Coccinellini Latreille, 1807

POORANI, J., 2023, An illustrated guide to lady beetles (Coleoptera: Coccinellidae) of the Indian Subcontinent. Part 1. Tribe Coccinellini, Zootaxa 5332 (1), pp. 1-307 : 12-26

Tribe Coccinellini Latreille, 1807

Diagnosis. Adults usually brightly coloured, reddish, orange or yellow with spots and other patterns ( Fig. 2 ), rarely pink or lime green. Body medium to large (2.50–18.00 mm), form circular, short oval, or elongate oval in outline, dorsum weakly to strongly convex, always glabrous (except for one genus, Singhikalia Kapur having distinct dorsal pubescence). Anterior clypeal margin normal, never laterally expanded, straight or shallowly to deeply arcuate between usually prominent lateral projections in predatory forms, without distinct lateral projections in mycophagous forms. Eyes prominent and large, usually finely facetted, weakly emarginate anteriorly. Antenna as long as or shorter or longer than head width, with 11 antennomeres, weakly or distinctly clubbed and inserted laterally in front of eyes. Mandibles usually apically bifid in predatory forms, apically multidentate, with a series of small serrations on the ventral side of apical tooth in mycophagous forms. Terminal maxillary palpomere typically securiform or axe-head shaped, elongate or more transverse in different degrees, apical margin strongly divergent. Prosternal intercoxal process moderately broad, usually with a pair of parallel carinae, occasionally absent. Middle and hind tibiae with or without a pair of apical spurs. Tarsal formula 4-4-4, tarsal claws usually appendiculate, rarely apparently double and apically cleft or almost simple with only a basal dilation. Abdomen with six ventrites; abdominal postcoxal lines on ventrite 1 complete or incomplete, occasionally with an associated oblique line, rarely absent; abdominal ventrites 5 and 6 posteriorly modified and sexually dimorphic in most cases. Male genitalia with tegmen symmetrical, parameres well developed with inner and / or apical margins densely setose; penis usually with a well-defined capsule, penis apex variously modified. Ovipositor broad, coxites of club and handle type, bearing short, or rarely prominent styli, colleterial glands always present; spermatheca with well-differentiated nodulus and ramus or only nodulus or only ramus distinct, infundibulum present or absent.

Adults of some Endomychidae ( Fig. 3c, d ) and Tenebrionidae ( Fig. 3e–h ) have bright red and black colour patterns and are often mistaken with Coccinellini and these can be easily separated from Coccinellini by the above character combination.

Immature stages. The immature stages of Coccinellini are briefly described below with representative illustrations. For detailed descriptions of the morphology of the larvae and pupae of Coccinellini , see Tomaszewska & Ślipiński (2010) and Ślipiński et al. (2020).

Eggs ( Fig. 4 ). Members of Coccinellini typically lay oval and spindle-shaped eggs in small or large groups. The eggs are usually orange yellow or bright golden yellow and rarely cream or white and the surface is smooth and shiny without any microsculpture.

Larvae ( Fig. 5 ). General appearance alligator-like (‘campodeiform’), ground colour slate grey to dark chocolate brown or black or yellow or reddish, with red, orange, yellow, or white spots / patterns, dorsal surface sometimes pruinose / covered with a whitish dust. Larvae of mycophagous species usually bright lemon yellow, or pale greyish with black and yellow spots and patterns.

Pupae ( Fig. 6 ). Coccinellini pupae are exarate, though legs and other appendages are not fully free, with some degree of partial fusion and are usually found attached to the substrate with the last larval skin attached to the caudal end. The pupae of most Coccinellini are often brightly coloured with spots and other patterns.

Biology / prey preferences. In general, members of the tribe Coccinellini are aphidophagous and also feed on other Hemiptera such as psyllids, leaf- and planthoppers, whiteflies and early-stage larvae of Lepidoptera and also mites. Among Indian species, Anegleis cardoni (Weise) , Phrynocaria unicolor (Fabricius) and Phrynocaria perrotteti (Mulsant) show a propensity to feed on whiteflies in preference to aphids (label data, personal observations). Aiolocaria hexaspilota (Hope) and some species of Calvia are notable for being predatory on the eggs, larvae and pupae of chrysomelid beetles (Booth 1997). Species of Synona Pope are predators of plataspidid bugs ( Hemiptera ) in India and Australia ( Ślipiński 2007; Poorani et al. 2008) and Indian Synona spp. are well known predators of Coptosoma ostensum Distant , a major pest of pigeonpea ( Poorani et al. 2008). In India and elsewhere, Bulaea lichatschovii (Hummel) and B. lividula bocandei Mulsant feed on pollen and may be occasionally phytophagous (Poorani et al. 2018). Micraspis spp. are abundant in the paddy ecosystems of the Indian region and feed primarily on rice pollen and also on insect prey ( Poorani et al. 2023).

