Cissococcus fulleri, Cockerell, 1902

Hodgson, C. J., Millar, I. M. & Gullan, P. J., 2011, Cissococcus Cockerell (Hemiptera: Coccoidea: Coccidae), a unique gall-inducing soft scale genus on Vitaceae from South Africa, with description of a new species, Zootaxa 2996 (1), pp. 1-32 : 7-25

CISSOCOCCUS FULLERI Cockerell

( Figs 1A–G , 4–12 )

Cissococcus fulleri Cockerell, 1902: 23 ; Ferris, 1919: 112; 1920: 63; Steinweden, 1929: 233; Ben-Dov, 1993: 63; Hodgson, 1994: 178; Beardsley, 1997: 337–338.

Type material: Material of this species is divided among institutions, as follows: BMNH: SOUTH AFRICA, “ Type ”, pres. Cockerell, 1920.153, 1/1ad ♀, poor (this is one of the specimens illustrated in fig. 34 in Hodgson (1994) as the “ type ”). BME: 6/1 ad ♀, 9 1 st -instar nymphs, 1 exuviae of immature male nymph, one box with 3 dry galls of females, 2 labels: “fulleri, Cissococcus / On Cissus TYPE, S. Africa ” and “Part of type / galls of Cissococcus / fulleri, Ckll S. Africa / On Cissus (Fuller) ”, the dry material and 2 slides are from the G.F. Ferris collection, 4 slides are recent mounts from the dry material by T. Kondo 2003. USDA: 2/2ad ♀ (very poor) labelled: Co-types: right label only: Cissococcus / fulleri, Ckll / Cotype / S. Africa .

Other material studied: SOUTH AFRICA, Eastern Cape (EC), East London, Orange Grove, roadside, 33° 02’S, 27° 49’E, on Rhoicissus tridentata , attended by ants of Crematogaster (Crematogaster) desperans , 16.xi.2005, P.J. Gullan (ANIC; BME; SANC: HC 6966 ): 23/10 ad ♀ (including DNA voucher TK0154), 3 3 rd -instar ♀, 53 1 st -instar nymphs, 9 2 nd -instar ♂♂, 1 ad ♂ pharate in pupa + 1 ad ♂ + associated dry galls; EC, Boesmansriviermond , on bank of Boesmansrivier , 33° 40’S, 26° 39’E, 25 m, ex Rhoicissus sp. , 26.xi.2005, P.J. Gullan ( ANIC; SANC: HC 6967 ): 22/5 ad ♀, 27 1 st -instar nymphs 10 2 nd -instar males + associated dry galls; EC, Uitenhage, 33° 46'S, 25° 25'E, on R. tridentata , -. III.1976, H.G. Zimmermann ( SANC: HC 5988 ): 24/19 ad ♀, 14 1 st -instar nymphs, 3 3 rd -instar ♀ ; Mpumalanga (MPU), Houtenbek Farm, NW of Dullstroom, Macabelel Lodge , 25°18’S 30°03’E, on leaves of Rhoicissus , 2.xii.2002, I. M. Millar ( SANC: HC 6915 ): 57/9 ad ♀, 10 3 rd -instar ♀ + 2 pharate 3 rd instar, 18 2 nd -instar + 1 pharate 2 nd instar, 8 2 nd -instar ♂♂, 7 1 st -instar nymphs + 1 pharate 1 st instar, 8 prepupae, 13 pupae and 3 ad ♂♂ + plant specimens with unopened galls; MPU, Goedehoop Farm, 15 km W of Lochiel, The Brook trail, 26°10’S 30°38’E, on leaves of Rhoicissus , 29.xi.2002, I.M. Millar ( SANC: HC 6916 ): 12/1ad ♀, 15 2 nd -instar ♂♂ + 2 pharate 2 nd instar, 33 1 st - instar nymphs; Northern Province , Lekgalameetse Nature Reserve , 24º10S 30º15E, 15.i.1997, R. tridentata, I.M. Millar ( SANC: HC 6726 ): 6/4 ad ♀, 4 2 nd -instar ♂♂, 15 1 st -instar nymphs .

Note. The type locality was specified by Cockerell (1902) as “Umquahumbi Valley, S. Africa” with Claude Fuller as the collector. We have been unable to locate Umquahumbi except that we believe it was a tributary of the Illovo River ( Russell, 1903). The closest town to that region is Richmond, KwaZulu-Natal (Grid references 29°52’S and 30°16’E). In 1916, Fuller also collected the material of the Cissococcus species that Brain (1918) mistook for C. fulleri and that we describe later in this paper as a new species. Both of Fuller’s collections came from KwaZulu-Natal; the new species that was confused with C. fulleri was from the Natal coast near Durban, whereas Fuller’s earlier collection was made farther inland.

Gall structure: Each gall of female ( Fig. 1A,B ) develops on stem of host plant. Initially, each gall appears as a small convexity with an apical opening. When fully mature, gall brownish, hairless, globular or cup-shaped, sometimes almost cylindrical, 5–6 mm in diameter, slightly less tall; basal attachment to plant usually broad (at least half diameter of gall); gall surface broken into polygonal, slightly raised brown ‘barky’ areas, each separated by greenish fissures. Top of gall flat and with a small concavity with a small central orifice, about 0.3–0.4 mm in diameter. When cut open, walls thick, fleshy and green. Adult female scale insect lying within a large, approximately round space, with body filling cavity and dorsum placed in gall opening. A short glassy white wax protrusion sometimes present, extending from anal plate area through orifice in gall; the glassy secretion is produced as a filament from each anal-plate seta and each dorsal marginal seta and these filaments combine to form a broad brush-like protrusion. Sometimes 2 or more galls coalesce.

ADULT FEMALE ( Fig. 4 ). Described from 6 specimens in good condition.

Unmounted material. When removed from gall, body dark burgundy pink. Body slightly dorsoventrally flattened, with a covering of white wax on one, only slightly rounded, surface, which shows strong signs of segmentation and so clearly referable to median areas of venter (i.e. lower venter (see below)). Other, upper surface, more convex, with two converging lines of white wax ( Fig. 1B ) in shallow grooves extending to small sclerotised dorsum – presumably wax secreted by bands of spiracular disc-pores which extend from spiracles to dorsum. Position of vulva possibly indicated by a small medial indentation on lower venter just anterior to dorsum.

Mounted material. See generic diagnosis. Body approximately round and apparently highly convex. Radius 1.5–5.5 mm.

