, Geoje-si, 15 IX 2021, Jaeil Shim ; JB: 1 ♀, Hoyja 3 - dong , Wansan-gu, Jeonju-si, 28 VIII 2014, Taeman Han ; 1 ♂, Deokjin Park , Deokjin-gu, Jeonju-si, 6 IX 2018, Junhee Park ; 1 ♀, Jeonju riverside , Jeonju-si, 15 IX 2018, Jaeil Shim ; 1 ♂, Mt. Moak , Gui-myeon, Wanju-gun, 6 X 2018, Hyeon-Ha Yoo ; 1 ♂, Jinan-Maisan Rest area of expressway , Jinan-gun, 22 VIII 2019, Jaeil Shim ; 1 ♀, Jeonju Univ. , Jeonju-si, 4 IX 2019, Jaeil Shim ; 1 ♂, Jangsu-eup , Jangsu-gun, 5 IX 2019, Jaeil Shim ; 11 ♂ 2 ♀, Mt. Moak , Gui-myeon, Wanju-gun, 7 IX 2019, Jaeil Shim ; 3 ♂, Temple Geumsansa , Gimje-gun, Gimje-si, 5 IX 2020, Jaeil Shim ; JN: 1 ♂, Myeongsasimni , Coastal Dune, Island Bigeumdo, Sinan-gun, 22 IX 2001, Haechul Park ; 1 ♀, Island Gogeumdo , Wando-gun, 3 IX 2003, Mikyung Ahn ; 1 ♂ 1 ♀, Bukyi-myeon , Jangseong-gun, 20 VIII 2021, Jaeil Shim;) ; 2 ♂ 1 ♀, Island Yeoseodo , Yeoseo-ri, Cheongsan-myeon, Wando-gun, VI 2023, Jaeil shim (reared from nymph) ; JJ: 1 ♀, Ihoteawu Beach , Jeju-si, 26 VIII 2014, Taeman Han, windbreak forest ; 2 Nymphs, Gwangchigi Beach , Goseong-ri, Seongsan-eup, Seogwipo-si, 16 V 2021, Jaeil Shim .

Redescription.

Measurements (mm): Total length (vertex to tip of abdomen) ♂ 54.2–89.1, ♀ 58.2–100.8; head width ♂ 5.9–6.5, ♀ 6.9–7.8; head length ♂ 4.5–4.9, ♀ 5.6–6.7; pronotum width ♂ 4.3–5.6, ♀ 4.9–7.4; pronotum length ♂ 18.0–24.3, ♀ 19.2–27.0; forewing (tegmina) length ♂ 38.4–55.2, ♀ 38.1–63.2. Male (Figs 88 , 90 , 92 , 95 – 101 ). Large to very large. Coloration (Figs 88 , 98 ): Body and forewing discoidal area green to greenish brown or brown. Head (Fig. 90 ): Triangular. Head width 1.2 × as long as head. Vertex slightly convex, brownish; apex with a bright brown transverse line (in live specimens). Ocelli large, oblong. Antenna nearly 1.5 × as long as pronotum. Epistomal sulcus slightly concave. Lower frons, clypeus, and labrum with two darkish longitudinal stripes. Prothorax (Fig. 92 ): Pronotum long but robust, flattened dorso-ventrally, dorsal surface smooth; pronotum length 3.8–4.0 × as long as maximum width. Prozone lateral margin and dorsal surface with minute denticles. Metazone (Fig. 92 ) often orangish; lateral margin smooth; 3.0–3.2 × as long as prozone. Between forecoxa basis membranous attachment surface yellow (Fig. 94 ). Postcervical plate reddish in brown morph (Fig. 94 ), anterior area of furcasternite with gradational dark pattern in brown morph, its pattern occasionally occurring in green morph. Forelegs (Prothoracic legs) (Figs 94–97 ): Coxa dorsal margin with 14–17 whitish conical spines; ventral surface of coxa proximal area with gradational dark pattern (Fig. 94 ); remaining surface with numerous small white spots. Tibial spur groove with faint dark spot (Fig. 94 ). Spination formula (Figs 95–97 ): Avts = 14–15; Pvts = 8–10; Avfs = 14–17; Pvfs = 4; Ds = 4. In 16 Avfs (Figs 95 , 96 ): spines 2, 4, 6, 8, 10, 12, and 16 larger than remaining Avfs. Tarsomere distal end dark brown. Tarsi 5 - segmented. Wings (Fig. 98 ): Forewing completely surpassing the end of abdomen. Forewing costal area green, discoidal area transparent. Hindwing venation brown, cross veins and cells of subcostal to radius area reddish to magenta; radius to cubitus proximal area, near cells of arculus veins dark brown (Fig. 98 ); anal area with dark brownish smoky pattern. Abdomen (Figs 99 , 100 ): Middle of the abdominal sternites with longitudinal yellow stripe pattern (Fig. 99 ). Cerci setose, not flattened, with 17–20 segments. Male subgenital plate (coxosternite IX) (Fig. 100 ) irregular rhomboidal, inter-stylar margin convex at the middle; ventral surface with numerous setae. Styli rather short. Male genitalia (Fig. 101 ): Right phallomere forming nearly V-shaped pva; pia sclerotized and weakly wrinkled; posterior surface of pia (Fig. 101 ) with membranous wide hump, surface with minute denticulation; fda elongate lobe shape. Left phallomere (Fig. 101 ) with elongated and curved paa, its apex round; aafa sclerotized, surface smooth, weakly bulbous spike-shaped basally, apically curved dorso-laterally; pafa sclerotized, nearly 90 ° angle, arched wide blade-shaped (apex expanded), posterior and apical margin of pafa with numerous spines; loa membranous, elongate finger-shaped, much longer than pafa; L 4 B curved spoon-shaped. Ventral phallomere irregular rhomboidal; posterior margin prominently expanded; sdpl hardly sclerotized, curved at ~ 45 ° at middle, its distal half slightly thicker and more sclerotized and melanized than basal, point of sdpl tips shallowly concave. Female (Figs 89 , 91 , 93 , 94 ). Similar to male, with following differences. Head (Fig. 91 ): width 1.1 × as long as head length. Vertex convex. Antenna as long as pronotum. Ocelli smaller than male. Prothorax (Fig. 93 ): Pronotum length 2.8–2.9 × as long as maximum width; lateral margin with numerous denticles. Prozone with numerous blunt denticles on dorsal surface. Medial keel protruded. Forelegs (Prothoracic legs): Coxa dorsal margin (Fig. 94 ) with 14–20 large conical spines (with sharp tips), small denticles located between them. Wings: Forewing occasionally reaching end of abdomen. Abdomen: Elongate oval, much broader than in male. Cerci with 15–17 segments. Ootheca (Fig. 102 ). Measurements (mm): Length 30.0–42.2; maximum width 22.2–27.6; maximum height 20.7–24.9; length of emergence area 18.4–32.8; width of emergence area 5.2–5.5. Identification: Barrel-like shape, mostly circular in cross-section. Ootheca attached to flat substrate by its ventral surface or fully encircling substrates such as sticks. External wall with very thick air-filled area, colored beige to bright brown. External coating covering entire surface of ootheca; beige in color. Emergence area depressed. Exhibiting ~ 15– 29 egg chambers; lips invisible due to very thick air-filled layer of ootheca. Distal end of ootheca obliquely truncate. Nymph. Mid to last instar nymph: Avfs 1 base (near femoral brush), last Avfs base to tibial spur groove area with dark spotted pattern; between forecoxa basis attachment membranous surface yellow.

