Waxiella egbara (Cockerell)

Hodgson, Chris J. & Peronti, Ana L. B. G., 2012, 3372, Zootaxa 3372, pp. 1-265: 203-211

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Waxiella egbara (Cockerell)


Waxiella egbara (Cockerell)  

( Figs 91, 94, 95; Map fig. 105; Table 8)

Ceroplastes egbarum Cockerell, 1899: 127   .

Ceroplastes egbarum Cockerell   ; Brain, 1920b: 28.

Ceroplastes africanus var. cristatus Green, 1899: 190   . Synonymised by Fernald, 1903: 151.

Waxiella africanus cristatus (Green)   ; Ben-Dov, 1986: 166.

Waxiella egbara (Cockerell)   ; De Lotto, 1971: 148.

Ceroplastes egbarum subsp. fulleri Cockerell & Cockerell   in Cockerell, 1902b: 113. Syn. nov.

Waxiella egbara fulleri (Cockerell & Cockerell)   ; De Lotto, 1971: 148.

Ceroplastes egbarum subsp. rhodesiensis Hall, 1931: 294   . Syn. nov.

Gascardia egbara rhodesiensis (Hall)   ; De Lotto, 1965: 181.

Waxiella egbara rhodesiensis (Hall)   ; De Lotto, 1971: 148.

Ceroplastes martinoi Almeida, 1969: 15   . Syn. nov.

Waxiella martinoi (Almeida)   ; Almeida, 1973: 7.

Ceroplastes ugandae Newstead, 1911a: 94   . Syn. nov.

Ceroplastes zonatus Newstead, 1917: 32   . Syn. nov.

Material examined. Ceroplastes egbarum Cockerell. Labelled   “Part of type material”: West Africa [ Nigeria], Egbaland , Abeokuta, on mimosa, Strachan (BMNH)   : 3/3 (poor, all old females, mainly bits. Also: as previous ( USNM) 1/1 (good condition) (remounted by Y. Ben-Dov, 1981).

Other material labelled W. egbara   : South Africa, Natal, on Acacia sp.   , May 1915, collector unknown ( SANC, Brain #87): 2/2, mounted from Brain's dried material by De Lotto (good but, due to denseness of anal cone, posterior end of abdomen hard to interpret).

Ceroplastes africanus var. cristatus Green.   South Africa: One slide labelled “ Ceroplastes africanus   (crossed out) var. cristatus Green   = egbarum Ckll   [Giant Insect Wax], Natal, ex coll. Brit. Mus.” (poor, 1 old female, mainly bits) ( BMNH).

Ceroplastes egbarum var. fulleri Cockerell   : Lectotype ♀ (here designated): South Africa: left label: Ceroplastes   / egbarum / fulleri Ckll / TYPE / (Fuller). right labels (top)   : Lectotype; (bottom) Remounted / Y. Ben- Dov, 1981 ( USDA): 1/1?ad (very poor; old female, venter and dorsum separate)   .

Paralectotype ♀: left label: 10592 / CO-TYPE / T.D.A. Cockerell, com. / Let. April 4, 1904; right labels (top): Ceroplastes   / egbarum / fulleri. / Cotype; (bottom) Remounted / Y. Ben-Dov, 1981 ( USDA): 1/1 (very poor)   .

Other material identified as Ceroplastes egbarum subsp. fulleri Cockerell & Cockerell   : front labels: left: Natal Coast / Umbilo and / Equeefa Rivers / on Monkey Rope / CKB 8a, and right: Ceroplastes   / egbarum fulleri (both labels in E.K. Hartwig’s handwriting); labels on reverse side of slide: left CKB no. 88a / S. Afr.: Transvaal / Pretoria / June 1918 / ex. Acacia sp.   / coll.? Delport and right label: Waxiella   / mimosae / (Sign.) (both labels in De Lotto’s handwiting) ( SANC, Brain #88a): 4/4 (good-fair) (see Discussion below).