Dorsum. Total length of dorsum 470–600 µm; greatest width 350–420 µm. Derm sclerotised throughout, with an irregularly fissured pattern; fissures arranged semi-longitudinally over posterior two-thirds of dorsum, tending to radiate from median to margin over anterior third. Derm with many pale pores of uncertain structure, about 3 µm wide. Dorsal setae hard to discern but each rather spinose, 5–7 µm long, when broken off position shown by small pore-like sockets, each about 1 µm wide; frequent throughout. Anal plates together oval, each 190–240 µm long; together 170–190 µm wide, dorsal surface of each plate with 75–90 spinose setae, covering almost entire surface; each seta 25–35 µm long, but with 1 or 2 near apex 45–65 µm long. Eyespots distinct, each slightly displaced from margin near anterior edge of dorsum, each 23–26 µm wide.

Margin. Marginal setae very similar to those on dorsal surface of anal plates but sometimes somewhat curved, each mainly sharply spinose but occasionally blunt or bifid, 25–50 µm long, in more or less a single line along margin, bases almost touching, with 100–120 on each side.

Venter. Becoming enormously swollen in mature specimens as described under generic diagnosis. Microducts frequent throughout, each about 2 µm wide. Loculate pores abundant medially on lower venter, generally appearing to be segmentally arranged (frequency of loculate pores and number of loculi somewhat variable between collections); each pore with mainly 5–10 loculi, but a few with 11 or 12 loculi, and each 6–10 µm wide; rarely (HC 5988) with a long inner ductule arising from central loculus, each ductule about 17 µm long. Similar loculate pores (but structure often less distinct) also associated with each spiracle on upper venter; number of pores somewhat variable between collections. Setae: lower venter setae abundant and long, each 17–40 µm long (up to 75 µm long on HC 6915); those medially on upper venter infrequent and short, mostly 8–12 µm long.

Antennae distorted, but each about 65 µm long; apical segment with 3 or 4 fleshy setae plus a few other setae; apical seta 30–45 µm long; two preapical segments each with 1 fleshy seta; other segments obscure; antennae located just anterior to anterior margin of dorsum on upper venter. Spiracles quite large, width of peritreme: anterior 60–65 µm, posterior 70–80 µm. Legs as per diagnosis.

Comment. For comparison with adult female of C. braini , see under that species below.

THIRD-INSTAR FEMALE ( Fig. 5 ). Described from about 8 specimens in good condition and with reference to another 11.

Unmounted material. Gall similar to that of adult female but much smaller.

Mounted material: obovate in shape and symmetrical; length 0.85–1.07 mm from posterior margin to folded anterior edge of venter, greatest width (measured between lateral edges of folded venter) 0.67–0.85 mm; anal cleft well-developed, about 1/7th length of dorsum; stigmatic clefts absent.

Dorsum: similar to that of adult female but smaller, length 0.51–0.63 mm, width 0.32–0.40 mm. Edges of dorsum next to anal plates heavily sclerotized, with a transverse bar just anterior to anal plates, measuring 90–112 µm wide and 5–7 µm long. Dorsal setae small, each seta 4–6 µm long, restricted to 2 mediolateral lines, each line with 5–7 setae, positions difficult to determine but probably a pair on head, 1 on each thoracic segment and 1 on several abdominal segments. Dorsal pores appearing as round clear pores, arranged submarginally in a loose line, double in a few places, with 21–31 pores on each side. Anal plates together obovate, outer margins rounded, each plate 143– 160 µm long, 55–65 µm wide, and with 3–8 thick setae on dorsal surface, pointing posteriorly, each 22–57 µm long, plus an apical seta, 33–40 µm. Anal fold possibly with 2 pairs of setae along anterior margin, very difficult to detect, longest about 15 µm; plus a seta ventrally on each anal plate, presence/absence hard to see. Anal ring with 3 pairs of setae, each seta 115–137 µm long; anal ring pores probably numbering 20 or more, difficult to discern due to folding of anal ring; anal tube 125–145 µm long.

Margin: marginal setae stoutly spinose, sharply pointed, each 13–36 µm long and with a well-developed basal socket; arranged in a single line around margin, with (on each side) 4–8 between apex of head and eyespot, 37–62 along remainder of margin; setae set close together along abdominal area, becoming more widely spaced on thorax and head; occasionally a few also present submarginally. Eyespots prominent, dome-shaped, each 18–23 µm wide, on margin.

Venter: considerably larger than dorsum, especially in head and thoracic areas, so that antennae located on upper venter on flattened, slide-mounted specimens; legs appear to be placed near edges of mounted body; derm membranous, with rows of fine spinules on abdomen; segmentation not evident. Preanal disc-pores absent. Spiracular disc-pores each with 5–8 outer loculi and a small, round central loculus; each pore band with 3–9 disc-pores. Ventral microducts each with a short broad outer ductule and a long narrow inner ductule about 20 µm long, present mainly anterior to mouthparts but also with a very few elsewhere. Ventral setae sparse; with a pair of long preanal setae on abdominal segment VII, each seta 33–75 µm long; about 6–8 short submarginal setae in a row on each side, each 8–13 µm long, most abundant along posterior edge of abdomen but also on thorax and head; other setae 6–24 µm long, positions difficult to determine owing to distortion of body; with 2 pairs of interantennal setae located some distance nearer mouthparts, longer pair 25 µm long, shorter pair 12 µm long.

Antennae well developed, each 98–118 µm long, 6-segmented with segment VI longest. Antennal segment lengths: I 18–26 µm, II 8–12 µm, III 15–22 µm, IV 10–12 µm, V 12–15 µm, VI 28–35 µm; setal distribution typical of Coccidae ; apical stiff seta 32–48 µm long. Clypeolabral shield normally developed, 200–217 µm long; labium 2- segmented, with 3 pairs of setae. Spiracles small, width of each peritreme 23–27 µm. Legs well developed but generally misshapen, data for metathoracic leg: coxa 25–30 µm long, with 5 setae, longest 20–35 µm; trochanter + femur 50–62 µm long, trochanter with 1 long and 1 shorter seta; each femur with 2 setae; tibia 45–57 µm long, with 2 setae; tarsus 68–75 µm long, with 4 setae; tibio-tarsal articulation sometimes obscure or absent; tarsal digitules extending beyond tip of claw, one digitule sometimes slightly thinner and shorter than other; claw 22–27 µm long, with a denticle; claw digitules sometimes dissimilar to each other, each extending beyond tip of claw.

Comments. Mature second- and third-instar females are distinctive and can be distinguished from all other immature instars of Cissococcus by their swollen, globular body shape, with a much enlarged venter and antennae on upper venter near dorsum. However, young teneral second-instar females are not swollen and have the venter approximately the same size as the dorsum. These two instars can be separated at all stages by the presence of the enlarged dorsal cone-shaped spines on the second-instar females (also present on second-instar males). Second- and third-instar females of C. fulleri can be separated from second-instar males by the absence of dorsal and ventral tubular ducts; and from adult females by the absence of multilocular disc-pores (frequent throughout much of the venter of the adult females) and in having much better developed legs and antennae. For differences from firstinstar nymphs, see under that instar below.

