Eriococcus melnikensis Hodgson & Trencheva, 2008

Hodgson, Chris & Trencheva, Katia, 2008, A new species of Eriococcus (Hemiptera: Sternorrhyncha: Coccoidea: Eriococcidae) and redescriptions of Eriococcus roboris Goux and E. thymelaeae Newstead, with a key to the Eriococcus species on Quercus in the western Palaearctic, Zootaxa 1959 (1), pp. 1-38 : 12-27

Eriococcus melnikensis Hodgson & Trencheva , spec. nov.

Material studied. Holotype f: BULGARIA, Pirin Mountains , Melnik, 10.iv.2008, on Quercus pubescens, K. Trencheva (BMNH) : 1 of three on slide, holotype specimen on left, nearest the data label.

Paratypes ff: remaining 2 specimens on holotype slide + data as above ( BMNH): 3/6 adff, 3 2 nd -instar males + 1 2 nd instar female ( USNM); as previous, but dated 23.vii.2006 ( BMNH, USNM, PPI): 12/3 young adf, 3 2 nd -instar f, 8 2 nd -instar m, 4 pupae + 12 adm (all in fair to good condition). Also, as previous but collected on 7.iv.2007 ( BMNH, USNM, PPI): 10/7 young adff, 6 2nd-instar ff, 5 2nd-instar mm, 3 1st-instar nymphs (all fair to good). Also nr. Kardzhali, Rodopi Mountains , on Quercus pubescens , 5.V.2008, K. Trencheva ( BMNH): 1/2 mature adff in good condition .

Non-paratype material: CYPRUS, Mylos Forest Stn., on Myrtus communis (Myrtaceae) 12.xii.1931, E.E. Green ( BMNH), (with manuscript name of Eriococcus myrtus Green ): 4/10 adf, in good to poor condition.

Other material: Hungary, Vászoly , on Quercus sp. , 14.vii.1969, Kozár ( PPI), 1/1ad in fair condition but not all characters visible – however, it lacks setae on antennal segment III and the limb measurements are all small and so it is probably referable to this species .

Adult female ( Fig. 5 ). Described from 5 specimens from Melnik, mainly in good condition, and 2 from Rodopi Mtns (data in brackets refers to Cypriot material where it differs).

Instar diagnosis. Young adults (from Melnik) small, mounted material in alcohol pale pink to reddish, long and narrow, pointed at both ends; about 1.25–1.8 mm long; venter slightly wider than dorsum: dorsum 475–700 µm wide, total width of mounted specimens about 575–800 (600–850) µm; older specimens (from Rodopi Mountains): 2.75–3.0 mm long; dorsum 1.35–1.70 mm wide, greatest width of mounted specimens 1.75–1.93 mm. Anal lobes sclerotised, comparatively large; median plate present, triangular to rather quadrate. Dorsum covered in truncate spinose setae; those along margin slightly larger. Dorsum also with numerous macrotubular ducts (of one size) and microtubular ducts (each with a divided dermal orifice); venter with macrotubular ducts similar to those on dorsum, restricted to near margin; also with small macrotubular ducts present ventrally on abdomen and metathorax. Quinquelocular pores abundant on abdomen, less frequent more anteriorly; cruciform pores present submarginally, mainly in groups on anterior abdominal segments and thorax. Legs comparatively well developed; metacoxae without pores but with spicules on anterior surfaces; claws with a strong denticle. Antennae 7 segmented; with frontal lobes.

