Heptodonta pulchella ( Hope, 1831 )

Görn, Sebastian, 2020, Revision of the Oriental tiger beetle genus Heptodonta Hope, 1838 (Coleoptera: Cicindelidae), Zootaxa 4875 (1), pp. 1-62 : 48-53

Heptodonta pulchella ( Hope, 1831)

Figs. 6 , 116 , 124–143

Cicindela pulchella Hope, 1831: 21 .

Heptodonta pulchella . W. Horn, 1898b: 193.

Odontochila (“group” Heptodonta) pulchella . W. Horn, 1910: 204.

Odontochila (Heptodonta) pulchella . W. Horn, 1926: 123.

Syn. Heptodonta Hopei Parry, 1844: 454 .

Heptadonta Hopei. Schaum, 1862: 174 .

Cicindela Hopei. Gemminger & Harold, 1868: 18 .

Heptodonta pulchella syn. Hopei. W. Horn, 1898b: 193.

Odontochila (“group” Heptodonta) pulchella syn. Hopei. W. Horn, 1910: 204.

Heptodonta pulchella syn. hopei . Fowler, 1912: 312.

Odontochila (Heptodonta) pulchella syn. Hopei. W. Horn, 1926: 124.

Syn. Aenictomorpha variipes Chaudoir, 1850: 11 .

Cicindela varipes . Schaum, 1862: 174.

Heptodonta Hopei syn. variipes . Chaudoir, 1865: 20.

Cicindela Hopei syn. variipes . Gemminger & Harold, 1868: 18.

Heptadonta Hopei syn. variipes . Fleutiaux, 1892: 128.

Odontochila (“group” Heptodonta) pulchella syn. variipes . W. Horn, 1910: 204.

Heptodonta pulchella syn. variipes . Fowler, 1912: 312.

Odontochila (Heptodonta) pulchella syn. variipes . W. Horn, 1926: 124.

Syn. Cicindela (Heptodonta) yunnana Fairmaire, 1887: 88 .

Heptadonta Yunnana. Fleutiaux, 1892: 128 .

Heptodonta yunnana . Fleutiaux, 1893: 496.

Odontochila (“group” Heptodonta) pulchella syn. yunnana . W. Horn, 1910: 204.

Odontochila (Heptodonta) pulchella syn. yunnana . W. Horn, 1926: 124.

Heptodonta pulchella syn. yunnana . Heynes-Wood & Dover 1928: 51.

Syn. Heptadonta Ferrarii Gestro, 1893: 366 . syn. nov.

Heptodonta Ferrarii. Fleutiaux, 1893: 496 .

Heotodonta Hopei var. Ferrarii . W. Horn, 1902, 66.

Odontochila (“group” Heptodonta) pulchella syn. Ferrarii. W. Horn, 1910: 204.

Heptodonta pulchella syn. ferrarii . Fowler, 1912: 312.

Odontochila (Heptodonta) pulchella syn. Ferrarii. W. Horn, 1926: 124.

Heptodonta pulchella var. ferrarii . Heynes-Wood & Dover, 1928: 51.

Heptodonta ferrarii. Naviaux, 1991: 238 .

Syn. Heptodonta freudei Mandl, 1969b: 152 .

Heptodonta ferrarii ferrarii syn. freudei . Sawada & Wiesner 2002a: 77.

Syn. Heptodonta shooki Wiesner, 1986a: 107 . syn. nov.

Heptodonta ferrarii shooki . Naviuax, 1991: 240.

Type locality. Nepal .

Type material of H. pulchella . Lectotype (designated here): ♂ in BMNH, labelled: “SYN- / TYPE” [round with blue frame, printed] // “Type” [round with red frame, printed] // “nepal / 3923 / a.” [round, handwritten] // “pulchella. Hope / Java.” [handwritten] // “Hope’s mss det. label / stuck into 8° mss. catalogue / of the collection no. 3923 / C. M. F. von Hayek / det. 1972.” [handwritten/printed] // “ LECTOTYPE / Cicindela / pulchella / Hope, 1831 / design. Sebastian Görn 2020” [red, printed] // “ Heptodonta / pulchella / ( Hope, 1831) / det. Sebastian Görn 2020” [printed].

