Heptodonta analis ( Fabricius, 1801 )

Görn, Sebastian, 2020, Revision of the Oriental tiger beetle genus Heptodonta Hope, 1838 (Coleoptera: Cicindelidae), Zootaxa 4875 (1), pp. 1-62 : 8-12

Heptodonta analis ( Fabricius, 1801)

Figs. 8–22

Cicindela analis Fabricius, 1801: 236 .

Heptodonta Analis. Hope, 1838: 11 , 25.

Heptadonta analis . Lacordaire, 1842: 27.

Euryoda analis . Lacordaire, 1842: 27.

Aenictomorpha analis . Chaudoir, 1850: 11.

Heptodonta analis. W. Horn, 1892: 94 .

Odontochila (“group” Heptodonta) analis . W. Horn, 1910: 203.

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

Heptodonta analis analis . Cabras et al. 2016: 193.

Type locality. Sumatra .

Type material. Lectotype (designated here): ♂ in ZMK, labelled: “analis” [handwritten] // “ LECTOTYPE / Cicindela / analis / Fabricius / by R. E. Acciavatti, ‘86” [red, printed/handwritten]. Paralectotypes: 1 ♂ in ZMK: “ Sumatra / Daldorff / Mus. Sig. P. L. / Cicindela / analis. F.” [handwritten]. 1 ♂ 1 ♀ in MNB: “243” [printed] // “Hist.-Coll. ( Coleoptera ) / Nr. 243 / Euryoda analis Fab. * / Sumatra, Daldorf / Zool. Mus. Berlin ” [yellow with black frame, printed] // “analis / F. Dej. / Sumatra Dald.” [yellow with black frame, handwritten] // “ SYNTYPE / Cicindela / analis / Fabricius, 1801 / labelled by MFNB 2018” [red, printed]. All type specimens labelled: “LECTO- TYPE (or PARALECTOTYPE respectively) / Cicindela / analis / Fabricius, 1801 / design. Sebastian Görn 2020” [red, printed] // “ Heptodonta / analis / ( Fabricius, 1801) / det. Sebastian Görn 2020” [printed].

Diagnosis. Distinguished by the mediolateral and anteroapical elytral impressions, in combination with the coarsely grooved pronotum, testaceous proximoapical protibiae, and testaceous apical sternites. Typical cleavershaped aedeagus.

Redescription. Body size: Length 10.8–14.3 mm (without labrum), width 3.2–4.4 mm ( Figs. 8–9 , 15–16 ).

Head with dorsal surface green and/or bronze, temples blue. Frons transversely to irregularly grooved, crumpled wrinkles in transition to vertex. Vertex median with irregular longitudinal striation, that converging in transition to the orbital plates. Orbital plates with fine, irregular to parallel converging striae, and with two setae on each side. Occiput with irregular transverse to completely irregular grooves. Genae glabrous and shallowly grooved, violetblue to yellowish-green, often with bronze anterior area. Clypeus glabrous, bronze-green. Labrum testaceous, with four setae, five apical teeth, and one lateral tooth on each side, shape variable, male labrum short (0.63–0.85 mm long, 1.30–1.70 mm wide, Fig. 10 ), median tooth often extended, female labrum with three protruding median teeth (0.78–1.18 mm long, 1.43–1.70 mm wide, Fig. 11 ). Mandibles testaceous, teeth dark rufous-brown to black. Labial and maxillary palpi testaceous, terminal palpomeres distally black in males, and almost entirely black in females. Antennae slender, extending back over the first third of the elytra, dark-brown, scape, pedicel, and apices of antennomeres 3–4 dorsal with faint green-violet lustre, pedicel testaceous at basal tip, scape with a single apical seta, antennomeres 3–4 with few scattered setae, antennomeres 5–11 finely and evenly pubescent.

Thorax entirely glabrous. Pronotum iridescent green and/or bronze, median line moderately pronounced with blue margin, proportions variable, usually slightly longer (2.08–2.95 mm) than wide (2.03–2.83 mm), transverse grooves coarse and irregular, pronounced anterior and posterior sulci, anterior lobe wider than posterior, lateral margins of the median lobe moderately converging to the base. Episterna iridescent bronze-green to blue. Sterna and epimera iridescent bronze-green to bluish-green, posterior mesosternum, lateroposterior metasternum, proepimeron, and mesoepimeron usually iridescent bronze.

