Waxiella subsphaerica (Newstead)

Hodgson, Chris J. & Peronti, Ana L. B. G., 2012, 3372, Zootaxa 3372, pp. 1-265 : 228-232

Waxiella subsphaerica (Newstead)

( Fig. 102; Map fig. 105)

Ceroplastes subsphaericus Newstead : 1911b: 166.

Waxiella subsphaerica (Newstead) ; De Lotto, 1971: 148.

Ceroplastes berliniae Hall, 1931: 291 . Syn. nov.

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

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

Ceroplastes berliniae var. enkeldoorni Hall : 1931: 292. Syn. nov.

Gascardia berliniae var. enkeldoorni (Hall) ; De Lotto, 1965: 181.

Gascardia enkeldoorni (Hall) ; Hodgson, 1969a: 25.

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

Material studied. Ceroplastes subsphaerica Newstead. Lectotype ♀ (here designated): East Africa ( Tanzania): left label: Tanzania / (D. Ost. Africa) / Ngambe / on Albizia lebbeck / 27.vi.1902 / Prof. A. / Zimmermann; right label: Ceroplastes / subsphaeroides / = subsphaericus / Newstead. / Mounted from / dried material / by / C. J. Hodgson, 2011 ( BMNH): 1/1 (old, poor, broken into many bits but with some venter). Although the vial was labelled C. subsphaeroides , this is clearly the material that Newstead later referred to as subsphaericus .

Paralectotype ♀: data as for lectotype ( ZMB): 2/2 (old, poor, dorsum broken into several bits, with almost no venter; mounted from spirit material by T.K. Qin, 1990). Also as previous ( ZMB); 2 / 2 specimens (probably immature but poor). Also 1 vial with 2 dried old females with above collection data but labelled C. subsphaeroides (BMNH) .

Ceroplastes berliniae Hall : Lectotype ♀ (here designated): Zimbabwe (Southern Rhodesia): top, scratched on glass: Ceroplastes / berliniae / Hall / Berlinia / globifera / Mazoe / WJH 14/8/27; bottom: TYPE / 12.iii.30 WJH ( BMNH): 1/2ff (lectotype top specimen, arrowed; in good condition; labelled on envelope as Holotype; see discussion on “type” specimens in Materials and methods above).

Paratype ♀, remaining specimen on lectotype slide, plus: as for lectotype ( BMNH): 1/4 (fair) .

Also identified as C. berliniae : Malawi, Mzuzu , no host, 17.ix.1970, E.S. Brown #2088 ( BMNH): 1/2 (fair) . Madagascar, Tsimbazaza , on Acacia dealbata , March 1950, A. Robinson #83 ( BMNH): 2/2 (fair-poor) . Democratic Republic of the Congo, Bukaru [Costermansville], on Albizia sp. , 7.iv.1938, J. Ghesquière #6700

( MNHN, TERV): 4/11 (good-poor); [Belgian Congo], on indigenous tree, no date, Ringoot ( BMNH): 1/5 (fair, labelled Ceroplastes congoensis Newstead , a m/s name). Zimbabwe [Southern Rhodesia], Mazoe, on Brachystegia sp. , 24.v.1932, W.J. Hall ( BMNH): 1/3 (good); also Rusape, on Brachystegia sp. , 11.ix.1927, W.J. Hall ( BMNH): 1/5 (fair); also McIlwaine, on unknown host, Aug. 1964, C.J. Hodgson ( BMNH): 1/1 (fair).

Ceroplastes berliniae var. enkeldoorni Hall : Lectotype ♀ (here designated): Zimbabwe [Southern Rhodesia]: top, scratched onto glass slide: Ceroplastes / berliniae var. / enkeldoorni / Hall / Enkeldoorn / 28.10.1929; bottom: ex. Coll. Dept. of / Agriculture, and on paper label: Cotype / 12.iii.30 / 3 WJH ( BMNH): 1/1 (poor, old, mainly bits).

Paralectotype ♀: as for lectotype ( BMNH): 1/1 (poor, old, mainly bits) .

Also identified as C. enkeldoorni : Cameroon, no site, on Bridelia sp. , 27.xi.1988, A. Dejean ( MNHN #11279): 2/5 (good). Côte d’Ivoire, Oumé, on Caesalpiniaceae , 12.xii.1979, F. Brunck ( MNHN #8070): 8/18 (fair-good). Mozambique [Portuguese East Africa], Musapa Gap, on Acacia (Albizia) adianthifolia , 7.vii.1967, C.J. Hodgson ( BMNH): 2/2 (good). Zimbabwe, Wankie Main Camp, on Baikiaea plurijuga , 3.ix.1967, C.J. Hodgson ( BMNH): 1/1 (good); as previous but on Acacia albida (BMNH) : 1/1 (poor).

Also: Gabon, Makokou, no host, 12.xi.1973, A.S. Balachowsky ( MNHN #5884): 5/5 (old, poor; identified as W. africana ); Makokou, Ivindo River, Loaloa, on leaves and woody twigs, 9.xii.1973, A.S. Balachowsky ( MNHN #5858; identified as Waxiella sp. ).