Type material of H. hopei . Lectotype (designated here): ♂ in BMNH, labelled: “TYPE” [with black frame, printed] // “Type” [round with red frame, printed] // “ Cicindela / Hopei Parry / India ” [handwritten] // “F. Bates Coll. / 1911-248” [printed] // “ LECTOTYPE / Heptodonta / Hopei / Parry, 1844 / design. Sebastian Görn 2020” [red, printed] // “ Heptodonta / pulchella / ( Hope, 1831) / det. Sebastian Görn 2020” [printed].

Type material of H. varipes . Lectotype (designated here): ♂ in MNB, labelled: “42453” [printed] // “Hist.- Coll. ( Coleoptera ) / Nr. 42453 / Cicindela varipes Chaud * / India bor., Coll. Schaum / Zool. Mus. Berlin” [yellow with black frame, printed] // “varipes / Chd * / Ind. bor.” [handwritten] // “ SYNTYPE / Aenictomorpha / variipes Chaudoir, 1850 / labelled by MFNB 2018” [red, printed] // “ LECTOTYPE / Aenictomorpha / variipes / Chaudoir, 1850 / design. Sebastian Görn 2020” [red, printed] // “ Heptodonta / pulchella / ( Hope, 1831) / det. Sebastian Görn 2020” [printed].

Type material of H. yunnana . Lectotype (designated here): ♀ in MNHN, labelled: “pulchella / Hope” [hand- written] // “ Yunnan / R.P.Delavay ” [with black frame, printed] // “ Heptodonta / Yunnane / Fairm.” [handwrit- ten] // “ExMusaeo / ARM.DAVID / 1900” [with black frame, printed] // “MUSÉUM PARIS / 1952 / COLL R. OBERTHÜR” [yellow with black frame, printed].

Type material of H. ferrarii . Lectotype (designated here): ♂ in SDEI, labelled: “Carin Chebà / 900–1100 m. / L. Fea V XII-88” [with black frame, printed] // “Type! / coll. W Horn” [with black frame, printed] // “ Syntypus ” [red, printed] // “Coll. W. Horn / DEI Eberswalde” [printed] // “f. / Ferrarii / Gestro” [yellow with black frame, handwritten] // “SDEI Coleoptera / # 301571” [printed]. Paralectotypes: 1 ♀ in SDEI: “Carin Chebà / 400– 900 m. / L. Fea IV. 88” [with black frame, printed/handwritten] // “Type! / coll. W Horn” [with black frame, printed] // “ Syntypus ” [red, printed] // “SDEI Coleoptera / # 301572” [printed]. 1 ♀ in MNB: “Carin Chebà / 900–1100 m. / L. Fea V XII-88” [with black frame, printed]. All type specimens labelled: “ LECTOTYPE (or PARALECTOTYPE) / Heptadonta / Ferrarii / Gestro, 1893 / design. Sebastian Görn 2020” [red, printed] // “ Heptodonta / pulchella / ( Hope, 1831) / det. Sebastian Görn 2020” [printed].

Type material of H. freudei . Holotype: ♀ in ZSM, labelled: “ Laos / Umg. Vientiane / III.–VI.1963 ” [printed] . Paratype: 1 ♀ in ZSM, with same label as holotype. All type specimens labelled: // “HOLO-Typus (or PARA-Ty- pus respectively) / Heptodonta / freudei m. / 1966 / det. Dr.K.Mandl ” [red, printed/handwritten] // “ Zool. Staatsslg. / München ” [blue, printed] // “ Heptodonta / pulchella / ( Hope, 1831) / det. Sebastian Görn 2020” [printed] .