Dorsum. Derm membranous. Dorsal setae all spinose, of approximately one size, each with a truncated apex, narrowing slightly, with a narrow basal socket; each mainly 20–30 (18–25) µm, range 13–50 (18–30) µm, longest medially on head but some specimens with median pair of spinose setae on each segment somewhat larger, up to 38–50 (35) µm long; present throughout although in distinct segmental bands on abdomen, as follows: abdominal segment VIII 1–3, VII 17–21 (13–14), VI 25–27 (15–22), V 31–37 (24–34). Dorsal pores absent. Dorsal ducts of two types: (i) macrotubular ducts, each quite large, cup-shaped invagination well developed and symmetrical, width subequal to width of base of dorsal seta; outer ductule broad, each 7–8 µm wide and about 19–28 (18–20) µm long; inner ductule narrow, widening towards distal end, length about 20–26 µm; present throughout, as abundant as or rather less abundant than dorsal spinose setae; and (ii) microtubular ducts, each small, dark and narrow, outer ductule about 8–10 (7) µm long, with an inner dark line; inner ductule thin, short, thread-like; dermal orifice oval, bifid when seen from side and triangular when seen end on; present throughout (and on dorsal surface of each anal lobe) and about as abundant as dorsal spinose setae. Anal lobes sclerotised throughout, finger like, each about 83–116 µm long, 40–54 (33–50) µm wide; inner margins serrated or nodulated; dorsal surface with 3 truncate setae (of rather variable length) and a long apical seta: anterior inner margin seta about 25–38 µm long; posterior inner margin seta about 33–55 (31–37) µm long; outer margin seta about 30–58 µm long; apical setose seta about 227–270 (190–210) µm long. Median plate triangular, with a serrate edge, about 25–30 µm long and 46–50 µm wide. Anal ring with 4 pairs of long setae, each about 85–155 µm long; with a complete outer ring of conical pores.

Margin. Not demarcated but probably represented by longer truncate spines, similar to but larger than those dorsally, each 33–50 (30–42) µm long; most abdominal segments with 4 (occasionally 5, rarely 3) on each side.

Venter. Membranous, apart from anal lobes which are sclerotised, with a short, sclerotised bar extending anteriolaterally from base of basal setae; bar about 28–34 µm long. Ventral setae: with a group of about 4–6 truncate spinose setae plus 4 sharply-pointed spinose setae near anterior margin of head; remaining setae mainly setose, most abundant anteriorly between antennae and clypeolabral shield, where 8–10 pairs of flagellate setae present; and medially on posterior 5 or 6 abdominal segments (no. setae across each segment approximately: II 7 , III 10 , IV 13 , V 9–13 , VI 10–13 , VII 12 or 13, and VIII 5 or 6); longest up to about 85 (55–60) µm long; also with sparse groups mesad to each coxa; submarginal setae setose along submargin of abdomen (where mainly 2 on each side of each segment, each 17–22 (16–20) µm long), becoming spinose on submargins of thorax and head, stoutest on head, longest on anterior thorax, where up to 33–42 (27–30+) µm long). Ventral surface of each anal lobe with a single long setose seta near apex, about 70–115 µm long, and a basal seta on outer margin about 30–40 µm long; suranal setae, on either side of anal ring, each about 85–112 µm long. Ventral pores of three types: (i) quinquelocular pores, each about 5 µm wide; abundant in a segmental band on posterior abdominal segments, becoming less abundant anteriorly and only sparse medially on thorax and head; few submarginally on thorax but with distinct concentrations near each spiracle; (ii) cruciform pores, each oval, about 3.5–4 µm long, and 2 µm wide, appearing to have an oval pore; in a sparse interrupted band along submargin of head and anterior thorax, but in larger submarginal groups (15–20+ pores) on meso- and metathorax, and in small groups (1–15 pores) on abdominal segments I– V, and (iii) 1 or 2 small preantennal pores present anterior to each scape. Ducts of four types: (i) large macrotubular ducts, similar to those on dorsum but restricted to a submarginal band 1–2 pores wide, tending to become (ii) a slightly smaller medially in a narrow band just ventral to type (i); also in a group anteriorly between antennae and with a pair on abdominal segment VIII; (iii) small macrotubular ducts, each narrow, with an asymmetric cup-shaped invagination; outer ductule 9–12 (10–13) µm long, inner ductule about 17 (?) µm long: distributed as follows: abdominal segment VIII 0, VII generally 0 (one specimen with 2), segments VI – IV 7–10 (8–10), III & II 4–8 (6–8) plus 2–4 (1–2) medially on metathorax; and (iv) microtubular ducts similar to those on dorsum but restricted to immediately ventrad of marginal spines .