Ceroplastes egbarum var. rhodesiensis Hall.   Lectotype ♀ (here designated): Rhodesia ( Zimbabwe): top label, scratched onto glass slide: Ceroplastes   / egbarum var. / rhodesiensis / Hall / Hunters / Road / 30/10/23; bottom label: TYPE / 14.iii.30 / (2) WJHall ( BMNH): 1/?1 (poor, old, broken into bits).  

Paralectotype ♀: as for lectotype slide, but labelled Ceroplastes egbarum var. rhodesiensis   ? Hall. Hunters Road, Bulawayo, 11/9/21, ex coll. Dept. Agriculture, COTYPE, 14.iii.30 (i) ( BMNH): 1/1 (poor, old, broken into bits)   .

Ceroplastes martinoi Almaeida   : Syntypes: Angola: Bruco , on undetermined plant ( CZLP): 1/ 1 in poor condition. Almeida did not designate a holotype in his original description of this species. The labels on this slide are as follows: I-Hom. Sintipos 3-4; Waxiella martinoi (Almeida)   comb. n. Dino, det., Lisboa- Portugal Direita- J.I.U. Centro de Zoologia Entomologia Reg. 5547 ( IICA); Angola (Bruco) Reg. 3828 Lisboa – Portugal   .

Ceroplastes ugandae Newstead.   Lectotype ♀ (here designated): 1 specimen on 2 slides, both with left label: Colonial Office / Ent. Res. Com. (T.A.) / A / Amakebe specimens; right label: Ceroplastes   / ugandae / Newstead / Co-type ♀♀ / 1912-152” ( BMNH): 2/1 (mainly poor; these slides refer to just one old specimen (despite being labelled ♀♀), with dorsum on one slide and venter on other).  

Other material labelled C. ugandae   : Kenya, Nairobi , on wattle, 18.v.1925, H. Wilkinson ( BMNH, IBE 1350): 1/1 (good, young). French West Africa [perhaps Burkino Faso], Upper Volta, Prosopis juliflora   , -. ix.1962, F. Brunck ( BMNH, CIE 8893 - A495): 1/3 (fair)   . Uganda, Entebbe , on Cajanus indicus   , 20.iii.1913, C.C. Gowdey ( BMNH): 1/2 (fairly young but poor); as previous but dated 30.iii.1913 ( BMNH, IIE 814 a): 2/3 (young, fair to good); as previous but dated 30.v.1913 ( BMNH, IIE 814 b): 1/1 (old, fair); no site, on unknown host, 1.vi.1913, C.C. Gowdey ( UCDC): 1/1 (fair)   .

Ceroplastes zonatus Newstead.   Lectotype ♀ (here designated): South Africa: left label: Dept. Agric., Mauritius / BM 1945, 121; right label “ Ceroplastes   / zonatus, / Newst. / Co-type ♀ / RN” ( BMNH)1/1 (fair). For explanation of Dept. Agric., Mauritius, see under C. bipartitus   above.  

Paralectotype ♀: As for lectotype ( BMNH): 1/1 (poor)   .

Other material labelled W. zonatus   . South Africa, ex collection Dept. Agric. Pretoria, det. Brain, no. 344 ( BMNH, received 24.x.1928): 1/2 (young, fair). Malawi, Chitedzi Res. Stn., on Albizia lebbeck   , 4.vii.1966, C.J. Hodgson ( BMNH): 2/2 (young, good). Tanzania [Tanganyika Terr.], no site, on Samanea saman   , no date, A.H. Ritchie ( BMNH): 2/2 (young, fair); as previous but on Enterolobium saman   (= Samanea saman   ) ( BMNH) 1/1 (young, fair; identified as C.? africanus   ). Zimbabwe [Southern Rhodesia], Inyangombe Falls, Brachystegia sp.   , 25.v.1964, C.J. Hodgson ( BMNH): 2/5 (mainly good-fair); Harare [Salisbury] Botanical Gardens, Acacia sp.   , 1.ix.1967, C.J. Hodgson ( BMNH): 1/1 (good, split into dorsum and venter); Harare [Salisbury], Melia sp.   , 8.ii.1924, W.J. Hall ( BMNH): 1/1 (youngish, good).