The molecular phylogenetic study on Coccinellini by Escalona et al. (2017) was focused on the reconstruction of the evolution of selected morphological traits and food preferences in Coccinellini . Their analysis supports the idea of aphidophagy being the ancestral state in Coccinellini . They suggested that evolution of host preferences in Coccinellini possibly involved multiple independent host switches from different insect orders to fungal spores and plant tissue and that prey diversification and switch to pollinivory, herbivory, mycophagy or predation on non-aphid hosts during periods of low aphid abundance in Coccinellini probably played a major role in their evolution.

Key to the genera of Coccinellini of Indian mainland

(Note. Schematic illustrations / photos of the major diagnostic characters used in the key are given in Figs 7–12 ).

1. Dorsum with distinct, short and dense pubescence, ground colour reddish with black maculae on pronotum and elytra ( Fig. 2a, b ). Abdominal postcoxal lines incomplete with an oblique associated line ( Figs 7b , 11e, f, k, m, p ). Distributed in the Himalayas (Sikkim) and Pakistan, rarely collected............................................................ Singhikalia

- Dorsum always glabrous, coloration variable ( Fig. 1a–h ). Abdominal postcoxal lines complete ( Fig. 11l ) or incomplete, with ( Fig. 11e, f, k, m, p ) or without ( Fig. 11g –j, n, o ) an oblique associated line. Distribution variable...................... 2

2. Mandibles apically bifid with ventral tooth / incisor edge having a row of small teeth ( Fig. 9g ). Anterior clypeal margin without distinct lateral projections or only barely produced. Antenna much longer than head width, with a loosely segmented club ( Fig. 9e ). Coloration predominantly yellow or creamy yellow with other patterns....................................... 3

- Mandibles large, apically bifid with a sclerotized basal mola bearing a prominent tooth ( Fig. 9h, i ). Anterior clypeal margin with distinct lateral projections ( Fig. 9b–d ). Antennal length and coloration variable.................................... 6

3. Terminal maxillary palpomere greatly transverse ( Fig. 10m ). Adults bright lemon yellow or creamy yellow, usually with and rarely without a pair of black spots on pronotum ( Fig. 1f ................................................... Illeis

- Terminal maxillary palpomere less strongly transverse ( Fig. 9f ). Adult coloration variable............................ 4

4. Adult very small (2.5 mm or less), creamy yellow, elytra with 18 black spots.Anterior margin of pronotum shallowly emarginate, covering only posterior half of eyes............................................................... Psyllobora

- Adult much larger (4.0 mm or longer), coloration and elytral pattern variable. Anterior margin of pronotum more or less straight, almost fully covering eyes....................................................................... 5

5. Male genitalia with parameres bilobed with a distinct subapical lobe, both lobes densely setose; penis capsule with inner process much reduced. Spermatheca c-shaped with distinct nodulus and ramus............................. Macroilleis

- Male genitalia with normal parameres, penis capsule with inner and outer processes almost subequal and distinct. Spermatheca c-shaped with nodulus and ramus distinct but produced to a lesser degree.................................... Halyzia

6. Size small (2.5–3.0 mm), orange- or yellow-coloured adults with black maculae. Anterior margin of prosternum medially prominent, forming a chin piece; prosternal intercoxal process at a distinctly elevated plane and carinate. Abdominal postcoxal line incomplete without an associated line. Distribution restricted to northeastern India, Myanmar, Southeast Asia...................................................................................................... Protothea

- Size larger (>3.0 mm), coloration variable. Anterior margin of prosternum not forming a chin piece; prosternal intercoxal process normal, T-shaped, with or without carinae. Abdominal postcoxal line complete or incomplete, with or without an associated line. Distribution variable...................................................................... 7

7. Form elongate oval, dorsum weakly to only moderately convex. Anterior margin of pronotum weakly excavated. Abdominal postcoxal lines complete and shallowly semicircular or absent. Legs long, distinctly projecting beyond outer margin of elytra. Tarsal claws double and cleft near apical one-third ( Fig. 11r ).......................................... Hippodamia

- Form broadly round, oval or elongate oval, dorsum moderately to strongly convex. Anterior margin of pronotum more strongly excavated. Abdominal postcoxal lines always present, complete or incomplete, with or without an associate line. Legs not unusually long. Tarsal claws appendiculate ( Fig. 11u, v ) or apically split ( Fig. 11t ), or almost simple ( Fig. 11q, s )......... 8

