An illustrated guide to lady beetles (Coleoptera: Coccinellidae) of the Indian Subcontinent. Part 1. Tribe Coccinellini Author POORANI, J. text Zootaxa 2023 2023-08-18 5332 1 1 307 http://dx.doi.org/10.11646/zootaxa.5332.1.1 journal article 264199 10.11646/zootaxa.5332.1.1 66e0ec51-d494-43d4-965e-a2cd1462ef54 1175-5326 8261502 424F7439-4095-46A5-93E3-C4130E3B6D9A Cheilomenes sexmaculata (Fabricius) ( Figs 46–49 ) Coccinella sexmaculata Fabricius, 1781: 96 ( Lectotype female, BMNH; Type locality: ‘Ind. Orient.’). Coccinella 4—plagiata Swartz in Scĥnherr, 1808: 195 ( Type locality: ‘ India Orientali’); Synonymized by Nakane & Araki 1959: 50 ; Sasaji & Akamatsu 1979: 16 . Cheilomenes sexmaculata : Dejean 1837: 435 ; Mulsant 1850: 444 ; 1866: 289 ; Crotch 1871: 8; 1874: 180 (as Chilomenes ); Poorani 2002a: 325 . Cheilomenes quadriplagiata : Dejean 1837: 435 ; Mulsant 1850: 447 ; 1866: 290 ; Crotch 1871: 8; 1874: 181 (as Chilomenes ). Menochilus sexmaculatus : Timberlake 1943: 40 ; Nakane & Araki 1959: 50 ; Sasaji & Akamatsu 1979: 2 ; Iablokoff-Khnzorian 1979: 58 ; 1982: 146 ; Pope 1989: 646 . Menochilus quadriplagiatus : Timberlake 1943: 41 . Cydonia triangulifera var. inops Mulsant, 1866: 286 ; Crotch 1874: 180 (as Chilomenes ); Bielawski & Chujo 1964: 228 (as Verania ). Micraspis inops Chunram & Sasaji, 1980: 488 ; Iablokoff-Khnzorian 1982: 511 . Diagnosis. Length: 3.30–6.20 mm ; width: 3.00– 5.30 mm . Form broadly oval to subrounded, dorsum moderately convex, glabrous and shiny. Ground colour orange, light red, yellow or pinkish with the following markings in the typical form ( Figs 47 , 49g –l ): head with a black transverse marking in posterior half ( Fig. 46a ); pronotum with a T-shaped median marking connected to a broad black band along posterior margin; elytra with two zigzag lines and a posterior black spot, suture with a black stripe. Ventral side uniformly yellow. Male genitalia ( Fig. 46f–j ), female genitalia ( Fig. 46k ) and spermatheca ( Fig. 46l, m ) as illustrated. This is the most ubiquitous Coccinellini of the Indian region but its polymorphic forms are widely misidentified due to superficial similarities with other common but unrelated species such as Micraspis discolor (Fabricius, 1798) , Propylea dissecta ( Mulsant, 1850 ) and Chilocorus nigrita (Fabricius, 1798) . The following forms are frequently seen: (a) the nominate form with two zigzag maculae and an apical spot ( Fig. 49g ); (b) the elytral markings coalesce and form broader bands ( Fig. 49h, i ); (c) elytra immaculate except for the sutural black stripe ( Fig. 49j ), (d) the elytra and pronotum are either partially black leaving only the margins ( Fig. 49k ), and (e) completely black ( Fig. 49l ). The pronotal marking is always constant and can be faintly seen even in completely black forms ( Fig. 49k, l ). All these forms freely intermate and several intermediate forms are found in the field ( Subramaniam 1924a ). Immature stages. Eggs ( Fig. 48a ) are yellow, spindle-shaped and laid in groups. The larva ( Figs 48b, c ; 49a–d ) is dark slaty grey to brown, with creamy yellowish markings. The pupa ( Figs 48g , h ; 49e, f ) is yellow or orange with black spots. Congregations of larvae and pupae are commonly seen on host plants like Calotropis gigantea and french bean ( Fig. 48d, e ). Distribution. Distributed almost throughout the Indian Subcontinent, the Oriental region and Asia; Iran ; Australo-Pacific region. Introduced and established in the Caribbeans (Assour & Behm 2019) and South America ( Chile ; Ecuador ; Peru ; Venezuela ; Colombia ) ( Ramírez et al . 