Also: Côte d’Ivoire, Rives du Nzo, Zoba (Man), no host, 21.vii.1978, G. Remaudiere ( MNHN #7463): 1/1 (good; identified as ‘ C. mimosae   group’). Gabon, Makokou Ivindo, no host, 20.x.1973, A.S. Balachowsky ( MNHN): 4/4 (fair; identified as W. mimosae   ). Kenya, Machakos, on Acacia seyal   , 4.viii.56, G. De Lotto ( BMNH): 1/1 (good). Mali, Barnako-Siguini, on Pterocarpus esculentus, Sept. 1918   , J. Vuillet ( MNHN #7647): 3/3 (good-fair; identified as Ceroplastes sp.   ). Namibia, Rundu   , 1.i.2002, on Acacia sp.   , P.J. Gullan ( DCBU): 1/2 (fair). Nigeria, Bauchi, on Acacia sp.   , -. iv.1990, J. Deeming ( BMNH): 1/1 (fairly young, fair; collected at same time as 2 specimens of W. senegalensis   ); Agege and at Moor Plantation, on pigeon peas, no date, W.A. Lamborn ( BMNH): 2/several (mainly poor, in bits; as W. vuilleti   ). Sierra Leone, Njala, on Dalbergia sexatilis   , -. ii.1932, Cockerell ( MNHN #7645): 1/1 (old, poor; identified as Ceroplastes sp.   ). South Africa, Limpopo Province [Northern Transvaal], Witbank, Acacia sp.   , 10.ii.1972, C.G.P. van Dyk ( SANC, 4502): 3/3 (young, good; identified as W. mimosae   ); Gauteng Province [Transvaal], Pienaarspoort, on unknown plant, 20.vii.1963, C.J. Cilliers ( SANC, 1318): 1/1 (young, good; identified as Gascardia mimosae   ); Gauteng Province, Pretoria, no host plant, 24.viii.1918, C.P. Lounsbury ( SANC, 516): 1/1 (old, good, mounted by De Lotto; identified as C. mimosae   ); Limpopo Province [Northern Province], Lekgagameetse Nature Reserve, on Dalbergia armata   , 15.i.1997, I.M. Millar ( SANC, 6727): 3/6 (young, good; unidentified); KwaZulu Natal, Greater St. Lucia Wetlands Park, Sta Lucia township, 15.vii.2008, G. Svenson ( DCBU): 1/1 (fair).

Democratic Republic of the Congo, Eala, on Legume, -. vii.1936, J. Ghesquière ( MNHN, TERV): 2/8 (fair, very young); Eala, on Entada sp.   , 18.iii.1936, J. Ghesquière #4020 ( MNHN, TERV): 2/8 (good-fair); Bitshumbi [Tshumbe?], on Acacia sp.   , 21.x.1933, de Witte ( MNHN #7648): 5/5 (good; identified as C. africanus   ); Rutshuru, on Albizzia   [ grandistipulata   ], 7.i.1938, J. Ghesquière #4186 ( MNHN, TERV): 4/4 (good-poor).

Note. The main description below is taken from Brain’s #87 material from South Africa. Data from a selection of the other material are given in (..) brackets. See also Table 8.

Unmounted material. “Female test broadly ovate in outline, highly convex; marginal plates very faintly indicated, but apparently without nuclear spots; dorsal plate very large, with a central nuclear spot of white wax; cephalic margin slightly clypeate; lateral margins in very old examples with a pronounced foot-like extension from each stigmata, from which extends a thick white waxen appendage. In the young forms, the foot-like extension is wanting, but the white waxen appendages are always present and always porrected. Colour creamy white, suffused with very pale brown; dorsal plates surrounded by a patch of dull flesh-colour. In very old examples, the zone of colour extends to the margins and is a shining madder-brown to piceous colour. On the removal of the outer surface of the test with chloroform, it is seen to be divided into seven areas by pale coloured lines: a central polygonal area, corresponding to the area occupied by the dorsal plate, from the angles of which radiate to the margin single lines marking off the areas of the lateral and cephalic plates.” “Female adult (denuded of the test). Ovate, cephalic margin clypeate; dorsum low and wrinkled; with 2 large bilateral extensions, both longitudinally striated; the space between these extensions of the body wall and the margin markedly constricted. Dorsum with a large keel-like process. Caudal process very short and conical. Stigmatic clefts deep.” (Description of W. zonata, Newstead 1917: 32   ). Hodgson (1969a, p. 38) illustrates the waxy test and the adult female with the wax removed (of W. zonatus   ).