SECOND-INSTAR FEMALE ( Fig. 6 ). Described from about 8 specimens in good condition and with reference to another 10.

Unmounted material: individually enclosed within a woody gall formed on twig; gall small initially but beginning to swell by end of instar. Body becoming rotund/globose when mature, with anterior and lateral parts of venter much larger than dorsum, with result that antennae on upper venter; mouthparts ventral, legs arising ventrally and projecting beyond edges of body. Venter not visible dorsally along posterior margins of body.

Mounted material: obovate in shape and symmetrical; length, measured from posterior margin to anterior folded edge of venter, 0.56–0.87 mm; width, measured across broadest part to each folded lateral edge of venter, 0.41–0.74 mm; anal cleft short, about 1/22nd of dorsum length; stigmatic clefts absent.

Dorsum: length 0.46–0.65 mm, width 0.35–0.42 mm; derm lightly sclerotized, faintly marked with an irregular cellular pattern, marginal area roughened by sclerotized low nodules arranged in several loose rows along body edge. A heavily sclerotized transverse bar present just anterior to anal plates, 55–70 µm wide and 8–10 µm long; dorsum without intersegmental lines. Dorsal setae minute, each 5–6 µm long, restricted to 2 mediolateral lines of 5–7 setae, exact positions difficult to determine but probably with pairs on head, on each thoracic segment, and on several abdominal segments. Dorsal secretory spines large, sunken and cone-shaped, each 8–16 µm high and 7–10 µm wide at the base, distributed in 4 longitudinal lines: 2 mediolateral lines each with 8–10 spines, and 2 submarginal lines each with 9 or 10 spines. Dorsal pores small, flat and simple, distributed more or less in 6 longitudinal lines: 2 submedially, each with about 2–6 pores, 2 lines (poorly-defined) mediolaterally (between sunken conical spines) each with about 5–7 pores, and 2 submarginal lines (between sunken conical spines and margin), arranged mostly as a single pore next to each large conical spine, each line with about 6–8 pores. Anal plates together quadrate, with antero- and posterolateral margins approximately equal in length, each plate 55–62 µm long, 28–40 µm wide; each with 4 setae: an anterior inner margin seta, 8–10 µm long, a posterior inner margin seta, 6–10 µm long; an apical seta, 28–35 µm long, and an outer marginal seta (actually located on dorsal surface), each 25–32 µm long. Anal fold with 2 pairs of setae along anterior margin, longest 13 µm, plus a seta ventrally on each anal plate. Anal ring with 3 pairs of setae, each seta 62–85 µm long; anal ring pores apparently few, difficult to discern due to folding of anal ring; anal tube 58–65 µm long.

Margin: marginal setae stout and spinose, mostly sharply pointed, each 17–32 µm long with a well-developed basal socket; arranged in a single line around margin, with (on each side) 7 or 8 setae extending from apex of head to anterior stigmatic spine, 3 between stigmatic spines, and 9 or 10 from abdominal apex up to posterior stigmatic spine. Stigmatic clefts absent; but each stigmatic area with a stigmatic spine, similar to marginal spines but smaller, each 10–15 µm long. Eyespots present on margin, each 18–23 µm wide.

Venter: except in very young specimens, venter considerably larger than dorsum, especially in head and thoracic areas, so that antennae on upper venter on flattened, slide-mounted specimens, and legs appear to be placed near body edges; derm membranous; segmentation visible only to some extent, indicated on some specimens by slight folding of derm and faint sclerotisation between segments; youngest specimens with venter subequal to dorsum in shape and size. With a submarginal row of 16–22 disc-pores on each side extending around entire margin, each with 4–6 outer loculi, similar to spiracular disc-pores, interspersed with 12–15 simple flat pores, similar to those on dorsum, but located between margin and disc-pores. Preanal disc-pores absent. Spiracular disc-pores each with 4 or 5 outer loculi and a small, round central loculus; each band with 1–3 disc-pores. Ventral microducts not detected but probably present between pro- and mesocoxae. Ventral setae sparse; with a pair of long preanal setae on abdominal segment VII, each 38–75 µm long, and shorter pairs on segments V & VI, each 5–12 µm long; with pairs of short submarginal setae on each abdominal segment plus 1 or 2 setae on thorax and head; with 2 pairs of interantennal setae located some way posterior to antennae, longest pair 25–32 µm long, shorter pair 7–12 µm long; other setae very sparse.

Antennae well developed, 6 segmented, each 111–132 µm long, with segment III longest. Antennal segment lengths: I 15–20 µm, II 13–18 µm, III 28–35 µm, IV 13–20 µm, V 12–20 µm, VI 17–30 µm; setal distribution typical of Coccidae ; apical stiff seta 43–50 µm long. Clypeolabral shield 145–155 µm long; labium 2 segmented, with 3 pairs of setae. Spiracles small, width of each peritreme 12–14 µm. Legs well developed, data for metathoracic leg: coxa 50–65 µm long, with 4 setae, longest 35–45 µm, trochanter + femur 75–100 µm long, trochanter with 1 long and 1 shorter seta; each femur with 2 setae; tibia 62–72 µm long, with 2 setae; tarsus 55–70 µm long, with 2 setae; tarsal digitules extending beyond tip of claw, one digitule often slightly thinner and shorter than other; claw 18–23 µm long, with a denticle; claw digitules usually similar to each other but occasionally dissimilar and extending beyond tip of claw.

Comment. Second-instar females differ from third-instar females of C. fulleri as follows (character-states on thirdinstar female in parentheses): (i) dorsal, sunken conical spines present (absent), (ii) dorsum not fissured and much less sclerotised (fissured and much more sclerotised); (iii) setae on anal plates mainly marginal (all dorsal), (iv) dorsum more or less the same size as venter on youngest specimens (dorsum very much smaller than venter), and (v) presence of a submarginal row of loculate disc-pores (absent).

Second-instar female nymphs have cone-shaped spines on the dorsum, similar to those on second-instar males, and so it is likely that both instars secrete the long glassy tubes or filaments from these spines (as described under the second-instar males below).

SECOND-INSTAR MALE ( Figs 7 , 8 ). Main description made from about 6 good specimens from Goedehoop Farm and Lekgalameetse Nature Reserve ( Fig. 7 ). Data in [..] parentheses for specimens from Boesmansriviermond ( Fig. 8 ), which differed slightly from other male nymphs.

