Aulacophora indica ( Gmelin, 1790 )

Reid, Chris, Halling, Luke & Beatson, Max, 2021, Revision of the Australopapuan and West Pacific species of plain pumpkinbeetles, the Aulacophora indica species-complex (Coleoptera: Chrysomelidae: Galerucinae), Zootaxa 4932 (1), pp. 1-73 : 31-34

Aulacophora indica ( Gmelin, 1790)

( Figs 5 , 13 , 21 , 28–29 , 38–39 , 49–52 , 70–76 , 87–89 , 100–102 , 114–115 , 127–129 , 141–144 , 155–158 , 170–175 , 185 )

Cryptocephalus indicus Gmelin, 1790: 1720 (nomen novum).

Rhaphidopalpa indica: Weise 1924: 8 .

Aulacophora indica: Wilcox 1972: 237 .

Crioceris testacea Fabricius, 1787: 87 (nec Cryptocephalus testaceus Fabricius, 1775 ).

Galeruca similis Olivier, 1808: 624 ; Baly 1886: 16 ( Aulacophora ); Kimoto 1989: 56 (synonymy).

Raphidopalpa femoralis Motschulsky, 1857: 37 ; Baly 1886: 16 ( Aulacophora ; synonymy).

Aulacophora flavipes Jacoby, 1883: 202 ; Baly 1886: 17 (synonymy).

Rhaphidopalpa pubescens Allard, 1888: 306 ; Lee & Beenen 2015: 164 (synonymy).

Rhaphidopalpa bengalensis Weise, 1892: 394 ; Weise 1924: 8 (synonymy).

Rhaphidopalpa ceramensis Weise, 1892: 394 ; Barroga & Mohamedsaid 2002: 103 (synonymy).

Rhaphidopalpa niasiensis Weise, 1892: 394 ; Barroga & Mohamedsaid 2002: 103 (synonymy).

Rhaphidopalpa chinensis Weise, 1892: 395 ; Gressitt & Kimoto 1963: 486 (synonymy).

Crioceris abdominalis sensu Fabricius 1787: 87 , nec Fabricius 1781.

Rhaphidopalpa foveicollis sensu Bechyné 1961: 256 , nec Lucas, 1849.