Mounted material. Body elongate oval, probably quite convex, with distinct, shallow, stigmatic clefts; dorsum without distinct tubercles. Stigmatic setae of 2 types, sharply-spinose setae along margin and conical setae in a roundly oval group on dorsum, each group 200–275 (425–850) µm long and 250–390 (375–700) µm wide. Caudal process short and stout, 0.72–0.8 (0.52–2.0) mm long, 0.75–0.83 (0.45–2.0) mm wide. Body length 2.88–3.20 (1.7–8.0) mm, width 2.25–2.5 (1.3–6.5) mm.

Dorsum. Derm membranous except for heavily sclerotised caudal process plus roundly-oval sclerotised areas associated with each group of conical stigmatic setae; old specimens with some sclerotisation developing along body margins (particularly ventrally along cephalic margin) and narrowly around each group of conical stigmatic setae. Derm: number of clear areas uncertain (8 distinct clear areas present, dorsal rather narrow). Dorsal setae each quite long and sharply spinose, often slightly bent, with more or less parallel sides towards base (narrowing slightly on longest); each 16–24 (15–24) µm long; basal socket width 6.5–7.0 (6.0–7.0) µm; present sparsely throughout except in clear areas (occasionally 1 or 2 in anterior clear area). Dorsal pores: (i) loculate microducts of complex type, each with 1–4 small, more or less round, satellite loculi; those with 1 satellite loculus mainly along margin; those with 2 & 3 loculi most abundant, each about 5 (4–6) µm wide; fairly evenly distributed throughout apart from clear areas; pores too few to detect wax-plate lines (but see comments below); and (ii) simple microducts: distribution uncertain but present in some clear areas. Preopercular pores present in a transverse line (13–37 in a band 1–3 pores wide). Anal plates each with 3 long dorsal setae (each 35–50 µm long); length of each plate 145 (145–155) µm, width of both plates combined 157 (150–175) µm. (Anal ring setae each 140–170 µm long).

Margin. Marginal setae similar to dorsal setae; each anal lobe with 3 (1–3) long setae, longest about 50 (30–40) µm long. Stigmatic clefts generally shallow and broad basally, each with 2 types of stigmatic setae: (a) sharply-pointed spinose setae in a group of 36 (14–65) on ventral surface associated with spiracular disc-pores, each 20–22 (13–23) µm long; group often wider than base of group of conical setae; and (b) roundly conical setae, in a large, generally almost round, group on dorsum; most conical setae about 11–14 µm long, 7.5–8 µm wide, but generally with 1 larger seta placed either rather centrally or slightly towards dorsal margin, each up to about 16 (17–33) µm long; each cleft with at least 250–300 (200+, but see discussion below) conical setae; each group with 8 (5–14) conical setae along basal margin, 48 (31–65) around dorsal margin and with about 22 (14–33) across widest part of group. Each group of conical setae on mature specimens surrounded by a narrow area of heavy sclerotisation, sclerotisation generally only slightly broader than width of group of conical setae. Eyespots each about 40 (30–40) µm wide.

Venter. Derm entirely membranous apart from anterior margin of head which becomes heavily sclerotised on older specimens. Pregenital disc-pores abundant around genital opening (segment VII) and across preceding 2 segments; with a group of about 11 (0–17) medially and 20 (0–13) mediolaterally on IV; 0 (0–3) medially and 0–2 (0–9) mediolaterally on III; and 0 (0–3) medially on II, absent more anteriorly. Spiracular disc-pores present in bands (narrower than width of peritreme) of at least 120–150 (80–150) pores, each band widening considerably near margin, with disc-pores intermingled among sharply-spinose stigmatic setae; with few or no disc-pores extending medially past peritremes. Ventral microducts present sparsely throughout most of venter (very sparse medially on anterior abdominal segments and perhaps absent on posterior segments). Ventral tubular ducts, some with a slender inner ductule, present in a group of about 30–35 (about 25–40) in cephalic area anterior to antennae but otherwise only rarely present laterally on head, thorax and abdomen; also present medially on abdominal segments: V 8 (0–5), VI? (1–11), VII? (0–3+ plus occasionally 1 or 2 associated with anogenital fold). Submarginal setae frequent, each 15–20 µm long; other setae infrequent.