Unmounted material. Tend to settle against midrib on lower surface of young leaves or on petioles, often in groups. Initially mottled reddish-brown marginally, with median to submedian area translucent and marked by pairs of dark red-brown spots arranged segmentally; on mounted specimens, these spots correspond to conical sunken spines. On older individuals, dorsum becomes covered in glassy-wax tubes ( Fig. 1D,E ) secreted from each dark spot; these wax tubes often curled and as long as or longer than body length. In addition, margin covered with short, glassy-wax extrusions, probably secreted by the fringe of marginal setae.

Mounted material: oval in shape, symmetrical; length 0.84–1.28 mm, width 0.45–0.71 mm; anal cleft very short, about 1/30th total length.

Dorsum: derm membranous apart from a transverse, heavily sclerotised bar anterior to anal plates, about 80– 100 µm wide and 8–16 µm long; derm without signs of segmentation. Dorsal setae small, each 4–8 µm long; restricted to 2 mediolateral lines each with 8–10 setae, probably with pairs on head, on each thoracic segment and about 6 pairs on abdominal segments. Dorsal secretory spines very large, sunken and cone-shaped, each 25–27 [17–20] µm tall and 17–20 [14–16] µm wide; distributed more or less in four longitudinal lines: 2 mediolateral lines, each with 5–7 pores and 2 submarginal lines, each with 10 or 11 pores [secretory spines more or less absent, 1 specimen with no spines, other specimens with 1–4 plus 0–9 “scars”, randomly placed]. Dorsal pores small, flat and simple with a dark centre, distributed more or less in six longitudinal lines: 2 median lines, each with 5–6 pores, 2 mediolateral lines (between conical spines) each with about 8–12 pores and 2 submarginal lines (between conical spines and margin) each with perhaps 17–19 pores. Rarely with spiracular disc-pores extending onto dorsum near stigmatic spines. Dorsal tubular ducts of one type, each with a short, broad outer ductule and a short, broad inner ductule with a large glandular end; each outer ductule 12–13 µm long, inner ductule 10–11 µm long; in two diagonal groups of 6–10 ducts mediolaterally on abdominal segment IV; tubular ducts absent from submargin. Anal plates together quadrate, each 68–75 [62–66] µm long and 100–125 [85–95] µm wide, with posterolateral margin shortest, each plate with four setae, a very short anterior inner margin setose seta, 8–10 [5–6] µm long; a posterior inner margin setose seta, 11–17 [12–14] µm long; an strongly spinose apical seta, 30–42 [26–32] µm long, and an outer margin strongly spinose seta, located on dorsal surface, 28–42 [25–28] µm long. Anal fold with 2 pairs of setae on anterior margin, longest 18–36 µm long; and with one seta subapically on ventral surface of each plate, each 15–17 µm long. Anal ring with 3 pairs of setae, each about 110–135 [85–115] µm long; presence of anal ring pores uncertain.

Margin: marginal setae finely spinose, sharply pointed, each 20–42 [28–40] µm long; each with a narrow, well-developed basal socket; present in a single line around margin, with (on each side) 5 [5–7] anteriorly between eyespots on head, 3–5 [5–7] between eyespots and anterior stigmatic spine, 3 or 4 [7–9] between stigmatic spines, and 20–29 [24–25] on abdomen - those on head and thorax subequally spaced but becoming much closer on posterior margins of abdomen. Each stigmatic area with 1 (rarely 2) [0–1] stigmatic spines, each rather similar to marginal spines but much smaller and slightly blunter, each 12–16 µm long. Eyespots present on margin, each 15– 17 µm wide.

Venter: derm membranous, segmentation fairly distinct on abdomen; derm between antennae appearing folded, perhaps forming a concavity. Preanal disc-pores absent. Spiracular disc-pores each with mainly 5 outer loculi and a small, round inner loculus; in bands with 1–8 [3 or 4] disc-pores between spiracles and margins; when many disc-pores present, pores also occasionally present laterally on head and thorax. Ventral microducts each with a short inner ductule, sparse; perhaps restricted to single pores between pro- and mesocoxae [none located]. Preantennal pores possibly occasionally present. Ventral tubular ducts similar to those on dorsum; frequent more or less throughout venter but most abundant submarginally. Ventral setae sparse: with a pair of long preanal setae on abdominal segment VII, each 80–120 [40–75] µm long; plus shorter pairs on segments V & VI, and with pairs of short setae medially on segments III–VII; with a single short seta near each pro- and mesocoxa; submarginal setae sparse, with 1 on each side between stigmatic areas; with 2 pairs of interantennal setae, longest 33–42 [20–25] µm long; also with a pair of longer setae near anterior margin, each about 30–35 [25] µm long.

Antennae well developed, 7 segmented, each 169–200 [170–185] µm long; setal distribution typical of Coccidae [flagellate seta on segment VI missing]; apical stiff seta 40–65 µm long. Clypeolabral shield 100–130 [90–100] µm long; labium 2 segmented, with 3 [4] pairs of setae; occasionally twisted sideways. Spiracles small, width of each peritreme 17–20 µm. Legs normally developed; lengths (metathoracic) in µm: coxa 83–100 [80–90]; trochanter + femur 130–140 [115–125]; tibia 86–95 [75–85], and tarsus 78–86 [62–66]; each coxa with 3 setae, longest 60–70 µm; trochanter with 2 long setae, longest 58–62 µm; femur with 3 setae; tibia with 2 setae and tarsus with 4 or 5 setae; tarsus without campaniform pores; tarsal digitules offset, similar [dissimilar, proximal digitules longer and slightly thicker] and each extending to tip of claw [well beyond]. Claws each 25–28 µm long, with a strong denticle; claw digitules similar [1 slightly slimmer] and slightly longer than claw.

Comment. Both male and female second-instar nymphs can be quickly recognised by the presence of the large, sunken conical spines on the dorsum, although some male nymphs from Boesmansriviermond lack them. Most specimens from Boesmansriviermond had 1 or more “scars” in the position where sunken conical spines might have been expected. Second-instar males can be easily separated from second-instar females by the presence of: (i) tubular ducts on the dorsum and venter (absent on females), and (ii) 7-segmented antennae (6-segmented on females) and (iii) absence of a submarginal row of loculate disc-pores (present).

The presence and position of the groups of tubular ducts on the dorsum is very similar to those on the secondinstar males of most Coccidae . The absence of tarsal campaniform sensilla is also typical of Coccidae ( Koteja, 1974) , as is the distribution of the interantennal setae (more or less in a transverse line on Coccidae ; in longitudinal lines on Eriococcidae ) and the unilocular appearance of the ventral microducts (more nearly bilocular or cruciform on Eriococcidae ). The absence of microtubular ducts (present on many Eriococcidae ) could be through secondary loss. However, the presence of the large sunken, conical spines on the dorsum is unlike any known second-instar soft scale male (although somewhat similar structures are present on Eriococcidae , but they are not sunken), and the anal plates and associated setae are not dissimilar from those on some South American and New Zealand Eriococcidae (see Miller & González, 1975; Hodgson & Henderson, 1996), although the inner margin setae on eriococcids are generally spinose

FIRST-INSTAR NYMPH (sex not determined) ( Fig. 9 ). Described from about 5 specimens in good condition but with reference to another 23.