Material examined (188): AUSTRALIA: 1♀ */ Broken Hill, NSW, 1924 FW Shepherd/ Fam. Chrysomelidae Aulacophora palmerstoni Blackburn / K463078/ ( AMS); 1♁*/ “Toyota Maru No 2” G30 Geraldton, 22.10.1976 / ( ANIC); BRUNEI: 1♁/ KBFSC, Ashton Trail, lowland rf, 7.viii.1995, PJ Gullan ( AMS); CHINA: 1♀ / China [Blackburn writing] ( SAM); GUAM: 1♁, 1♁*, 2♀ / Nimitz Hill, Guam, on pumpkin, 8 March 1983, MF Kirby ( ANIC); INDONESIA: Java: 33, 1♁*, 1♀ */ Buitenzorg [Bogor], Java, 1931 ( ANIC); 1♁/ Buitenzorg [Bogor], Jan, AM Lea & wife ( SAM); 1♁/ Dieng Plateau, Java, 1929 G Windred ( ANIC); 1♁, 2♀ / Java, Jan–Feb 1925 C.T. McNamara ( SAM); Kalimantan: 2♁, 1♁*/ E. Kalimantan, Wanariset Res. Cen. of dipterocarp, 1–30.iii.1992, Dar- ling, Rosichon & Sutriano ( ROM); Nusa Tenggera Timor: 2♁ 1♀ / Flores, Endeh, 4.vi.1929 I Mackerras ( ANIC); 1♁, 1♁*, 1♀, 1♀ */ West Timor, Kabuna, 9: 4.2S 124: 54.54E, ex sweeping Cucurbita , 30.iii.1998 G Bellis & Supandi ( AMS); 2, 1♀ / West Timor, Nonbes, Amarasi, Kupang, 10: 12.31S 123:49:45E, ex sweeping Cucurbita sp, 27.iii.1998, Supandi ( AMS, DPID); Papua: 1♀ */ Biak, ex choko, 27.iii.1992, JW Turner ( NAQS); Sulawesi: 1♀ / Wawolemo, 50k W Kendari, 1983, R De Keyzer ( ANIC); Sumatra: 1♁*, Lampung, Desa Kedaton, Kalianda, 5:43S 105:35E, c 25m, arable (rice/maize etc), on cucurbit, ii.2010, C Reid ( AMS); 1♁/ E Sumatra, Riau, Bukit Tigapuluh NP, 0:50S 102:26E, 18–25.i.2000, J Bezdek ( AMS); JAPAN: 1♁/ Yachiyodai, Chiba Prefecture, 21 Dec 1960 ( SAM); LAOS: 1♀ / Champasak, Pakse, 15.1226N 105.8003E, 106m, watermelon seedling trial, 4.iii.2016, I Smith ( AMS); 1♀ / Champasak town, Phoxai watermelon farm, 14.911N 105.8849E, 100m, yellow pan traps, 12.ii.2016, I Smith ( AMS); 3♀, 1♀ */ Pha Song, Champasak, on jackfruit, 17.vi.2014 L Grieve ( AMS); 1♁, 1♀ / Salavan, Lakhonpheng, 15.8012N 105.6977E, 178m, watermelon farm, swept, 12.ii.2016 I Smith ( AMS); 1♀ / Sa- lavan, Lakhonpheng, Lak92, 77k N Pakse, 15.8073N 105.6914E, watermelon farm, watermelon, 12.i.2016 I Smith ( AMS); 1♁/ Savannakhet, Song Khong, Nong Hai, 16.25907N 105.17234E, 165m, watermelon farm, sweepnet, 28.i.2016, I Smith ( AMS); PHILIPPINES: 1♀ / Banaue, Luzon, 23.vii.1982 R De Keyzer ( ANIC); 22♁, 2♁*, 21♀, 1♀ */ Luzon, Tanay, Rizal, 1–5.vi.1992 R Gallardo ( AMS); 1♁, ditto, vii.1992, Melo family ( AMS); 2♁, ditto, v–vi.1992, A & E Melo ( AMS); 3♁, 4♀, ditto, v.1992 ( AMS); 1♁/ Luzon, Pleasant Village, 15:50N 121:00E, v.1985, J. Montagnes ( ROM); 1♁/ Negros Oriental, Cuernos de Negros, 7k W Valencia, 9:17N 123:15E, 700m, mal- aise trap, forest edge 15–23.xi.1987, DC Darling, E Mayordo ( ROM); TAIWAN: 3♁*, 1f/ 25.05N 121.33E, Taipei, Shulin, 4.iv.1987, DCF Rentz & YM Chiang, stop T-8 ( ANIC); TIMOR LESTE: 4♁, 8f/ 8.71S 125.58E, Aileu, Sabouria, on zucchini, 4.iii.2017, A Guterres ( NAQS); 2♁, 1♁*, 5f/ Aileu, Seloi Karik, Tasoni, 8.70S 125.53E, cucumber, 3.iii.2017, A Guterres ( NAQS); 2♀ / Becora, Dili, 8: 33.842S 125: 36.52E, ex sweep Cucurbita moschata , 9.v.2002, G Bellis ( AMS); 1♁, 1♀ / Desa Ililai, Kec. Moro, Kab. Lautem, 8: 24.665S 126: 44.236E, ex sweep Cucurbita moschata , 14.vii.2000 G Bellis ( DPID); 6, 2♀ / Desa Rahi Tahu, 8: 54.596S 126: 23.667E, defoliating Cucurbita moschata , 18.vii.2000, G Bellis ( AMS, DPID); 4♁, 13♀ / Ermera, Gleno, Talimara, 8.72S 125.43, cucumber & eggplant, 28.ii.2017, D Britton & D Ximenes ( NAQS); 1♁*, 1♀ */ Manatuto Agr Office, 8.53181S 126.00462E, sweep & hand coll from Cucurbita pepo , 18.iv.2012, S Anderson ( NAQS); VIETNAM: 1♀ */ Dong Nai: Cat Tien NP, Crocodile Lake, 11:27: 31.5N 107:20: 42.5E, edge field/forest, 120m, malaise trap, 26–31.viii.1998, B Hubley ( ROM); 1♀ / Nghe An, c25k SW Con Cuong, Khe Moi River forestry camp, 18:56N 104:49E, 308m, tropical forest, 3–10.vi.1995, B Hubley, J Swann ( ROM).