Type material of H. shooki . Holotype: ♂ in JWCW, labelled: “ THAILAND, Nakhon Ratchasima, / Khao Yai National Park / 11–13 June 1983 / Coll. G.A. Shook ” [with yellow frame, printed] . Paratypes: 1 ♂ 1 ♀ in JWCW, 1 ♀ in ZSM, with same labels as holotype. All type specimens labelled: “ HOLOTYPUS (or PARATYPUS respec- tively) / HEPTODONTA / shooki / Wiesner 1985” [red, printed/handwritten] // “ Heptodonta / pulchella / ( Hope, 1831) / det. Sebastian Görn 2020” [printed] .

Diagnosis. This species is extremely variable. The only distinct character of H. pulchella is the typical hookshaped anteromedian sclerite of the aedeagus (best seen in right lateral aspect, Figs. 130 , 134–137 ). Usually specimens from the western population ( India, Nepal) are large and green ( Figs. 124–125 , 131 , 139–140 ), individuals from China large to medium sized and copper-green ( Figs. 132–133 , 143 ), and specimens from Myanmar, Yunnan, and Thailand medium to small sized and black or green ( Figs. 138, 141–142 ). Western specimens are similar to H. schuelei sp. nov. and H. tempesta sp. nov., but possess neither the slender, elongated elytral apex of H. schuelei sp. nov. nor the coarse zigzag pattern on the vertex of H. tempesta sp. nov. From H. posticalis it is distinguishable by the uniform silky shine of the elytra owing to its pronounced regular elytral microsculpture ( Fig. 116 ), and the usually bronze-green, not violet, mesepisternum.

Redescription. Body size: Length 11.4–16.6 mm (without labrum), width 3.2–5.0 mm ( Figs. 124–125 , 131– 133 , 138–143 ).

Dorsal surface of head metallic green and bronze, copper, or black with some bluish reflections. Frons transversely to irregularly grooved. Vertex with coarse, irregular longitudinal striation or almost completely irregular grooves, in transition to orbital plates with converging irregular striation ( Fig. 6 ). Orbital plates with fine irregular striation and two setae on each side. Occiput with wrinkled transverse grooves, anteromedian irregularly rugose. Genae glabrous (except for three specimens from Fujian with one or two setae on each side), rather shallowly or conspicuously grooved, colouration variable, often metallic yellowish-bronze and green, copper, bluish-green and violet, or blackish with bronze-blue lustre. Clypeus glabrous, colouration variable, frequently metallic bronze and green with two anterior blue spots, copper, or blackish with some bluish-green and bronze lustre. Labrum testaceous, with four setae, five apical teeth, and one lateral tooth on each side, three median teeth acute, third apical teeth reduced, triangular shape, male labrum shorter (0.78–1.35 mm long, 1.35–2.10 mm wide, Fig. 126 ) than elongated female labrum (1.10–1.80 mm long, 1.48–2.20 mm wide, Fig. 127 ). Mandibles testaceous, teeth black to rufous-brown, basal inner margin frequently with dark-brown spot. Labial and maxillary palpi testaceous, terminal palpomeres distally black in males, and almost entirely black in females, some females with dark-testaceous preterminal maxillary palpomeres or just distally black terminal palpomeres. Antennae slender, extending back over the first third of the elytra, metallic black, scape and pedicel frequently with iridescent green lustre, antennomeres 3–4 distally often with golden-violet or golden-green shine, pedicel testaceous basally, scape with a single apical seta, antennomeres 3–4 with few scattered setae, antennomeres 5–11 finely and evenly pubescent, in males terminal antennomere slightly dilated.

Thorax entirely glabrous. Pronotum with variable colouration, metallic bronze-green, bronze and bluish-green, copper, reddish, or blackish with bluish-green and violet-bronze lustre, moderate median line blue, approximately as long (2.18–3.30 mm) as wide (2.05–3.20 mm), but more elongated in smaller males, irregular transverse grooves wrinkled on disc, pronounced anterior and posterior sulci, anterior lobe wider than posterior, convex lateral margins of the median lobe slightly converging to the base. Episterna, epimera, and sterna metallic bronze and green, copper, reddish, bluish-green and yellowish-copper, or yellowish-blue, frequently mesepisternum anterior with bluish aspect and proepimeron with iridescent copper colouration.

