Candezea Chapuis, 1879

Wagner & Kurtscheid, A., 2005, Revision of Candezea Chapuis, 1879 (Coleoptera, Chrysomelidae, Galerucinae) from continental Africa, Journal of Natural History 39 (28), pp. 2591-2641 : 2594-2601

publication ID	https://doi.org/ 10.1080/00222930500102611
persistent identifier	https://treatment.plazi.org/id/C13187E1-C46B-FF9F-FE3E-FAEFB0C1E8E9
treatment provided by	Felipe
scientific name	Candezea Chapuis, 1879
status

Treatment

Type species: Monolepta occipitalis Reiche, 1847 by monotypy.

Redescription

Total length. 5.7–8.1 mm (adults), means of single species 6.09–7.14 mm. Usually females slightly longer and broader than males. Larvae and pupae unknown.

Head. Eyes convex, ovate. Labial and maxillary palpi slender ( Figure 2B View Figure 2 ) yellow to brownish yellow, mandible yellow, apically dark brown or entirely black. Clypeus and frons yellow, brown or black, labrum black, without any significantly incised sutures. Antenna with 11 slender and elongated antennomeres ( Figure 1 View Figure 1 ). Mean antennal length 4.1 mm, antenna reaching towards apical quarter of the elytra. Three basal antennomeres yellow to brownish yellow, antennomeres 4–11 yellow, brownish yellow, black ( C. occipitalis ), or antennomeres yellow at base and black at apex, i.e. antennae appear annulate ( C. haematura ). Third antennomere on average about twice as long as second, relative length of second to third antennomere 0.55–0.56, range of means for single species ( Figure 2A View Figure 2 ), third antennomere about two-thirds as long as fourth, length of third to fourth antennomere 0.62–0.64).

Thorax. Pronotum yellow or brownish yellow, rarely black. Pronotal width 1.8–2.5 mm (mean for single species 1.98–2.17 mm), length to width of pronotum 0.42–0.59. Pronotum sub-rectangular, posteriorly significantly extended in the middle ( Figure 2C View Figure 2 ). Procoxal cavities nearly closed ( Figure 2D View Figure 2 ), prosterna and hypomera usually yellow, sometimes brown or black. Meso- and metathorax mostly dark brown or black, only in C. flaveola usually yellow. Metasternum very broad ( Figure 2E View Figure 2 ). Length of elytron 4.3–6.3 mm (mean for single species 4.82–5.79 mm), elongated, maximum width in most species in the last third. Maximum elytral width 2.8–4.2 mm (mean for single species 3.12–3.90 mm), maximum width to length of elytron 0.59–0.73. Elytra yellow to brownish yellow, in some species with reddish brown apex, or irregularly arranged circular black spots. Males of all species have sexual dimorphic structures on the elytra. Males of C. occipitalis , C. atomaria , C. haematura , and C. flaveola have small, heart-shaped elytral protrusions beyond scutellum only ( Figures 6A View Figures 6–9 , 15A View Figures 15–18 , 29A View Figures 29–32 , 34A View Figures 34–37 ), males of C. costatipennis have these protrusions with protruded elytral ridges from humerus towards the apical third of elytra ( Figure 20A View Figures 20–23 ), males of C. bicostata have two pairs of protruded elytral ridges ( Figure 24A View Figures 24–27 ), and males of C. irregularis and C. franzkrappi sp. nov. have deep, ovate depressions beyond scutellum ( Figures 11A View Figures 11–14 , 39A View Figures 39–42 ). Alae fully developed ( Figure 3 View Figure 3 ). Scutellum triangular, yellow, brown, or black. Legs usually yellow to brownish yellow throughout, sometimes darker towards base of femora, in C. bicostata coxae and trochantera black, in C. haematura coxa, trochantera and femora black, in C. occipitalis legs black throughout. Basi-metatarsus elongated and slender ( Figure 4A View Figure 4 ). All tibiae with single apical spur, longest on metatibiae.

Abdomen. Yellow, brown, or black, significantly wider in females. Five visible sternites, anal sternite in males apically with a pair of deep incisions ( Figure 5A View Figure 5 ), in females rounded ( Figure 5B View Figure 5 ).

Female genitalia. Spermatheca 0.40–0.55 mm long, with small nodulus and long, usually strongly curved cornu (cf. Figure 8 View Figures 6–9 ). Bursa sclerites larger than spermathecae (0.69– 1.10 mm), strongly sclerotized, with strong spines ( Figures 7 View Figures 6–9 , 12 View Figures 11–14 , 16 View Figures 15–18 , 21 View Figures 20–23 , 25 View Figures 24–27 , 30 View Figures 29–32 , 35 View Figures 34–37 , 40 View Figures 39–42 ), number of these spines varies between species from four to eight, number within single species sometimes slightly variable.

