Hierodula patellifera ( Audinet-Serville, 1838 )

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

Hierodula patellifera ( Audinet-Serville, 1838)

Figs 121–132 , 133–136

Mantis patellifera Audinet-Serville, 1838: 185.

Mantis bipapilla Audinet-Serville, 1838: 188.

Hierodula assamensis Mukherjee et al., 1995: 185.

Hierodula manillensis Saussure, 1870: 233.

Hierodula raptoria Stål, 1877: 38.

Hierodula dispar Kirby, 1900: 146.

Hierodula saussurei Kirby, 1904: 245.

Hierodula manillana Giglio-Tos, 1912: 96.

Hierodula (Hierodula) manillana Giglio-Tos, 1927: 448.

Hierodula yunnanensis Wang, 1993: 137.

Hierodula xishaensis Wang, 1993: 140.

Hierodula multispina Wang, 1993: 141.

Hierodula daqinshanensis Wang, 1993: 143.

Hierodula patellifera ( Audinet-Serville, 1838): Jeon et al. 1999: 226 ( South Korea); Kim 2010: 31; Kim 2021: 65; Shim et al. 2021 b: 149. Korean record.

Specimens examined.

[ NASIC] South Korea: GG: 1 ♀, Seodun-dong , Suwon-si, 2 IX 1999, Taewoo Kim ; 1 ♂, Seodun-dong , Suwon-si, 13 IV – 17 VIII 2001, Taewoo Kim (reared) ; 4 ♂ 1 ♀, SETEC, Daechi-dong , Gangnam-gu, Seoul, 8 VI 2023, Jaeil Shim (reared from nymph) ; CB: 1 ♂ 5 ♀, Parking area of Cheongju Airport , Ipsang-ri, Naesu-eup, Cheongwon-gu, Cheongju-si, 15 IX 2023, Jaeil Shim ; CN: 1 ♂, Rest area , Geumsan-gun, 10 IX 2013, Haechul Park ; 1 ♂, Mt. Gubong , Gwanjeo-dong, Seo-gu, Daejeon, 1 IX 2014, Taekyu Kim ; 1 ♀, Chungnam Nat. Univ. , Yuseong-gu, Daejeon, 27 IX 2015, Taeman Han ; 4 ♂ 2 ♀, Samsong-ri , Haemi-myeon, Seosan-si, 24 IV 2023, Jaeil Shim (reared from oothecae) ; 4 ♂ 9 ♀, Chungnam Nat. Univ. , Yuseong-gu, Daejeon-si, 19 VIII 2023, Jaeil Shim ; GB: 2 ♀, Street near Gomo station (Gomo-ro) , Suseong-gu, Daegu, 14 IX 2019, Jaeil Shim ; 8 ♀, Hyomok Elementary Scholl , Hyomok-dong, Daegu-si 29 X 2023, Jaeil Shim ; GN: 1 ♀, Mt. Mang , Island Geojedo, Geoje-si, 23 VII 2019, Jun-Gi Lee, Jun-Ho Lee ; JB: 2 ♂ 2 ♀, Jeonbuk Nat. Univ. , Jeonju-si, 21 VIII 2017, Jaeil Shim ; 3 ♀, National Institute of Agricultural Sciences , Iseo-myeon, Wanju-gun, 4 VII 2018, Jaeil Shim (reared from nymph) ; 2 ♂, Jangsu-eup , Jangsu-gun, 5 IX 2019, Jaeil Shim ; 2 ♂ 3 ♀, Mt. Moak , Gui-myeon, Wanju-gun, 7 IX 2019, Jaeil Shim ; 2 ♂, Jeonbuk Art Museum , Gui-myeon, Wanju-gun, 21 IX 2019, Jaeil Shim ; 4 ♂ 6 ♀, Jeonbuk Art Museum , Gui-myeon, Wanju-gun, 22 IX 2019, Jaeil Shim ; 2 ♂ 10 ♀, Iseo-myeon , Wanju-gun, 10 IX 2021, Jaeil Shim ;: 5 ♂ 3 ♀, Eunpa Lake , Gunsan-si, VI 2022, Jaeil Shim, JuHyeong Sohn (reared from nymph) ; 2 ♀, Byeonsan-myeon , Buan-gun, VIII 2022, Jeonbuk Jaeil Shim ; 1 ♂ 1 ♀, Hyangga-ro , Pungsan-myeon, Sunchang-gun, 26 VII 2023, Jaeil Shim (reared from nymph) ; JN: 1 ♂ 1 ♀, Near the Korea Coast Guard Academy (KCGA), Yeosu-si, 1 IX 2019, Byeongmin Jeong ; 1 ♂ 1 ♀, Dal-dong , Mokpo-si, VIII 2020, Jaeil Shim ; JJ: 1 ♀, Jeju Airport , Jeju-si, Jeju-do, 22 IX 2023, Jaeil Shim ; 1 ♂ 3 ♀, Donnaeko , Seogwipo-si, Jeju-do, 22 IX 2023, Jaeil Shim ; Guam: 1 ♀, Guam , USA, 19 VII 2017, Yeong-Hun Kim ; Japan: 1 ♂ 1 ♀, Yanabaru , Okinawa, Japan, 1–4 I 2020, Wonjun Sung ; Java: 1 ♂ 1 ♀, Mt. Argopuro , East Java, Indonesia, V 2019, Jaeil Shim (purchase) Vietnam: 1 ♂, Quang Trung , Bao Loc, Lam Dong, 20 II 2012, Lam Dong Agro-Foresty Research and Experiment Center .

