Cassida Linnaeus 1758
Creators: name: Borowiec, Lech
name: Świętojańska, Jolanta
Publisher: name: Zenodo
Year: 2022
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Coleoptera
Chrysomelidae
Cassida

description: Genus Cassida Linnaeus, 1758
Cassida Linnaeus, 1758: 362 (type species: Cassida nebulosa Linnaeus, 1758, designated by Spaeth, 1914 b: 92); Chapuis, 1875: 388; Spaeth, 1914 b: 92; Gressitt, 1952: 484; Hincks, 1952: 339; Gressitt & Kimoto, 1963: 963; Brovdii, 1983: 74; Lopatin & Kulenova, 1986: 180; Chen et al., 1986: 456, 472; Gruev & Tomov, 1986: 344; Riley, 1986: 102; Borowiec, 1990 b: 4, 1994 a: 17, 1999 a: 234.
Deloyala Redtenbacher, 1858: 952 (type species: Cassida seraphina Ménétries, 1836, designated by Hincks, 1950: 508); Spaeth, 1914 b: 93, not Deloyala Duponchel & Chevrolat, 1843. Palearctic Region.
Cassidula Weise, 1889: 260 (type species: Cassida nobilis Linnaeus, 1758, designated by Hincks, 1952: 339); Spaeth, 1914 b: 97, not Cassidula de Blainville, 1830. Palearctic Region.
Pseudocassida Desbrochers, 1891: 15 (as subgenus, type species: Cassida murraea Linnaeus, 1768, by monotypy); Hincks, 1952: 339; Seeno & Wilcox, 1982: 177, subgenus. Palearctic Region.
Mionycha Weise, 1891: 204 (type species: Cassida azurea Fabricius, 1801, by monotypy); Spaeth, 1914 b: 95; Gressitt, 1952: 503; Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Brovdii, 1983: 90; Chen et al., 1986: 468, subgenus. Palearctic Region.
Odontionycha Weise, 1891: 204 (type species: Cassida viridis Linnaeus, 1758, designated by Hincks, 1952: 339); Spaeth, 1914 b: 93; Gressitt, 1952: 521; Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Brovdii, 1983: 84; Chen et al., 1986: 457, subgenus. Palearctic Region.
Crepidaspis Spaeth, 1912 b: 119 (type species: Crepidaspis varicornis Spaeth, 1912, by monotypy), 1914 b: 129; Hincks, 1952: 340 (as syn. of Thlaspidosoma Spaeth, 1901); Seeno & Wilcox, 1982: 177 (in syn. of Thlaspidosoma); Borowiec, 1990: 4 (as subgenus of Cassida). Oriental Region.
Taiwania Spaeth, 1913: 47 (type species: Taiwania sauteri Spaeth, 1913, by monotypy), 1914 b: 144; Gressitt, 1952: 486 (as subgenus); Hincks, 1952: 339 (as subgenus); Seeno & Wilcox, 1982: 177 (as genus); Chen et al., 1986: 490, 504 (as genus); Borowiec, 1990: 4 (as syn. of Crepidaspis). Oriental Region.
Eremocassis Spaeth in Spaeth and Reitter, 1926: 15 (type species: Eremocassis transcaspica Spaeth, 1926, by monotypy); Hincks, 1952: 338; Lopatin, 1977: 247; Seeno & Wilcox, 1982: 176; Lopatin & Kulenova, 1986: 178; Borowiec, 1994 b: 157 (as syn. of Tylocentra Reitter, 1926). Palearctic Region.
Lordicassis Reitter in Spaeth & Reitter, 1926: 23, 27 (type species: Cassida undecimnotata Gebler, 1841, by monotypy); Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Chen et al., 1986: 462, subgenus. Palearctic Region.
Tylocentra Reitter in Spaeth & Reitter, 1926: 24, 57 (type species: Cassida turcmenica Weise, 1892, designated by Hincks, 1952: 339); Gressitt, 1952: 525; Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Medvedev, 1982: 286 (as genus); Chen et al., 1986: 469, subgenus. Palearctic Region.
Lordiconia Reitter in Spaeth & Reitter, 1926: 23, 26 (type species: Cassida canaliculata Laicharting, 1781, by monotypy); Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Brovdii, 1983: 82, subgenus. Palearctic Region.
Onychocassis Spaeth in Spaeth & Reitter, 1926: 23, 26 (type species: Cassida brevis Weise, 1884, designated by Hincks, 1952: 339); Hincks, 1952: 339; Seeno & Wilcox, 1982: 177, subgenus. Palearctic Region.
Cassidulella Strand, 1928: 2 (new name for Cassidula Weise, 1889 not de Blainville, 1830; type species: Cassida nobilis Linnaeus, 1758, designated by Hincks, 1952: 339); Gressitt, 1952: 523; Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Brovdii, 1983: 156; Chen et al., 1986: 464, subgenus. Palearctic Region.
Alledoya Hincks, 1950: 508 (new name for Deloyala Redtenbacher, 1858 not Duponchel and Chevrolat, 1843; type species: Cassida seraphina Ménétries, 1836, designated by Hincks, 1950: 508), 1952: 339; Seeno & Wilcox, 1982: 177; Lopatin & Kulenova, 1986: 186 (as genus); Chen et al., 1986: 547 (as genus). Palearctic Region.
Mionychella Spaeth in Hincks, 1952: 346 (type species: Cassida hemisphaerica Herbst, 1799, by monotypy); Hincks, 1952: 339; Seeno & Wilcox, 1982: 177; Brovdii, 1983: 88, subgenus. Palearctic Region.
Lasiocassis Gressitt, 1952: 485 (new name for Deloyala Redtenbacher, 1858 not Duponchel & Chevrolat, 1843; type species: Cassida vespertina Boheman, 1862), proposed as subgenus. Palearctic Region.
Cyclocassida Chen & Zia, 1961: 442 (type species: Taiwania (Cyclocassida) variabilis Chen & Zia, 1961, by original designation), proposed as subgenus of Taiwania; Seeno & Wilcox, 1982: 177 (as subgenus of Taiwania); Chen & al., 1986: 491 (as subgenus of Taiwania); Borowiec, 1990: 4 (as syn. of Crepidaspis). Oriental Region.
Yunocassis Chen & Zia, 1961: 442 (type species: Cassida appluda Spaeth, 1926, by original designation), proposed as subgenus of Taiwania; Seeno & Wilcox, 1982: 177 (as subgenus of Taiwania); Chen & al., 1986: 500 (as subgenus of Taiwania); Borowiec, 1990: 4 (as syn. of Crepidaspis). Oriental Region.
Cyrtonocassis Chen & Zia, 1961: 446 (type species: Cyrtonocassis tumidicollis Chen & Zia, 1961, by original designation); Seeno & Wilcox, 1982: 177 (as genus); Chen & al., 1986: 546 (as subgenus of Taiwania), subgenus. Oriental Region.
Dolichocassida Günther, 1958: 568 (type species: Cassida pusilla Waltl, 1839 = Dolichocassida veselyi Günther, 1958, by monotypy); Seeno & Wilcox, 1982: 177, subgenus. Palearctic Region.
Pseudocassis Steinhausen, 2002: 24 (type species: Cassida flaveola Thunberg, 1794, by original designation), subgenus. Palearctic Region.
Betacassida Steinhausen, 2002: 26 (type species: Cassida nebulosa Linnaeus, 1758, by original designation), proposed as subgenus, objective synonym of Cassida Linnaeus, 1758. Palearctic Region.
Coptocycla auct., part; Spaeth, 1914 b: 130, part (Old World species).
Metriona auct., part; Spaeth, 1914 b: 136, part (Old World species).
Description. Very heterogenous genus. Length from 2.5 to 11.6 mm, body varying from parallel sided to circular, from depressed to strongly convex, without or with postscutellar gibbosity or angulation, rarely with conical postscutellar tubercle. Base of elytra not to distinctly wider than pronotum. Pronotal sides from angulate to rounded, usually without distinct basal corners, occasionally with basal corners protruding posterad, disc of pronotum more or less bordered from explanate margin, surface of disc from smooth and shiny to distinctly punctate, or with granulate or striate sculpture with shiny, or microreticulate, dull background. Explanate margin of pronotum from narrow to very broad, from smooth to distinctly punctate or sculptured, often transparent with honeycomb structure. Scutellum more or less triangular, always well visible. Punctation of elytral disc from regular to completely irregular, intervals, if present, or interspaces with surface from smooth and shiny or microreticulate to punctate, dull. Explanate margin of elytra from strongly declivous to subhorizontal, from very narrow to very broad, with surface from smooth and shiny to punctate or sculptured with microreticulate and dull interspaces, often transparent with honeycomb structure. Clypeus from narrow to broad, usually flat to slightly convex, occasionally with distinct apical angulation. Clypeal grooves usually faint, sometimes deep, complete, converging in triangle or arch, or more or less shortened and visible only at base of clypeus. Surface of clypeal plate from smooth and shiny to punctate, or microreticulate and dull. Antennae from short to moderately long, the third antennomere usually longer than the second. Venter of pronotum without antennal grooves along sides of head at most with shallow channels. Sides of prosternal collar without excavation. Prosternal process usually narrow but more or less expanded apically, with or without median channel, its surface from smooth and shiny to punctate or sculptured, especially expanded part with more or less developed sculpture. Meso- and metathorax without gibbosities or angulations. Legs short, last segment of tarsi not or insignificantly longer than third segment, tarsomeres usually with distinct, hairy sole. Claws from simple to appendiculate, often apparently appendiculate due to distally projecting flanks of claw segments.
Diversity. With 458 described species it is the most speciose and most diverse genus in the subfamily Cassidinae. It is distributed in almost the entire Old World (except New Zealand) but with only one native species in North America. Such a large genus is difficult to characterize with a short diagnosis. Several subgenera have been proposed for the genus Cassida but exclusively for Palearctic and Oriental taxa. Borowiec (2007) discussed all subgeneric names described in the genus Cassida and proposed rejection of all names. He proposed grouping species into morphologically similar species-groups, not necessarily monophyletic but practical in identifying and creating comparative diagnoses that allow easier understanding of the species richness of this large genus. Below, based on morphology, we propose the following species groups for African members of the genus Cassida:
Cassida aethiopica species-group: Cassida aethiopica Boheman, 1854 and C. amorifica Boheman, 1862 — small species, length below 6 mm, body elongate oval; pronotal and elytral disc yellow with black pattern, explanate margin always yellow, ventrites usually partly brown to black, only the palest specimens completely yellow; pronotum elliptical, sides broadly rounded, no basal corners, surface of disc with very fine and sparse punctation; elytral punctation arranged in regular rows, but regularity of rows sometimes disturbed by yellow relief, disc without special sculpture, sometimes yellow parts sligthly elevated foms a relief but elytra in profile depressed, without hump or H-shaped postscutellar elevation; explanate margin narrow, moderately to strongly declivous, with fine and very dense punctation; claws simple.
Cassida andreinii species-group: Cassida andreinii Spaeth, 1933 and Cassida heroni sp. nov. —small cassids, length below 4.6 mm; dorsum predominantly green to yellow, at most with small patches in postscutellar point, behind humeral callus and in posterolateral part of disc, ventrites yellow; body oval to broadly-oval; pronotum elliptical, sides broadly rounded, no basal corners; base of elytra not or only slightly wider than pronotum, disc regularly convex, without hump, at most with short transverse fold in postscutellar point, elytral rows regular; explanate margin narrow, strongly declivous, surface with shallow but coarse and moderately dense punctation, appears more or less irregular; claws with large basal tooth.