Description (based on material examined). Colour ( Fig. 5 ). There are two colour forms which may be clinal ( Lee & Beenen 2015). Almost all of the material we have examined is of the pale, southern form. Head brownish-yellow, except apical half to two-thirds of labrum and apices of mandibles dark brown to black, eyes black; antennae usually entirely brownish-yellow, occasionally antennomeres 4–11 may appear feebly darker (yellowishbrown), rarely darkening from pale base to dark brown apex; pronotum and elytra entirely brownish-yellow; venter of prothorax entirely brownish-yellow or prosternum partly brown (Borneo and eastern Sumatra only); scutellum brownish-yellow or disc brown (Borneo and eastern Sumatra only); mesanepisternum, mesepimeron and mesoventrite brownish-yellow or partly brown with mesanepisternum often dark brown ( Guam, Borneo and eastern Sumatra only); metaventrite dark brown to black, occasionally with yellowish anterior margins; procoxae brownish-yellow; mesocoxae anteriorly yellowish-brown and posteriorly dark brown; metacoxae usually almost entirely dark brown to black; legs variable in colour, usually entirely brownish-yellow, sometimes with meso- and metafemora medially brown, darkest forms with mid and hind legs almost entirely black (trochanters, extreme bases of tibiae and third tarsomeres brown) and forelegs brownish-yellow with narrow dorsal edge of femora and broad dorsal edge of tibiae dark brown ( Guam, Borneo); median tergites pale brown, propygidium usually darker; pygidium variable, usually entirely brownish-yellow or with brown base, to almost entirely dark brown with apical margins brown (one Bornean specimen); abdominal ventrites 1–4 dark brown to black, sometimes with apical margins pale brown; ventrite 5 with dark brown to black basal band, extending apically at sides, remainder brownish-yellow, or entirely dark brown (one Bornean and one Guam specimen).

Male: length 6–8 mm; frontoclypeus without arcuate ridges or densely setose patches; first antennomere expanded, oval flat area in apical half defined by a ridge; antennae about 0.6x body length; antennomere 2 shortest, variable in size, less than to greater than one third length of 1, antennomere 1 longest, comparative lengths: 1> 11> 3 = 4 = 5 = 6 = 9> 7 = 8> 10> 2 ( Philippines specimen), 1>11> 5 = 9> 4 = 6 = 7 = 8 = 10> 3> 2 (Timorese specimen); antennomere 5 length 2.7 (Timorese specimen)—3.3 (Philippine specimen) x width; antennomeres 3–7 slightly expanded to apices; antennomeres 3–11 each with 3–6 erect lateral setae, not forming a row; pronotal transverse depression posteriorly shallowly arcuate, deep at sides and middle, broadest at middle; in lateral view convexity of anterior half of pronotum similar to posterior half and median depression at sides with anterior and posterior slopes similar; disc with pair of large pits anterior to transverse groove; elytra less shiny than pronotum, shallowly microreticulate; elytral humeri with elongate oval patch of 100–200 erect setae; apical lobe of ventrite V asymmetrically sculptured, cavity abruptly ridged on left (more sharply in southeast Asian and Indonesian specimens, less so in Philippines and Timor), ridge often with slight basal swelling, and gradually elevated on right without a basal swelling; elongate cavity deepened from base almost to apex and deepest at left side before apex (more so in southeast Asian and Indonesian specimens, less so in Philippines and Timor), then gradually elevated to apical margin; tergite VIII bicornual with two prongs separated by a deep U-shaped concavity, prongs elongate and narrow with upturned or straight apices and left prong usually slightly shorter and straighter than right; prominant acute or rounded projection lateral to the central prongs; penis thin and almost parallel-sided in lateral view, with small sharply angulate tubercle at tip; sides without punctures, smooth and unridged; narrow and almost parallel-sided in dorsal view, usually slightly constricted at about middle, sides abruptly attenuated in apical 1/7 to acute apex.