8. Scutellar shield small to very small (about 1/15 times as broad as elytra at shoulders or less), triangular................. 9

- Scutellar shield regular triangular or broader, transverse triangular, not very small (at least 1/10 times as broad as elytral width at shoulders or wider)................................................................................. 11

9. Form distinctly elongate oval. Antenna much longer than width of head capsule. Elytral epipleuron narrow. Prosternal intercoxal process narrow, without carinae ( Fig. 11a ). Mesoventrite anteriorly barely emarginate. Mid- and hind tibiae with a pair of apical spurs ( Fig. 11d ). Distributed mainly in the colder parts of northwestern India, Himalayas, Pakistan, Nepal and Bhutan. Adalia

- Form broadly rounded or oval.Antenna shorter than or at most only slightly longer than width of head capsule. Elytral epipleuron distinctly broader. Prosternal intercoxal process broader, with ( Fig. 11b ) or without carinae ( Fig. 11a ). Mesoventrite anteriorly distinctly but shallowly emarginate. Mid- and hind tibiae lacking apical spurs ( Fig. 11c ). Widely distributed............ 10

10. Form circular. Prosternal intercoxal process without carinae. Spermatheca vermiform without well differentiated cornu, nodulus and ramus..................................................................................... Anegleis

- Form broad or elongate oval to almost circular. Prosternal intercoxal process with a pair of carinae. Spermatheca not vermiform, with regular c-shaped cornu but ramus and nodulus often not well demarcated.............................. Micraspis

11. Antenna short, scape greatly enlarged and club spindle shaped, distinctly shorter than width of head capsule ( Fig. 10e ). Abdominal postcoxal lines incomplete with an associated oblique line. Male genitalia with penis apex thread-like and very long. Female genitalia with a very long, intricately coiled sperm duct................................... Cheilomenes

- Antenna distinctly longer than width of head capsule, scape not unusually enlarged ( Fig. 10b, d, i ). Abdominal postcoxal lines complete or incomplete with or without an associated line. Male genitalia with penis apex not thread-like, modified otherwise. Female genitalia with sperm duct not unusually long or coiled................................................. 12

12. Medium to large adults (7–15 mm) with explanate elytral margins. Elytral base distinctly much broader than pronotum... 13

- Medium to small adults (usually <7 mm), with or without explanate elytral margins. Elytral base not unusually broader than pronotum........................................................................................... 18

13. Anterior clypeal margin semicircularly emarginate between lateral projections. Abdominal postcoxal line incomplete with an associated line....................................................................................... 14

- Anterior clypeal margin straight between lateral projections. Abdominal postcoxal line incomplete without an associated line. .................................................................................................. 16

14. Medium sized adults (6.5–8.0 mm long), elytra with a prominent marginal bead. Head and pronotum fully or partially orange-yellow to reddish, elytra fully black ( Fig. 1g ). Spermatheca with a well-developed ramus but much reduced nodulus. Synona

- Large adults (9.0 mm– 15 mm long), elytra without a marginal bead. Coloration variable. Spermatheca with ramus much reduced, nodulus elongate tubular and snout-like........................................................... 15

15. Pronotum with steep and almost straight lateral sides, antero- and posterolateral corners acutely angulate. Spermatheca with a tubular and straight nodulus. Distributed in northwestern and eastern Himalayas, Pakistan; rarely collected...... Palaeoneda

- Pronotum with strongly rounded sides, posterolateral corners rounded. Spermatheca with a tubular but distinctly curved nodulus. Distributed in northwestern and northeastern India, Pakistan, Nepal and Bhutan; more commonly collected................................................................................................... Aiolocaria

16. Form broad around middle of elytra and distinctly narrower towards both ends. Ground colour bright carmine red with black maculae on pronotum and elytra. Sides of pronotum steep but gradually narrowed towards anterior, widest at posterior margin, posterolateral corners broadly rounded. Distribution restricted to northwestern and eastern Himalayas........... Callicaria

- Form broad oval to almost circular. Ground colour reddish or orange-yellow with black maculae. Sides and anterolateral corners of pronotum rounded. Distributed more widely, commonly associated with bamboo aphids.................... 17

17. Tarsal claws stout, double and apically split ( Fig. 11t )................................................ Synonycha

- Tarsal claws appendiculate ( Fig. 11u, v ).......................................................... Megalocaria