Cassida coagulata species-group: Cassida coagulata Boheman, 1854 and C. irrorata Weise, 1898 —large, length above 6.4 mm; body strongly convex, hemispherical to cylindrical; colour from yellowish brown to brown, elytra with dark brown to black pattern of small spots; pronotum broadly rounded on sides without basal corners, pronotal disc impunctate, at most with fine pricks; base of elytra not or only slightly wider than pronotum, surface of elytra with hardly marked postscutellar H-shaped elevation, without postscutellar hump, elytral sculpture hardly marked of partly elevated intervals on top of elytra, elytral rows regular, only slightly disturbed by elytral sculpture, explanate margin of elytra narrow, strongly declivous; claws simple.
Cassida flavosignata species-group: Cassida benguelica Spaeth, 1933, Cassida flavosignata Spaeth, 1932 and C. leleupi sp. nov. —small species, length below 5 mm; pronotal and elytral disc mostly black with yellow pattern of numerous spots, sometimes posterior spots marked with pink to red, explanate margins and ventrites always yellow; body short-oval to almost circular; pronotum elliptical, sides broadly rounded, no basal corners; base of elytra distinctly wider than pronotum, disc slightly, regularly convex, without hump, without H-shaped elevation, only yellow spots more or less elevated, elytral rows regular but regularity disturbed by elevated yellow spots; explanate margin broad, only slightly declivous, shallowly and densely punctate, appears slightly irregular; claws simple or with moderate basal tooth.
Cassida lacrymosa species-group: Cassida bamendana sp. nov., C. callosicollis Spaeth, 1926, C. camerunensis (Spaeth, 1903), C. delenifica Boheman, 1862, C. depicta Boheman, 1862, C. elgonensis sp. nov., Cassida eluta Boheman, 1862, C. fuscosignata Boheman, 1854, C. humerosa Spaeth, 1902, C. innotata Boheman, 1854, C. lacrymosa Boheman, 1854, C. namibiensis Borowiec, 2005, C. natalensis Spaeth, 1932, C. irregularis Boheman, 1854, C. ghesquieri Spaeth, 1943, C. schoutedeni Spaeth, 1932, C. setosa Chapuis, 1880, C. spartea Shaw, 1961 and C. sparteiformis sp. nov. —small to moderate species, length below 5.9 mm; colouration varies within and between species, from uniformly yellow to maculate, with reddish, brown, black pattern, or disc of elytra black with yellow relief spots, in extreme case entire pronotal and elytral disc black, ventrites from uniformly yellow to mostly black; base of elytra slightly to moderately wider than pronotum, disc from slightly depressed to slightly convex, without hump, but with more or less marked H-shaped elevation, punctation coarse and dense, mostly regular, sometimes regularity disturbed by elytral relief, intervals narrow, often linear, usually inner intervals slightly elevated, in species with black and yellow pattern yellow spots elevated, form a relief; claws variable, from simple to simple but appearing slightly appendiculate due to distally projecting flanks of last tarsal segment or with distinct basal tooth. This group is close to several species of Cassida described from the Oriental Region in the genus Taiwania Spaeth, 1913 (sensu Chen et al. 1986 and Chen & Zia 1961). Borowiec (1990) synonymized the genus Taiwania Spaeth, 1913 with the genus Crepidaspis Spaeth, 1912 (included only species close to Taiwania sauteri Spaeth, 1913 —the type species for the genus Taiwania) and later (Borowiec 2007) proposed to treat all species from the Taiwania circle (including Taiwania s. str., Crepidaspis Spaeth, 1912, Cyclocassida Chen & Zia, 1961 and Yunocassis Chen & Zia, 1961) as only single species-group within the large and heterogenous genus Cassida.
Cassida litigiosa species-group: Cassida chrysanthemoides Borowiec & Świętojańska, 2001, C. cordula Boheman, 1854, C. distinguenda Spaeth,1928, C. diversepunctata Borowiec&Świętojańska,2001, C. foveolatipennis Borowiec & Świętojańska, 2001, C. limpopoana Borowiec & Świętojańska, 2001, Cassida litigiosa Boheman, 1854, C. lycii Borowiec & Świętojańska, 2001, C. melanophthalma Boheman, 1854, C. oxylepiformis sp. nov., C. pudens Boheman, 1854, C. reticulipennis Borowiec & Świętojańska, 2001, C. spatiosa Spaeth, 1928, C. spatiosiformis Borowiec & Świętojańska, 2001 and C. wittmeri sp. nov. — small to medium-sized cassids up to 6.8 mm in length; dorsal colouration uniformly green or yellow or occasionally green with small yellow spots, ventrites uniformly yellow; body from elongate-oval to almost circular, elytra always regularly convex or slightly depressed with no tubercles or impressions; elytral punctation always irregular but with distinct and regular marginal row, explanate margin of elytra indistinctly bordered from disc, strongly to moderately declivous, never horizontal, punctate. This group is restricted to the southern Africa, the northern most distributed species is known from Malawi, most of them are restricted to the Cape Province of the Republic of South Africa and Namibia.
Cassida sphaerula species-group: Cassida guttipennis Boheman, 1862, C. quatuordecimsignata Spaeth, 1899, C. sphaerula Boheman, 1854 and C. vespertilio Boheman, 1862, small species, length below 5.6 mm—pronotum usually yellow, only in the darkest forms mostly black, elytral disc usually with black pattern of small spots, sometimes also stripes along sides, explanate margin of elytra yellow or partly black; pronotal sides subangulate to narrowly rounded, no basal corners; base of elytra not to moderately wider than pronotum; elytral disc regularly convex, without hump, without H-shaped elevation; elytral rows regular; explanate margin narrow to moderately broad; claws simple; associated with Asteracae plants; distribution restricted to South Africa.
Cassida successiva species-group: Cassida stipidosa sp. nov. and C. successiva Spaeth, 1924 —small species, length below 4.8 mm; dorsum yellow, pronotum at most with small brown spot in front of scutellum, elytra with brown to black pattern limited to punctures or at most punctures and areolae around punctures, no spots or reticulate pattern, ventrites partly brown to black; pronotum elliptical, sides broadly rounded, no basal corners; base of elytra moderately to distinctly wider than pronotum, rows of punctures partly regular, party irregular, regularity disturbed by elytral sculpture; postscutellar H-shaped elevation distinctly marked, intervals on top of elytra more or less elevated but surface of elytra appears regular; claws with large basal tooth.
Cassida sulphurago species-group: Cassida imitatrix Spaeth, 1916, C. sulphurago Boheman, 1854 and C. sulphurea Boheman, 1854 —body moderately large to large, length 5.0–7.5 mm; dorsum and ventrites always uniformly yellow; pronotum narrowly to broadly rounded on sides without basal corners, surface of disc shiny, top of disc and sides with fine and sparse punctation; base of elytra moderately to distinctly wider than pronotum, rows of punctures completely regular, sculpture absent or indistinct, limited to slightly elevated intervals on top of disc; claws with or without basal tooth.
Cassida tosta species-group: Cassida altiuscula Spaeth, 1916, C. inaequalis Thomson, 1858, C. overlaeti sp. nov. and C. tosta Klug, 1835 —body large, length always above 6.5 mm, usually above 7.0 mm; colour dark, red to black including explanate margin; pronotum broadly rounded on sides without basal corners, pronotal disc impunctate; base of elytra distinctly wider than pronotum, surface of elytra with strong sculpture of folds, tubercles and ridges, postscutellar hump always well marked, regularity of elytral rows disturbed by sculpture; claws simple but with widened base.
Cassida troglodytes species-group: Cassida satanas sp. nov. and C. troglodytes Boheman, 1854 —small species, length below 4.6 mm; body colouration variable, dorsum from uniformly yellow to mostly black, often mostly purple - red, postscutellar elevation often with dark spot, sometimes posterior part of elytral disc with few very small dark spots, explanate margins from uniformly yellow or purple - red to mostly black with yellow spots, ventrites from mostly yellow to mostly black; body almost circular; pronotum with subangulate to angulate sides but without basal corners; base of elytra as wide as pronotum, disc convex, with well marked postscutellar hump, more or less distinct H-shaped elevation and slightly elevated inner intervals, elytral rows regular but regularity slightly disturbed by elytral sculpture; explanate margin moderately broad, moderately declivous, shallowly and densely punctate, appears slightly irregular; claws simple.
Cassida turcmenica species-group: Cassida pellegrini Marseul, 1868 and C. rothschildi Spaeth, 1922 —small species, length below 5.3 mm; body strongly convex, subtriangular in outline, with highest point in the postscutellar area but without hump; dorsal colour mostly to completely green or yellow, pattern, if present, reddish forms small spot in postscutellar point, sometimes also base of elytra with few small reddish spots, occasionally suture behind postscutellar point with very small reddish spots or only sutural elevation marked with red, ventrites yellow; pronotum elliptical, with maximum width in the middle, sides subangulate to angulate, pronotal disc distinctly punctate; base of elytra only slightly wider than pronotum, surface of elytra without postscutellar H-shaped elevation, elytral rows completely regular, intervals flat wider than rows, explanate margin of elytra narrow, strongly declivous; claws simple. Species from this group in former classification belong to the subgenus Tylocentra Reitter, 1926. The group comprises several Palaearctic species associated with desert habitats and plants of the genus Lycium L. (Solanaceae).
Cassida unimaculata species-group: Cassida drakensbergensis Borowiec, 2005 C. emontinensis sp. nov. and C. unimaculata Boheman, 1854 —moderately sized species, length 5.2–6.2 mm; dorsal colouration yellow, dark pattern limited to a small spot or stripe in postscutellar point, sometimes part of elytral punctures with dark centre, ventrites yellow; pronotum elliptical, sides broadly rounded, no basal corners; base of elytra slightly wider than pronotum; body almost circular in outline; elytra strongly convex with peak of convexity in postscutellar area but without hump and without special sculpture, surface of elytra completely regular; elytral punctation always regular; claws simple. The group is restricted to southern Africa.
Cassida viridipennis species-group: Cassida devylderi Spaeth, 1928, C. franklinmuelleri Spaeth, 1925, C. granula Boheman, 1856, C. kantnerorum sp. nov., C. mashonensis Spaeth, 1928, C. rhodesiaca Spaeth, 1928, C. sublesta (Weise, 1904) and C. viridipennis Boheman, 1854 —moderately sized species, length 4.65–5.80 mm; dorsum green or yellow, without pattern or with reddish to brown pattern of small spots and lateral stripes but never with black spots, ventrites always yellow; pronotum with more or less rounded sides, without basal corners, pronotal disc usually distinctly punctate or at least with pricks; base of elytra not or only slightly wider than pronotum, elytral disc always with more or less marked H-shaped elevation but without hump, punctation coarse, from mostly regular to almost completely irregular, sometimes only postscutellar impressions and area behind H-shaped elevation irregularly punctured, intervals usually very narrow, linear; explanate margin narrow to moderately broad, moderately declivous, surface with coarse and moderately dense punctation, appears irregular to rugose; claws with large basal tooth.