Redescription.

Measurements (mm): Total length (vertex to tip of abdomen) ♂ 44.3–57.8, ♀ 54.3–74.2; head width ♂ 6.6–7.5, ♀ 8.4–9.7; head length ♂ 5.1–5.7, ♀ 7.3–8.1; pronotum width ♂ 5.1–6.0, ♀ 6.7–8.6; pronotum length ♂ 12.9–15.1, ♀ 16.1–20.2; forewing (tegmina) length ♂ 33.8–40.9, ♀ 36.8–47.5. Male (Figs 121 , 123 , 125–126 , 129 , 133–135 ). Medium to large sized, body robust. Coloration (Fig. 121 ): Bright green to green or bright brown to darkish brown. Head (Fig. 123 ): Triangular. Head width 1.3 × as long as length. Vertex flat. Compound eye globular, inverse drop-shaped. Ocelli large, oblong. Antenna filiform; length nearly 1.2 × as long as pronotum. Lower frons with two very weakly protruding parallel vertical ridges, lower frons width 1.5 × as long as height. Epistomal sulcus transverse. Lateral margin of compound eye, mandible, and gena yellow. In live specimens, labrum posterior margin orangish. Prothorax (Figs 125 , 126 ): Pronotum short, clavate, flatted dorso-ventrally; pronotum dorsal surface smooth and covered in waxy secretion; pronotum length 2.5 × maximum width. Prozone lateral margin with numerous denticles. Metazone lateral margin weakly expanded. Medial keel very weakly protruding. Furcasternite (Fig. 128 ) with two thick transverse and purple markings, larger one at furcasternite medial area and smaller one at posterior one-fourth of furcasternite. Forelegs (Prothoracic legs) (Figs 128–132 ): Coxa dorsal margin (Figs 128 , 129 ) with 2–5 large spines, rounded triangular or round tooth-like shape, yellow; occasionally 1–3 very minute, white, blunt spines located between large spines. Dorsal and ventral coxal lobes continuous, lacking space between them. Spination formula (Figs 130–132 ): Avts = 13–15; Pvts = 10–12; Avfs = 14–16; Pvfs = 4; Ds = 4. In 15 Avfs (Figs 130 , 131 ): spines 2, 4, 6, 8, 10, 12 and 15 size larger than remaining Avfs; spines 1, 2, 4, 6, 8, 10 and 12 black. Ds (Fig. 130 ) 1–3 interior surface black. Tarsomere distal end black. Meso- and metathorax and their legs: Mesothorax sternite anterior area with purple pattern. Meso- and metathoracic legs simple, long, and slender. Tarsi 5 - segmented. First tarsomere of midleg slightly shorter than remaining segments combined, first tarsomere of hindleg slightly longer than remaining segments combined. Wings (Figs 121 , 133 ): Forewing completely surpassing end of abdomen; costal area thick, discoidal area transparent; discoidal area occasionally with brightly colored (yellow, beige, brown) mottled pattern. Stigma (Fig. 133 ) triangular, white to whitish yellow; rimmed with dark pattern. Hindwing hyaline. Abdomen: Fusiform. Tergites bright yellow to green. Cerci setose, not flattened, with 17 or 18 segments. Male subgenital plate (coxosternite IX) (Fig. 134 ) irregularly rhomboidal; inter-stylar margin slightly convex and protruding dorsally, margin with 14–20 black spines; ventral surface of subgenital plate with numerous setae; left margin with 41–56 black spines, right margin with 16–28 black spines. Styli rather short. Male genitalia (Fig. 135 ): Pia sclerotized and weakly wrinkled; fda triangular; maa short and stout, surface covered by minute spines. Left phallomere (Fig. 135 ) with elongate and curved paa, its surface smooth, distal area curved dorso-laterally, apex round; afa (aafa + pafa) weakly sclerotized, wide trapezoidal, surface densely covered in minute denticles; anterior margin area of afa, basal one-third with stout dark decumbent projection (aafa); on dorsal surface of afa with long longitudinal groove; posterior apex mostly rounded; loa membranous, weakly humped; L 4 B curved spoon-shaped. Ventral phallomere (Fig. 135 ) irregular rhomboidal; sdpl hardly sclerotized, spear and hook-like in shape, curved at basal one-sixth by little more than 90 °, remaining distal area long and straight. Female (Figs 122 , 124 , 127 – 133 ). Similar to male, with following differences. Body (Fig. 122 ) robust. Head (Fig. 124 ): Vertex slightly convex. Head width 1.1 to 1.2 × as long as length. Ocelli smaller than male. Antenna slightly shorter than pronotum. Lower frons width 1.3–1.4 × as long as height. Prothorax (Fig. 127 ): Pronotum length 2.3–2.4 × as long as maximum width; lateral margin with numerous denticles. Medial keel protruding. Forelegs (Prothoracic legs) (Figs 128–132 ): Large forecoxal spines more pronounced than in male. Number of Avfs = 14–17. Wings (Figs 122 , 133 ): Forewing occasionally reaching end of abdomen. Ootheca (Fig. 136 ). Measurements (mm): Length 21.4–29.0; maximum width 11.2–15.6; maximum height 11.3–13.5; length of emergence area 15.1–21.3; width of emergence area 3.0–3.9. Identification: Oblong, nearly elliptical (mostly dorso-ventrally compressed) in cross-section. Ootheca attached to flat substrate by ventral surface or fully encircling substrate such as sticks. External wall (Fig. 136 ) dark green to brown. External coating comes off easily in the wild, colored beige. Exhibiting ~ 15– 27 egg chambers clearly delimited by visible slightly curved lips. Distal end of ootheca truncate and surface rough.