Unmounted material. Small, orange to rusty brown in life and rather flat ( Fig. 1D ); eyespots distinct.

Mounted material: oval but often rather more pointed posteriorly, with no signs of asymmetry; 0.5–0.8 mm long and 0.27–0.44 mm wide; stigmatic clefts absent; anal cleft very shallow.

Dorsum: derm membranous apart from a heavily sclerotized transverse bar just anterior to anal plates, about 60–65 µm wide and 7–10 µm long. Dorsal setae sparse, with 4 pairs in two parallel lines medially, with pairs on head and pro-, meso- and metathorax. Dorsal pores simple, flat, apparently with a small dark centre, each 2.5–3.0 µm wide; distributed more or less in 4 longitudinal lines: each submarginal line with 9–10 pores, each submedial line with about 10 pores. Trilocular pore near anterior margin absent. Anal plates each more or less triangular but with postero-lateral margin shortest; each plate about 45–60 µm long and 78–100 µm combined width; each plate with 4 setose setae around margin: anterior inner margin seta about 10–12 µm long, posterior inner margin seta 17– 20 µm long; long apical seta 80–100 µm long, and outer margin seta 25–30 µm long. Anogenital fold with 2 setae along anterior margin, longest 25–33 µm; plus 1 seta subapically on ventral surface of each plate. Anal ring with 3 pairs of setae, each about 55–70 µm long; anal ring pores not noted; anal tube short.

Margin: marginal setae strong and bluntly spinose, each 13–19 µm long; basal sockets only slightly broader than seta; with (on each side) 5 between apex and eyespot, and (on each side) 3 between eyespot and anterior stigmatic spine; 3 between stigmatic spines and 9 along abdominal margin, each more or less subequally spaced but posterior two close together. Stigmatic areas each with a shorter, spinose stigmatic spine, each 11–13 µm long [generally absent on Boesmansriveirmond specimens, represented by a larger gap between marginal setae]. Anal cleft with a short sclerotised protuberance where it meets outer margin. Eyespots on margin, each 10–12 µm wide.

Venter: venter apparently slightly broader than dorsum on mounted material; derm membranous; segmentation on abdomen just visible. Preanal disc-pores absent. Spiracular disc-pores each with mainly 5 outer loculi and a small central loculus; with 3 (rarely 2) in each anterior stigmatic band and 4 (rarely 3) in each posterior band. Ventral microducts not detected and possibly absent. A small pore, sometimes appearing to be trilocular, and sometimes convex, present between each scape and margin. Ventral setae few, with a pair of longer setae on each posterior 3 abdominal segments; with a submedial and a submarginal band of short setae on abdomen; other submarginal setae very few; with one pair of longer interantennal setae and another pair of long setae near anterior margin.

Antennae well developed, 6 segmented, each 105–140 µm long; setal distribution typical of Coccidae ; apical stiff seta 56–60 µm long. Clypeolabral shield 85–100 µm long; 2 segmented, with 3 pairs of setae. Spiracles particularly small, width of each peritreme 9–10 µm. Legs well developed, measurements (µm) and data for metathoracic leg: coxa 56–63, with 5 setae; trochanter + femur 96–108, each trochanter with 1 long and 1 shorter seta on ventral margin and 1(?) short seta on dorsal margin; each femur with 2 setae; tibia 50–65 with 2 setae; presence of microspines on distal end of tibia uncertain; tarsus 63–68, with 3 setae; tarsal campaniform pores absent; tarsal digitules on meso- and metathoracic legs offset, so that digitules of different lengths; one digitule on prothoracic legs rather short and almost spinose, other normal; claws quite long, each 27–29 µm long, with a distinct denticle; claw digitules more or less similar and longer than claw.

Comment. The 1st-instar nymphs of Cissococcus appear to be reasonably typical coccid crawlers. The main unusual characters are: (i) the presence of what appears to be a trilocular pore anteriorly on the venter (this is usually present on the dorsum; otherwise only known on the venter of Ceroplastodes dugesii (Signoret) (Rainato & Pellizzarri, 2008)) ; and (ii) the presence of a sclerotised bar just anterior to the anal plates, otherwise only known on first-instar nymphs of Cryptostigma (Kondo, 2010) . First-instar nymphs differ from other instars in having the very long apical anal plate seta; they also differ from second-instar nymphs in lacking the sunken cone-shaped pores.

Four lots of material have been seen (see under material studied above). Those of C. braini (described as new below) ( SANC: CKB 40) were similar to those described above except that the claw digitules were more dissimilar, with one clearly narrower than the other.

PREPUPA ( Fig. 10 ). Described from about 5 specimens in good condition plus reference to another 3.

Test. Appears to be composed of loose white wax strands, with longer wax tubes emerging through woven texture. No distinct operculum-like structure noted.

Mounted material: elongate oval; rather pointed posteriorly and rounded anteriorly. Division into head, thorax and abdomen obscure; segmentation obscure apart from on abdomen. Derm membranous, with small dermal spinules ventrally. Ducts and pores absent and setae few. Quite large: length 1.5–1.63 mm; head width about 380 µ m.

Head: lacking mouthparts and simple eyes, although mouthparts perhaps indicated by some folded derm. Antennae each moderately long, obscurely 10 segmented, pointing posteriorly, short, only reaching to prothoracic spiracle: total length 320–370 µm (ratio of total body length to antennal length 1:0.22). Setae: with 2 pairs of setae on anterior margin on both dorsum and venter, ventrally with 1 mesad to each scape, 1 pair medially on either side of mouth area, and another pair dorsally.

Thorax: unsclerotised, segmentation unclear; with 3 pairs of moderately well-developed legs; coxa and trochanter slightly sclerotised; prothoracic legs directed anteriorly but short; metathoracic legs extending posteriorly to about abdominal segment IV or V; tarsal campaniform pores absent; each tarsus with a small triangular finger on apex, probably an incipient claw; length of metathoracic legs 250–275 µm. Wing-buds (wb) on each side, extending to about abdominal segment I; mildly sclerotised; length 445–450 µm, width 160–205 µm (ratio length to width 1:0.4). With 2 pairs of spiracles, anterior pair (sp 2) each with peritreme 27–30 µm wide and with 2–5 disc-pores, each disc-pore with a highly variable number of loculi; posterior spiracles (sp 3) without disc-pores. Setae 3 or 4 dorsally and ventrally.