An additional 29 species (even 28.7%) each has a unique morphology and it is difficult to assign them to any of the proposed species-groups.
The biology of African members is poorly known. Host plants are known only for 29 species (28%). Most biological data refer to South African members of Cassida. The most preferred host plant families are Asteraceae, Amaranthaceae, Chenopodiaceae and Solanaceae (Muir & Sharp 1904, Heron & Borowiec 1997, Heron 2003, Rice 2003).
Key to species
1. Large species, length always above 6.4 mm, usually above 7.0 mm (up to 9.35 mm). Dorsum dark coloured, often partly to mostly black (figs. 51 –54, 57, 62) or partly reddish to reddish brown (figs. 55, 58, 61, 63–66, 72–74), if mostly or completely yellow to ochraceous (figs. 60, 67, 68, 69–71) then elytra with strong sculpture of numerous folds, wrinkles or costae and well marked postcutellar H-shaped elevation or hump (figs. 60, 67, 68) or elytra with pattern of numerous brown to black spots (figs. 69 –74, 176, 177).................................................................................... 2.
- Usually smaller species, length below 6.0 mm, if larger (up to 7.6 mm) then dorsum mostly uniformly yellow (figs. 218, 256 –261), green or yellowish green (figs. 317–319, 331, 332), or yellow with small black postscutellar spot (fig. 327) and elytra without distinct sculpture or hump..................................................................... 20.
2. Elytra mostly to completely black (figs. 51 –54, 57, 62)...................................................... 3.
- Elytra mostly to completely ochraceous yellow to red (figs. 58, 60, 61, 63 –74, 176, 177)........................... 6.
3. Elytral disc without (fig. 56) or with low postscutellar hump (fig. 59), elytra widest at 1/3 length then gently converging posterad (figs. 54, 62)................................................................................ 4.
- Elytral disc with high postscutellar hump (fig. 53), elytra widest behind humeri then strongly converging posterad (figs. 51, 52). Central Africa from Cameroon to Rwanda (fig. 2).............................................. altiuscula Spaeth
4. Postscutellar elevation very low, not forming hump (figs. 56, 65). Elytral disc behind humeral callus shallowly impressed, folds on elytral slope low, straight........................................................................... 5.
- Postscutellar elevation low but distinct, forming an obtuse hump (fig. 59). Elytral disc behind humeral callus deeply impressed, folds on elytral slope high, often S–shaped or irregular. Forest regions of West and Central Africa (fig. 47)................................................................................................. tosta Klug, black form
5. Punctation of explanate margin of elytra coarse and dense, surface appears strongly irregular. Sculpture on elytral slope higher thus surface appears distinctly irregular (figs. 54-56). Democratic Republic of Congo (fig. 32)..................................................................................................... overlaeti sp. nov., black form
- Punctation of explanate margin of elytra moderately coarse and dense, surface appears moderately irregular. Sculpture on elytral slope higher thus surface appears moderately irregular (figs. 63-66). Western and Central Africa from Guinea to Uganda (fig. 22)................................................................... inaequalis Thomson, black form
6. Surface of pronotal disc smooth or minutely punctate....................................................... 7.
- Surface of pronotal disc granulate (fig. 218). South Africa and southern Mozambique, probably introduced and established in Kenya (fig. 19).......................................................... granulicollis Spaeth, large specimens
7. Surface of elytra with strong sculpture of folds or/and costae, H–shaped postscutellar elevation distinct, dorsum often dark, reddish, brown to partly black (figs. 58, 60, 61, 63, 64, 66, 67, 69, 177) or disc with hump (fig. 68), if elytra without distinct sculpture then with distinct pattern of small dark spots (figs. 69 –74)............................................ 8.
- Surface of elytra without strong sculpture, without or with hardly marked postscutellar H–shaped elevation, without hump, dorsum mostly or completely yellow or yellowish green (figs. 256 –261, 317–319, 326–328, 331, 322)............... 16.
8. Dorsum uniformly yellow, ochraceous or red (figs. 60, 63, 67, 68)............................................. 9.
- Dorsum partly black (figs. 55, 57, 58, 61, 66) or yellow to brown with dark pattern (figs. 69 –74, 177, 178)............ 11.
9. Elytra without or with low postscutellar hump (figs. 59, 65). Elytral base distinctly wider than pronotum, sides of pronotum broadly rounded (figs. 60, 63)......................................................................... 10.
- Elytra with high, tuberculate postscutellar hump (fig. 68). Elytral base only slightly wider than pronotum, sides of pronotum narrowly rounded (fig. 67). Tanzania (fig. 2).................................................... blanda Spaeth
10. Postscutellar elevation very low, not forming hump (fig. 65). Elytral disc behind humeral callus shallowly impressed, folds on elytral slope low, straight (fig. 63). Western and Central Africa from Guinea to Uganda (fig. 22).................................................................................................. inaequalis Thomson, red form
- Postscutellar elevation low but distinct, forming obtuse hump (fig. 59). Elytral disc behind humeral callus deeply impressed, folds on elytral slope high, often S–shaped or irregular (fig. 60). Forest regions of West and Central Africa (fig. 47)................................................................................ tosta Klug, red or ochraceous form
11. Elytra strongly sculptured, with more or less marked H–shaped postscutellar elevation, longitudinal and irregular folds, colouration partly black (figs. 55, 57, 58, 61, 66).......................................................... 12.
- Elytra gently sculptured, with very low H–shaped postscutellar elevation, surface less distinct sculptured sometimes without folds but with longitudinal costae, background colouration mostly yellow to ochraceous–reddish, dorsum with pattern of small, brown to black numerous spots (figs. 69 –74, 176–178)..................................................... 14.
12. Postscutellar elevation very low, not forming hump (figs. 56, 65). Elytral disc behind humeral callus shallowly impressed, folds on elytral slope low, straight.......................................................................... 13.
- Postscutellar elevation low but distinct, forming an obtuse hump (fig. 59). Elytral disc behind humeral callus deeply impressed, folds on elytral slope high, often S–shaped or irregular. Explanate margin of elytra often mostly black with red spot in the middle (figs. 57, 58, 61). Forest regions of West and Central Africa (fig. 47).............. tosta Klug, black and red form
13. Punctation of explanate margin of elytra coarse and dense, surface appears strongly irregular. Sculpture on elytral slope higher thus surface appears distinctly irregular. Pronotum mostly black with reddish spot anteriorly, disc of elytra mixed red with brown to black (fig. 55). Democratic Republic of Congo (fig. 32).................. overlaeti sp. nov., black and red form
- Punctation of explanate margin of elytra moderately coarse and dense, surface appears moderately irregular. Sculpture on elytral slope higher thus surface appears moderately irregular. Pronotum and elytral disc red, explanate margin of elytra black (fig. 66). Western and Central Africa from Guinea to Uganda (fig. 22).................. inaequalis Thomson, black form
14. Body strongly convex (figs. 71, 74), elytral sculpture low and obtuse, pronotal sides broadly rounded (figs. 69, 70, 72, 73)................................................................................................... 15.
- Body moderately convex (figs. 178), elytral sculpture at least with one sharp, longitudinal carina, pronotal sides subangulate (figs. 176, 177). Southern Africa north to Malawi (fig. 46)....................... thomsoni Boheman, large specimens
15. Explanate margin of elytra with posterolateral and sutural spots. Body stouter, more rounded on sides, L/W ratio 1.24–1.36, surface of elytra distinctly sculptured, with distinct H–shaped elevation and lateral folds (figs. 69, 70). Antennae uniformly yellow to ochraceous. Kenya, Tanzania, Malawi, Zambia, Zimbabwe, Mozambique, and South Africa (fig. 12)............................................................................................... coagulata Boheman
- Explanate margin of elytra without posterolateral and sutural spots. Body slimmer, less rounded on sides, L/W ratio 1.34–1.46, surface of elytra indistinctly sculptured, only with very low and obtuse H–shaped elevation (figs. 72, 73). Apical antennal segments 8–11 brown to black. Kenya, Rwanda, Tanzania and Zambia (fig. 24)........................ irrorata Weise
16. Punctation of two sutural and two or three lateral rows mostly regular or all rows regularly punctate (figs. 256 –261, 326–328)................................................................................................. 17.
- Punctation of elytra completely irregular (figs. 317, 319, 331, 332)........................................... 19.
17. Dorsum uniformly yellow, elytra widest at anterior third? then elytral sides more or less distinctly converging posterad (figs. 256, 257, 259, 260)................................................................................. 18.
- Elytra in postscutellar area with small, black spot, body almost circular, widest in the middle (figs. 326, 327). South Africa (fig. 49).............................................................................. unimaculata Boheman
18. Pronotum widest in the middle with broadly rounded sides (fgs. 256, 257). Claws with large basal tooth. Sierra Leone, Central Africa east to Ethiopia (fig. 21)................................................ imitatrix Spaeth, large specimens
- Pronotum widest at anterior third, sides subangulate (fgs. 259, 256). Claws simple. Botswana, Nambia, and South Africa (fig. 45).................................................................. sulphurago Boheman, large specimens
19. Pronotum almost semicircular, with maximum width at base, sides angulate. Punctation of elytra moderate, quite dense, distance between punctures from twice narrower to as wide as diameter of a puncture (figs. 317–319). Malawi, Mozambique and sounthern Zimbabwe (fig. 18)........................... limpopoana Borowiec & Świętojańska, large specimens
- Pronotum elliptical, with maximum width in the middle, sides subangulate. Base of elytra as wide as base of pronotum. Punctation of elytra very coarse, dense, distance between punctures mostly twice narrower than diameter of a puncture (figs. 331, 332). South Africa: Western Cape (fig. 11).............. foveolatipennis Borowiec & Świętojańska, large specimens
20. Entire surface of elytra covered with dense, short, white, erect setae (figs. 97, 99, 109)............................ 21.
- Surface of elytra without erect setae, at most elytral disc with sparse, short, adpressed hair......................... 22.
21. Base of elytra distinctly wider than pronotum (fig. 109). Surface of pronotal disc without semierect setae. Elytral setae very long, longer than puncture with areola. Ethiopia, Eritrea, Tanzania (fig. 42)............................ setosa Chapuis
- Base of elytra slightly to moderately wider than pronotum (figs. 97, 98, 100–102). Surface of pronotal disc covered witht semierect setae. Elytral setae short, not longer than puncture with areola. East, Central, and South Africa (fig. 23).............................................................................................. innotata Boheman
22. Explanate margin of elytra with dark markings, red, brown to black, forming humeral, or both humeral and posterolateral spots or occupying entire or large parts of margins (figs. 75, 76, 78, 83, 84, 86, 87, 91–95, 103, 105, 107, 136, 161, 221, 231, 329)................................................................................................. 23.
- Explanate margin of elytra without dark markings, yellow, green or brown, at most with narrow sutural spot (figs. 155, 156).................................................................................................. 36.
23. Surface of pronotal disc smooth or punctate, sometimes with irregular sculpture but never granulate. Explanate margin of elytra never completely black.............................................................................. 24.
- Surface of pronotal disc granulate. Explanate margin of elytra completely black (fig. 221). South Africa and S Mozambique, probably introduced and established in Kenya (fig. 19)..................... granulicollis Spaeth, the darkest specimens
24. Explanate margin of elytra with reddish markings forming spots or entire surface of margins red (figs. 86, 103, 105, 231, 329)................................................................................................. 25.