Female as male, except: length 6–8 mm; pygidium variable in colour, as male; abdominal ventrites 1–4 often with pale brown apical margins; antennomeres slightly thinner than male, antennomere 5 length 3.2 –3.3 x width, antennomere 8 length 2.9 ( Philippines)—3.5 (Timor) x width; transverse pronotal depression shallower; pygidium almost flat, but shallowly medially keeled, with flat apical extension in most southeast Asian and Indonesian specimens, apical half slightly raised and thickened in Philippines specimens, Timorese specimens mostly intermediate or flat; apex pygidium in dorsal view elongate-triangular (apical angle 50–80̊) with straight sides but the tip vari- able, acute, rounded, truncate or narrowly concave; pygidial apex without tubercle; venter of pygidial apex slightly concave, or flat; apex ventrite V deeply and narrowly concave, concavity rarely asymmetric (U-, or V-shaped), and usually as deep as broad (depth: width ratio 1:1.8 or less, shallowest in Vietnamese specimens examined); apex of ventrite V with preapical lateral depressions defined by lateral ridges, apical margin flat, not reflexed at middle; vaginal palpi elongate ovate, length 3–4.3x width, with 7–8 pairs of setae in apical half; basal apodemes straight to slightly curved, 0.43–0.48 mm long; sternite VIII with tignum separated from weakly sclerotised posterior margin of the sternite by a transparent membranous area or this area slightly pigmented, and posterior margin truncate, not produced; tignum 1.2–1.5 mm long, apex membranous, rounded to almost triangular and usually separated from shaft by a band of deeper pigmentation; spermathecal shape variable, falcate, collum abruptly demarkated from receptaculum and reflexed relative to receptaculum but variable in shape and insertion point of gland (ramus) flat to produced; receptaculum strongly hook-shaped with angulate to evenly curved interior bend; appendix variable from large and beak-like to almost absent.

Diagnosis. Male: pronotal disc with pair of glands ( Fig. 10 ), pronotal depression almost symmetrical in lateral view ( Figs 27 ), humeral setal patch present ( Fig. 21 ), scutellum usually pale ( Fig. 5 ), tergite 8 deeply arcuate ( Figs 87–89 ), penis thin and almost parallel-sided with blunt apex ( Figs 100–102 ). Female: frontoclypeus medially keeled, scutellum usually pale, apex of abdominal venter pale ( Figs 49–52 ), apical margin of ventrite 5 deeply concave with width at most 1.8x depth and with subparallel ridges at sides ( Figs 49–52 , 70–76 ).

Notes. The justification of validity of the name of Aulacophora indica ( Gmelin 1790) , has not been made explicit, so we provide the rationale here. It is a junior subjective synonym of A. testacea ( Fabricius 1787) , which was formerly a junior subjective homonym. Gmelin’s name was a substitute because he had placed both Crioceris testacea Fabricius 1787 and Cistela testacea Fabricius 1775 in the genus Cryptocephalus ( Gmelin 1790) and had recognised the consequent homonymy of these names. Gmelin therefore renamed the former as Cryptocephalus indica . Crioceris testacea is now placed in Aulacophora and Cistela testacea in Exosoma Jacoby 1903 . The lectotype and identity of Crioceris testacea was fixed recently ( Lee & Beenen 2015). However, although the two species names are no longer bound by the same genus, Article 59.3 (ICZN 1999) states: “the junior secondary homonym is permanently invalid unless the substitute name is not in use AND [our emphasis] the relevant taxa are no longer considered congeneric”. In this case the substitute name, A. indica ( Gmelin 1790) , is well established and is therefore valid, even though the relevant taxa are no longer congeneric.