Eyespots distinctly displaced ventrally from margin, each about 20–27 µm wide. Antennae each 7 segmented, about 252–315 µm long; scape with 4 setae, pedicel with 3 setae + a campaniform sensillum; III without setae; IV 1 long seta + a shorter setae; V 1 fleshy seta; VI with 1 fleshy seta + 3 setose setae; and VII with 3 stout fleshy setae + 6 or 7 other setae, some stiff; apical seta each about 40–50 (50–60) µm long; each antenna with a frontal lobe antero-medially to scape, each about 30–35 µm wide and 30–55 µm long when expanded (not obvious on either Rodopi or Cypriot material). Clypeolabral shield about 150–175 (175–186) µm long; labium about 145–165 (165–170) µm long, 3 segmented, basal segment with 2 pairs of setae (one specimen off Myrtus sp. with 3 on one side), middle segment 1 pair and apical segment with 5 or 6 (5) pairs; stylet loop reaching posteriorly to metacoxae. Spiracles quite small, width of peritremes about 26–33 µm. Legs: data for metathoracic legs: lengths (µm) coxa 100–135; trochanter + femur 153–195; tibia 100–125; tarsus 125–150; with few spinulae on anterior surfaces of procoxae but frequent on meso– and metacoxae; each coxa with 5 or 6 setae; each trochanter with 2 long setae, longest about 65–75 µm, + 1 short seta; each femur with 3–6 setae; tibia with 4 setae and each tarsus with 4 or 5 setae; tarsal campaniform sensillum present; tarsal digitules with relatively narrow apices and subequal to length of claw; claw about 30–35 µm long, with a distinct denticle; claw digitules capitate and longer than claw. Vulva opening between segments VII and VIII, with a complex area of sclerotisation associated with opening, possibly with inner lining of oviduct sclerotised .

Comment. Adult female E. melnikensis sp. nov. differ from those of E. roboris in having (character-states on E. roboris in brackets): (i) all measurements rather smaller; (ii) few or no small macrotubular ducts ventrally on abdominal segment VII (8–12); (iii) fewer than 10 small macrotubular ducts present across abdominal segment VI–II (mainly more than 15 and sometimes over 30 across some segments); (iv) segment III of the antennae without setae (setae apparently always present on segment III on E. roboris ).

The material from Cyprus off Myrtus communis is very similar to the type series and clearly belongs to this species, despite being off a different host and away from the type site .

Second-instar female ( Fig. 6 ). Described from 5 specimens in fair to good condition.

Instar diagnosis. Small, oval but more pointed posteriorly; about 0.87–1.05 mm long; venter clearly wider than dorsum: dorsum 385–525 µm wide, total width of mounted specimens about 500–675 µm. Anal lobes sclerotised, comparatively large; quadrate median plate present. Dorsum with truncate spinose setae; those along margin slightly larger, those submedially on abdominal segments V–VII (sometimes III and IV also) smaller. Dorsum also with numerous microtubular ducts (each dermal orifice with two large wing-like extensions); macrotubular ducts absent from both surfaces. Quinquelocular pores absent from dorsum but frequent on venter; cruciform pores present submarginally in groups on ventral anterior abdominal segments and thorax. Legs comparatively well developed; claws with a strong denticle. Antennae 6 segmented, with welldeveloped frontal lobes.

Dorsum. Derm membranous. Dorsal setae all spinose, each with a truncate apex, narrowing slightly from a broad base; each with a narrow basal socket; of two sizes: most quite large, mainly 30–40 µm long, but with short setae of similar structure but only 10–18 µm long submedially on abdominal segments V–VII (occasionally II–IV); those medially on segments II–V also slightly smaller; setae in 4 longitudinal lines on abdominal segments VI & VII but six lines on more anterior abdominal segments and more randomly distributed on thorax and head. Dorsal pores absent. Dorsal microtubular ducts each small, with outer ductule about 7–8 µm long, dark and narrow, with an inner dark line; inner ductule thin, short, thread-like; dermal orifice oval, with two wing-like extensions; usually with 1–3 in general area of each dorsal seta; present throughout dorsum plus 2 or 3 on dorsal surface of each anal lobe. Anal lobes sclerotised throughout, finger-like, each about 60–66 µm long, 25–33 µm wide; inner margins slightly serrate; dorsal surface with 3 truncate setae and a long apical seta: anterior inner margin seta about 16–22 µm long; posterior inner margin seta about 33–38 µm long; outer margin seta about 30–35 µm long; apical setose seta about 180–205 µm long. Median plate quadrate, with four lightly sclerotised projections along posterior margin; about 30 µm wide and 18 µm long. Anal ring with 3 pairs of long setae, each about 80–100 µm long.