- Explanate margin of elytra with brown to black markings forming spots, bands or occupying large part of margins (figs. 75, 76, 78, 83, 84, 87, 91–95, 107, 136, 161, 221)............................................................... 29.
25. Explanate margin of elytra with reddish humeral or both humeral and posterolateral spots (figs. 86, 103, 105, 329). Elytral disc regularly convex, or only slightly elevated in postscutellar area (figs. 88, 104, 106), occasionally distinctly elevated (fig. 330)................................................................................................. 26.
- Explanate margin of elytra completely purple–red (fig. 231). Elytral disc with distinctly elevated postscutellar hump (fig. 228). West and Central Africa south to Zimbabwe, east to Ethiopia (fig. 48)................... troglodytes Boheman, red form
26. Explanate margin of elytra with both humeral and posterolateral red spots (figs. 103, 105, 329)..................... 27.
- Explanate margin of elytra only with reddish humeral spots (fig. 86). Equatorial part across Africa (fig. 20)......................................................................... humerosa Spaeth, form with reddish humeral spots
27. Thorax and abdomen partly black. Dorsum, except spots on explanate margin of elytra, yellow to pale yellowish red (figs. 103, 104, 329, 330)..................................................................................... 28.
- Thorax and abdomen yellow. Dorsum, except spots on explanate margin of elytra, at least partly deep red to purple–red (figs. 105, 106). Eastern and southern Democratic Republic of Congo (fig. 28)............................... pernix Spaeth
28. Body elongate, L/W ratio 1.78 (fig. 103); postscutellar elevation low (fig. 104). Kenya: E slope of Mt. Elgon (fig. 13).......................................................................................... elogonensis sp. nov.
- Body stout, almost circular, L/W ratio 1.12–1.18 (fig. 329); postscutellar elevation high (fig. 330). Democratic Republic of Congo (fig. 19)........................................................................ ghesquieri Spaeth
29. Explanate margin of elytra with only humeral dark spot (figs. 87, 91 –95, 107)................................... 30.
- Explanate margin of elytra with both humeral and posterolateral dark spots (figs. 76, 84, 136), or only with posterolateral spot (fig. 83), or elytral margins largely dark (figs. 75, 78, 161).................................................. 32.
30. Explanate margin of pronotum yellow to reddish, at most pronotal disc partly to completely black. Base of elytra only slightly to moderately wider that pronotum (figs. 87, 91 –95)....................................................... 31.
- Explanate margin of pronotum largely dark, disc completely black only anterior part of explanate margin yellow. Base of elytra distinctly wider that pronotum (fig. 107). Only Mt. Cameroon in Cameroon (fig. 39)................ rogozinskii sp. nov.
31. Elytral sculpture less marked, second interval only in posterior third obtusely convex, punctation smaller, interspaces not convex, surface of disc appears mostly regular; punctation of explanate margin of elytra very shallow. Elytral pattern constant, brown to black forms humeral spots, several small, partly coalescent spots around, disc forming more or less distinct U–shaped figure, stripe at postscutellar elevation, small spots at top of disc, also some punctures with dark centre (figs. 95, 96). Cameroon (fig. 7)............................................................................. camerunensis Spaeth
- Elytral sculpture distinctly marked, second interval on entire length more or less convex, punctation coarser, interspaces partly convex, surface of disc appears more or less irregular, especially on sides of disc; punctation of explanate margin of elytra deeper than in previous species. Elytral pattern extremely variable, disc from yellow to reddish, without pattern (fig. 86), or with brown to black pattern (figs. 87, 91) to completely black (figs. 93, 94). Equatorial part across Africa (fig. 20).............................................................. humerosa Spaeth, form with brown to black humeral spots
32. Explanate margin of pronotum yellow (figs. 136, 161)..................................................... 33.
- Explanate margin of pronotum partly brown or black, sometimes dark spot reduced to small triangular patch at base of explanate margin (figs. 75, 76, 78, 83, 84)....................................................................... 34.
33. Explanate margin of elytra with broad humeral and narrow posterolateral spots (fig. 136). Elytral disc variable coloured, in the most common form mostly yellow with black pattern as in fig. 136, in the darkest form mostly black except few small yellow spots on top of disc. South Africa (fig. 50)................................................. vespertilio Boheman
- Explanate margin of elytra in anterior 3/4 length black, except yellow humeral angle, yellow extreme margin and yellow fenestrate spot in the middle, close to border of disc. Elytral disc mostly black with two small reddish spots at top of disc and yellow extreme apex (fig. 161)........................................... guttipennis Boheman, the darkest form
34. Elytral disc with high postsutellar hump and distinct sculpture in posterior half. Dark colour of explanate margin of elytra variable, forms regular humeral and posterolateral spots (fig. 76) or semicircular figure, inside with large fenestrate yellow spot (fig. 75), or margins mostly black with small yellowish fenestrate spots in the anterior third and yellow apex (fig. 78). Surface of disc without granulate sculpture, finely punctate, shiny between punctures................................... 35.
- Elytral disc depressed, without postscutellar hump and without sculpture in posterior half. Entire surface of pronotal disc with more or less minutely granulate sculpture, dull, appears shiny. Dark colour of explanate margin of elytra constant, forms regular posterolateral or both humeral and posterolateral spots (figs. 83, 84). Tanzania and Uganda (fig. 46).......... tarda Weise
35. Elytral margins behind humeral angles shallowly emarginate thus humeri more or less acute. Dorsal pattern brown to dark brown, never black. Dark colour of explanate margin of elytra variable, forms regular humeral and posterolateral spots (fig. 76) or semicircular figure, inside with large fenestrate yellow spot (fig. 75). Elytral sculpture, except postscutellar hump, also forms high folds on slope (fig. 77). Eastern Democratic Republic of Congo, Uganda, Tanzania and Zambia (fig. 7).................................................................................................... calvaria (Weise)
- Elytral margins behind humeral angles not emarginate, humeri subangulate. Dorsal pattern black. Dark colour covers large part of pronotum except yellow anterior part of explanate margin, entire elytral disc, and large part of explanate margin of elytra, except small yellow fenestrate spots in anterior third and yellow apex (fig. 78). Elytra without high folds on slope (fig. 79). Uganda (fig. 40).......................................................................... satanas sp. nov.
36. Dorsum never uniformly yellow, green or reddish, at least elytra with darker pattern or mostly dark coloured. Sometimes pattern limited to small red or black spot or stipe in postscutellar area (figs. 232, 322, 326), or small reddish spots on humeral callus and posterolateral part of disc (figs. 236, 239), or elytral disc partly yellow partly reddish (figs. 250, 274, 285), or partly green and partly yellow (fig. 280)...................................................................... 37.
- Dorsum uniformly yellow, green or ochraceous, without pattern, spots or figures with different colour than pale background.................................................................................................. 98.
37. Ground colour of dorsum ochraceous, yellowish brown to brown, elytral disc with marble pattern of mixed ochraceous and yellow or with numerous small, brown to black spots (figs. 176 –181). Surface of elytra appears irregular due to slightly convex yellow relief or elevated folds, at least second interval forms longitudinal costa. Moderately large species, length 5.2–6.8 mm, elongate oval (L/W ratio 1.29–1.49), elytral disc regularly convex without postscutellar elevation or hump............ 38.
- Different combination of characters. Dorsum ground colour usually yellow or green, sometimes reddish but never dark ochraceous or brown, dorsal pattern usually forms spots, bands, vittae or lyriform figures, sometimes large red or black areas but only occasionally marble pattern, length often below 5 mm, body often subcircular to short–oval................ 39.
38. Elytral disc with marbled pattern of yellow and ochraceous. Pronotal sides angulate (figs. 176–178). Claws with large basal tooth. Southern Africa north to Malawi (fig. 46).............................................. thomsoni Boheman
- Elytral disc with numerous small, dark brown to black spots on ochraceous to brown background. Pronotal sides rounded (figs. 179–181). Claws simple. Ethiopia and Eritrea (fig. 32)........................................ weinmanni Chapuis
39. Elytral disc surrounded by thin to broad black ring of regular borders, other parts of disc, except vittae and bands, yellow (figs. 168, 170, 174)..................................................................................... 40.
- Elytral disc with various pattern but it never forms regular black ring. If black pattern suroounds elytral disc then borders of pattern irregular or other parts of disc bicoloured (figs. 150, 336, 338)......................................... 42.
40. Elytral pattern except black ring with bands or spots. Pronotal disc with large black spot or M–shaped figure (figs. 168, 170)................................................................................................. 41.
- Elytral pattern forms only broad, black ring without bands and spots inside. Pronotal disc at base only with five small, brown spots (fig. 174). Kenya (fig. 33)........................................................... rabaiensis sp. nov.
41. Elytral disc with narrow black band across the middle. Scutellum and elytral suture black, pronotal disc with black M–shaped figure (fig. 168). Mozambique and Zimbabwe (fig. 12)........................................ circumflexa Spaeth
- Elytral disc without band only with small black spots in postscutellar area and 2/3 length of disc, posterior spots tends to form transverse band. Scutellum yellow, pronotum on disc with large, black, trapezoidal spot (fig. 170). South Africa: Eastern Cape (fig. 31)................................................................................. wanati sp. nov.
42. Elytral disc almost completely black, at most with few very small reddish spots of diffused borders (figs. 80, 81, 92, 118, 132, 152)............................................................................................. 43.
- Elytral disc never completely black, with various dark pattern............................................... 47.
43. Elytral punctation fine to moderate, intervals mostly wider than rows. Pronotal disc with black spots in the middle and yellow sides (figs. 118, 152)................................................................................ 44.
- Elytral punctation coarse, intervals narrower than rows, often linear. Pronotal disc completely black (figs. 80, 81, 92, 132)................................................................................................... 45.
44. Base of elytra only slightly wider than pronotum. Pronotal disc punctate, with two black spots, small triangular anteriorly and large, transverse at base (fig. 118). Ventrites mostly black. Body oval, L/W 1.33–1.43. South and Central Africa north to Rwanda (fig. 3)......................................................... amorifica Boheman, the darkest form
- Base of elytra distinctly wider than pronotum. Pronotal disc impunctate, with single cross–shaped black spot (fig. 152). Ventrites completely yellow. Body almost circular, L/W 1.16–1.22. Central Africa from Cameroon to Ethiopia, south to Mozambique and Zimbabwe (fig. 4)...................................................... ancorifera Boheman, the darkest form
45. Scutellum mostly to completely black. Last one or two antennal segments distinctly infuscate dorsally............... 46.