Aulacophora indica was clearly identified and redescribed by Barroga (2002). This species has at least 8 synonyms (Barroga 2002; Lee & Beenen 2015). We are not able to check all of them, but since the synonymy has been determined recently by two independent authors we see little need to. However we note that photographs of a female syntype or holotype of the Sulawesian species A. flavipes Jacoby, 1883 , are available online at the time of writing, confirming Baly’s synonymy of this species with A. indica ( Perkins et al. 2019) .

Gressitt’s description of female A. similis in Micronesia fits this species ( Gressitt 1955) and we can confirm that A. indica occurs on Guam. Specimens from other Micronesian islands have not been examined as this area is outside the region of our study. Gressitt’s note on variation in the shape of female apical abdominal ventrite ( Gressitt 1955: 30) suggests that both A. abdominalis and A. indica occur in Micronesia.

Aulacophora indica may have been confused with A. foveicollis in southern Asia, where both species are supposed to occur. The latter clearly belongs to the indica species-complex, and has a penis even more sinuate than those of A. abdominalis and A. relicta ( Mann 1985; Berti 1990; Borowiec 2013).

Lee & Beenen (2015) noted the colour variation in A. indica , suggesting it was geographical (clinal), with dark specimens in the northern and eastern part of the range ( Taiwan, Japan, Philippines), pale specimens in the south ( India, southeast Asia) and both forms present in China and Indonesia. There are colour photographs of A. indica in reviews of the chrysomelids of Taiwan ( Lee & Cheng 2007) and Japan ( Hayashi, Morimoto & Kimoto 1984) showing the dark legs of the typical northern colour forms. Our study partly corroborates the observations of Lee & Beenen (2015), but all the Philippines material we examined was of the pale form. The few specimens we examined from Micronesia and nearby islands belong to the dark form. Gressitt (1955) noted colour variation in A. indica (as A. similis ) in this area. Our colleague Al Samuelson has examined Micronesian material in BPBH and has confirmed that only A. indica is involved, although some specimens have dark apical antennal segments. The two specimens we have examined from Australia belong to each colour variety. The specimen intercepted on a Japanese boat in Perth belongs to the dark form typical of Japan. The specimen labelled from Broken Hill belongs to the pale southern Asian form (see below, under distribution).

The male penis is diagnostic for A. indica and shows little variation ( Figs 100–102 , 127–129 ). The female abdominal ventrite 5 is diagnostic ( Figs 70–76 ) but other female genitalic characters are variable and not useful for diagnosis of this species: sternite 8 ( Figs 141–143 ; vaginal palpi ( Figs 155–158 ) and spermatheca ( Figs 170–175 ). The genitalic variation does not seem to correlate with the colour variation noted above.