Abdomen: segmentation distinct ventrally, anterior-most segment considered to represent segment II. Setae: with pairs of short (5–7 µm long) dorsal abdominal setae present (ads) on segments III–VI; ante-anal setae (aas) absent; with single pairs of longer (20 µm long) ventral abdominal setae (avs) on segments III–VII; with a broken line of 19 or 20 fleshy dorsopleural seta (dps) on each side on segments I–VII; ventropleural setae (vps): with 1 minute ventropleural seta on each side from segment II to segment VII. Lobes of segment VII rounded. Lobes of segment VIII small, each with 2–4 small fleshy setae. Anus present dorsally at anterior end of penial sheath, about 21–25 µm wide. Penial sheath (ps) lightly sclerotised, slightly wider than long (about 95–125 µm long, 130–158 µm wide at base; ratio length to width 1:1.3); with a pair of minute setae on dorsal surface; genital opening present medially on ventral surface, with a pair of minute setae on either side.

Comment. The prepupa of Cissococcus is similar to that of Crystallotesta ornatella Henderson & Hodgson ( Hodgson & Henderson, 2004) in having no obvious lobes on abdominal segment VII and a line of fleshy pleural setae along the abdominal margins.

PUPA ( Fig. 11 ). Described from 5 specimens in good condition and with reference to another 8.

Mounted material: elongate oval, rounded anteriorly, rather pointed posteriorly. Segmentation obscure apart from on abdomen. Derm membranous, with small dermal spinules. Ducts and pores absent and setae few. Quite large: length 1.4–1.63 mm; head width about 340–380 µm.

Head: lacking mouthparts and simple eyes. Antennae moderately long, 10 segmented, pointing posteriorly and reaching mesocoxae: length 735–780 µm (ratio of total body length to antennal length 1:0.5); with a series of short fleshy fingers on apex; basal segments slightly sclerotised. Setae: with a group of longer setae anteriorly, mesad to each scape; with 3 small setae anterior to each procoxa, plus a seta medially on venter and a pair on dorsum.

Thorax: unsclerotised, segmentation unclear. With 3 pairs of moderately well-developed legs; coxa and trochanter slightly sclerotised; prothoracic legs C-shaped, directed anteriorly and curving round in front of anterior margin of head; metathoracic legs extending posteriorly past abdominal segment VIII; coxae each with 1 or 2 minute seta; tarsal campaniform pores absent; each tarsus with a small triangular finger on apex, probably an incipient claw; length of metathoracic legs 615–685 µm. Wing-buds (wb) extending to about abdominal segment III or IV; mildly sclerotised; length 670–690 µm, width 235–240 µm (ratio length to width 1:0.35). Anterior spiracles (sp 2): width of peritreme 34–37 µm, each with a group of 6–8 spiracular disc-pores, each with a highly variable number of loculi; posterior spiracles without disc-pores. Setae: with 2 pairs on dorsum and on venter.

Abdomen: segmentation distinct ventrally; anterior-most segment considered to represent segment II, so that there are 7 visible segments ventrally (segments II to VIII) anterior to penial sheath. Setae: dorsal abdominal setae all short, with a pair present submedially on segments IV–VII; ante-anal setae absent; ventral abdominal setae (avs) rather longer, each about 20 µm long, present on segments IV–VII; dorsopleural seta (dps) more or less in a line on each side between segment II and VII, with 15–20 fleshy setae on each side, each 12–22 µm long; ventropleural setae (vps) with 1 small seta on each side of segments III–VII. Lobes of segment VII rounded or absent, not even extending posteriorly to base of penial sheath. Lobes of segment VIII inconspicuous, each with 2 or 3 fleshy setae, longer two each 18–33 µm long plus one about 12 µm long. Anus present dorsally at anterior end of penial sheath, 20–26 µm wide. Penial sheath (ps) sclerotised, slightly longer than wide (about 155–175 µm long, 135–140 µm wide at base; ratio length to width 1:0.83); with 2 pairs of minute setae on dorsal surface; genital opening present medially on ventral surface.

Comment. Although the basic pattern of this pupa is similar to those of other Coccidae , it can be separated from all others known to the authors by the following combination of character states: (i) very short and rounded lobes on abdominal segment VII (usually extremely well developed on most species, extending posteriorly to at least half the length of the penial sheath), and (ii) apparent absence of pairs of small dorsal setae on abdominal segments V, VI and VII (usually present on one or more segments but sometimes present on all segments).

ADULT MALE ( Fig. 12 ). Described from 3 specimens in fair to good condition.

Unmounted material. In life, orange-brown body with darker thorax ( Fig. 1F,G ), wings whitish. Wings extending past end of long, thin penial sheath. Glandular pouch setae with long white wax extrusions (tail filaments), which also extend past apex of penial sheath. Simple eyes dark.

Mounted material: moderately large, total body length about 1.8–1.85 mm, width at mesothorax about 360– 380 µm; antennae slightly longer than half body length; body rather bald, with few body setae, mostly hair-like setae (hs) but these rather similar to fleshy setae (fs); appendages, however, hirsute, with numerous fs easily differentiated from hs. Wings large, about 9/10th total body length.

Head: rounded in dorsal view; length about 207–210 µm; width across genae 210–240 µm. Almost hairless. Median crest (mc) reticulated, extending posteriorly to about level with postocular ridge (pocr); with 1–3 hs dorsal head setae (dhs); postoccipital ridge absent. Mid-cranial ridge absent dorsally; ventrally (vmcr) narrow and poorlydefined, extending posteriorly as far as ocular sclerite; with a broad, lightly reticulated margin; with 0 or 1 hs ventral mid-cranial ridge setae (vmcrs) on each side. Postoccipital ridge absent. Preocular ridge (procr) distinct and extending posteriorly about 2/3rds of way to midcranial ridge. Genae (g) large and membranous, with large polygonal reticulations, each reticulation occasionally with an inner microridge; without genal setae. Eyes: 2 pairs of large, round, simple eyes, 1 pair dorsally (dse) and 1 pair ventrally (vse), subequal in size, each 33–42 µm wide. Ocelli quite large, touching postoccipital ridge (pocr), each about 25 µm wide. Ocular sclerite (ocs) strongly reticulated, most reticulations with a slightly curved inner microridge. Postocular ridge (pocr) extending medially almost to median crest dorsally. Dorsal ocular setae absent. Ventral head setae few, perhaps 4–6 on each side. Preoral ridge (pror) well developed. Cranial apophysis rather bluntly bifurcated; length about 33 µm.