Note. The following description was made from the material of W. berliniae Hall. Where data were available for the type series of W. subsphaerica , these are in […] brackets.

Unmounted material. ( W. berliniae ) "Test of adult female white, very highly convex and approximately oval in outline, the short axis being the longitudinal axis of the insect. A very deep mediodorsal longitudinal depression is usually present giving a bilobed appearance. In the centre of this depression is a small, flattened, transversely striated whitish boss which often carries indications of longitudinal striations in addition. In very young individuals, this boss very nearly covers the entire insect. Opercular plates black, or very dark, and sunken; in some examples, this and the mediodorsal depression are continuous. The wax is soft in fresh specimens and unusually deep in relation to the size of the insect. Fusion frequently takes place where individuals are contiguous. A localised tinge of pink is seen in some specimens. Denuded of its wax test, the adult female is oval in outline, being longer than broad. The central portion is raised into a conical dome which has a tendency to be directed posteriorly. The margin is flattened and rounded, and it appears as though the central dome is set on a cushion. The cephalic lobe is comparatively well developed. Stigmatic clefts deep and conspicuous (from the under surface of the insect). A cephalic and three bilateral tubercles are present; these are small and conical, but in some specimens they are very much better developed than others. Caudal process short, conical, with its axis at approximately 45˚ with that of the main body. Length of test of adult female 5–6.5 mm; breadth 8–9.5 mm, length of adult female 4 mm; breadth 2.5 mm ” ( Hall, 1931: 291).

Unmounted material (of W. subsphaerica ). “Test of old female thin, divided into large lateral plates with nuclear centres; dorsum forming a large hemisphaerical or dome-shaped mass, at the sides of which the wax is so thin that the dark colour of the insect shows through.” ( Newstead, 1911b: 166). (This description almost certainly refers to old dried material that had lost much water from the waxy test).

Mounted material. Body almost round; stigmatic clefts distinct and quite broad; derm membranous apart from caudal process; caudal process well developed and conical; dorsum with small, distinct lateral tubercles. Length 2.6–3.3 mm, width 2.4–3.1 mm.

Dorsum. Derm membranous but becoming heavily sclerotised on old specimens; caudal process heavily sclerotised, but rather short and conical. Clear areas all small but distinct, although dorsal clear area often appearing absent. Dorsal setae each small, about 3–5 µm long, usually rather longer than width of basal socket (about 3 µm wide) but variable between collections, shortest near margins, generally with sides converging to a moderately sharp or fairly blunt apex, only rarely parallel-sided; about half as abundant as dorsal pores but present throughout, although very sparse or absent from more posterior and dorsal clear areas. Dorsal pores: (i) loculate microducts of complex type, each 4–5 µm wide with 1–3 satellite loculi; sparsely scattered over dorsum but absent from all clear areas; pores too few to detect wax-plate lines, and (ii) simple microducts present throughout. Preopercular pores small, rather few, sometimes possibly absent but generally 6–10 around anterior margin of anal plates [few, perhaps 10]. Anal plates each with 1 long, robust dorsal seta, perhaps 50 µm long [only 1 present], plus 2 smaller setae nearer apex; also with a small subapical seta about 25 µm long; length of each anal plate 105–125 [108] µm; width of both plates combined 110–140 µm. Anal tube short, subequal to length of anal plates; anal ring setae each about 100 µm long.

Margin. Marginal setae rather variable, with sharply-spinose setae in a line of about 12–18 anteriorly between eyespots, each about 12–14 µm long; setae becoming less spinose along rest of margin and some tending to be finely setose along margin of abdomen; sparse along margins of thorax and abdomen; anal lobe setae in a group of 3, longest 60–66 µm long. Stigmatic clefts distinct, shallow, each with 2 types of stigmatic setae: (i) those along margin on venter each sharply spinose, often slightly curved and slightly larger than marginal setae, each 13–18 µm long and 2.5–4.0 µm wide at base, with 6–27 in each cleft; and (ii) roundly conical setae in an elongate group at least twice as long as wide extending radially onto dorsum; setae rather uniform in size, most 5.5–7.5 µm wide at base and 7–8 µm long but generally with a single larger spinose seta set near dorsal end of group; with 20–55 [31–36] setae in each cleft. Eyespots each about 30–32 µm wide.

Venter. Derm entirely membranous. Pregenital disc-pores abundant around genital opening and in preceding segment ( VI), plus a few in segment V [certainly present in VI, perhaps also in V]. Spiracular disc-pores in narrow bands near peritreme but broadening to slightly narrower than group of sharply-spinose stigmatic setae near margin; each band with about 50–60 disc-pores. Ventral microducts small, each about 2.5 µm widest, rather sparse and only slightly more abundant submarginally; present medially on abdomen. Ventral tubular ducts entirely absent. Submarginal setae sparse but more abundant than marginal setae.