Distribution ( Fig. 185 ) and biology. This is the dominant PPB species in south-east Asia and the central western Pacific. Records of A. indica in the southwestern Pacific (eg. Anonymous 2013, 2018) almost certainly belong to A. abdominalis . Besides the material listed above, the following western Pacific specimens in BPBM have been identified by our colleague Al Samuelson, using our diagnostic key: GUAM: 1m, 1f, Talofofo, viii.1952, NHL Krauss; 1m, ditto, x.1957; 1m, 1f, Yigo, x.1957, NLH Krauss; 1f, Yona, x.1952, NHL Krauss; MICRONE- SIA: 1, Yap, vii–viii.1950; NORTHERN MARIANA ISLANDS: 10, Saipan, ii–iii.1958; 2, Tinian, x.1952; PA- LAU (“many”): Angaur, ii.1948; Babelthaup, vii.1946; Garakayo, viii.1945; Kayangel, x.1952; Koror, vii.1940 & iii–v.1954;; Melokeiok, vi.1936; Ngeremlengui, iv.1936; Pelelieu, vii.1945; Sonsorol, ix.1952. The mapped distribution shows that this is the dominant and usually the only species of PPB in the central west Pacific north of the Equator, including Palau, the Mariana Islands and western Caroline Islands. The records for this area are all from the 1930s onwards and may be due to importation of cucurbits under the Japanese occupation (Palau was the first of the islands annexed by Japan, in 1914). Aulacophora indica may be native to the area and simply not previously collected, but it is regarded as an invasive species in Guam where Gressitt failed to find it in 1945 ( Gressitt 1955). Most invasive chrysomelid species to Guam appeared to have come from the Philippines ( Gressitt 1955). However Gressitt noted slight morphological differences amongst the material of A. indica from Micronesia and suggested that it might have had two or more origins. Aulacophora indica has only been reliably recorded in New Guinea from the far northwest, at Biak Island, but is otherwise common in the Indonesian Archipelago from Timor westwards. Its western limits in the Indian Subcontinent are obscure as this species seems to have been confused with A. foveicollis .

We have only seen two specimens of Aulacophora indica labelled as Australian. One was a biosecurity interception off a Japanese ship (the “Toyota Maru 2”) in a Western Australian port, with no evidence that it represented an established population. The other is labelled from Broken Hill, a central Australian mining town, and it was supposedly collected in 1924 by F.W. Shepherd who provided material to the Australian Museum from the Broken Hill area, from 1905 to 1925. We were therefore concerned that this notifiable pest might be established in Broken Hill. However more recent surveys of cucurbits there have only found the native species A. relicta (OAI; Peter Gillespie, pers. com. April 2017). We suggest that if A. indica was established in Broken Hill it would have become extinct quite quickly as the annual rainfall is less than 300mm and mean July minimum temperature is 5̊ C ( Anonymous 2020). These figures lie well outside the range of temperature and rainfall experienced by A. indica in its native range in south-east Asia. More likely is that this specimen is wrongly labelled. In the Australian Museum collection there are several specimens of another exotic pest species, Basilepta subcostata (Jacoby, 1889) , appended with exactly the same label. Basilepta subcostata is a serious pest in Southeast Asia on banana ( Mishra et al. 2015), a plant which does not grow in Broken Hill, and these specimens are also the only record of the beetle in Australia. They must be incorrectly labelled. The label for the Broken Hill specimen of A. indica is therefore also almost certainly incorrect.

Material we have examined was collected on the following plants (number of collection events in brackets): Cucurbitaceae : Citrullus lunatus (2), Cucumis sativus (2), Cucurbita maxima (3), Cucurbita moschata (3), Cucurbita pepo (2), Sechium edule (1); Moraceae : Artocarpus heterophyllus (1). Most of these are exotic to the area and the likely native hosts in Asia are species of Cucumis These records suggest that A. indica has a greater host range within Cucurbitaceae than A. abdominalis .

Figures

FIGURES 2–5. Aulacophora species, male habitus: 2, A. abdominalis (Fabricius, 1781) (Guadalcanal); 3, A. barrogae Reid, Halling & Beatson, sp. nov.; 4, A. cornuta Baly, 1879 (New Ireland); 5, A. indica (Gmelin, 1790) (Luzon).

FIGURES 10–15. Aulacophora species, male head, anterior view: 10, A. abdominalis (Fabricius, 1781) (Guadalcanal); 11, A. barrogae Reid, Halling & Beatson, sp. nov.; 12, A. cornuta Baly, 1879 (New Ireland); 13, A. indica (Gmelin, 1790) (Luzon); 14, A. mbabaram Reid, Halling & Beatson, sp. nov.; 15, A. relicta (Boisduval, 1835) (Mount Molloy).

