Coccidula Kugelann, 1798

Coccidula Kugelann, 1798: 421. Type species: Chrysomela scutellata Herbst, 1783, by subsequent designation by Crotch 1874.

Strongylus Panzer, 1813: 114.

Cacidula Dejean, 1821: 132. Type species: Chrysomela pectoralis Fabricius, 1792 (= Dermestes rufus Herbst, 1783).

Cacicula Stephens, 1831: 397.

Diagnosis.

Representatives of the genus Coccidula with its general body shape may resemble Tetrabrachys Kapur, however, it can be separated based on the structure of the tarsi which are tetramerous in both genera but in Coccidula the first tarsomere is sub-triangularly broadened apically and the second is elongate and distinctly lobbed, while in Tetrabrachys both the first and second are narrow, elongate and without lobes. Moreover, in Tetrabrachys the apical maxillary palpomere is widely securiform, and beetles are brachypterous, while in Coccidula the apical maxillary palpomere is only slightly widened and the second pair of wings is functional. Coccidula is also externally similar to European species of Rhyzobius but it can be separated based on the following characters: body almost parallel sided, elytra covered with punctures of two sizes, larger punctures arranged in nine rows (in C. litophiloides some of them are reduced), base of the pronotum not bordered, while in Rhyzobius the lateral body outline is broadly rounded, the elytra are covered with single sized, randomly arranged punctures, and base of the pronotum with distinct bordering line.

Description.

Body elongate-oval, with sides parallel (Fig. 1C-H), body flattened in lateral view, convex in cross-section; dorsum covered with setiferous punctures of two sizes (Figs 7A, 9A), hairs directed forwards on pronotum, backwards on elytra.

Head partially withdrawn into prothorax (Fig. 1C-H); ventral antennal grooves shallow and moderately long, extending to posterior border of an eye (Fig. 3E). Eyes prominent, coarsely facetted (7-8 ommatidia per eye width), ocular canthus distinct, about as long as 4-5 ommatidium diameters; interocular distance about 3 × as eye diameter; interfacetal setae present only in basal part; temple behind eye distinctly longer than eye (Fig. 3E). Antennal insertion placed laterally, invisible from above, distance between antennal insertions about same as between eyes; frons around antennal insertions slightly expanded, covering antennal insertions, anterior tentorial pits placed ventrally below antennal insertions. Antennae (Figs 5A, E, 7A, D) longer than maximum head width including eyes, composed of 11 antennomeres (AN); scape simple, without projections, slightly curved; pedicel distinctly narrower than scape, elongate (1.5 × longer than wide); AN 3-8 elongate (AN3 ≈ 3.5 ×; AN8 ≈ 1.3 × longer than wide); AN 9-11 forming a loose, asymmetric club, ultimate AN truncate apically. Frontoclypeus short, transverse, anterior margin straight. Labrum entirely exposed, transverse, anterior margin straight. Mandibles asymmetric, bifid apically (Fig. 10G), molar part with basal tooth; prostheca distinct. Maxillary stipes (Figs 2B, 5C, 9C, 10H) with distinct groove for reception of maxillary palp in repose; palpomere 2 shorter than terminal (4th) one, slightly broadened apically; palpomere 3 about 2.3 × shorter than terminal one, subtriangular; terminal palpomere slightly securiform; lacinia with stiff setae on outer margin in apical half, with several additional spurs on surface (Fig. 10H). Labial palps (Figs 3E, 9C) with 3 palpomeres, inserted ventrally on prementum; palpomere 1 very small, apical palpomere as long as and about as broad as penultimate; distance between palp insertions about 1.5-2 × as its width. Prementum subquadrate, transverse apically. Mentum trapezoidal, broadest in anterior part, with horseshoe impression at base (Figs 7C, 9C). Submentum broad, transverse, with suture invisible.

Anterior margin of pronotum weakly, broadly emarginate (Figs 3B, 5B) with anterior corners broadly rounded; lateral margins with moderately (Figs 7B, 9B) to distinctly expanded lateral beads (Fig. 3B), distinctly margined; hind corners sharply pointed; hind margin not bordered. Prothoracic hypomeron smooth, without delimited foveae (Figs 3C, 7C). Prosternum in front of coxae about as long as longitudinal length of procoxal cavity; anterior margin straight or slightly emarginate with distinct border. Prosternal process about 0.4 times of coxal diameter, surface smooth (Fig. 3C) or with lateral carinae (Figs 7C, 9E). Procoxal cavity oval, distinctly bordered anteriorly.

Mesoventrite 1.3 × longer than its width at the level of mid coxae (Figs 1D, 5D, 7D); mesal surface with deep emargination for receiving tip of prosternal process (Fig. 2C); anterior margin with completely raised border. Meso-metaventral process narrow (Figs 1D, 2C, 5D, 7E), about 0.5 times of mesocoxal diameter, junction slightly arcuate (Figs 2C, 3D, 5D, 7E, 9D), with suture visible. Metendosternite with stalk sub-quadrate, tendons long, separated by a distance of about width of stalk and situated closer to center (Fig. 10I). Scutellar shield pentagonal (Figs 7B, 9B). Elytra at base wider than pronotum, lateral margins clearly visible from above throughout (Figs 2D, 3A, 5A, 7A) (except C. scutellata where it is obscured in basal part, Fig. 9A), surface covered with punctures of double size, smaller irregularly distributed, larger punctures arranged in nine irregular longitudinal rows. Sutural stria absent. Elytral epipleuron narrow, incomplete, reaching base of ventrite 4 (Fig. 1D), with complete bordering line, epipleural foveae absent. Hind wings fully developed or missing (in C. litophiloides). Metaventral postcoxal lines roundly joined medially, complete laterally, straight or descending (Figs 2C, 3D, 5D, 7E, 9D). Metaventrite with discrimen visible in posterior 2/3.

Trochanters simple, subtriangular, without projection (figs 7E, 9D). Tibiae slightly expanded apically with one apical spur on forelegs, and two in mid and hind legs. Tarsi consisting of four tarsomeres, second tarsomere truncate apically; tarsal claws cleft apically (Fig. 9G) with single empodial seta present.

Abdomen in both sexes with 6 ventrites (Fig. 1D); ventrite 1 about as long as ventrites 2-4 combined, ventrite 2 longer than ventrite 3, ventrites 3-5 subequal in length. Abdominal postcoxal lines (Figs 7E, 9D) separate medially, recurved and complete, reaching anterior margin of ventrite, posteriorly reaching about half length of ventrite 1. Ventrite 5 in female posteriorly rounded (Fig. 7F), in male truncate (Fig. 9F). Ventrite 6 rounded in both sexes.

Male terminalia. Tegmen (Figs 4A, B, 8A, B, 10A, B) symmetrical; parameres articulated with penis guide. Penis (Figs 4C, 8C, 10C) slender, pointed apically; penis capsule asymmetrical with outer arm reduced, inner arm well developed. Apodeme of male sternum IX simple, not broadened apically (Figs 8D, 10D). Tergite X broadly rounded, semicircular (Figs 8D, 10D).

Female terminalia. Coxites (Figs 10F) distinctly elongate, subtriangular; styli small but visible, bearing several short setae; infundibulum absent (Figs 4D, 8E, 10E); sperm duct simple. Spermatheca (Figs 4D, 8E, 10E) worm-like, without clear ramus or nodulus; spermathecal accessory gland small, elongate. Proctiger elongate, rounded apically (Fig. 10F).

Immature stages.

Larva as in Fig. 1A, pupa as in Fig. 1B.

Distribution.

Holarctic: Asia, Europe, Africa (North), North America.