Male genitalia. Aedeagus without parameres, with short tegmum, attached in the basal third (cf. Figure 9 View Figures 6–9 ). Outer shape and endophallic structures symmetrical, length 2.3– 3.0 mm. Median lobe straight, elongated and usually slender, apex carinated ventrally. Tectum long, subapically widened, often lanceolate, apically slightly incised. Endophallus with three distinct pairs of strong, usually curved spiculae, and weakly sclerotized sac structure. Orifice usually small. For terminology see Figure 9 View Figures 6–9 .

Distribution. This study includes data of 10,735 specimens of Afrotropical Candezea for which locations could be sufficiently identified from the label data, and thus distribution of most species is now known very well. Candezea mainly occur in humid savanna and forest biomes south of the Sahara to northern South Africa (cf. Table II). No species is known from Mali, Burkina Faso, Benin, Togo, Niger, Chad, Botswana, and most parts of other countries with predominantly dry savannas and deserts such as Nigeria, Sudan, Somalia, Namibia, and western South Africa.

Diagnosis. A general overview of the generic characters of most taxa mentioned in this paper was given recently ( Wagner 2003b). A comparison of Candezea with phylogenetically related species-rich taxa is given here. In comparison to Monolepta , Afrocrania , and Afrocandezea , Candezea are large Galerucinae. Total length varies from 5.7 to 8.1 mm, average of single species from 6.09 to 7.14 mm, while total length of Monolepta is 3.0– 7.5 mm ( Wagner 1999, 2000a, 2000b, 2001, 2002, 2003a), Afrocrania is 4.5–6.8 mm ( Middelhauve and Wagner 2001), and Afrocandezea is 4.2–6.7 mm ( Wagner and Scherz 2002). Afrocrania has much narrower elytra (width of both elytra to maximal length of elytron 0.57–0.62), and Afrocandezea has on average broader elytra (0.65–0.85) than Candezea (0.59–0.73). Pronotum is very broad in Candezea (length to width 0.42–0.59) but narrower in most species of Monolepta (0.52–0.65), Afrocandezea (0.52–0.72), and Afrocrania (0.59–0.67), the latter has also a pronotum significantly narrowed at base. The third antennomere is much longer than second in Candezea (length of antennomere 2– 3 0.49–0.63) which is similar in Afrocrania (0.57–0.80), and Afrocandezea (0.53–0.80), but different from Monolepta where second and third antennomeres are approximately of the same length (0.82–1.10).

Basic coloration of pronotum and elytra of Candezea is yellow to brownish yellow, while most Afrocandezea and Afrocrania are brown, reddish brown, or pale yellow. Most Monolepta species have no uniform coloration and show combinations of red and black, yellow and black, or red, yellow and black often forming large spots or transverse bands on the elytra. Neither in Afrocandezea and Afrocrania , nor in Monolepta , do irregularly scattered elytral spots occur, which is a unique character of several Candezea species.

External sexual dimorphic patterns of males do not occur in Monolepta , but are characteristic for Candezea , Afrocrania , and Afrocandezea . Males of several Afrocrania have unique patterns and sculpturation like head cavities sometimes combined with horned basal antennomeres, or complex folded elytral extrusions (cf. Middelhauve and Wagner 2001), but there are also some species of Afrocrania and all species of Afrocandezea where males show small, heart-shaped elytral protrusion beyond scutellum as in some species of Candezea .

Outline of median lobe and endophallic structures are also different from those of Monolepta , Afrocrania , and Afrocandezea . The median lobe of Afrocrania is more strongly curved in lateral view, tectum is very slender and much shorter than the median lobe ( Middelhauve and Wagner 2001). Afrocandezea has a longer tectum, and is characterized by one pair of strong fixed spiculae with apices protruding laterally outside the median lobe ( Wagner and Scherz 2002). In contrast to Candezea , tectum and apex of median lobe in Monolepta and Afrocrania are not incised. Endophallic structures of Afrocrania and Afrocandezea are relatively similar to those of Candezea , but Afrocrania differs remarkably by having only two pairs of slender spiculae, one of them curved the other straight. In Monolepta three different types of distinct spiculae occur which do not occur in any species of Candezea , Afrocandezea , and Afrocrania ( Wagner 2000a) . Also outline and size of the spermathecae and bursa sclerites of Afrocrania , Candezea , and Afrocandezea are more similar in comparison to Monolepta , but the long curved cornu of Candezea is an easily recognizable character to distinguish females of Candezea from those of Afrocandezea and Afrocrania .