Elytra elongate, length 6.5–8.9 mm, slightly dilated laterally, maximal width posterior, colouration green with yellow-bronze lustre to almost entirely bronze, lateral margin and apex violet to bluish-green, juxtahumeral, discal, mediolateral, anteroapical, and apical impression considerably pronounced, posterior gibbosity and basodiscal convexity significant, deep blue punctures slightly decreasing in size from base to apex, punctures irregularly fused to transverse lines, converging to apex on anteroapical impression, basal and apical irregularly punctured.

Coxae iridescent bronze-green to bluish-green, pro- and mesocoxae anteriorly and metacoxae laterally densely covered with long white setae, mesocoxae posteriorly with few white setae. Trochanters and femora testaceous. Femoral distal apices, tibiae, and tarsi metallic black, often with some gently violet lustre. Tibiae proximoapically testaceous or at least slightly lightened. Claws rufous-testaceous to black.

Abdominal sternites glabrous except for a few long setae at posterior margins, some scattered minute setae on the last four sternites of males, and some single minute setae on sternites 4–5 of females, first four sternites darkbrown with bronze, green, blue, and violet reflections, last two sternites and some lateral margin aspects of sternites 3–4 testaceous ( Fig. 12 ).

Aedeagus sigmoid-trapezoidal, length 2.45–2.80 mm, aedeagal base sigmoid, basoventrally convex, round apical tip slightly directed ventrally, dorsal apex straight diagonal, with dorsobasal concavity and dorsomedian convexity (usually a straight slope in dry material). Inner sac in left lateral aspect basally and apically with broad scleriteplates, apical plate sigmoid, medioventrally with diagonal arciform sclerite, and median with vertical sclerite ( Figs. 13–14 , 17–22 ).

Variability. Due to its fragmented distribution over several islands of the Malay Archipelago, speciation seems to be ongoing in H. analis . While specimens from the Malay Peninsula and Sumatra are usually coloured iridescent green, bluish-green, or bronze-green, specimens from Java and especially Borneo are for the most part characterized by a dull bronze-green to mainly bronze colouration. The population of the Bako National Park on Borneo is unique, in which many individuals are entirely blue ( Fig. 16 ). However, other individuals from this locality share the characters of other localities.

Distribution. Western INDONESIA (Sumatra, Borneo, Java), entire MALAYSIA, most southern THAI- LAND (Yala), BRUNEI, SINGAPORE.