- Scutellum yellow. Antennae yellow or last antennal segments indistinctly infuscate dorsally. Equatorial part across Africa (fig. 20)........................................................ humerosa Spaeth, dark form without humeral spots
46. Surface of pronotal disc between punctures distinctly microreticulate, appears more or less alutaceous. Dark elytral disc not deep black, at least with reddish diffused borders, often on dorsal surface and elytral relief with small reddish spots of diffused borders (figs. 80, 81). Mountain regions of Kenya, Rwanda, Uganda, and Eastern Democratic Republic of Congo (fig. 41)........................................................................ schoutedeni Spaeth, the darkest form
- Surface of pronotal disc between punctures less distinctly microreticulate, appears shiny. Dark elytral disc deep black, without small reddish spots (fig. 132). Forest species known from Central Africa south to Zimbabwe, few localities also in Guinea, Sierra Leone and Togo (fig. 14).............................................. depicta Boheman, the darkest form
47. Explanate margin of elytra without apical, sutural spot..................................................... 48.
- Explanate margin of elytra with apical, sutural spot (figs. 155, 156). Cameroon and Nigeria (fig. 13).... delenifica Boheman
48. Elytral disc without adpressed hair..................................................................... 49.
- Elytral disc covered with short, sparse adpressed hair (sometimes difficult to see, best viewed under oblique lighting), with several small, brown to black spots (figs. 214, 216). Kenya, Tanzania, Malawi, Mozambique, Namibia, and Transvaal province in South Africa (fig. 6).................................................................. callosicollis Spaeth
49. Elytral pattern forms black, isolated spots on yellow background (figs. 111, 115, 117, 158, 160, 162, 163, 172), sometimes this pattern is reduced to two small spots on sides of disc or single spot or stripe in postscutellar point (figs. 165, 166, 226, 227, 322, 324, 326, 327)..................................................................................... 50.
- Elytral pattern variable but never forms black, isolated spots on yellow background but yellow elevated spots on black background (figs. 119, 120, 122, 124, 126, 131, 151) or mixed yellow and brown to black marble pattern (figs. 128, 129, 133, 134, 140, 141, 146, 189, 190, 212, 336, 338), or only punctures with brown to black centre and areola (figs. 199, 200, 202), or disc with reddish spots (232, 234, 237, 239), or with red, brown and yellow figures (figs. 244, 245, 249, 250, 274, 279, 280, 285)................................................................................................................................................................................................. 59.
50. Elytral disc with only a single spot at postscutellar area, sometimes with minute additional two spots on elevated second interval, or two spots on side of disc (figs. 165, 166, 226, 227, 322, 324, 326, 327)............................... 51.
- Elytral disc with more than three dark spots (figs. 111, 115, 117, 158, 160, 162, 163, 172)......................... 55.
51. Elytral disc with single dark spot or stripe at postscutellar area, sometimes with minute additional two spots on elevated second interval. Body stout, almost circular (figs. 226, 227, 322, 324, 326, 327)....................................... 52.
- Elytral disc only in the middle of eighth interval with small, brown to black spot. Body short–oval (figs. 165, 166). South Africa (fig. 35).......................................................... sphaerula Boheman, typical maculate form
52. Elytral disc without sculpture, rounded in profile, elytral punctation moderately coarse and dense, intervals well marked, mostly wider than rows, sides of pronotum rounded (figs. 322–328)................................................. 53.
- Elytral disc with distinct sculpture and well-marked postscutellar hump, elytral punctation coarse and dense, intervals narrow, partly linear, sides of pronotum angulate (figs. 226–228). West and Central Africa south to Zimbabwe, east to Ethiopia (fig. 48)....................................................................... troglodytes Boheman, pale form
53. Elytral punctures without dark centres, dark spot in postscutellar area round (figs. 326, 327) or in form of short stripe (fig. 322), interspaces on sides of elytra flat and surface of elytral sides appears completely regular, surface of explanate margin of elytra slightly irregular................................................................................... 54.
- Elytral punctures at top of disc with dark centre, dark spot in postscutellar area elongate, punctation of elytra coarser than in both relatives, interspaces on sides of elytra slightly convex, forming a low fold, surface of elytral sides appears slightly irregular, surface of explanate margin of elytra strongly irregular (fig. 324). North-eastern part of South Africa (fig. 9)................................................................................... drakensbergensis Borowiec
54. Elytral spot in form of short, narrow stripe, punctation of sides of disc slightly coarser and denser (fig. 322). East Cape Province in South Africa (fig. 8)......................................................... emontinensis sp. nov.
- Elytral spot more or less round, occasionally in form of stripe, punctation of sides of disc slightly finer and sparser (figs. 326, 327). South Africa (fig. 49)........................................................... unimaculata Boheman
55. Pronotum with black spots (figs. 111, 115, 117)........................................................... 56.
- Pronotum without black spots (figs. 158, 160, 162, 163, 165, 172)............................................ 57.
56. Pronotal disc at top with two round or oval spots (fig. 111). Elytral suture on slope without black spot, prosternum yellow. Western, eastern and central Africa west to Mali, north to Sudan, and south to N Tanzania (fig. 1).............................................................................................. aethiopica Boheman, typical form
- Pronotal disc at base with mushroom–shaped or trapezoidal black spot and often with small, traingular or rhomboidal spot in area above head (figs. 115, 117). Elytral suture on slope with black spot, prosternum yellow. South and Central Africa north to Rwanda (fig. 3)............................................................ amorifica Boheman, typical form
57. Smaller species, length below 5.6 mm. Body slimmer, L/W ratio 1.18–1.51. Postscutellar area without black spot (figs. 158, 160, 162, 163). Claws simple......................................................................... 58.
- Larger species, length 6.0 mm. Body stouter, circular, L/W ratio 1.08. Postscutellar area with black spot (fig. 172). Claws simple but appearing distinctly appendiculate due to projecting flanks of claw segment. Tanzania (fig. 2).. procurva Spaeth
58. Body larger and stouter, L 4.65–5.50, L/W ratio 1.18–1.31. Explanate margin of elytra broader and more explanate (figs. 158, 160). South Africa (fig. 11)................................................ guttipennis Boheman, maculate form
- Body smaller and slimmer, L 3.85–4.85, L/W ratio 1.35–1.51. Explanate margin of elytra narrower and less explanate, declivous (figs. 158, 160). South Africa (fig. 37).............................................. quatuordecimsignata Spaeth
59. Pronotal and elytral disc predominantly black, both or only elytral disc with small yellow spots of distinct borders (figs. 119, 120, 122, 124, 131, 151)............................................................................. 60.
- Pronotal and elytral disc not predominantly black, usually with marble pattern of yellow and brown to black, if black predominate then yellow spots with irregular shape and borders (figs. 90, 126, 128, 133, 134, 138, 142), or only punctures with brown to black centre and areola (figs. 199, 200, 202), or disc with reddish spots (figs. 232, 234, 237, 239), or with red, brown and yellow figures (figs. 150, 244, 245, 249, 250, 274, 279, 280, 285).................................................. 65.
60. Pronotal black spot with two small, more or less round spots at base (figs. 119, 120, 131).......................... 61.
- Pronotal black spot without or with U or V–shaped yellow spot at base (figs. 122, 124, 151)....................... 63.
61. Elytral punctation moderately fine and sparse, distance between punctures mostly wider than puncture diameter, rows interrupted by elytral relief and partly appear irregular (figs. 122, 124). Surface of pronotal disc smooth and shiny. Ventrites uniformly yellow........................................................................................... 62.
- Elytral punctation coarse and dense, distance between punctures mostly narrower than puncture diameter, rows slightly interrupted by elytral relief but appear mostly regular (fig. 131). Surface of pronotal disc punctate. Ventrites partly brown to black. Forest species known from Central Africa south to Zimbabwe, few localities also in Guinea, Sierra Leone and Togo (fig. 14).............................................................................................................................................................................. depicta Boheman, dark form
62. Elytral margins distinctly emarginate behind humeral angle thus humeri appear acute. Behind scutellum two small yellow spots. Elytral spots not elevated and impunctate (fig. 120). Black ground colour behind humeral callus extending to marginal row (fig. 121). Surface of disc more shiny with mirror brilliance. Eastern Democratic Republic of Congo and Rwanda (fig. 15).............................................................................................. leleupi sp. nov.
- Elytral margins shallowly emarginate behind humeral angle thus humeri appear angulate. No yellow spots behind scutellum. Central elytral spots elevated and with 1–2 coarse punctures. Black ground colour behind humeral callus extending only to submarginal row. Surface of disc less shiny and slightly alutaceous. Angola (fig. 1).................. benguelica Spaeth
63. Elytral punctation coarse and dense, distance between punctures mostly narrower than puncture diameter. At base of elytra close to scutellum two small, yellow spots (figs. 122, 124). At least prosternum brown............................ 64.
- Elytral punctation fine to moderate, distance between punctures from as wide as to thrice wider than puncture diameter. Base of elytra close to scutellum without yellow spots (fig. 151). Ventrites completely yellow. Central Africa from Cameroon to Ethiopia, south to Mozambique and Zimbabwe (fig. 4).............................. ancorifera Boheman, dark form
64. Slimmer species, L/W ratio 1.23–1.26. Pronotal black spot with U or V–shaped yellow spot at base (fig. 124). Marginal interval in front and behind lateral fold mostly black (fig. 125). Ventrites mostly black, including coxa, only abdomen surrounded by yellow. Claws with large basal tooth. North–western Cameroon (fig. 6)........................... bamendana sp. nov.
- Stouter species, L/W ratio 1.13–1.18. Pronotal black spot without additional yellow spot at base (fig. 122). Marginal interval in front and behind lateral fold mostly yellow (fig. 123). Prosternym brown, metasternum and abdomen yellow. Claws with small basal tooth. Eastern Democratic Republic of Congo, Rwanda, and N Tanzania (fig. 13).............. flavosignata Spaeth
65. Only elytral punctures with black centre and areola, sometimes areoles of adjacent punctures merge with each other but the pattern on the elytra does not create clear spots or a marbled pattern. Pronotum immaculate or with very small V–shaped brown spot (figs. 199 –203)................................................................................. 66.
- Dark markings on elytral disc form distinct pattern, sometimes it is reduced to small spots in postscutellar area or figures on slope or only punctures on sides of disc with dark centre. Pronotum immaculate or with basal spots (figs. 205, 224, 285)..................................................................................................... 67.
66. Elytral punctures denser, tends to form more or less regular rows, surface except postscutellar H–shaped elevation appears regular, second interval does not form a dinstinct elevated costa (figs. 199, 200). East Africa from Kenya to Eastern Democratic Republic of Congo (fig. 44)............................................................... successiva Spaeth
- Elytral punctures sparser, punctation appears mostly irregular, second interval almost on entire length elevated (figs. 202, 200). Ethiopia (fig. 44)........................................................................ stipidosa sp. nov.
67. Elytral pattern dark, dark brown to black, occupies large part of disc (figs. 80, 126–131, 133–136, 138–144, 150, 185–187, 336–339)......................................................................................... 68.
- Elytral pattern pale, reddish to brown, occasionally black, occupies small part of disc, the pale background predominate (figs. 89, 145, 146, 147, 148, 153, 138–144, 150, 182–184, 188 –198, 205, 210–213, 220, 224, 232, 234, 236–240, 249–251, 274, 279, 280, 285)......................................................................................... 75.
68. Pronotum with distinct brown to black basal spot (figs. 126, 128, 129, 133, 134, 138–143, 150, 185, 186)............. 70.
- Pronotum immaculate or at most with a fuzzy darker spot in the centre (figs. 90, 336, 338, 339)..................... 69.
69. Larger, L 4.25–5.80 mm. Elytral pattern regular, yellow spots spread regularly on top of disc, H– shaped elevation indistinct (fig. 90). Ventrites mostly black. Equatorial part across Africa (fig. 20)... humerosa Spaeth, form with black elytral pattern
- Smaller, L 3.90–4.60 mm. Elytral pattern less regular, H–shaped elevation distinct and mostly yellow, yellow spots tend to form transverse irregular spot on slope (figs. 336, 338, 339). Ventrites uniformly yellow or only central part of metathorax infuscate. Kenya, Tanzania, Mozambique and South Africa (fig. 42)......................................... shimba sp. nov.