Margin. Marked by longer truncate spinose setae, similar to but slightly larger than those dorsally, each up to about 40 µm long; each abdominal segment with 2 on each side.

Venter. Membranous. Ventral setae mainly setose but with a group of 6–8 truncate spinose setae anteriorly on head, similar to those on dorsum, but smaller, each about 28 µm long; with 5 pairs of long flagellate setae between antennae and clypeolabral shield (longest about 33–45 µm long); also medially on each thoracic segment and with 1–3 on abdominal segments II – VIII (no. setae across each segment: I 2, II – V 4 , VI 6 , and VII 2 ); longest on segment VI 35–40 µm long; also with submedial and submarginal short setae on most abdominal and thoracic segments, latter becoming significantly more spinose anteriorly, where up to 25 µm long. Ventral surface of each anal lobe with 1 setose seta, 45–66 µm long; suranal setae on either side of anal ring, each about 45–60 µm long. Preantennal pores: single pores present anterior to each scape, each about 3 µm tall. Other ventral pores of two types: (i) quinquelocular pores, each about 4 µm wide, sparse throughout venter, and (ii) cruciform pores, each oval, about 3 µm long, in submarginal groups each associated with a submedial seta, with (on each side): meso- and metathorax 2–4, abdominal segments I– III 1–4 , and IV & V 0–1; absent along head submargin. Macrotubular and microtubular ducts absent .

Eyespots distinctly displaced ventrally from margin, each about 16–18 µm wide. Antennae each 6 segmented, about 165–170 µm long; scape with 4 setae, pedicel with 3 setae + a campaniform sensillum; segment III 2 setae; IV 1 fleshy seta; V 1 fleshy seta + 2 setose setae; and VI with 3 stout fleshy setae + 6 or 7 other setae, mainly flagellate; each antenna with a small frontal lobe at base of scape. Clypeolabral shield about 85–115 µm long; labium about 100–115 µm long, 3 segmented, basal segment with 2 pairs of setae, middle segment with 1 pair and apical segment with 4 pairs of setae; stylet loop extending posteriorly as far as metacoxae. Spiracles quite small, width of each peritreme about 15 µm. Legs: lengths metathoracic legs (µm): coxa 63–73; trochanter + femur 100–112; tibia 60–65; tarsus 85–98; with a few spinulae on anterior surfaces of all coxae; each coxa with 6 setae; trochanter 3 setae, longest about 63–66 µm; femur 3 setae; tibia with 3 setae, and each tarsus with 6–8 setae; tarsal campaniform sensillum present; tarsal digitules slightly longer than claw, with small capitate apices; claw about 25 µm long, with a distinct denticle; claw digitules longer than claw, with small capitate apices.

Comment. For differences with 2nd-instar female E. roboris , see under that species above.

Second-instar male ( Fig. 7 ). Described from 5 specimens in fair to good condition.

Instar diagnosis. Small, oval but more pointed posteriorly; about 0.67–0.98 mm long; venter clearly wider than dorsum: dorsum 305–350 µm wide, total width of mounted specimens about 355–415 µm. Anal lobes sclerotised, comparatively large; rectangular median plate present. Dorsum with truncate spinose setae; those along margin slightly larger, those submedially on abdominal segments V–VII somewhat smaller (sometimes also on segments II–IV). Dorsum also with numerous microtubular ducts, each dermal orifice with two large wing-like extensions; macrotubular ducts present throughout dorsum and marginally on venter; quinquelocular pores absent from dorsum but frequent on venter. Cruciform pores present submarginally in groups on anterior abdominal segments and thorax. Legs comparatively well developed; coxae without pores; claws with a strong denticle. Antennae 7 segmented, with well-developed frontal lobes.