Additional material examined. 312 specimens. INDONESIA : Sumatra, Sereda Radjah, 19.I.1980 ( JWCW) ; Sumatra, Dolok Merangir, 13.XI.1978 ( JWCW) ; N Sumatra, Huta Padang, 300m, 30km SE P. Siantar, 20.I.1990 ( JWCW) ; N Sumatra, 40km W Sidikalang , 8.II.1981 ( JWCW) ; N Sumatra, Aceh, 27.V.1979 ( JWCW) ; West Sumatra, Kerinci, Seblat NP, 24km NE Tapan, Muara Sako env., 2°05’S 101°15’E, 400–550m, 4.–18.III.2003 ( JWCW) ; Sumatra Barat, Harau Valley , 500–800m, IX.2008 ( JWCW) ; W Sumatra, Lubuk Sikaping, 28.X.1991 ( JWCW) ; W Sumatra, Harau, VIII.1992 ( JWCW) ; Sumatra, Jambi, Bukli Tigapuluh NP, 140m, 01°09’S 102°33’E, 10.– 17.IX.2003 ( JWCW) ; Java, 1872 ( SMNS) ; Java Barat, Sukabumi, 1893 ( SNMF, MNB, SDEI) ; N Sumatra, Dolok, Merangir ( SNMF) ; Sumatra Barat, Merapi env., XII.1995 ( PSCH) ; Sumatra Utara, Medan ( MNB, ZSM) ; Java Barat, Mount Gede ( MNB) ; Java 1895, 1898 ( MNB); Borneo , Kalimantan Barat, Mt. Bawang , X.1989, IV.1991, VI–VII.1991 ( ZSM) ; Borneo, Kalimantan Barat, Mt. Bawang , 250–300m, VII–VIII.1990, X.1990 ( ZSM) ; Borneo, Kalimantan Barat, Bengkayang, X.1993 ( ZSM) ; Sumatra, Riau, Akar river , 300m, IV.1999 ( ZSM) ; Sumatra Utara, Bukit Lawang, X.1990 ( ZSM) ; Java ( ZSM) ; Sumatra ( SDEI) ; Sumatra Utara, Nias Is., Lahago, 4.II.–10.III.1896 ( SDEI) ; Java Barat, Sukabumi, 1894 ( SDEI) ; W Java, Palaboehan, Ratoe, Wynkoopsbaai ( SDEI) ; MALAYSIA: Borneo, Sabah, Mt. Kinabalu , ca. 1500m, Headquarters, 25.VIII.1990 ( JWCW) ; Borneo, Sabah, Bukit Monkobo, 700m, 51°48’N, 116°58’E, 7.VIII.1987 ( JWCW) ; Borneo, Sarawak, Kapit , Rajang, Pelagus, 18.VIII.1994 ( JWCW) ; Borneo, Sarawak, Bako NP, 1987 ( JWCW) ; Borneo, Sarawak, Kuching, Bako, 7.X.1994 ( JWCW) ; Pahang, 30km NE Raub, Lata Lembik , 3°56’N, 101°38’E, 200–400m, 22.IV.–1.V., 8.–15.V.2002 ( JWCW) ; Kelantan, 90km N Gua Musang, Gunung Basor , 1700m, Kampong Kubur Datu, 10.IV.–5.V.2016 ( JWCW) ; Pahang, Cameron Highlands, Tanah Rata, edge of degraded rainforest at light, 21.III.–2.IV.1995 ( JWCW) ; Pahang, Cameron Highlands, Gunung Jasar, montane rainforest, 23.III.1995 ( JWCW) ; Pahang, Cameron Highlands, 1500m, 4°29’01N / 101°22’95E, 25.III.2006 ( JWCW) ; Johor, Endau Rompin NP, 120–300m, N2°62‘, E103°35‘, 28.–31.V.2013 ( JWCW) ; Johor, Gunung Panti, forest, 40m, N1.866, E103.88, 23.–27.V.2013 ( JWCW) ; Borneo, Sarawak, Kuching, Santubong, 8.–9.III.1990 ( SMNS) ; Borneo, Sarawak, Bako NP, 6.–7.XII.2010 ( SMNS) ; Borneo, Kinabalu ( SNMF) ; Selangor, Gombak, 28.IV.1998 ( SNMF) ; Johor, 30km N of Kota Tinggi, Kangkar Dohol , 35m, 7.–11.IV.2007 ( PSCH, SGCH) ; Borneo, Sabah, Keningau, V.1993 ( PSCH) ; Penang Is., IV.1989 ( MNB) ; Perak ( MNB) ; Pahang, Cameron Highlands Mts. , Tanah Rata, 1000–1500m, 2.–9.IV.1997 ( MNB) ; Borneo, Sarawak, Bako NP, IV.1987 ( ZSM) ; Borneo, Sarawak, Kuching, Mt. Serapi , 27.–29.III.1994 ( ZSM) ; Borneo, Sarawak, Kuching, Mt. Penrissen , 1000m, 24.–26.III.1994 ( ZSM) ; Borneo, Sarawak, Katang, Gunung Serapi, 900m, 110°12’E 1°36’N, 13.V.1981 ( ZSM) ; Borneo, Sarawak, Kapit , 112°56’E 2°1’N, 18.VIII.1978 ( ZSM) ; Borneo, Sarawak, Bako NP, 110°27’E, 1°43’N, 11.–14.VIII.1978, 11.–17.IV.1981 ( ZSM) ; Borneo, Sarawak, Mt. Bakak ,> 1000m, 20km SW Kapit, IV–V.1997 ( ZSM) ; Borneo, Sarawak, Mt. Bakak , 900m, 20km SW Kapit, III.1998 ( ZSM) ; Borneo, Sarawak, Long Linau, III.1990 ( ZSM) ; Borneo, Sarawak, Santubong, III.1990 ( ZSM) ; Borneo, Sarawak, Gunung Serapi, Matang, 700m, 25.IV.1987 ( ZSM) ; Borneo, Sabah, Crocker Mts. , Gunung Emas, IV.1993 ( ZSM) ; Borneo, Sabah, Crocker range, N of Keningan, Apin-Apin en., 1.V.1999 ( ZSM) ; Borneo, Sabah, Crocker Range, V.1987, IV–V.1988, VII.1988 ( ZSM) ; Borneo, Sabah, Kinabalu ( ZSM) ; Perak, Kuala-Kangsar ( ZSM) ; Penang Is., VII.1989, IV.2000 ( ZSM) ; Pahang, Cameron-Highl., Tanah Rata, 4.IV.1990 ( ZSM) ; Johor, Muar-Segamat, Gunung Ledang, 15.IV.1990 ( ZSM) ; Perak, Taiping, Nal, 1978–80 ( ZSM) ; Perak ( SDEI) ; BRUNEI: Waterstradt ( SDEI) ; SINGAPORE: Singapore ( MNB) ;? JAPAN: Iriomote, Funaura, 28.IV.1996 ( MNB) ;? PHILIPPINES: Philippines ( MNB) ; Manila ( MNB) .