Antennae each with 8 (8) segments, all segments distinct (but Brain (1920b) suggests that they might be 6–8 segmented, presumably due to variation in pseudo-articulation in segment III); total length 410–460 (323–465) µm. Clypeolabral shield about 230 (200–270) µm long. Spiracles: width of peritremes 95–100 (75–130) µm. Legs well developed, each with a well-developed tibio-tarsal articulatory sclerosis; claw denticle obscure or absent; claw digitules both very broad and slightly shorter than tarsal digitules; dimensions of metathoracic legs (µm): coxa 180–210 (152–220); trochanter + femur 225–250 (190–265); tibia 173–185 (150–182); tarsus 93–102 (80–105), and claw 35 (30–36).

Discussion. At least eight species/subspecies in the africana   -group of Waxiella   have been described from central and southern Africa, namely: W. africana   , W. africana var. cristata   , W. egbara   , W. egbara var. fulleri   , W. egbara var. rhodesiensis   , W. martinoi   , W. ugandae   and W. zonatus   . It is here considered that these represent only 2 species, W. africana   and W. egbara   , differing in the number of stigmatic setae, the basic shape of the conical setal group and in the width of sclerotisation around each group of conical setae on mature specimens. Ceroplastes (Waxiella) africana var. cristata   was synonymised with W. egbara   by Fernald (1903) and this synonymy is here accepted. In addition, the following are here synonymised with W. egbara   : Ceroplastes (Waxiella) egbarum var. fulleri Cockerell & Cockerell   , Ceroplastes (Waxiella) egbarum var. rhodesiensis Hall   , Ceroplastes (Waxiella) martinoi (Almeida)   , Ceroplastes (Waxiella) ugandae (Newstead)   and Ceroplastes (Waxiella) zonatus Newstead.  

The specimens of W. (Ceroplastes) egbara fulleri   listed above in SANC are not part of the type material of this subspecies. According to De Lotto’s notes in his inventory (see under C. eucleae   above), the “original slide labels” were those of E.K. Hartwig, who copied the data from Brain’s paper (1920b: 30). However, De Lotto points out that the data given in Brain’s accession book indicates that the specimens were collected “in Pretoria in June 1918 by a certain Delport on Acacia sp.   ” De Lotto considered that the specimens were W. mimosae   but here we believe that W. mimosae   does not occur in southern Africa and consider these specimens to be W. egbara (Ckll)   .

The distinction between the details (number of setae, basic shape of conical setal group, width of sclerotisation around group of conical setae, etc) which are here used to separate W. africana   and W. egbara   are fairly obvious on most specimens. However, a few young specimens can be difficult to place, particularly in northern South Africa, because, on young specimens, the group of conical setae are unsclerotised and tend to be tightly packed and so cannot be easily counted. Occasional specimens have about 150–200 conical setae (i.e., intermediate between W. africana   and W. egbara   ) and/or a broad area of sclerotisation around each conical group, as on W. africana   . Most of these specimens can usually be placed by counting the number of conical setae along the basal margin – when less than about 5 or 6, almost certainly W. africana   but, if more, probably W. egbara   .

Similarly, the distinction between mature W. egbara   and mature W. senegalensis   is reasonably easy based on the characters suggested here. However, immature adult females, in which the sclerotisation has yet to set in, are extremely difficult to place with confidence. Of the specimens studied here, many from West Africa remain uncertain ( Guinea, Foulaya, host unknown, 15.ix.1947, no collector (MNHN # 7644); Senegal, Matam, on Acacia sp.   , 17.viii.1979, B. Sigwalt (MNHN # 7263); similar data but on A. nilotica   , 18.viii.1979 (MNHN # 7865); similar data but from Richard Toll, 16.viii.1979 (MNHN # 7862)). These can look very similar to W. mimosae   , but lack the multilocular disc-pores posterior to each metacoxa and medially on the metathorax. Perhaps it should be noted here that Sigwalt, who collected some of the above Senegal specimens, also collected W. senegalensis   in Senegal at about the same time. See under W. senegalensis   below.