Biological notes.

Tenodera sinensis occurs throughout the Korean peninsula and has adapted well to urban, suburban, and riverside environments. It spawns ootheca in various locations, such as on stones, tree trunks, and branches. First instar nymphs hatch from mid-April to mid-May, while adult mantises typically emerge in mid-August.

Distribution.

China, Nepal, Japan, Russia, Thailand, South Korea. Invasive in Canada and the USA.

Remarks.

Tenodera sinensis was originally described as a variation of the widely distributed species Tenodera aridifolia (Stoll, 1813) , and treated as a subspecies for quite a long time ( Saussure 1871 b; Rehn 1903; Shiraki 1932; Beier 1932; Tinkham 1937; Bazyluk 1977; Iwasaki 1996; Bruins 1999), but already Giglio-Tos (1927) and Ehrmann (2002) considered T. sinensis as a valid species. Tenodera aridifolia and T. sinensis are extremely close and morphologically very similar ( Ehrmann and Borer 2015), but Jensen et al. (2010) provide differences in male genitalia of T. sinensis and T. aridifolia . According to Jensen et al. (2010), it is a distinctly divided sister species pair based on phylogeny using nuclear genes (histone III, wingless gene) and mitochondrial genes (large and small rRNA, cytochrom oxidase II). Tenodera aridifolia is found in tropical and subtropical regions but is replaced by T. sinensis in temperate habitats. Our study used one male West Javan T. aridifolia specimen to comparatively examine and recovered remarkable characters from T. sinensis including the narrow pronotum (length / maximum width = 5.1) and forewing, a smaller head proportional to body size, short and stout aafa, pafa, and sdpl curved at ~ 45 ° ( Mukherjee et al. 1995; Jensen et al. 2010).

Subfamily Hierodulinae Brunner de Wattenwyl, 1893

Figures

Figures 88–97. Habitus, head, pronotum and foreleg of Tenodera sinensis 88 male dorsal aspect 89 female dorsal aspect 90 male face 91 female face 92 male pronotum 93 female pronotum 94 ventral aspect of foreleg and furcasternite (live specimens) 95 foreleg ventral aspect 96 foreleg interior aspect 97 foreleg tibia and tarsus (dorsal aspect). Abbreviation: Tsg = tibial spur groove. Red arrows = dark spot near the tibial spur groove.

Figures 98–102. Wings, abdomen, male genitalia and ootheca of Tenodera sinensis 98 wings 99 male abdomenal sternites (above: green morpho-type below: brown morpho-type) 100 subgenital plate 101 male genitalia (small box = lateral aspect of aafa) 102 ootheca (left: dorsal aspect right above: lateral aspect right below: distal aspect). Red arrow = arculus area.