Biological notes.

Hierodula patellifera occurs throughout the Korean peninsula. This species is well-adapted to urban and suburban environments, and can be easily found near mountains, expressway rest areas, and parks. It typically lives under tree leaves throughout its life cycle and lays its oothecae on tree branches, trunks, and building walls near trees. Nymphs fold their abdomens back to camouflage themselves. First instar nymphs hatch in early June and adult mantises emerge in mid-August.

Distribution.

China, Guam, India, Japan, Java, New Guinea, Philippines, Sumba, Taiwan, Thailand, Vietnam, Korea. Invasive in France, Italy, and Hawaii.

Remarks.

Hierodula patellifera is a widely distributed species ( Ehrmann and Borer 2015; Patel and Singh 2016; Shcherbakov and Anisyutkin 2018; Battiston et al. 2019; Moulin 2020). The species exhibits a high degree of morphological variation, which has led to the recognition of numerous synonyms ( Audinet-Serville 1838; Wang 1993, Schwarz et al. 2018). Morphological variation, including differences in the number of forecoxal spines, and shapes of forewing stigma and afa, have been observed among H. patellifera populations, including those in Korea. Shim et al. (2021 b) discuss the challenges of species delimitation in H. patellifera in detail and emphasize the need for a comprehensive analysis of both morphological and molecular data to resolve taxonomic uncertainties.

Figures

Figures 121–132. Habitus, head, pronotum and foreleg of Hierodula patellifera 121 male dorsal aspect 122 female dorsal aspect 123 male face 124 female face 125 male pronotum (small sized) 126 male pronotum (large sized) 127 female pronotum 128 ventral aspect of foreleg and furcasternite (live specimens) 129 foreleg coxa ventral aspect (right: female with 4 forecoxal spines left: male with 2 forecoxal spines) 130 foreleg ventral aspect 131 foreleg interior aspect 132 foreleg dorsal aspect.

Figures 133–136. Wings, male genitalia and ootheca of Hierodula patellifera 133 stigma of forewing (3 variations) 134 male subgenital plate 135 male genitalia (small box = variation of afa) 136 ootheca (left: dorsal aspect right: lateral aspect). Red arrows = stigma.