Antennae: filiform, 10 segmented, each about 1.2 mm long; most segments with both fs and hs. Scape (scp): 48–60 µm long and 65 µm wide, with 3 hs. Pedicel (pdc): 60–67 µm long and 53–58 µm wide; reticulated or ridged throughout, with 4–8 fs, 4–6 hs + a campaniform pore. Segments III–VI each about 23–31 µm wide; length of fs 36–42 µm: lengths (µm): III 140–157; IV 163–193; V 202–212; VI 214–220; VII 170–177; VIII 124–128; IX 96– 99; X 64–72; approximate number of setae per segment: III 9–10 fs + 1 hs; IV 17–26 fs + 3 or 4 hs; V 25–29 fs + 3 or 4 hs; VI 29–31 fs + 4 or 5 hs; VII 20–32 fs + 2 hs; VIII 11–21 fs, 1–3 hs + a bristle (ab); IX 11–14 fs, 1 hs + 1 ab; X 6–10 fs, 1 or 2 hs, 3 capitate setae (caps), 4 or 5 ab + 0 or 1 basiconic sensilla (bacs) near apex; apex not constricted. Antennal bristles barely differentiated from fs.

Thorax. Prothorax: pronotal ridge (prnr) well developed and meeting medially on dorsum; with broad, striated, lateral pronotal sclerites (prn); without lateral pronotal setae. Post-tergite (pt) not detected; without post-tergital setae. Medial pronotal setae absent. Proepisternum + cervical sclerite (pepcv) well developed. Sternum (stn 1) with a strong, more or less straight transverse ridge; median ridge represented by a few small areas of sclerotisation; with a few lateral striations; prosternal setae, anteprosternal and antemesospiracular setae absent.

Mesothorax: possibly rather convex in life; prescutum (prsc) about as long as broad (144–160 µm long, 155– 195 µm wide); with a nodulated surface; prescutal ridges (pscr) and prescutal sutures (pscs) well developed. Scutum (sct): median membranous area (sma) about four times as wide as long, perhaps 33–55 µm long, 175–200 µm wide; scutal setae (scts) absent; lateral margins of scutum with polygonal nodulations laterad to prescutum but not laterad to scutellum. Scutellum (scl) 62–70 µm long, 178–200 µm wide; probably with a large foramen; without scutellar setae. Mesepisternum (eps 2) with polygonal nodulations. Basisternum (stn 2) large, about 195–215 µm long, 290–320 µm wide; with a rather weak median ridge (mdr), generally not complete; without setae; bounded by strong marginal (mr) and precoxal ridges (pcr 2); lateropleurite (lpl) rather large, with an extension from marginal ridge anteriorly; furca (f) well developed and extending anterior to point where marginal ridge (mr) and precoxal ridges (pcr 2) meet laterally. Mesothoracic spiracle (sp 2): width of peritreme 34–36 µm; postmesospiracular setae absent. Postalare (pa) reticulated at anterior end; without postalare setae. Tegula (teg) well developed with generally 3–5 tegular setae (tegs) (one specimen with 10 on one side). Subepisternal ridge (ser) well developed. Antemetaspiracular setae absent. Mesopostnotum (pn 2) with large, shallow polygonal nodulations.

Metathorax: metapostnotum (pn 3) sclerotised; with 1 pair of hs metatergal setae. Metapleural ridge (plr 3) well developed; suspensorial sclerite (ss) probably present; metepisternum (eps 3) sclerotised, with 4–6 fs + 0–2 hs postmetaspiracular setae (pmss) on each side; metepimeron (epm 3) well developed but only mildly sclerotised; without setae. Metathoracic spiracle (sp 3): width of peritreme 37–40 µm. Dorsospiracular setae absent. Metasternum (stn 3) slightly sclerotised but not reticulated; with no anterior metasternal setae but with 0–2 posterior metasternal setae.

Wings: hyaline; of moderate length and width, each 1.60–1.65 mm long and 0.72–0.78 mm wide (ratio of length to width 1:0.46); with a single alar seta near base of each wing. Hamulohalteres present, length 120 µm, width 23–25 µm; with a single hooked hamulus, each 60–65 µm long.

Legs: long and hairy; subequal in length. Coxae (cx): length (µm): I 124–135, II 132–140, III 135–150; with 13–15 fs + 6–8 hs; longest seta 65–70 µm; without coxal bristles. Trochanter (tr) + femur (fm) (µm): I 320–330; II 290–310, III 290–306; each trochanter (tr) with 2 small campaniform sensilla on each side, and 8–14 fs + 1–4 hs; long seta barely differentiated, each 44–60 µm long; each femur with 29–35 fs + 22–25 hs. Tibia (ti) length (µm) I 350–360; II 345–350; III 340–360; each with many setae, mainly fs and hs but with increasing numbers of spurlike setae on distal third; apical spur (tibs) stout and strong, each 24–27 µm long. Tarsi (ta) one segmented, lengths (µm): I 153–155; II 145–158; III 145–162; each with numerous setae, fs, hs and spur-like setae; tarsal campaniform pore absent; distal tarsal spur (tars) barely differentiated, each 19–22 µm long; tarsal digitules (tdt) shorter than claw. Claws (c) moderately long, slight longer than width of tarsi; with a distinct denticle (cd); length 34–39 µm; claw digitules (cdt) slightly longer than claw.

Abdomen. Segments I–VII: tergites (at) barely sclerotised on segments (V)–VII; sternites (as) slightly more distinct, on segments V–VII; pleurites absent. Caudal extension of segment VII absent. Dorsal abdominal setae (ads) almost absent, only represented by 1 pair of hs on segments III–VII. Pleural setae: dorsopleural setae (dpls): with single setae on each side of segments V–VII; ventropleural setae (vpls) present more or less in a line along margins of segments III–VI; segment VII with about 10–12 setae; IV–VI 3 or 4 and segment III 0 or 1. Ventral abdominal setae very few, with 1 pair on segment VII, 1 or 2 pairs on VI and V, 1 pair on IV and III. Segment VIII: tergite and sternites sclerotised; tergite with 0–2 ante-anal setae (aas); sternite without setae; caudal extension absent, each with 3 hs pleural setae. Glandular pouches (gp) present, each divided into an inner and outer chamber, each with different types of pore; each pouch with two capitate setae (gls), each 125–150 µm long.

Genital segment (IX + X). Penial sheath (ps) sclerotised, long, narrowing to a sharp apex; basal area membranous on both dorsal and ventral surfaces; length 465–475 µm, width at base 110–118 µm, about 1/4th total body length (ratio of total body length to penial sheath length 1:0.25). Anus oval, 28–30 µm long and 21–27 µm wide, on dorsal surface of penial sheath, surrounded by sclerotisation of penial sheath (probably segment IX). Basal rod (bra) short, about 78–85 µm long, anterior to aedeagus, with an extension of about 12–25 µm within aedeagus; basal rod not nearly reaching basal membranous area (bma) anteriorly. Aedeagus (aed) more than half length of penial sheath, 275–290 µm long, of approximately uniform width. Apex of penial sheath without a membranous extension; with about 13 minute setae (gts) along each margin of penial sheath and with a cluster of small sensillae (gtp) near apex.