2018 ). Prey/associated habitat. Mainly aphidophagous, but feeds on a diverse range of prey that includes psyllids, whiteflies, mealybugs, tingids, leaf-and planthoppers, mites, early instar lepidopteran larvae and isopods. Agarwala & Yasuda (2000) gave a worldwide checklist of prey of this species. Some of the common hosts are listed below. Aleyrodidae : Aleurodicus dispersus Russell , Aleurolobus barodensis (Maskell) , Bemisia tabaci (Gennadius) , Lipaleyrodes euphorbiae David & Subramaniam , Neomaskellia bergii (Signoret) , Rusostigma eugeniae (Maskell) , Siphoninus phillyreae (Haliday) ; Aphididae : Acyrthosiphon pisum (Harris) , Aphis affinis Del Guercio , Aphis craccivora Koch , A. craccivora pseudoacaciae Takahashi , Aphis cytisorum Hartig , Aphis fabae Scopoli , Aphis gossypii Glover , Aphis longisetosa Basu , Aphis nerii Boyer de Fonscolombe , Aphis pomi De Geer , Aphis spiraecola Patch (as A. citricola van der Goot ), Aphis umbrella (Boerner) , Aulacorthum nipponicum (Essig & Kuwana) , Aulacorthum solani (Kaltenbach) , Brachycaudus helichrysi (Kaltenbach) , Brevicoryne brassicae (Linnaeus) , Capitophorus himalayensis Ghosh et al . , Cavariella simlaensis Chowdhuri et al . , Cerataphis brasiliensis (Hempel) (as C. palmae (Ghesquiere)) , Ceratovacuna lanigera Zehntner , Cervaphis rappardi indica Basu , Cervaphis schouteniae van der Goot , Chaitophorus himalayensis (Das) , Coloradoa rufomaculata (Wilson) , Cryptosiphum artemisiae Buckton , Dreyfusia (as Adelges ) knucheli (Schneider-Orelli & Schneider) , Greenidea heeri Raychaudhuri et al . , Greenideoida ceyloniae van der Goot , Hyadaphis coriandri (Das) , Hyalopterus pruni (Geoffroy) , Hyperomyzus lactucae (Linnaeus) , Hysteroneura setariae (Thomas) , Lipaphis pseudobrassicae (Kaltenbach) , Liosomaphis atra Hille Ris Lambers , Macrosiphoniella sanborni (Gillette) , Macrosiphum rosae (Linnaeus) , Melanaphis sacchari (Zehntner) (as M. indosacchari (David)) , Myzus nicotianae Blackman , Myzus obtusirostris David et al . , Myzus persicae (Sulzer) , Pemphigus ?napaeus Buckton, Pentalonia nigronervosa Coquerel , Pineus sp. , Rhopalosiphum maidis (Fitch) , Rhopalosiphum nymphaeae (Linnaeus) , Rhopalosiphum padi (Linnaeus) , Rhopalosiphum sp. , Schizaphis graminum (Rondani) , Shivaphis celti Das , Sinomegoura citricola (van der Goot) , Sitobion akebiae (Shinji) , Sitobion avenae (Fabricius) , Sitobion graminis Takahashi , Sitobion ibarae (Matsumura) , Sitobion rosaeiformis (Das) , Sipha maydis Passerini , Therioaphis trifolii (Monell) , Tinocallis kahawaluokalani (Kirkaldy) , Aphis (Toxoptera) aurantii Boyer de Fonscolombe , Aphis (Toxoptera) citricidus (Kirkaldy) , Aphis (Aphis) odinae (van der Goot) , Tuberculatus nervatus Chakrabarti & Raychaudhuri , Uroleucon compositae (Theobald) (as Dactynotus solidaginis (Fabricius)) , Uroleucon carthami (Hille Ris Lambers) , Uroleucon (as Dactynotus ) formosanus (Takahashi), Uroleucon (as Dactynotus ) nigrotuberculatum (Olive); Cicadellidae : Amrasca biguttula biguttula (Ishida) , Amritodus atkinsoni (Lethierry) , Empoasca kerri Singh-Pruthi , Hishimonus phycitis (Distant) , Empoasca sp. , Exitianus indicus (Distant) , Deltocephalus sp. , Idioscopus clypealis (Lethierry) , Nephotettix nigropictus (Stål) ; Coccoidea: Drepanococcus chiton (Green) , Drosicha mangiferae