Elytra elongate to extremely elongate, length 6.7–10.5 mm, slightly dilated laterally to almost parallel, maximal width at or slightly anterior to the middle, colouration mostly green with bronze lustre, but also copper, reddish, bluish-green with bronze shine, or blackish with gently bronze or bluish lustre, rather dull shine due to pronounced microsculpture ( Fig. 116 ), lateral margin violet with bluish-green transition; apical impression and posterior gib- bosity slightly pronounced, juxtahumeral impression, discal impression and basodiscal convexity moderately to slightly marked, violet-blue punctures slightly deeper anterior, punctures forming irregular transverse lines that are sometimes fused to short transverse grooves.

Coxae testaceous and dark-testaceous to black with green, reddish, bronze, bluish-green, or violet lustre, procoxae anterior, mesocoxae anteriorly and posteriorly, and metacoxae laterally densely covered with long white setae. Trochanters, femora, and tibiae testaceous. Femoral distal apices with black tip, followed always by a testaceous ring in profemora, mostly in mesofemora, and sometimes in metafemora, and a blackish (often green or reddish shining) area extended posteriorly and covering up to entire profemora, two-thirds of mesofemora, and half of metafemora. Tarsi and one to three-quarters of distal tibiae metallic black. Proximal tarsomere regions variably dark-testaceous, from terminus only to all tarsomeres. Claws testaceous, rufous-testaceous, or dark-brown. Some specimens lacking any dark femoral and tibial colouration besides distal black tip.

Abdominal sternites glabrous (apart from few long setae at posterior margins), with iridescent green to violet lustre, margin of or entire terminal sternite dark-brown to dark-testaceous ( Fig. 128 ).

Aedeagus shape slender but variable, arched or angled, 2.70–4.45 mm, ventrally concave to almost straight, sometimes with marginal medioventral convexity, acute aedeagal tip, dorsoapical usually moderate ascent, but also specimens with short steep or flat elongated dorsoapical slope, dorsal anteromedian rather straight to slightly convex, usually posterodorsal even curvature, aedeagal base rather short and moderate to steeply sloping. Inner sac in right lateral aspect with short diagonal testaceous to dark-testaceous spine row, dark-testaceous anteromedian sclerite posterior wide and anterior ending in one elongate diagonal spine ( Figs. 129–130 , 134–137 ).

Variability. The large variability of H. pulchella led to the description of six synonyms and the misinterpretation of reddish medium-sized specimens as H. vermifera ( Fig. 132 ). Actually, there occur three local variations in the northwestern, northeastern, and southern part of the distribution area: in Nepal and northern India specimens are usually large and green ( Figs. 124–125 , 131 , 139–140 ), individuals from China large to medium-sized and frequently copper-green ( Figs. 132–133 , 143 ), and specimens between Myanmar, Yunnan, and Thailand medium to small-sized and black or green ( Figs. 138, 141–142 ). Nevertheless, all possess the typical hook-shaped anteromedian sclerite of the aedeagus (best seen in right lateral aspect, Figs. 130 , 134–137 ). Also, the aedeagal shape differs considerably in H. pulchella , but rather within than between the local groups ( Figs. 134–137 ). However, on average aedeagi of northwestern males are considerably larger compared to northeastern and especially southern males, due to the overall larger body size. Further, assumed locally variable group characters also occur occasionally in other groups: e.g. large robust specimens are not only recorded from Nepal and India, but also from Fujian (eastern China); small green specimens typical for southern Thailand can also be found in Khammouane (central Laos); and the almost parallel elytral shape, characteristic for southern specimens, appears also in individuals from Darjiling (northeastern India).