Waxiella martinoi Almeida   is here considered to be a synonym of W. egbara   . The specimen depicted in Almeida (1969) is clearly an old adult. The single slide of this species seen during this study (see above) was poor but it is clear that the group of conical stigmatic setae is about as wide as long, with a broad base, and the margin shows little sign of a distinct cleft. Also, there are about 27 & 32 sharply-spinose setae along the base of 2 of the clefts, and 1 group has about 55 conical setae around the dorsal margin. These characters are typical of W. egbara   , even though the vignette Almeida provides of the stigmatic area (1969: p. 16) suggests a wide outer sclerotised margin, not present on the only available specimen.

Newstead (1917) mentioned that the inner wax of the test (when the outer layers have been removed with chloroform) was divided into areas (plates) by lines. From his description (quoted above under Unmounted material), it is clear that these lines relate to the “wax-plate lines” noted on some Ceroplastes species   above. However, on Ceroplastes species   , the lines are apparent on the mounted specimens due to a slightly denser grouping of loculate microducts, particularly pores with a larger number of satellite loculi. Wax-plate lines were not detected on any mounted specimens of Waxiella   .

Based on the above records, there have been many misidentifications in the past and the list of countries and host plants given in ScaleNet ( Ben-Dov et al., 2011) needs to be revised. Our study suggests that W. egbara   has been recorded from Angola, Côte d’Ivoire, Democratic Republic of the Congo, Gabon, French West Africa [Burkino Faso?], Kenya, Malawi, Namibia, Nigeria, South Africa, Tanzania, Uganda and Zimbabwe within the Afrotropical Region and from Saudi Arabia. The above records are all on Fabaceae   : Acacia spp.   , Albizia lebbek   , Brachystegia spiciformis   , Cajanus indicus   , Dalbergia armata   , Mimosa sp.   , Samanea saman   , Prosopis juliflora   and wattle, but Hodgson (1969a) also included Bidens bipinnata   ( Asteraceae   ) and Hall (1931) mentioned Melia azedarach   ( Meliaceae   ).


Smithsonian Institution, National Museum of Natural History


Agricultural Research Council-Plant Protection Research Institute


United States Department of Agriculture


R. M. Bohart Museum of Entomology


Museum National d'Histoire Naturelle


Universidade Federal de Sao Carlos














Waxiella egbara (Cockerell)

Hodgson, Chris J. & Peronti, Ana L. B. G. 2012

Waxiella africanus cristatus (Green)

Ben-Dov, Y. 1986: 166

Waxiella martinoi (Almeida)

Almeida, D. M. de 1973: 7

Waxiella egbara (Cockerell)

De Lotto, G. 1971: 148

Waxiella egbara fulleri (Cockerell & Cockerell)

De Lotto, G. 1971: 148

Waxiella egbara rhodesiensis (Hall)

De Lotto, G. 1971: 148

Ceroplastes martinoi Almeida, 1969: 15

Almeida, D. M. de 1969: 15

Gascardia egbara rhodesiensis (Hall)

De Lotto, G. 1965: 181

Ceroplastes egbarum subsp. rhodesiensis

Hall, W. J. 1931: 294

Ceroplastes egbarum Cockerell

Brain, C. K. 1920: 28

Ceroplastes zonatus Newstead, 1917: 32

Newstead, R. 1917: 32

Ceroplastes ugandae

Newstead, R. 1911: 94

Ceroplastes egbarum subsp. fulleri

Cockerell, T. D. A. 1902: 113

Ceroplastes egbarum Cockerell, 1899: 127

Cockerell, T. D. A. 1899: 127

Ceroplastes africanus var. cristatus

Fernald, M. E. 1903: 151
Green, E. E. 1899: 190