Identification key for Afrotropical Galerucinae with elongated basi-metatarsi, lacking pronotal depressions; valid only for recently revised taxa

1 At least parts of head, pronotum, and elytra metallic blue or green; in most species dorsum entirely metallic; median lobe broad, less than 4× longer than broad at orifice...................... 2

– Without any metallic coloration; median lobe slender, more than 5× longer than broad at orifice.................... 3

2 Legs brownish yellow, antennomeres very slender, pronotum very broad (pronotal length to width 0.39–0.49); endophallus without spiculae; widely distributed in Guineo-Congolian forests...................... Barombiella Labossière, 1931 (one species; Wagner and Freund 2003)

– Legs brown or black, antennomeres shorter, pronotum narrower (ratio pronotal length: width 0.54–0.58); endophallus with two or three asymmetrical endophallic spiculae; in forests and savannas of sub-Saharan Africa............. Bonesioides Laboissière, 1925 (21 species; Freund and Wagner 2003)

3 Second and third antennomere subequal in length (ratio length of antennomere 2:3 0.80–1.20); endophallus with three different types of spiculae or with two pairs of spiculae carrying rows of spines.............. 4

– Third antennomere much longer than second (ratio length of antennomere 2:3 0.45–0.80); endophallic armature absent or different......... 5

4 Small, total length 3.2–4.1 mm, elytra yellow with symmetrically arranged small black spots, endophallus with two pairs of spiculae having rows of spines; occurs only in savannas and semi-deserts, most species restricted to southern Africa..... Afromaculepta Hasenkamp and Wagner, 2000 (six species)

– Most species much larger, total length 3.0– 7.5 mm, elytral coloration very variable, in some species with transverse black bands or cross-like markings, never with symmetrically arranged small black spots; in all biomes throughout Africa............... Monolepta Chevrolat, 1837 (ca 100 species; Wagner 2000a, 2000b, 2001, 2002, 2003a, further parts in preparation)

5 Very large, total length 8.8–9.9 mm; elytra and abdomen black, head and prothorax red to reddish brown; in Central, East, and South Africa.............. Afromegalepta Schmitz and Wagner, 2001 (one species)

– Total length less than 8.5 mm ............... 6

6 Small, total length 2.4–4.9 mm, pronotum trapezoidal, elytra usually slender (ratio width of elytra together: length of elytron 0.39–0.75), legs short (ratio length of basi-metatarsi: metatibiae 0.34–0.50); median lobe without endophallic spiculae, deeply incised at apex; throughout Africa................. Galerudolphia Hincks, 1949 (15 species; Bolz and Wagner 2005)

– Total length 4.2–8.1 mm, pronotum rectangular or significantly narrowed in the basal third, elytra usually broader (ratio width of elytra together: length of elytron 0.57–0.75), legs usually much longer (ratio length of basi-metatarsus: metatibia more than 0.50); median lobe with endophallic spiculae, apex of median lobe not incised...................... 7

7 Elytra very narrow (ratio width of elytra together: length of elytron 0.57–0.61), pronotum very narrow (ratio pronotal length: width 0.59–0.67); males with complex sexual dimorphic structures on frons, basal antennal articles or elytra; median lobe very slender, slightly enlarged at apex, with very short tectum and two pairs of endophallic spiculae; throughout Africa............ Afrocrania Hincks, 1949 ( Middelhauve and Wagner 2001 (partim); ca 20 species)

– Elytra broader (width of elytra together to length of elytron 0.62–0.86), males without those complex sexual dimorphic structures at frons, antenna, or elytra...................... 8

8 Tibiae and tarsi black, femora contrasting yellow, elytra pale yellow; median lobe very slender, tectum very small, endophallus with many small, short spiculae; in savannas of eastern and southern Africa........................ Afrotizea Stapel and Wagner, 2001 (one species)

– Femora, tibiae, and tarsi concolorous, if contrasting, femora black and tibiae yellow; one to three pairs of strong endophallic spiculae........ 9

9 Large, total length 5.7–8.1 mm; elytra comparatively slender (ratio width of elytra together: length of elytron 0.62–0.70), all species with irregularly scattered spots belong to this group; median lobe with three pairs of strong endophallic spiculae; throughout Africa.... Candezea Chapuis, 1879 (eight species, this study)

– Smaller, 4.2–6.7 mm, if total length more than 5.5 mm elytra much broader.. 10

10 Males with deep cavity on vertex; median lobe arrow-like apically; one pair of curved endophallic spiculae, uniformly reddish brown; only known from forest of western Central Africa...................... Monoleptocrania Laboissière, 1940 (one species; Stapel and Wagner 2001)

– Males without head cavity; median lobe parallel-sided at apex; two or three pairs of endophallic spiculae, and one pair of fixed spiculae protruding outside the median lobe; not uniformly reddish brown; throughout Guineo-Congolian forests....... Afrocandezea Wagner and Scherz, 2002 (15 species)