Distribution. Southern CHINA (Fujian, Guangdong, Guangxi, S Sichuan, SE Tibet, Yunnan), northeastern and northern INDIA (? Meghalaya,? Sikkim, Himachal Pradesh, Uttarakhand, N West Bengal), entire LAOS, centraleastern MYANMAR (Karen Hills, Shan State), eastern and central NEPAL, entire THAILAND, entire VIET- NAM.

Additional material examined. 294 specimens. CHINA: SW Yunnan, Jinggu county, Mt. Longtanshan , 1.– 20.VI.2009 ( JWCW) ; NW Yunnan, Mt. Gaoligongshan , Sedang, Gongshan County, 1.–10.VI.2008 ( JWCW) ; SE Yunnan, Wenshan , Qiu Bei, Xin Guo, 1350m, 16.VI.2005 ( JWCW) ; W Yunnan, Tongbiguan , Yingjiang, VIII.2003 ( JWCW) ; SW Yunnan, 300 km SW Kunkin, Lintsan city env., 2200m, 15.VI.2000 ( JWCW) ; S Yunnan, Xishangbana, VII.1994 ( JWCW) ; C Yunnan, 100 km W Kunming, Diaolin Nat. Reservation , 22.V.–2.VI.1993 ( JWCW, ZSM) ; SE Tibet, Zayü County, Xia-Cha-Yu, 1900–2600 m, 1.–28.VII.2004, 22.–28.VI.2006 ( JWCW) ; SW Guangdong, Baishui Pubu , 21°53’N 111°23’E, 500–750m, 5.VI.2015 ( JWCW) ; SE Guangdong, Lianhuashan , 23°02’N 115°15’E, 250–570m, 3.VI.2014 ( JWCW) ; W Guangxi, Leitashan , 23°44–45’N 106°16–17’E, 800–1350m, 31.V.–1.VI.2013 ( JWCW) ; S Fujian, Longyan , forest park, 24°57’N 117°01’E, 680m, 28.VI.2007 ( JWCW) ; S Sichuan, Xichang env., 1700m, 18.VII.2001 ( JWCW) ; S Sichuan, S Xichang, Lushan Mt. , 1800m, 25.VII.1998 ( JWCW) ; S Sichuan, S Xichang, Lushan Mt. , 1600–1800m, 25.–28.VII.1996 ( JWCW) ; Yunnan, Vallis flumin, Soling-ho ( SNMF, MNB) ; Sichuan, Mts. Yunling ( MNB) ; Yunnan ( MNB, SDEI); S Sichuan, Xichang , 1900 m, VII.1996 ( ZSM) ; NW Yunnan, Haba mts. NE slope, 2800–3000m, 18.VI.1998 ( ZSM) ; Yunnan, Lou-Nan, 1931 ( SDEI) ; INDIA: West Bengal, env. Darjiling ( ZSM) ; Uttarakhand, Mussoorie , Kampty-Falls, 1500m, 8.VII.1989 ( ZSM) ; Uttarakhand, Mussoorie V. P., VII.1989 ( ZSM) ; West Bengal, Mungpoo ( SDEI) ; Sickim ( SDEI) ; N. Ind., Ianson ( SDEI) ; LAOS: Khammouane, Nakai env., 17°43’N 105°09’E, 500–600m, 22.V.–8.VI.2001 ( JWCW) ; Luang Namtha, 20 km NW Luang Namtha city, 21°09.2N 101°18.7E, 900± 100m, 24.–30.V.1997 ( JWCW) ; Khammouane, Nakai env., Route No. 8, N 17°42.8’ E 105°09.1’, 560m, 4.–8.V.1998 ( JWCW) ; Khammouane, Nakai env., Route No. 8, N 17°42.8’ E 105°08.9’, 560± 20m, 4.–8.V.1998 ( JWCW) ; Xiangkhouang, 17.V.1994 ( JWCW) ; Oudomxay, 17km NEE Oudomxay city, 20°45’N 102°09’E, 1100m, 1.–9.V.2002 ( JWCW) ; Phôngsali, Ban Bun Neua (4 km E), 21°38’N 101°57’E, 1100m, 20.VI.2003 ( JWCW) ; Khammouane, Ban Khoun Ngeun env., 18°07’N 104°29’E, 250m, 10.–13.VI.2001 ( JWCW) ; Bolikhamsai, Ban Nok env., Route No. 8, N18°08.7 E104°28.1, 220± 50m, 9.–14.V.1998 ( JWCW) ; Phôngsali, Phôngsali city env., 21°41–2’N 102°06–8’E, 1500m, 28.V.–20.VI.2003 ( JWCW) ; Khammouane, Ban Khoun Ngeun , 18°07’N 104°29’E, 200m, 19.–31.V.2001 ( JWCW) ; Houaphanh, 25km SE Vieng Xai (by road), Ban Kangpabong env., 20°19’N 104°25’E, 14.–18.V.2001 ( SMNS) ; Houaphanh, 38km (road) S of Sam Neua, Ban Saleui, 1350–1850m, 9.