70. Elytral punctation coarse and dense, distance between punctures mostly narrower than puncture diameter. Pronotal spot M– shaped, V–shaped or or forms oblique spots on sides of disc, occasionaly mushroom–shaped (figs. 142, 143), black pattern spread over the entire surface of disc (figs. 126, 128, 129, 133, 134, 138–143, 185, 186)........................... 71.
- Elytral punctation fine to moderate, distance between punctures from as wide as to thrice wider than puncture diameter. Pronotal spot mushroom–shaped. Black pattern surrounds elytral disc and forms incomplete band in 2/3 length of disc, other parts of disc reddish and yellow (fig. 150). Ventrites completely yellow. Central Africa from Cameroon to Ethiopia, south to Mozambique and Zimbabwe (fig. 4)........................................................ ancorifera Boheman, dark form
71. Smaller species, L 4.05–5.20 mm. Body short–oval to oval (figs. 126, 128, 129, 133, 134, 138, 140, 141, 185, 186), slightly convex (figs. 130, 135, 139, 187). Pronotal spots never mushroom–shaped, pronotal sides usually narrowly rounded (figs. 126, 128, 129, 185), or broadly rounded (figs. 133, 134, 138, 140, 141). Ventrites often partly brown to black.............. 72.
- Large species, L 5.80–7.00 mm. Body almost hemispherical, strongly convex (fig. 144). Pronotal spot mushroom–shaped, pronotal sides broadly rounded (figs. 142, 143). Ventrites uniformly yellow. Sierra Leone, Cameroon, Democratic Republic of Congo, Rwanda, Uganda, and Kenya (fig. 26).............................................. oberlaenderi Spaeth
72. Pronotal pattern forms M– or V–shaped basal spot (figs. 126, 128, 129, 133, 134, 185). Ventrites often partly brown to black. Here three species difficult to identification.............................................................. 73.
- Pronotal pattern forms broad, oblique spots on sides of disc and usually surrounds yellow lateral lobes (figs. 138, 140, 141). Ventrites uniformly yellow. Central Mozambique and South Africa (fig. 24)...................... irregularis Boheman
73. Pronotal sides narrowly rounded. Pronotal spot usually small, V–shaped, only in the darkest forms M–shaped (figs. 128, 129, 185, 186). Species widely spread in tropical Africa........................................................ 74.
- Pronotal sides broadly rounded. Pronotal spot usually large, M–shaped (figs. 133, 134). Distribution restricted to South Africa (fig. 30)............................................................................... natalensis Spaeth
74. In pattern of elytral disc yellow colour predominate, dark spots usually brown to dark brown, occupy mostly sides (figs. 185, 186). East and South Africa north to Somalia, south to Cape province in South Africa (fig. 25)................................................................................................... lacrymosa Boheman, dark form
- In pattern of elytral disc black colour predominate (fig. 129), dark spots always black, occupy almost entire surface of disc (figs. 128, 129). Central Africa south to Zimbabwe, few localities also in Guinea, Sierra Leone and Togo (fig. 14)......................................................................................... depicta Boheman, typical form
75. Surface of pronotal disc smooth or punctate, sometimes with irregular sculpture but never granulate................. 76.
- Surface of pronotal disc granulate. Elytral pattern brown to black, forms oblique line on sides, few small spots on slopes, also some punctures with dark centre (fig. 220). South Africa and S Mozambique, probably introduced and established in Kenya (fig. 19)................................................................ granulicollis Spaeth, maculate form
76. Smaller species, L 4.05–5.20 mm. Body from short–oval to distinctly narrowing posterad. Pronotal from angulate to broadly rounded. Ventrites uniformly yellow to partly brown or black................................................ 77.
- Large species, L 5.8–7.00 mm. Body almost hemispherical, strongly convex (fig. 144). Pronotal sides broadly rounded, base of elytra distinctly wider than pronotum (fig. 146). Ventrites uniformly yellow. Sierra Leone, Cameroon, Democratic Republic of Congo, Rwanda, Uganda, and Kenya (fig. 26)............................ oberlaenderi Spaeth, brown maculate form
77. Dark spots on elytral disc form various pattern but never reddish oval figure on slope............................. 78.
- Dark spot on elytral disc form large, reddish, oval figure on slope (fig. 285). Small species, L 4.25–4.60, oval, with completely regular elytral punctation and elytral sculpture resticted only to obtuse H–shaped figure, and slightly convex second interval (figs. 265, 286). Cameroon, Democratic Republic of Congo, and Kenya (fig. 24)......... luxuriosa Spaeth, maculate form
78. Central part of elytral disc yellow to red in relation to green or yellow sides (figs. 244, 245, 249, 250, 279, 280), the central area sometimes bordered by red to brown forming lyriform figure (figs. 153, 245, 249, 279) or with few small brown spots (fig. 244)............................................................................................. 79.
- Central part of elytral disc the same colour as rest of background, yellow to green, sometimes with reddish stripes along sides (figs. 147, 274), pattern of various small, red or brown spots or marble stripes (figs. 189, 191, 212, 224, 232, 234, 237, 239, 264)............................................................................................. 83.
79. Claws simple. Elytral punctation moderate, completely regular, intervals never linear (figs. 153, 244. 245)............ 80.
- Claws with basal tooth. Elytral punctation coarse, partly irregular, intervals vary narrow, partly linear (figs. 249, 250, 279, 280)............................................................................................. 81.
80. Elytral disc with well marked H–shaped postscutellar elevation, pronotal disc at base with two short, brown stripes (figs. 153, 154). Central Africa from Cameroon to Ethiopia, south to Mozambique and Zimbabwe (fig. 4).............................................................................................. ancorifera Boheman, the palest form
- Elytral disc without or indistinctly marked H–shaped postscutellar elevation, pronotal disc immaculate (figs. 243–245). Entire tropical Africa (fig. 16)................................................... dorsovitata Boheman, maculate form
81. Central part of elytral disc yellow with reddish brown or brown spots, especially on sides of yellow spot (figs. 279, 280), or the central spot margined by brown lines (fig. 249). Species from eastern and southern Africa (figs. 28, 32).............. 82.
- Central part of elytral disc partly red due to reddish centres of punctures and red lateral stripes (figs. 250, 251), sometimes red is reduced only to lateral stripes or punctures with red centre. Species from West and Central Africa (fig. 18).................................................................................................. granula Boheman
82. Pronotal disc moderately coarse and moderately dense punctate, distance between punctures from slightly narrower to twice wider than puncture diameter. Body more regularly oval, regularly rounded on sides (figs. 279, 280). Kenya, Zimbabwe, and Namibia (fig. 28)..................................................................... mashonensis Spaeth
- Pronotal disc impunctate or at most finely punctate, distance between punctures always wider than puncture diameter. Body less regularly oval, sides slightly converging posterad (fig. 249). South and East Africa north to Somalia (fig. 32).............................................................................. viridipennis Boheman, maculate form
83. Body distinctly converging posterad (figs. 232, 234). Elytral disc strongly convex, slightly gibbous in profile, elytral pattern of red spots limited to postscutellar impression and sutural part of disc (figs. 233, 235). Claws simple. Distribution area limited to north–eastern Africa and southwestern Saudi Arabia (figs. 33, 40)............................................ 84.
- Body from almost circular to elongate–oval but not distinctly converging posterad (figs. 182, 183, 189 –191, 193. 196, 210, 212, 224, 236, 237, 239). Elytral disc moderately, regularly convex or depressed, never gibbous in profile, elytral pattern of red spots not limited to postscutellar impression and sutural part of disc (figs. 192, 195, 198, 211, 213). Claws with basal tooth, only in two species simple.................................................................................. 85.
84. Larger: length 4.40–5.20 mm and slightly more elongate L/W ratio 1.39–1.49 (fig. 232). Elytral profile less convex and less angulate (fig. 233). Pronotal punctation coarser and denser. Cyprus, Israel, Lebanon, Saudi Arabia and Tunisia (fig. 33)............................................................................. pellegrini Marseul, maculate form
- Smaller: length 3.70–4.40 mm and slightly less elongate L/W ratio 1.29–1.39 (fig. 234). Elytral profile more convex and more angulate (fig. 235). Pronotal punctation finer and sparser. Saudi Arabia, Yemen, Sudan, and Kenya (fig. 40)...................................................................................... rothschildi Spaeth, maculate form
85. Claws with basal tooth............................................................................... 87.
- Claws simple...................................................................................... 86.
86. Elytral pattern forms reddish, oblique stripe on sides of disc, a moderately broad band in 2/3 length of disc and thin band on slope, also postscutellar area partly reddish (figs. 212, 213). Nigeria (fig. 26)....................... nigeriana sp. nov.
- Elytral pattern forms six pale brown spots on top of disc, oblique stripe in humeral area, and spot in posterolateral part of disc (fig. 114). Western, eastern and central Africa west to Mali, north to Sudan, and south to N Tanzania (fig. 1)........................................................................................ aethiopica Boheman, pale form
87. Elytral pattern never forms large reddish spots in humeral and posterolateral parts of disc.......................... 88.
- Elytral pattern forms large reddish spots in humeral and posterolateral parts of disc, sometimes these spots connected and form red stripe along side of disc(fig. 274). Body elongate–oval, with maximum width in the middle: L/W 1.53. Punctures in postscutellar impressions and red marked punctures in humeral and posterolateral part of disc partly irregular. Angola and Zambia (fig. 39)........................................................................ rhodesiaca Spaeth
88. Length above 3.5 mm, elytral pattern variable but never form narrow ring surrounding disc, occasionally with thin ring and band (fig. 189)..................................................................................... 89.
- Very samll, length 3.20–3.40 mm. Eytral pattern reddish brown, tends to form narrow ring around disc and incomplete band in 2/3 length of disc (figs. 210, 211). Somalia (fig. 28)......................................... minutissima sp. nov.
89. Elytral pattern limited to small spots on postscutellar elevation, humeral callus and posterolateral part of disc, in pale forms the pattern limited only to spot on postscuttellar elevation or stripes on humeral callus and posterolateral parts of disc (figs. 207, 236, 237, 239)..................................................................................... 90.
- Elytral pattern more widespread, forms a marbled pattern on the sides of the disc and/or small spots at top of disc (figs. 89, 188 –191, 193–195, 197, 198, 205, 206, 209, 224, 225)..................................................... 92.
90. Ground colour of dorsum yellow to yellowish green, elytra at least with four spots or stripes. Species generally from eastern and southern Africa (figs. 5, 20)....................................................................... 91.
- Ground colour of ochraceous yellow, elytra disc with only small, brown spot on postscutellar elevation. Species from Burkina Faso and Niger (fig. 44)........................................................ villiersi sp. nov., spotted form
91. Slimmer, L/W ratio 1.33–1.47, explanate margin of elytra narrower and less explanate (figs. 236–238). Almost entire eastern Africa, single locality from north–eastern Nigeria (fig. 5)........................................ andreinii Spaeth
- Stouter, L/W ratio 1.22–1.30, explanate margin of elytra broader and more explanate (figs. 239, 240). Mozambique and South Africa (Natal) (fig. 20)..................................................................... heroni sp. nov.