Comment. The structure of the penial sheath and associated structures on Cissococcus are very similar to those on other Coccidae , particularly (i) the structure of the glandular pouches with two chambers, each with different pores (see fig. 70 of Crystallotesta ornatella in Hodgson & Henderson (2004)), and (ii) the absence of tarsal campaniform pores. However, the position of the anus within the sclerotised part of the penial sheath is unlike any known coccid male, and the presence of alar setae is known only in Eulecanium tiliae (L.) and Nemolecanium abietis Borchsenius ( Giliomee, 1967) . It seems likely that Cissococcus evolved from early ancestors in the evolution of the Coccidae .

The "reticulations" present on the thorax have a different structure to those on the head and are here being referred to as polygonated nodulations. When the reticulations on the head are viewed under high-power, it is clear that they are shallow ridges, whereas those on the thorax (particularly on the prescutum, scutum and mesepisternum) are composed of shallow grooves between polygonally-shaped convex areas. This was noted previously by Hodgson & Foldi (2005) on male margarodoids and probably applies to most (all?) male Coccoidea with "reticulations" on sclerotised structures posterior to the head. Those on more membranous areas, such as the abdomen and mesopostnotum, may be similar to those on the head but this has still to be checked.

Figures

FIGURE 1. Cissococcus fulleri Cockerell on Rhoicissus tridentata, East London: A, galls of adult females on main twig and immature females on twig leading to leaf; B, two galls of adult females with the one on right cut open to reveal the anal area and stigmatic wax of a female, and one on left showing gall apex (in middle of gall); C, gall cut open to show middle venter of adult female; D, first- and second-instar male nymphs on the underside of a leaf; E, close-up of two second-instar male nymphs (note glassy-wax tubes emanating from cone-shaped pores on dorsum); F, an adult male recently emerged from one of the cocoon-like tests underneath him; G, adult male a few hours after eclosion. Galls of young adult females of Cissococcus braini Hodgson & Millar sp. n. on Rhoicissus digitata: H, immature gall viewed from side with large orifice visible, at Boesmanriviermond; I, same gall as in H viewed from top, with atrium below orifice visible; J, mature gall at Gxulu River (Photos by PJG).

FIGURE 4 Cissococcus fulleri Cockerell, adult female. The greatly expanded venter comprises most of the main figure. Unless otherwise stated, on this and the following figures: A = section of dorsum showing fissures; B = dorsal seta; C = dorsal pore; D = dorsal cone-shaped secretory spine; E = dorsal tubular duct; F = dorsal view of anal plate; G = ventral view of anal area; H = dermal nodulations; J = marginal spinose seta; K = stigmatic spine; L = ventral multilocular pore; M = ventral seta from medially on surface of lower venter; N = ventral seta from more lateral areas of upper surface of venter; O = preantennal pore; P = ventral microduct; Q = antenna; R = ventral tubular duct; S = spiracle; T = tarsus plus claw; V = spiracular disc-pores; X = dorsal view of venter of old specimen showing enlarged apodemes.

FIGURE 5. Cissococcus fulleri Cockerell, third-instar female. Lettering as for Figure 4.

FIGURE 6. Cissococcus fulleri Cockerell, young second-instar female. Lettering as for Figure 4 except that X = shape and limb distribution on older specimen.

FIGURE 7. Cissococcus fulleri Cockerell, second-instar male with sunken cone-shaped secretory spines. Lettering as for Figure 4.

FIGURE 8. Cissococcus fulleri Cockerell, second-instar male from Boesmansriviermond, without sunken cone-shaped secretory spines. Lettering as for Figure 4.

FIGURE 9. Cissococcus fulleri Cockerell, first-instar nymph (sex unknown). Lettering as for Figure 4 except T = tibia, tarsus + claw.

FIGURE 10. Cissococcus fulleri Cockerell, prepupa. Where ads = dorsal abdominal setae; an = anus; avs = ventral abdominal setae; dhs = dorsal head setae; dpls = dorsal pleural setae; ps = penial sheath; spds = spiracular disc-pores; vhs = ventral head setae; vpls = ventral pleural setae, and wb = wingbud.

FIGURE 11. Cissococcus fulleri Cockerell, pupa. Lettering as in Figure 10.

FIGURE 12. Cissococcus fulleri Cockerell, adult male. A = reticulations on gena; B = reticulations on ocular sclerite; C = cranial apophysis; D = apical antennal segment; E = distal end of metathoracic leg; G = dorsal view of penial sheath, H = ventral view of penial sheath. Also: aas = ante-anal setae; ab = antennal bristle; ads = dorsal abdominal setae; aed = aedeagus; als = alar setae; an = anus; as = abdominal sternite VIII; at = abdominal tergite VIII; avs = ventral abdominal setae; bma = basal membranous area; bra = basal rod of aedeagus; c = claw; caps = capitate setae; cd = claw denticle; cdt = claw digitule; cx = coxa; dhs = dorsal head setae; dpls = dorsal pleural setae; dse = dorsal simple eye; epm3 = metepimeron; eps2 = mesepisternum; f = furca; fem = femora; fs = fleshy seta; g = gena; gls = glandular pouch setae; gp = glandular pouch; gpp = glandular pouch pores; gts = penial sheath setae; gtp = penial sheath pores; h = hamulus; ham = hamulohaltere; lpl = lateropleurite; mc = median crest; mdr = median ridge; mr = marginal ridge; mts = metatergal setae; o = ocellus; ocs = ocular sclerite; pa = postalare; pcr2 = precoxal ridge of mesothorax; pdc = pedicel; pepcv = proepisternum + cervical sclerite; plr3 = metapleural ridge; pmss = posterior metasternal setae; pn2 = mesopostnotum; pn3 = metapostnotum; pocr = postocular ridge; prn = lateral pronotal sclerite; prnr = pronotal ridge; procr = preocular ridge; prsc = prescutum; ps = penial sheath; pscr = prescutal ridge; pscs = prescutal sclerite; s = spur-like seta; scl = scutellum; scp = scape; sct = scutum; ser = subepisternal ridge; sma = membranous area of scutum; sp2 = anterior spiracle; sp3 = posterior spiracle; stn1 = prosternum; stn2 = basisternum (mesosternum); ta = tarsus; tdt = tarsal digitules; teg = tegular; tegs = tegular setae; ti = tibia; tr = trochanter; vhs = ventral head setae; vmcr = ventral midcranial ridge; vmcrs = ventral midcranial ridge setae, and vpls = ventral pleural setae.