Biology. Horn (1908b) noted that H. analis occurs on sunny trails, and was reported on vertical slopes shaped by water or bare piles of gravel and clay ( Naviaux 1991). Damken et al. (2017) found it on forest roads. According to Wiesner (1986b) specimens are frequently attracted to light at night.

Remarks. While this species was frequently collected, the distribution is clearly restricted to the area between the Malay Peninsula, Sumatra, Java, and Borneo ( Fabricius 1801; Lacordaire 1842; Chaudoir 1865; Fleutiaux 1892; Horn 1897, 1910, 1926, 1927; Mandl 1981; Stork 1986; Wiesner 1986b, 1989, 1992, 1996, 2007, 2013, 2019; Sawada & Wiesner 2003; Cassola 2009; Damken et al. 2017). Hence, records from Iriomote ( Japan), Philippines ( Mandl 1970; Cabras et al. 2016), and “Makassar” (Sulawesi, in Cassola 1991) are doubtful. Although a few tiger beetle species cross the Wallace Line, the low number of old records, one specimen labelled “Makassar” ( Cassola 1991) and three specimens labelled “ Philippinen / leg. Böttcher”, suggest that this labelling was erroneous. Even more improbable is the record from Iriomote, especially since numerous amateurs and specialists have explored southern Japan extensively for Heptodonta (Michio Hori pers. comm.). Schaedla (2005) wrote that H. analis can be “found throughout Thailand ”, but a reliable record is only known from most southern Thailand ( Naviaux 1991; Naviaux & Pinratana 2004). Completely unreliable is the record from Bosnia by Fleutiaux (1892).

All recent records of H. analis from the Philippines refer to H. halensis sp. nov. or H. abasileia sp. nov.

Figures

FIGURES 8–14. Heptodonta analis (Fabricius, 1801), Indonesia, Sumatra, Bukit Tigapuluh NP (JWCW): 8, 10, 12–14—♂: 8—habitus, 10—head, 12—lateral body, 13—aedeagus in left lateral aspect, 14—aedeagus in right lateral aspect; 9, 11—♀: 9—dorsal habitus, 11—head. Scales: 8–12 = 1 mm, 13–14 = 0.5 mm.

FIGURES 15–22. Heptodonta analis (Fabricius, 1801): 15—habitus, ♂, lectotype, Sumatra (ZMK); 16, 19–20—♂, Malaysia, Borneo, Bako NP (JWCW): 16—habitus, 19—aedeagus in left lateral aspect, 20—aedeagus in right lateral aspect; 17–18—♂, paralectotype, Sumatra (MNB): 17—aedeagus in left lateral aspect, 18—aedeagus in right lateral aspect; 21–22—♂, Malaysia, Johor, Endau Rompin NP (JWCW): 21—aedeagus in left lateral aspect, 22—aedeagus in right lateral aspect. Scales: 15–16 = 1 mm, 17–22 = 0.5 mm.