92. Larger, length 5.20–5.80............................................................................. 93.
- Smaller, length at most 5.00 mm....................................................................... 94.
93. Pronotal sides angulate, elytral disc depressed (figs. 224, 225). Angola (fig. 43).................. sparteiformis sp. nov.
- Pronotal sides rounded, elytral disc convex (figs. 88, 89). Equatorial part across Africa (fig. 20)................................................................................................. humerosa Spaeth, the palest form
94. Stouter species, L/W 1.17–1.32 (figs. 190, 191)........................................................... 95.
- Slimmer species, L/W 1.32–1.45 (figs. 193, 194, 196, 197, 204, 205).......................................... 96.
95. Brown, small spots group mostly on sides of elytral disc (figs. 188, 190). East and South Africa north to Somalia, south to Cape province in South Africa (fig. 25)........................................... lacrymosa Boheman, the palest form
- Brown, small spots group mostly on top of elytral disc (figs. 191, 192). Senegal (fig. 17).......... fuscosignata Boheman
96. Elytral pattern of numerous small brown to black spots spread over the entire surface of disc, sometimes only most of punctures with dark centre (figs. 193–195, 205, 206). Prosternum at least partly brown to black............................. 97.
- Elytral pattern of several small reddish to brown spots on sides of disc and a single spot on postscutellar elevation, top of disc without spots (figs. 197–198). Prosternum yellow. Kenya (fig. 47)................................. voiensis sp. nov.
97. Elytral punctation coarser, with distance between punctures in lateral rows almost twice smaller than puncture diameter. Sides of elytra slightly more rounded (figs. 193–195). Ethiopia, Eritrea and Tanzania (fig. 17)................. eluta Boheman
- Elytral punctation finer, with distance between punctures in lateral rows mostly as wide as or only slightly smaller than puncture diameter. Sides of elytra less rounded. Namibia (fig. 27)..................................... namibiensis Borowiec
98. Pronotal disc smooth or punctate but never granulate....................................................... 99.
- Surface of pronotal disc granulate (fig. 218). South Africa and southern Mozambique, probably introduced and established in Kenya (fig. 19)...................................................... granulicollis Spaeth, the palest specimens
99. Punctation of elytra completely regular, if partly or mostly irregular then punctures very coarse and dense and surface of elytra appears more or less irregular and at least first two sutural and two marginal rows regular. Tarsal claws simple or toothed................................................................................................... 100.
- Punctation of elytra completely irregular. Tarsal claws usually simple or with slightly widened base, occasionally with basal tooth............................................................................................ 126.
100. Claws simple..................................................................................... 101.
- Claws with basal tooth, sometimes small, or simple but appear appendiculate due to distally projecting flank of last tarsal segment......................................................................................... 109.
101. Body distinctly converging posterad (figs. 232, 234). Elytral disc strongly convex, slightly gibbous in profile (figs. 233, 235). Distribution area limited to north–eastern Africa and southwestern Saudi Arabia (figs. 33, 40)
- Body from almost circular to elongate–oval but not distinctly converging posterad. Elytral disc moderately, regularly convex or depressed, never gibbous in profile.................................................................... 103.
102. Larger: length 4.40–5.20 mm and slightly more elongate L/W ratio 1.39–1.49 (fig. 232). Elytral profile less convex and less angulate (fig. 233). Pronotal punctation coarser and denser. Cyprus, Israel, Lebanon, Saudi Arabia and Tunisia (fig. 33)........................................................................... pellegrini Marseul, immaculate form
- Smaller: length 70–4.40 mm and slightly less elongate L/W ratio 1.29–1.39 (fig. 234). Elytral profile more convex and more angulate (fig. 235). Pronotal punctation finer and sparser. Saudi Arabia, Yemen, Sudan, and Kenya (fig. 40).................................................................................... rothschildi Spaeth, immaculate form
103. Smaller species, length usually below 5.5 mm, if slightly larger (up to 5.95 mm) then body almost parallel sided, elytral punctation completely regular and pronotal sides broadly rounded (fig. 113)................................... 104.
- Larger, length 5.90–7.50 mm. Body slightly converging posterad, elytral punctation mostly regular but with additional irregular punctures, pronotum trapezoideal, with maximum width at anterior 1/3 length, sides narrowly rounded. Botswana, Nambia, and South Africa (fig. 45)................................................................. sulphurago Boheman
104. Pronotum subtrapezoidal, widest in front of the middle, sides narrowly rounded to angulate (figs. 264, 287).......... 105.
- Pronotum elliptical, widest in or behind the middle, sides broadly rounded (figs. 113, 241) to subangulate (figs. 165, 222)................................................................................................... 106.
105. Elytra with deeply impressed postscutellar impressions and well marked H–shaped postscutellar elevation (fgs. 287, 288). Ventrites mostly black. Western Cape Province in South Africa (fig. 8)........................... capensis Borowiec
- Elytra without postscutellar impressions and without H–shaped postscutellar elevation (figs. 264, 265). Ventrites uniformly yellow. South Africa: Cape Province (fig. 45)............................................... sulphurea Boheman
106. Elytral punctation moderate, intervals well marked, usually as wide as to slightly wider than rows (figs. 113, 165, 241). Explanate margin of elytra narrow to moderately broad, more or less declivous (figs. 112, 167, 243)................ 107.
- Elytral punctation coarse and very dense, intervals linear, narrower than rows (fig. 222). Explanate margin of elytra very broad, strongly explanate, almost horizontal (figs. 222, 223). Southern part of Democratic Republic Of Congo (fig. 42)................................................................................................... spartea Shaw
107. Elytral disc in profile forms regular arch (fig. 167) or is depressed anteriorly and regularly rounded on slope (fig. 112). Pronotal disc at top finely punctate........................................................................... 108.
- Elytral disc irregularly convex in profile, the highest in postscutellar area (fig. 243). Pronotal disc at top impunctate, at most with fine pricks. Entire tropical Africa (fig. 16)....................... dorsovittata Boheman, typical immaculate form
108. Pronotal sides broadly rounded. Elytral punctation fine, intervals 1 and 2 distinctly wider than rows (fig. 113). Elytral disc in profile depressed anteriorly and regularly rounded on slope (fig. 112). Western, eastern and central Africa west to Mali, north to Sudan, and south to N Tanzania (fig. 1)............................... aethiopica Boheman, rare immaculate form
- Pronotal sides subangulate to narrowly rounded. Elytral punctation moderately coarse, intervals 1 and 2 as wide as to slightly wider than rows (fig. 165). Elytral disc in profile forms regular arch (fig. 167). South Africa (fig. 35)........................................................................................ sphaerula Boheman, immaculate form
109. Body elongate–oval, almost parallel sided, elytral disc with distinct H–shaped elevation and convex second interval (figs. 252, 320). Length above 5.3 mm.......................................................................... 110.
- Body of various shape, from circular (figs. 282, 289) to oval (figs. 269, 277) but never appears parallel sided. Length often below 5.0 mm.................................................................................... 111.
110. Sides of pronotal disc with irregular sculpture of punctures and wrinkles. Punctation of explanate margin of pronotum and elytra deep, surface appears distinctly irregular. Posterolateral parts of elytral disc with mostly irregular punctction. Ethiopia, Oromia Region (fig. 23)............................................................... oromiaensis sp. nov.
- Sides of pronotal disc punctate but without wrinkles. Punctation of explanate margin of pronotum and elytra shallow, surface appears slightly irregular. Posterolateral parts of elytral disc with mostly regular punctation. Namibia and South Africa (fig. 34)................................................................................. penelope Boheman
111. Punctation of elytra coarse and dense, at least partly irregular, intervals mostly narrower than rows, especially on sides of disc, elytral disc often with well marked and sharp H–shaped elevation. Dried specimens often retain their natural green colour. Length always above 4.6 mm (figs. 246–248, 266–273, 276–278)........................................... 112.
- Punctation of elytra from fine to coarse but usyually moderately dense, often mostly regular, if partly irregular than intervals partly as wide or wider than rows, especially on sides of disc, elytral disc usually with obtuse H–shaped elevation. Dried specimens never retain their natural green colour. Length often below 4.5 mm.................................. 116.
112. Elytral punctation mostly regular, the few extra punctures at intervals do not give the impression of irregularity of rows and surface irregularities. Punctation of pronotal disc from fine to moderately coarse, from scarce to moderately dense but surface of disc appears mostly regular to only slightly irregular (figs. 246–248, 271–273)............................... 113.
- Elytral punctation only partly regular, several extra punctures at intervals give the impression of irregularity of rows and surface irregularities. Punctation of pronotal disc from moderately coarse to coarse, dense and surface of disc appears more or less irregular (figs. 266 –270, 276–278).................................................................... 114.
113. Body stouter, L/W ratio 1.17–1.24, base of elytra distinctly wider than pronotum, sides of elytra distinctly converging posterad. Sides of pronotum broadly rounded, surface of pronotal disc finer and sparser punctate (figs. 246–248). South and East Africa north to Somalia (fig. 32)........................................ viridipennis Boheman, typical immaculate form
- Body slimmer, L/W ratio 1.29–1.34, base of elytra slightly wider than pronotum, sides of elytra slightly converging posterad. Sides of pronotum less broadly rounded, surface of pronotal disc coarser and denser punctate (figs. 271–273). Botswana, Malawi, Namibia, and Zimbabwe (fig. 15).................................................... devylderi Spaeth
114. Elytral disc irregularly convex with the highest point in postscutellar elevation, H–shaped elevation usually well marked and high (figs. 266–268, 276–278)....................................................................... 115.
- Elytral disc almost regularly convex, H–shaped elevation usually hardly marked and obtuse (figs. 269, 270). South Africa and Zimbabwe (fig. 17)................................................................. franklinmuelleri Spaeth
115. Stouter, L/W ratio 1.23–1.29, explanate margin of elytra slightly broader and less declivous (figs. 266–268). East Africa from Ethiopia to Zimbabwe (fig. 43)............................................................... sublesta Weise
- Slimmer, L/W ratio 1.31–1.40, explanate margin of elytra slightly narrower and more declivous (figs. 276–278). Malawi (fig. 22)............................................................................... kantnerorum sp. nov.
116. Large species, length 5.05–6.60 mm................................................................... 117.
- Smaller species, length 3.90–4.95 mm................................................................. 118.
117. Base of elytra distinctly wider than pronotum. Pronotum with broadly rounded sides, widest in the middle. Punctation of elytra less regular, intervals on slope and posterolateral parts of disc with several additional punctures (figs. 256–258). Sierra Leone, Central Africa east to Ethiopia (fig. 21)....................................................... imitatrix Spaeth
- Base of elytra slightly wider than pronotum. Pronotum with less regularly rounded side, widest slightly in front of the middle. Punctation of elytra more regular, intervals on slope and posterolateral parts of disc without additional punctures (figs. 262, 263). Ethiopia and Somalia (fig. 6)...................................................... benaadirensis sp. nov.
118. Body almost circular, pronotum widest at base (figs. 282, 289).............................................. 119.
- Body not circular (figs. 196, 204, 207, 284), if more or less circular then pronotum not widest at base (figs. 239, 254, 340).................................................................................................. 120.