Dorsum. Derm membranous. Dorsal setae all spinose, each with a truncate apex, narrowing slightly from a broad base; with a narrow basal socket; of two sizes, most quite large, mainly 25–35 µm long, in four (abdominal segments VI & VII) or six (segments V–III) longitudinal lines but more randomly distributed on thorax and head; with short setae of similar structure but only 10–18 µm long, present submedially on abdominal segments V–VII (and sometimes II–IV); also those medially on segments II–VI slightly smaller. Dorsal pores absent. Dorsal microtubular ducts each small, with outer ductule about 7–8 µm long, dark and narrow, with an inner dark line; inner ductule thin, short, thread-like; dermal orifice oval, with two wing-like extensions; present throughout and on dorsal surface of each anal lobe; usually with 1–3 associated with each dorsal seta and often macrotubular pore. Dorsal macrotubular pores abundant, each with a membranous outer ductule about 12–14 µm long and 6–7 µm wide, with a well-developed symmetric cup-shaped invagination; inner ductule narrow; distributed fairly evenly over dorsum. Anal lobes sclerotised throughout, finger like, each about 55–66 µm long, 25–30 µm wide; inner margins slightly serrate; dorsal surface with 3 truncate setae and a long apical seta: anterior inner margin seta about 10–16 µm long; posterior inner margin seta about 25–28 µm long; outer margin seta about 20–33 µm long; apical setose seta about 180–210 µm long. Median plate quadrate, about 26–32 µm wide and 16–18 µm long; with 4 or 5 projections along posterior margin. Anal ring with 3 pairs of long setae, each about 65–85 µm long.

Margin. Marked by longer truncate spinose setae, similar to but slightly larger than those dorsally, each up to about 35 µm long; abdominal segments I–VII with 2 on each side.

Venter. Membranous. Ventral setae mainly setose but with a group of 8 anteriorly on head similar to those on dorsum, but smaller, each about 25–28 µm long; with 5 pairs of long flagellate setae between antennae and clypeolabral shield (longest about 40–50 µm long); medially on thoracic segments and with 2 or 3 on each side of abdominal segments I– VII (no. setae across each segment: I 2, II 4 , III – VI 6 , and VII 2 ); longest on segment VII 30–35 µm long; also with submedial and submarginal short setae on most abdominal and thoracic segments, latter becoming more spinose anteriorly where up to 25 µm long. Ventral surface of each anal lobe with 1 long setose seta, 50–65 µm long; suranal setae on either side of anal ring, each about 50–65 µm long. Preantennal pores: single pores present anterior to each scape, each about 3 µm tall. Other ventral pores of two types: (i) quinquelocular pores, each about 4 µm wide, frequent throughout venter, and (ii) cruciform pores, each oval, about 3.0–3.5 µm long; in submarginal groups associated with a submedial seta and a macrotubular pore, with (on each side): mesothorax 1–8, metathorax of 1–4, abdominal segments I– III 1–4 , and IV 0–1; absent along head submargin. Microtubular ducts absent. Macrotubular ducts of similar structure to those on dorsum, but smaller, each about 6–7 µm wide, present in a narrow submarginal band .

Eyespots distinctly displaced ventrally from margin, each about 15–16 µm wide. Antennae each 7 segmented, each about 160–185 µm long; scape with 4 setae, pedicel with 3 setae + a campaniform sensillum; segment III 0 setae, IV 2 setae; V 1 fleshy seta; VI 1 fleshy seta + 2 setose setae; and VII with 3 stout fleshy setae + 6 or 7 other setae, mainly stiff setae; small frontal lobes present just anterior to each scape. Clypeolabral shield about 105–110 µm long; labium about 100–105 µm long, 3 segmented, basal segment with (on each side) 2 setae, middle segment with 1 seta and apical segment with 4 setae; stylet loop extending posteriorly to metacoxae. Spiracles quite small, width of peritremes each about 15 µm. Legs well developed; lengths metathoracic legs (µm): coxa 69–88; trochanter + femur 110–135; tibia 78–83; tarsus 90–105; with a few spinulae on anterior surfaces of all coxae; each coxa with 5 or 6 setae; trochanter 3 setae, longest about 40–45 µm; femur 5 setae; tibia with 4 setae and each tarsus with 4 or 5 setae; tarsal campaniform sensillum present; tarsal digitules with small capitate apices, slightly longer than claw; claw about 24–25 µm long, with a distinct denticle; claw digitules with small capitate apices, longer than claw.