Green , Hemiberlesia lataniae (Signoret) , Maconellicoccus hirsutus (Green) , Orthezia insignis Browne , Parlatoria blanchardi (Targioni Tozzetti) , Phoenicococcus marlatti Cockerell , Pulvinaria psidii Maskell ; Delphacidae : Nilaparvata lugens (Stål) , Peregrinus maidis (Ashmead) , Sogatella furcifera (Horvath) ; Lophopidae : Pyrilla perpusilla (Walker) ; Psyllidae : Diaphorina citri Kuwayama , Heteropsylla cubana Crawford , Paurocephala psylloptera Crawford , Phylloplecta sp. ; Lepidoptera : Chilo partellus (Swinhoe) (Crambidae) ; Pectinophora gossypiella (Saunders) (Gelechiidae) ; eggs and/or early instar larvae of Chilades lajus (Stoll) (Lycaenidae) ; Earias spp. , Helicoverpa armigera (Ḩbner), Spodoptera litura (Fabricius) (Noctuidae) ; Papilio demoleus Linnaeus (Papilionidae) ; Cydia leucostoma Meyrick (Tortricidae) ; Opisina arenosella Walker (Xylorictidae) . Found in association with Diptera : Atherigona soccata Rondani (Anthomyiidae) ; Dasyneura lini Barnes (Cecidomyiidae) and Acanthiophilus helianthi (Rossi) (Tephritidae) . Acari: Oligonychus coffeae (Nietner) on tea, Oligonychus mangiferus (Rahman & Sapra) , Tetranychus urticae Koch , Tetranychus sp. ( ludeni -group) ( Tetranychidae ). FIGURE 46. Cheilomenes sexmaculata (F.): a. head; b. antenna; c. maxilla; d. prosternal process; e. abdominal postcoxal line; f–j. male genitalia: f. tegmen, lateral view; g. tegmen, ventral view; h, i. penis, variants; j. penis apex; k. female genitalia; l, m. spermatheca. FIGURE 47. Cheilomenes sexmaculata (F.), live adult. FIGURE 48. Cheilomenes sexmaculata , life stages: a. eggs; b, c. larva; d, e. larva and pupa; f. parasitized larvae; g, h. pupa. FIGURE 49. Cheilomenes sexmaculata , life stages: a–d. larva; e, f. pupae; g–l. adult, polymorphic forms. Smartweed ( Polygonum hydropiper ) and sunnhemp ( Crotalaria juncea ) are common reservoir plants that harbour the beetles in large numbers (label data). Collected on a variety of agricultural crops including maize, sorghum, rice, finger millet, cowpea, cotton, bhendi, brinjal, cabbage, cauliflower, groundnut, and several other plants such as gliricidia (label data). Seasonal occurrence. Active almost throughout the year in India , with many overlapping generations. Natural enemies. Dinocampus coccinellae (Schrank) , Nothoserphus mirabilis Brues , Homalotylus eytelweinii Ratzeburg , Homalotylus flaminius (Dalman) , Homalotylus terminalis (Say) , Oomyzus scaposus (Thomson) (= Tetrastichus coccinellae Kurdjumov ), Pediobius foveolatus (Crawford) , Tetrastichus sexmaculatus Kurian , Tetrastichus sp. , Eocanthecona furcellata (Wolff) ( Hemiptera : Pentatomidae ), Coccipolipus sp. , Parasitylenchus coccinellae Iperti & Waerebeke (Nematoda: Allantonematidae ), and Wolbachia sp. Notes. This is probably the best-known species of Coccinellini of this region with numerous papers on its hosts, biology, population dynamics, susceptibility to insecticides, etc. Stebbing (1903) , Lefroy (1909) , Subramaniam (1924a) , Rahman (1940) , and Puttarudriah & Channabasavanna (1953 , 1956 ) provided accounts of its bionomics, associated host plants / insects, etc. Sasaji (1971) , Pope (1989) , Booth et al. (1990) and Ślipiński et al. (2020) described the genus / species with illustrations. Sasaji & Akamatsu (1979) carried out breeding experiments and illustrated the elytral pattern variations. Agarwala & Yasuda (2000) reviewed and provided a worldwide checklist of host insects.