119. Ventrites mostly black. Punctation of elytral disc coarser and denser, punctures on slope as coarse as in anterior half of disc, in rows distance between punctures often smaller than puncture diameter. Elytral disc slightly convex with only slightly marked postscutellar elevation (fig. 283). Pronotal sides narrowly rounded (fig. 282). Democratic Republic of Congo (fig. 19)......................................................................................... garambana sp. nov.
- Ventrites completely yellow. Punctation of elytral disc finer and sparser, punctures on slope distinctly smaller than in anterior half of disc, in rows distance between punctures often larger than puncture diameter, especially in posterior half of disc. Elytral disc strongly convex with high postscutellar elevation (fig. 290). Pronotal sides angulate (fig. 289). Senegal (fig. 40).......................................................................................... senegalensis sp. nov.
120. Ventrites uniformly yellow. Body oval to almost circular (figs. 196, 207, 239, 254, 284, 340)...................... 121.
- Thorax black. Body oval, almost parallel sided (figs. 204, 205). Namibia (fig. 27)...................................................................................................... namibiensis Borowiec, immaculate form
121. Punctation of elytra coarser and denser, distance between punctures mostly smaller than puncture diameter, intervals on sides of disc partly narrower than rows to linear (figs. 196, 198, 207, 209, 239, 240, 254, 255, 340, 341)................. 122.
- Punctation of elytra finer and sparser, distance between punctures at least on top of disc partly as large as or larger than puncture diameter, intervals on sides of disc partly as wide as to wider than rows (figs. 284, 286). Cameroon, Democratic Republic of Congo, and Kenya (fig. 24)................................................. luxuriosa Spaeth, immaculate form
122. Punctation of elytra on entire disc appears regular, at most in postscutellar impressions punctures can appear irregular and few additional punctures in median intervals do not disturb regularity of rows (figs. 204, 207, 239, 340)................ 123.
- Punctation along middle of elytron appears completely irregular, also postscutellar impressions irregularly punctate, only first two sutural and five lateral rows regularly punctate (figs. 254, 255). Angola, South Democratic Republic of Congo, and Zambia (fig. 33)................................................................................ patruelis Spaeth
123. Postscutellar impressions deep, well visible, H–shaped elevation complete (figs. 196, 207, 340)................... 124.
- Postscutellar impressions absent or very shallow, hardly visible, H–shaped elevation rudimental or completely absent (figs. 239, 240). Mozambique and South Africa (Natal) (fig. 20).......................... heroni sp. nov., immaculate form
124. Body slimmer, oval, L/W ratio 1.27–1.42 (figs. 196, 207). Claws with large basal tooth.......................... 125.
- Body stouter, circular, L/W ratio 1.24 (fig. 340). Claws with small basal tooth. Southern Ethiopia (fig. 48)..................................................................................................... somalica Spaeth
125. Explanate margin of pronotum punctate. Elevated second interval of elytral disc complete (fig. 207). Slightly larger and stouter, length 4.10–4.85 mm, L/W ratio 1.27–1.37. Burkina Faso and Niger (fig. 44).......... villiersi sp. nov., immaculate form
- Explanate margin of pronotum impunctate. Elevated second interval of elytral more or less interrupted or absent, occasionally complete (fig. 196). Slightly smaller and slimmer, length 3.75–4.55 mm, L/W ratio 1.32–1.42. Kenya (fig. 47).................................................................................... voiensis sp. nov., immaculate form
126. Claws simple, sometimes base of claw slightly widened and forms small tooth. Species from southern Africa north to southern Malawi .......................................................................................... 127.
- Claws with large basal tooth. Small species, length 3.95–4.90 mm, stout, with base of elytra not or only slightly wider than pronotum, elytral disc strongly convex (figs. 303, 304). Subsaharian Africa south to southern Tanzania (fig. 36)................................................................................................ praetimida Spaeth
127. Larger, length above 5.6 mm......................................................................... 128.
- Smaller, length below 5.6 mm........................................................................ 133.
128. Body slimmer, L/W ratio above 1.20. Punctation of elytra finer (figs. 301, 309, 315, 317, 318, 334)................. 129.
- Body stouter, almost circular, L/W ratio 1.17–1.19 (fig. 331). Elytral disc moderately convex (fig. 332). Punctation of elytra very coarse, punctures almost touching each other. South Africa: Western Cape (fig. 11)................................................................................................ foveolatipennis Borowiec & Świętojańska
129. Punctation of pronotal disc coarse and dense, distance between punctures mostly smaller to as wide as puncture diameter (figs. 301, 309, 317).................................................................................... 130.
- Punctation of pronotal disc fine and sparse, distance between punctures wider than puncture diameter. Base of elytra distinctly wider than pronotum, elytral disc distinctly convex (figs. 315, 316). South Africa: Eastern Cape (fig. 9)............................................................................. diversepunctata Borowiec & Świętojańska, 2001
130. Eyes small, distance between lower margin of eye and lateral angle of labrum longer than half length of eye.......... 131.
- Eyes larger, distance between lower margin of eye and lateral angle of labrum shorter than half length of eye......... 132.
131. Base of elytra as wide as base of pronotum (figs. 317, 318). Antennal segment 7 only slightly longer than wide. Larger, length 6.1–7.6 mm, and very convex (fig. 319). S Zimbabwe, Malawi and Mozambique (fig. 18)............................................................................................ limpopoana Borowiec & Świętojańska, 2001
- Base of elytra slightly wider than base of pronotum (fig. 309). Antennae slimmer, segment 7 more than twice longer than wide. Smaller, length 5.8–6.2 mm, and slightly convex (fig. 310). South Africa: Cape (fig. 9)............. distinguenda Spaeth
132. Pronotum regularly semicircular, with maximum width at base (figs. 333, 334). Body stouter, L/W ratio 1.20–1.33, sides regularly convex (figs. 333, 334). Elytral disc feebly convex (fig. 335). South Africa: Cape (fig. 30)............................................................................................ pudens Boheman, large specimens
- Pronotum inversely trapezoidal, with maximum width in the middle (figs. 300, 301). Body slimmer, L/W ratio 1.35–1.65, sides distinctly converging posterad (figs. 300, 301). Elytral disc strongly convex (fig. 302). Namibia and South Africa (fig. 29).................................................................. melanophthalma Boheman, large specimens
133. Explanate margin of elytra extremely declivous, almost perpendicular to surface................................ 134.
- Explanate margin of elytra moderately to strongly declivous but never appears perpendicular to surface............. 135.
134. Smaller, L 4.15 mm, body slimmer, L/W 1.66 (fig. 293). Elytra more convex (fig. 294). Punctation of elytral disc homogenous. South Africa (Western Cape) (fig. 35).................................................... oxylepiformis sp. nov.
- Larger, L 4.75–5.30 mm, body stouter, L/W 1.46–1.51 (fig. 307). Elytra less convex (fig. 308). Punctation of elytral disc heterogenous, between large punctation numerous fine punctures. South Africa: Cape (fig. 31).......... wittmeri sp. nov.
135. Punctation of pronotum sparse to dense, finer than punctation of elytral disc, basal part of pronotal disc never appears strongly rugose. Elytra slightly converging posterad (figs. 295, 298, 300, 311, 313, 333)................................. 136.
- Punctation of pronotum more extremely coarse and dense than punctation of elytral disc, basal part of pronotum appears strongly rugose. Elytra strongly converging posterad, disc strongly convex (figs.291, 292). South Africa: Cape (fig. 27)............................................................................... lycii Borowiec & Świętojańska
136. Body slimmer, L/W ratio 1.39–1.65, sides distinctly converging posterad (figs. 295, 298, 300)..................... 137.
- Body stouter, L/W ratio below 1.39, sides regularly convex or only slightly converging posterad (figs. 311, 313, 333)...................................................................................................... 139.
137. Punctation of pronotum fine and sparse, distance between punctures mostly wider than puncture diameter; punctures shallow, often hardly visible................................................................................ 138.
- Punctation of pronotum coarse and dense, especially on basal part of disc, distance between punctures mostly narrower than puncture diameter. Namibia and South Africa (fig. 29)................ melanophtkalma Boheman. small, slim specimens
138. Slimmer, L/W ratio 1.46–1.65. Punctation of elytral disc slightly coarser and denser, microreticulation between punctures finer (figs. 295–297). Wide spread from Zambia to South Africa (fig. 26)............................... litigiosa Boheman
- Stouter, L /W ratio 1.38–1.45. Punctation of disc slightly finer and sparser, microreticulation between punctures stronger (figs. 298, 299). Southern Africa: Western Cape only (fig. 37)...................... reticulipennis Borowiec & Świętojańska
139. Explanate margin of elytra more declivous, distinctly narrowed posterad. Pronotum with maximum width in front of base, if close to base then sides slightly rounded (figs. 300, 301, 305, 311, 313)....................................... 140.
- Explanate margin of elytra less declivous, broad in apical part. Pronotum regularly semicircular, with maximum width at base, sides angulate, elytral disc slightly convex (figs. 333, 334). South Africa: Cape (fig. 30).................................................................................................. pudens Boheman, rare small specimens
140. Punctation of basal part of pronotal disc fine and shallow, often hardly visible, if distinctly visible and quite dense then elytra with regularly rounded sides (fig. 311)................................................................. 141.
- Punctation of basal part of pronotal disc coarse and dense. Elytra distinctly converging posterad (figs. 300, 301). Elytral disc strongly convex (fig. 302). Namibia and South Africa (fig. 29)......... melanophthalma Boheman, small, stout specimens
141. Punctation of elytra dense, punctures disposed regularly, no impunctate intervals. Base of elytra usually distinctly wider than base of pronotum (figs. 305, 313). Species outside Namibia ................................................ 142.
- Punctation of elytra sparser, punctures disposed irregularly, having a tendency to form two impunctate intervals, especially in area homologous to third and anterior part of fifth interval. Base of elytra as wide as or only slightly wider than pronotum (fig. 311). Elytral disc slightly convex (fig. 312). Species from Namibia (fig. 34).......................... spatiosa Spaeth
142. Punctation of elytra finer and sparser, surface of elytra does not appear rugose (fig. 313)......................... 143.
- Punctation of elytra coarse and extremely dense, punctures almost touching each other (fig. 305), surface of elytra appears rugose, especially on slope. Elytral disc strongly convex (fig. 306). South Africa: Western Cape (fig. 8)...................................................................................................... cordula Boheman
143. Punctation of elytra finer, but appears more dense (figs. 313, 314). Surface between punctures very reticulate, appearing slightly dull. Base of elytra distinctly wider than base of pronotum (fig. 313). Cape Province in South Africa (fig. 10)............................................................................. chrysanthemoides Borowiec & Świętojańska
- Punctation of elytra coarser, but appears less dense. Surface between punctures less reticulate, appearing slightly glabrous. Base of elytra only slightly wider than base of pronotum. South Africa: Eastern and Western Cape Provinces (fig. 34)............................................................................. spatiosiformis Borowiec & Świętojańska
description: Published as part of Borowiec, Lech & Świętojańska, Jolanta, 2022, A monograph of the Afrotropical Cassidinae (Coleoptera: Chrysomelidae). Part 6. Revision of the tribe Cassidini 3, the genus Cassida L., pp. 1-250 in Zootaxa 5171 (1) on pages 8-22, DOI: 10.11646/zootaxa.5171.1.1, http://zenodo.org/record/6966207