Antennae well developed, each with 7 or 8 [7] segments, extra segments clearly due to subdivision of segment III; total length 227–305 µm. Clypeolabral shield 175–185 [215] µm long. Spiracles: width of each peritreme 44–55 [57] µm. Legs well developed; each with a distinct tibio-tarsal articulatory sclerosis; each claw sometimes with a minute denticle; claw digitules both broad; tarsal digitules subequal to length of claw digitules; dimensions of metathoracic leg (µm): coxa 115–145 [128–132]; trochanter + femur 150–185 [145–160], tibia 103–135 [114–116] and tarsus 66–83 [65–75], claw 23–28 [25].

Discussion. C. subsphaericus Newstead was described from material collected in German East Africa ( Tanzania) as part of an expedition from the Berlin Zoological Museum. It was sent to Prof. Newstead to identify. However, the mounted type specimens, which should have been present in the BMNH, could not be located. Nonetheless, a small box labelled ‘ Ceroplastes subsphaeroides’ [sic] with exactly the same collection data was found. This is clearly the only material of this species in the BMNH. The box contained 3 heavily sclerotised old females, one of which was mounted. In addition, 4 slides were located in the Berlin Zoological Museum which had been mounted by Dr. T. Qin in 1990, obviously also from old, heavily sclerotised adults. The specimen mounted from the dried material in the BMNH has been chosen as the Lectotype because it has some venter with most structures visible (the venter is missing from all BZM material). All of the mounted adult specimens are poor, but the stigmatic clefts can be seen on all of them and the arrangement and frequency of the conical stigmatic setae falls within the range for W. berliniae . Unfortunately, none of the marginal setae in the cephalic region could be located but, nonetheless, almost all that can be gleaned from these 3 adult specimens falls well within the range for W. berliniae . It is therefore considered that these 2 species are the same and so W. berliniae is here synonymised with W. subsphaerica .

In addition, with the availability of more specimens and with some collections having several specimens, it has been possible to check the frequency of certain character states used to separate C. enkeldoorni from C. berliniae , particularly by Hodgson (1969a). The possession of a small claw denticle is not here thought to be a diagnostic characteristic and the frequency of stigmatic setae is now shown to vary considerably even within single collections (and on single specimens) and so, despite the differences discussed by Hodgson (1969a) in the structure of the waxy test, it is here considered that C. berliniae var. enkeldoorni is a synonym of C. berliniae and therefore also a synonym of C. subsphaerica .

The available material of W. subsphaerica (as shown by specimens of C. berliniae ) differs from W. subdenudata in having the marginal setae along the anterior margin of the head all long and sharply spinose, whereas on W. subdenudata they are short and bluntly pointed. In addition, the dorsal setae on W. subsphaerica are mostly clearly longer than the width of the basal socket (particularly away from the margin), and mostly narrowing to a moderately sharp apex, whereas on W. subdenudata they are all short and blunt, subequal in length to the width of their basal sockets and with more or less parallel sides. However, the material from Gabon (for instance) has dorsal setae very similar to those of C. subdenudata and so this may not be a good differentiating character. In addition, W. subdenudata appears to have fewer spinose and conical stigmatic setae but, as little material of W. subdenudata has been seen, it is here considered that the name C. subdenudata could also be a junior synonym of C. subsphaericus .

Qin and Gullan (1995), in their morphological cladistic study of the wax scales, recorded only one difference for the characters scored between W. enkeldoorni and W. berliniae , but considered that W. subsphaerica was sufficiently different to record it separately, scoring 3 differences: (i) Qin considered that a dorsal clear area was absent on W. berliniae but present on W. subsphaerica — we considered that the specimens of the latter were too poor to be sure but that a small dorsal clear area was sometimes present on W. berliniae ; (ii) Qin scored more than 10 marginal setae between lateral stigmatic clefts on W. berliniae and 5 or less on W. subsphaerica — we recorded very few on both, and (iii) Qin scored a claw denticle as present on W. berliniae and absent on W. subsphaerica . We considered that a minute claw denticle was present or absent on W. berliniae but was absent on W. enkeldoorni . None was detected on W. subsphaerica .

Based on our study and on the available literature, it appears that this species has now been recorded in the Afrotropical Region from: Angola ( Almeida, 1969, 1973), Cameroon, Côte d’Ivoire, Democratic Republic of the Congo, Gabon, Malawi, Mozambique, Tanzania, Zambia and Zimbabwe and also from Madagascar. It has been recorded on the following hosts: Annona sp. (Annonaceae) ; Bridelia sp. (Euphorbiaceae) , Caesalpiniaceae , Acacia dealbata , A. albida , Albizia sp. , A. adianthifolia , Baikiaea plurifuga , Berlinia sp. , B. globiflora , Brachystegia sp. , B. boehmii , B. flagristipulata , B. longifolia , B. spiciformis , B. tamarinoides , B. utilis , Guibourtia coleosperma and Pericopsis angolensis (Fabaceae) ; and Manilkara macaulayae (Sapotaceae) .