Comment. The 2nd-instar male is immediately separable from the 2nd-instar female by the presence of macrotubular ducts. It differs from the 2nd-instar male of E. roboris in being rather smaller, in having fewer cruciform pores, in the shape of the median plate, and in generally having small submedial truncate dorsal setae only on abdominal segments VI and VII (rather than II–VII).

First-instar nymph ( Fig. 8 ). Described from 3 specimens in fair to good condition.

Instar diagnosis. Oval in outline, rather more pointed posteriorly, 720–740 µm long, dorsum 280–290 µm wide (venter wider); anal lobes short but with 3 pairs of truncate spinose setae dorsally in addition to long flagellate apical seta; median plate indicate by a small membranous outgrowth with about 3 protuberances. Dorsal setae truncate, of two sizes. Antennae 6 segmented; frontal lobes absent; dorsal microtubular ducts present, each outer orifice with two large wing-like extensions; quinquelocular pores present ventrally on head, thorax and abdomen; cruciform pores present submarginally on venter of thorax; marginal and dorsal setae all spinose and truncate, each with a slightly swollen base; dorsal spinose setae in 4 longitudinal lines + marginal setae; claw with a well-developed denticle.

Dorsum. Derm membranous. Dorsal setae all spinose, with pairs medially and submedially on most segments, of two sizes: most setae large (each mostly 16–20 µm long) but with short setae (each mainly 6.5–13 µm long) medially on abdominal segments II–V and submedially on abdominal segments II–VII; medial pairs on abdominal segments VI and VII large. Marginal setae similar to large spinose setae on dorsum, but perhaps a little larger, each mainly 23–27 µm long, with about 21 on each side. Dorsal microtubular ducts, each with a long dark outer ductule about 6.5–7.0 µm long and a large outer orifice with two wing-like extensions, sparsely distributed, submedially on abdomen (1 per segment) and submarginally and medially on thorax and head. Other pores absent. Anal lobes short, each about 25 µm long, unsclerotised, with (on dorsal surface) two spinose setae on inner margin (anterior about 13–14 µm long, posterior about 28 µm long), a long flagellate apical seta about 175–180 µm long, and a spinose seta on outer margin about 25 µm long. Anal ring with 6 setae, each about 65 µm long.

Venter. Derm membranous. Setae: spinose setae, similar to those on dorsum but slightly smaller, in a group of 4 on anterior margin of head; long flagellate setae distributed as follows: with 3 pairs between antennae, longest about 35 µm; 1 associated with each meso- and metacoxa; and medially on abdominal segments II and III, and medially and submedially on abdominal segments IV–VII; short submedial and submarginal setae present on most thoracic and abdominal segments; suranal setae about 33–40 µm long; ventral seta on each anal lobe about 60–65 µm long. Preantennal pores: single pores present anterior to each scape, each about 4 µm tall; frontal lobes absent. Quinquelocular pores few, each about 5 µm wide, present on head, thorax and medially/submedially on abdomen. Cruciform pores, each about 4–4.5 µm wide, present submarginally on thorax, with 3 between spiracles. Eyespots each about 13–15 µm wide.

Antennae 6 segmented, each 105–125 µm; scape with 4 setae, pedicel with 3 setae + campaniform sensillum, segment III with 2 flagellate setae, IV with 1 fleshy seta, V with 1 fleshy seta + 2 flagellate setae, and VI with 3 fleshy setae, and about 6 other setae, mainly flagellate. Clypeolabral shield about 90–95 µm long; labium about 80–90 µm long, with 7 pairs of setae, 2 on basal segment and 4 on apical segment; length of stylet loop unknown. Spiracles: width peritremes each about 8 µm. Metathoracic legs: lengths (µm): coxa 48–50; trochanter + femur 78–85; tibia 45–48; tarsus 63–70; claw 22–24; without spinulae on anterior surfaces of any coxa; number of setae: coxa 7; trochanter 3 (longest about 50 µm); femur 5; tibia 4 (tibial spurs not differentiated); tarsus 6. Tarsal digitules both capitate and subequal to length of claw; claw digitules slightly longer than claw and both slightly capitate; claw with a distinct denticle.