–22.V.2009 ( PSCH) ; Oudomxay, Ban Kheun , 6.–15.VI.2014 ( PSCH) ; Houaphanh, Mt. Pan , 1400m, VI.2013 ( JWCW, SGCH) ; Luang Namtha, Namtha-> Muang Sing , 21°09’N 101°19’E, 900–1200m, 5.–31.V.1997 ( ZSM) ; near Vientiane, 1995 ( ZSM) ; Xayabuli , env. Paklay, 1963 ( ZSM) ; Attapeu, Annam Highland Mts., Dong Amphan NBC-Area, Nong Fa (crater Lake) env., 15°05.9’N 107°25.6’E, 1160 m, 2.V.2010 ( CKCE) ; MYANMAR: Shan Highland , Monghkok, 1.–22.V.2005 ( JWCW) ; Shan State, env. Lashio, Namsham, 1500m, VI.2000 ( JWCW) ; SE-Burma, 1.VII.1990 ( JWCW) ; Burmese border, Tralea Huika, V.1981 ( ZSM) ; North Shan States , IV.–V.1914 ( SDEI) ; NEPAL: Bagmati, Sollenta-Dhading , 1000–1300m, 21.–22.VI.1993 ( JWCW, ZSM) ; Bagmati, Bhota- Koahi-Khola , Lamosangu, 700m, 28.VI.1987 ( JWCW) ; Janakpur, Jriri-Shivalaya , Khimti-Khola, 2500– 1800m, 11.VI.1987 ( JWCW) ; Arun V., Num, 1550m, 5.–6.VI.1983 ( JWCW) ; Arunthan-Chichila , 1300–1950m, 23.V.1980 ( JWCW) ; Dolakha, lower Khara Khola , 1200– 1050m, 5.VI.2000 ( SMNS) ; Bhojpur, Gothe to Majwa, 600–800m, 27.V.1997 ( SMNS) ; Dhading, W Samari, Banjyang / Topal Khola, agricultural land, 1000–1200m, 23.VII.1983 ( SNMF) ; Rasuwa, Trisuli Bazaar , 11.–14.VI.1999 ( SGCH) ; Lamabagar , 1000–1550m, 4.VI.1980 ( ZSM) ; Kaski , Gandaki, Chandrakot-Hille , 1000–1600m, 9.VI.1986 ( ZSM) ; Bothe-Kosi canyon, 14.VI.1964 ( ZSM) ; Kathmandu , 1400 m, 8.VIII.1968 ( ZSM) ; Likhu Khola valley , 1700m, 1.VIII.1962 ( ZSM) ; THAILAND: Kanchanaburi, Pilok, 24.V.1988 ( JWCW, ZSM) ; Mukdahan, Phu Phan, 1989 ( JWCW) ; Chiang Mai, Samoeng, 20.IV.1991 ( JWCW) ; Chiang Rai, Wiang Pa Pao, 15.V.1991 ( JWCW) ; Nakhon Ratchasima, Khao Yai, 9.V.1986 ( JWCW) ; Nakhon Ratchasima, Corat, 20.V.1990 ( JWCW) ; Chumphon, 5 km S Phato , 155m, lowland rainforest, NF, N 09°46.075’ E 98°46.992’, 06.VII.2008 ( PSCH) ; Mae Hong Son, Ban Huai Po, 800–1600m, 1-15.1991 ( ZSM) ; Mae Hong Son, Ban Huai Po, 1600–2000m, 9.–16.V.1991 ( ZSM) ; Phetchabun, Khao Kho, 1000m, V.2001 ( ZSM) ; Surat Thani, Khao Sok NP, 2.–8.V.1995 ( ZSM) ; Nakhon Ratchasima, Corat, 5.V.1991 ( ZSM) ; VIETNAM: Vĩnh Phúc, Tam Dao, 27.V.–2.VI.1986 ( JWCW, ZSM) ; Bắc Giang, Tay Yen Tu Nat. Res. , 21°11’10’’N 106°43’25’’E, 7.– 11.VII.2013 ( JWCW) ; Bắc Giang, Tay Yen Tu Nat. Res. , Van Danh (near Dong Ri), N21°09.965’ E106°49.557’, 415m, 20.V.2015 ( JWCW) ; Bắc Giang, Tay Yen Tu Nat. Res. , 6 km SW Thanh So’n, N21°10.830’ E106°43.427’, 200m, 18.–21.V.2015 ( JWCW) ; Vĩnh Phúc, Tam Dao, Tam Dao Nat. Park , 1000m, 2.VII.2011 ( PSCH) ; Phú Thọ, Xuan Son NP, 200–300m, 15.6.2004 ( SGCH) ; Vĩnh Phúc, Tam Dao NP, 22.V.2008 ( SGCH) ; Lạng Sõn, Mount M ẫu Sõn, April–Mai 2-3000‘ ( MNB, SDEI) ; Vĩnh Phúc, Tam Dao, 26.V.–5.VI.1986 ( ZSM) ; Lào Cai, Sa Pa, 1560m, VI.1990 ( ZSM) ; Vĩnh Phúc, 70 km NW Hanoi , Tam Dao, 21°27’N 105°39’E, 900–1200m, 3.–7.VI.1996 ( ZSM) ; Lâm Đ ồng, Bao Loc, V.–VI.1994 ( ZSM) ; Ngh ệ An, Vinh, 1910 ( SDEI);