18. Prothoracic hypomera with distinct foveae near anterolateral corners ( Fig. 8c ).................................... 19

- Prothoracic hypomera without foveae near anterolateral corners............................................... 23

19. Male genitalia with penis consisting of more than one sclerite, one basal and a smaller apical piece, distinctly separated by membranes......................................................................................... 20

- Male genitalia with penis not having more than one sclerite, with a distinct capsule and a modified apex............... 21

20. Form broadly rounded. Ground colour bright lemon yellow to creamy yellow, pronotum with a pair of black spots, elytra with 10 large, black maculae. Anterolateral corners of pronotum sinuate / weakly emarginate before apex. Male genitalia with penis guide distinctly shorter than parameres. Distribution restricted to Himalayas and North-eastern India, Nepal and Bhutan.................................................................................................. Alloneda

- Form broad oval to almost circular. Color pattern variable, not as above. Anterolateral corners of pronotum normally rounded, rarely sinuate before apex. Male genitalia with penis guide usually elongate and as long as parameres. Widely distributed............................................................................................... Coelophora

21. Form broad to elongate oval. Female genitalia with spermatheca having distinct but short nodulus and ramus, infundibulum distinctly sclerotized. Distribution restricted to NE India, Nepal, Bhutan.................................. Microcaria

- Form elongate oval to almost circular. Female genitalia with spermatheca not having well differentiated nodulus and ramus, infundibulum less prominent. Widely distributed........................................................... 22

22. Form broadly rounded, dorsum strongly convex. Eyes large, narrowly separated with strongly divergent inner margins, interocular distance about one-third as wide as head including eyes, ocular canthus narrow and deep. Mesoventrite usually with a short longitudinal median carina not reaching beyond middle. Middle and hind tibiae without apical spurs ( Fig. 11c ). Female genitalia with coxites having distinct styli ( Fig. 12a )................................................. Phrynocaria

- Form distinctly more elongate oval, dorsum only moderately convex. Eyes smaller, more broadly separated with inner margins subparallel, ocular canthus moderately deep. Mesoventrite without a median carina. Middle and hind tibiae with a pair of apical spurs ( Fig. 11d ). Female genitalia with coxites having reduced and barely visible styli......................... Propylea

23. Tibial spurs absent. Abdominal postcoxal line incomplete laterally, usually with an oblique, curved associated line connected at its end ( Fig. 11f ); antenna short with a compact, broad club, terminal antennomere expanded, strongly transverse ( Fig. 10c ).............................................................................................. Harmonia

- Tibial spurs present. Abdominal postcoxal line incomplete with an associated line distinctly separated from basal line ( Fig. 11 m, p ) or incomplete without associated line ( Fig. 11g –j, n, o ). Antenna variable................................... 24

24. Abdominal postcoxal line incomplete with an oblique associated line distinctly separated from basal line ( Fig. 11m, p ).... 25

- Abdominal postcoxal line incomplete without an associated line, occasionally very short ( Fig. 11g, n )................. 26

25. Form elongate oval. Head black, always with a pair of yellow or white frontal spots. Mesoventrite anteriorly barely emarginate. Female genitalia with spermatheca always having wart-like projections in the proximal half. Male genitalia with penis capsule usually very large............................................................................. Coccinella

- Form usually broad oval to almost circular. Head yellow or black, without frontal spots. Mesoventrite distinctly but shallowly emarginate anteriorly. Female genitalia with spermatheca differentiated into a nodulus and often elongate and distally bulbous ramus, and a sclerotized infundibulum, without wart-like projections. Male genitalia with penis capsule regular, not unusually large......................................................................................... Oenopia

26. Eyes small, widely separated, interocular distance about 3x as wide as an eye ( Fig. 9a ). Antenna with a compact club, terminal antennomere distinctly transverse and quadrate ( Fig. 10g ). Terminal maxillary palpomere elongate securiform. Prosternal process without carinae. Abdominal postcoxal line apically recurved and incomplete. Tarsal claws simple ( Fig. 11q ) or almost simple, at most with a slight median projection ( Fig. 11s ). Female genitalia without infundibulum................ Bulaea

- Eyes much larger, interocular distance less than thrice eye width, with inner margins usually strongly divergent at both ends. Antenna long and loosely jointed, terminal antennomere elongate oval. Terminal maxillary palpomere regular securiform. Prosternal process with carinae. Abdominal postcoxal line incomplete, parallel to posterior margin of ventrite 1, occasionally very short. Elytra often with deep, distinctly dual punctation. Tarsal claws appendiculate. Female genitalia with infundibulum............................................................................................... Calvia

Figures

FIGURE 1. Map of the geographic spread of the Indian Subcontinent.