FIGURES 18–25. Aulacophora species, male anterior and antenna: 18, A. abdominalis (Fabricius, 1781) (Guadalcanal); 19, A. barrogae Reid, Halling & Beatson, sp. nov.; 20, A. cornuta Baly, 1879 (New Ireland); 21, A. indica (Gmelin, 1790) (Luzon); 22, A. mbabaram Reid, Halling & Beatson, sp. nov.; 23, A. relicta (Boisduval, 1835) (Mount Molloy); 24, A. wallacii Baly, 1886; 25, A. wilsoni Baly, 1888.

FIGURES 26–31. Aulacophora species, pronotum, lateral view: 26, male A. abdominalis (Fabricius, 1781) (Vanuatu); 27, male A. barrogae Reid, Halling & Beatson, sp. nov.; 28, male A. indica (Gmelin, 1790) (Borneo); 29, female A. indica; 30, male A. relicta (Boisduval, 1835); 31, male A. wallacii Baly, 1886.

FIGURES 38–43. Aulacophora species, male, apex of abdominal venter: 38, A. indica (Gmelin, 1790) (Sumatra); 39, A. indica (West Timor); 40, A. mbabaram Reid, Halling & Beatson, sp. nov.; 41, A. relicta (Boisduval, 1835) (Mount Molloy); 42, A. wallacii Baly, 1886; 43, A. wilsoni Baly, 1888.

FIGURES 44–49. Aulacophora species, female, apex of abdominal venter and pygidium: 44, A. abdominalis (Fabricius, 1781) (Vanuatu); 45, A. abdominalis (New Ireland); 46, A. abdominalis (Mer Island); 47, A. barrogae Reid, Halling & Beatson, sp. nov.; 48, A. cornuta Baly, 1879 (Sulawesi); 49, A. indica (Gmelin, 1790) (Laos).Dotted lines indicate depth and width of apical concavity.

FIGURES 50–55. Aulacophora species, female, apex of abdominal venter: 50, A. indica (Gmelin, 1790) (Biak Island); 51, A. indica (Vietnam); 52, A. indica (Broken Hill); 53, A. mbabaram Reid, Halling & Beatson, sp. nov., Almaden; 54, A. relicta (Boisduval, 1835); 55, A. wallacii Baly, 1886.

FIGURES 70–76. Apical margin of female sternite VII, Aulacophora indica (Gmelin, 1790) (locality sequence from west to east): 70, Laos, 71, Vietnam, 72, Luzon, 73, east Borneo, 74, west Timor, 75, east Timor, 76, west New Guinea (Biak).

FIGURES 83–88. Aulacophora species, male sternite 8, ventrolateral view: 83, A. abdominalis (Fabricius, 1781) (Vanuatu); 84, A. abdominalis (New Ireland); 85, A. barrogae Reid, Halling & Beatson, sp. nov.; 86, A. cornuta Baly, 1879 (Guadalcanal); 87, A. indica (Gmelin, 1790) (Timor); 88, A. indica (Luzon).

FIGURES 89–93. Aulacophora species, male sternite 8, ventrolateral view: 89, A. indica (Gmelin, 1790) (Sumatra); 90, A. mbabaram Reid, Halling & Beatson, sp. nov.; 91, A. relicta (Boisduval, 1835) (Coonabarabran); 92, A. wallacii Baly, 1886; 93, A. wilsoni Baly, 1888.

FIGURES 98–101. Aulacophora species, penis, dorsal view: 98, A. barrogae Reid, Halling & Beatson, sp. nov.; 99, A. cornuta Baly, 1879 (Guadalcanal); 100, A. indica (Gmelin, 1790) (Laos); 101, A. indica (Luzon).

FIGURES 102–105. Aulacophora species, penis, dorsal view: 102, A. indica (Gmelin, 1790) (Timor); 103, A. mbabaram Reid, Halling & Beatson, sp. nov.; 104, A. relicta (Boisduval, 1835) (Mount Molloy); 105, A. relicta (Mount Mulligan).