Biology. Occurs on steep slopes, roads in forests, road cuts, bare soil in open areas, old gravel pits, and excavation sites ( Wiesner 1986a; Probst 1996; Shook & Wu 2007). Recorded from lowland rainforest in southern Thailand (155 m) up to elevations of 2900 m in Yunnan.

Remarks. Most widely distributed species of the genus Heptodonta ( Hope 1831; Parry 1844; Chaudoir 1850; Fairmaire 1887; Fleutiaux 1892, 1893; Gestro 1893; Horn 1910, 1926; Fowler 1912; Heynes-Wood & Dover 1928; Mandl 1969b; Wiesner 1986a, 1992, 2006; Naviaux 1991; Probst 1996; Sawada & Wiesner 1999b, 1999c, 1999d, 2000b, 2000c, 2001, 2002a, 2004, 2006c; Cassola & Klícha 2002; Naviaux & Pinratana 2004; Shook & Wiesner 2006; Shook & Wu 2007; Uniyal & Bhargav 2007; Matalin 2015; Wiesner & Geiser 2016; Putchkov & Matalin 2017; Wiesner et al. 2017). Specimens recorded from Balphakram National Park in Meghalaya ( Sawada & Wiesner 1997) refer to H. tempesta sp. nov., and records from Cherrapunjee in Meghalaya and Jamiri in Arunachal Pradesh ( Sawada & Wiesner 2006a) relate to H. schuelei sp. nov. Records from Mizoram in northeastern India ( Harit 2013) may belong to H. schuelei sp. nov. The record from Sikkim needs to be verified as the old record probably refers to the area around Darjeeling, which nowadays belongs to the state of West Bengal. Parry (1845) remarked in his second description of H. hopei that this species was collected in the Khasi Hills. However, this exceptionally large specimen is the only individual of H. pulchella recorded in the distribution area of H. schuelei sp. nov., between Darjeeling and Yunnan; this old record needs to be confirmed.