FIGURE 2. Representative images of Coccinellini (adult): a. Coccinella transversalis Fabricius; b. Megalocaria dilatata (Fabricius); c. Oenopia kirbyi (Mulsant); d. Synonycha grandis (Thunberg); e. Halyzia tschitscherini Semenow; f. Illeis cincta (Fabricius); g. Synona melanopepla (Mulsant); h. Harmonia octomacutata (Fabricius).

FIGURE 3. a, b. Singhikalia spp., the only Coccinellini genus with dorsal pubescence; c–h. Beetles similar to Coccinellini: c, d. Cyclotoma sp. (Endomychidae); e–h. Derispia coccinelloides (Westwood) (Tenebrionidae).

FIGURE 4. Eggs of Coccinellini: a. Cheilomenes sexmaculata (Fabricius); b. Megalocaria dilatata (Fabricius); c. Synonycha grandis (Thunberg) (image credit: H.M. Yeshwanth); d. Anegleis cardoni (Weise); e. Coccinella transversalis Fabricius; f. Illeis cincta (Fabricius); g. Micraspis tenuilinea (Walker); h. Propylea dissecta (Mulsant).

FIGURE 5. Larvae of Coccinellini: a. Calvia flaveola Booth; b. Propylea dissecta (Mulsant); c. Harmonia dimidiata (Fabricius); f.Coelophora bissellata (Mulsant); e. Illeis cincta (Fabricius); f. Coccinella transversalis Fabricius; g. Cheilomenes sexmaculata (Fabricius); h. Micraspis tenuilinea (Walker).

FIGURE 6. Pupae of Coccinellini: a. Cheilomenes sexmaculata (Fabricius); b. Coccinella transversalis Fabricius; c. Illeis bielawskii Ghorpade; d. Coelophora bissellata Mulsant; e. Calvia explanata Poorani; f. Harmonia octomaculata (Fabricius); g. Oenopia sauzeti (Mulsant); h. Anegleis cardoni (Weise).

FIGURE 7. Schematic illustration of the major diagnostic characters of Coccinellidae / Coccinellini:A. Dorsal view; B. Ventral view (Abbreviations: Em2—mesepimeron; Em3—metepimeron; Es2—mesanepisternum; Es3—metanepisternum; V1–V6: abdominal ventrite 1–6).

FIGURE 8. Diagnostic characters in Coccinellini: a. head, dorsal aspect; b. head, ventral aspect; c. prothorax, ventral view. (Abbreviations: AN—antenna; SC—scape; PD—pedicel; EY—eye; CL—clypeus; FR—frons; LB—labrum; ST—stipes; CA—cardo; MX—maxilla; MT—mentum; HP—prothoracic hypomeron; PST—prosternum; PSC—prosternal carina; FO— hypomeral fovea).

FIGURE 9. Key characters in various Coccinellini: a–f. Head: a. Bulaea; b. Anegleis; c. Coelophora; d. Harmonia; e. Illeis; f. Halyzia; g–i. mandibles: g. Illeis; h. Coccinella; i. Bulaea.

FIGURE 10. Representative illustrations of diagnostic characters: a–i. antenna: a. Adalia; b. Coelophora; c. Harmonia; d. Propylea; e. Cheilomenes; f. Hippodamia; g. Bulaea; h. Coccinella; i. Illeis; j–o. maxilla: j. Coccinella; k. Oenopia; l. Bulaea; m. Illeis; n. Adalia; o. Cheilomenes.

FIGURE 11. Representative illustrations of key characters in Coccinellini: a. prosternal process without carinae; b. prosternal process with carinae; c. tibia without apical spur; d. tibia with a pair of apical spurs; e–p. abdominal postcoxal line: e, f, k, m, p. incomplete with an associated line; g–j, n, o. incomplete without associated line; l. complete; q–v. tarsal claw types: q, s. simple; r, t. bifid / double; u, v. appendiculate.

FIGURE 12. Genitalia in Coccinellini: a. female genitalia; b. spermatheca; c–e. male genitalia: c. tegmen, lateral view; d. tegmen, ventral view; e. penis. (Abbreviations: bp—basal piece; bs—bursa; cx—coxite; cg—colleterial gland; co—cornu; inf—infundibulum; no—nodulus; rm—ramus; pa—penis apex; pc—penis capsule; pg—penis guide; pm—paramere; sp— spermatheca; sd—sperm duct; sg—spermathecal gland; ts—tegminal strut / trabes).