FIGURES 112–115. Aulacophora species, penis, ventral view: 112, A. barrogae Reid, Halling & Beatson, sp. nov.; 113, A. cornuta Baly, 1879; 114, A. indica (Gmelin, 1790) (Laos); 115, A. indica (W Timor).

FIGURES 125–128. Aulacophora species, penis, lateral view: 125, A. barrogae Reid, Halling & Beatson, sp. nov.; 126, A. cornuta Baly, 1879; 127, A. indica (Gmelin, 1790) (Laos); 128, A. indica (Luzon). Arrow indicates small spine on dorsal surface.

FIGURES 129–132. Aulacophora species, penis, lateral view: 129, A. indica (Gmelin, 1790) (W Timor); 130, A. mbabaram Reid, Halling & Beatson, sp. nov.; 131, A. relicta (Boisduval, 1835) (Mount Molloy); 132, A. relicta (Mount Mulligan).

FIGURES 135–142. Aulacophora species, female sternite 8: 135, A. abdominalis (Fabricius, 1781) (Mer); 136, A. abdominalis (Fuiloro); 137, A. abdominalis (Vanuatu); 138, A. abdominalis (New Ireland); 139, A. barrogae Reid, Halling & Beatson, sp. nov.; 140, A. cornuta Baly, 1879 (Sulawesi); 141, A. indica (Gmelin, 1790) (Luzon); 142, A. indica (Laos).

FIGURES 143–149. Aulacophora species, female sternite 8: 143, A. indica (Gmelin, 1790) (Manatuto); 144, A. indica (Biak); 145, A. mbabaram Reid, Halling & Beatson, sp. nov.; 146, A. relicta (Boisduval, 1835) (Ashford); 147, A. wilsoni Baly, 1888 (Eungella); 148, A. wallacii Baly, 1886; 149, A. wilsoni (Lamington).

FIGURES 150–155. Aulacophora species, female ovipositor: 150, A. abdominalis (Fabricius, 1781) (Mer); 151, A. abdominalis (Vanuatu); 152, A. abdominalis (New Ireland); 153, A. barrogae Reid, Halling & Beatson, sp. nov., 154, A. cornuta Baly, 1879; 155, A. indica (Gmelin, 1790) (Luzon). Sensilla not shown on Fig. 154.

FIGURES 156–160. Aulacophora species, female ovipositor: 156, A. indica (Gmelin, 1790) (Laos); 157, A. indica (Manatuto); 158, A. indica (Biak); 159, A. mbabaram Reid, Halling & Beatson, sp. nov.; 160, A. relicta (Boisduval, 1835) (Ashford). Sensilla not shown on Figs 156, 158, 160.

FIGURES 164–172. Aulacophora species, spermatheca: 164, A. abdominalis (Fabricius, 1781) (Mer); 165, A. abdominalis (Fuiloro); 166, A. abdominalis (Vanuatu); 167, A. abdominalis (New Ireland); 168, A. barrogae Reid, Halling & Beatson, sp. nov., 169, A. cornuta Baly, 1879 (Sulawesi); 170, A. indica (Gmelin, 1790) (Luzon); 171, A. indica (Laos); 172, A. indica (Manatuto).

FIGURES 173–181. Aulacophora species, spermatheca: 173, A. indica (Gmelin, 1790) (Biak); 174, A. indica (Vietnam); 175, A. indica (Timor); 176, A. mbabaram Reid, Halling & Beatson, sp. nov. (Almaden); 177, A. mbabaram (Almaden); 178, A. wilsoni Baly, 1888 (Eungella); 179 A. relicta (Boisduval, 1835) (Ashford); 180, A. wallacii Baly, 1886; 181, A. wilsoni (Lamington).

FIGURES 184–185. 184, distribution of Aulacophora cornuta Baly, 1879, east of Indonesia; 185, distribution of Aulacophora indica (Gmelin, 1790) (specimens examined only).