The distribution gap between large specimens from Nepal or India and medium-sized individuals from Myanmar or Yunnan indicate that there might be some ongoing speciation between these variation groups.

Figures

FIGURES 1–7. 1–2—anterior elytra: 1—Heptodonta melanopyga (Schaum, 1862), ♀, Philippines, Luzon, Banaue (PSCH); 2—H. mindoroensis Cassola, 2000, ♂, Philippines, Mindoro, Mt. Halcon (JWCW). 3–4—pronotum: 3—H. arrowi W. Horn, 1900, ♂, Myanmar, Magway Region, Pyon Village (SGCH); 4—Heptodonta horii sp. nov., ♂, holotype, Myanmar, Bago Re- gion, Chanthakwin (JWCW); 5–7—head: 5—Heptodonta tempesta sp. nov., ♂, holotype, India, Meghalaya, Balpakram NP (JWCW); 6—Heptodonta pulchella (Hope, 1831), ♂, Nepal, Bagmati, Sollenta-Dhading (JWCW); 7—Heptodonta schuelei sp. nov., ♂, holotype, India, Meghalaya, Cherrapunjee (JWCW). Scales: 1 mm.

FIGURES 113–116. 113–115—Heptodonta posticalis (White, 1844): 113—habitus, ♂, lectotype, China, Hong Kong (BMNH); 114–115—♀, China, Guangdong, Qimuzhang (JWCW): 114—mesepisternum, 115—detail elytra. 116—detail elytra, H. pulchella (Hope, 1831), ♀, Nepal, Bagmati, Sollenta-Dhading (JWCW). Scales: 113–114 = 1 mm, 115–116 = 0.5 mm.

FIGURES 124–130. Heptodonta pulchella (Hope, 1831), Nepal, Bagmati, Sollenta-Dhading (JWCW): 124, 126, 128–130—♂: 124—habitus, 126—head, 128—lateral body, 129—aedeagus in left lateral aspect, 130—aedeagus in right lateral aspect; 125, 127—♀, paratype: 125—habitus, 127—head. Scales: 124–128 = 1 mm, 129–130 = 0.5 mm.

FIGURES 131–137. Heptodonta pulchella (Hope, 1831): 131, 134—♂, lectotype, Nepal (BMNH): 131—habitus, 134—ae- deagus in right lateral aspect; 132, 135—♂, China, Sichuan, Xichang (JWCW): 132—habitus, 135—aedeagus in right lateral aspect; 133, 136—♂, China, Fujian, Longyan (JWCW): 133—habitus, 136—aedeagus in right lateral aspect; 137—aedeagus in right lateral aspect, ♂, lectotype of syn. H. ferrarii Gestro, 1893, Myanmar, Carin Cheba [Karen Hills] (SDEI). Scales: 131–133 = 1 mm, 134–137 = 0.5 mm.

FIGURES 138–143. habitus, Heptodonta pulchella (Hope, 1831): 138—♂, lectotype of syn. H. ferrarii Gestro, 1893, Myanmar, Carin Cheba [Karen Hills] (SDEI); 139—♂, lectotype of syn. H. variipes (Chaudoir, 1850), India bor. (MNB); 140—♂, lectotype of syn. H. hopei Parry, 1844, India (BMNH); 141—♀, holotype of syn. H. freudei Mandl, 1969b, Laos, Vientiane (ZSM); 142—♂, holotype of syn. H. shooki Wiesner, 1986a, Thailand, Nakhon Ratchasima, Khao Yai NP (JWCW); 143—♀, lectotype of syn. H. yunnana (Fairmaire, 1887), China, Yunnan (MNHN). Scales: 1 mm.