Comment. The first-instar nymphs of E. melnikensis and E. roboris are extremely similar, perhaps only differing in the size of the median dorsal spinose setae, E. melnikensis having the posterior two setae quite large, whereas only one is large on E. roboris .

PUPA ( Fig. 9 ). Described from 4 specimens in fair to good condition, one with pharate adult .

Instar diagnosis: quite small, total body length about 1.05–1.27 mm, width across head 250–290 µm. Body not clearly divided into tagmata; antennae 10 segmented, extending posteriorly to about anterior spiracles. Legs well developed, with tibio-tarsal articulation usually distinct. Body with small setose setae, each mainly 10–14 µm long, longest setae on posterior abdomen (about 23 µm long); multilocular pores present ventrally on prothorax and abdomen. Other pores absent. Anal lobes (caudal extensions) present on segment VIII, each with a long apical seta and small setae. Anterior wingbuds moderately long, extending posteriorly to metacoxa; posterior (hamulohaltere) wing buds small, on margin just posterior to posterior spiracles. Penial sheath triangular, with two pairs of long setae dorsally.

Dorsum. Derm membranous, apart from a sclerotised plate on posterior margin of head. Derm with frequent short setose setae, subequal in length to those on venter, in a group on head, singly on prothorax, but in transverse bands across other thoracic and abdominal segments, with 2–4 pairs per segment. All pores absent. Anal lobe distinct, each with a long pleural seta, 90–100 µm long, plus 1 short setose seta. Position of glandular pouch visible. Anus present at anterior end of penial sheath, 25–33 µm wide. Penial sheath about as long as wide, 90–95 µm long and 90–105 wide, with two pairs of long setae on dorsal surface, each 25–35 µm long.

Margin. Anterior wingbuds quite long and narrow, each about 360–380 µm long and 85–135 µm wide; posterior (hamulohaltere) wingbuds small but distinct, represented by small lobes, each about 20 µm long, just posterior to posterior spiracles. Margin of abdomen with segmentally arranged groups of longer setae, with mainly 3 dorsal pleural setae on all abdominal segments.

Venter. Derm membranous. Ventral setae few, with 1 anteriorly on head and medially on meso- and metathorax and abdominal segments II–VII; also submedially on segments IV–VI and submarginally on all abdominal segments. Multilocular pores each with mainly 10 loculi, with a pair laterad to each anterior spiracle, and then across meso- and metathorax and abdominal segments II–VII, and with groups marginally on segments VI–VIII. Other pores absent. Frontal lobes absent. Eyespots absent.

Antennae 10 segmented, each about 310–380 µm long; segments without any short setae. Mouthparts entirely absent, although position of mouth visible. Spiracles: width of all peritremes about 23–26 µm. Metathoracic legs: lengths (µm): coxa 70–87; trochanter + femur 140–155; tibia 140–185; tarsus 82–100; claw absent; setae only indicated by sockets on coxae.

Comment. Relatively few pupae of Eriococcidae have been described ( Stibococcus cerinus Miller & Gonzáles (Miller & Gonzáles, 1975) ; Eriochiton armatus Brittin. E. dugdalei Henderson & Hodgson and E. hoheriae Hodgson ( Hodgson & Henderson, 1996) ; Pseudotectococcus rolliniae Hodgson & Gonçalves ( Hodgson et al., 2004) , Eriococcus koelreuterius Wei & Wu ( Wei & Wu, 2004) , and Cryptococcus ulmi Tang & Hao ( Wu, 2000) . The last paper has not been seen but the pupae of the present species and the others listed above all share: (i) loculate pores ventrally and in pleural areas (and dorsally on Eriochiton spp and P. rolliniae ); (ii) absence of membranous lobes (caudal extensions) on abdominal segment VII; (iii) lobes on abdominal segment VIII quite distinct and rounded; (iv) penial sheath quite long, length probably at least equal to width, and (v) a mouth opening just posterior to head. The pupa of the present species differs from those of Eriochiton spp and P. rolliniae in lacking loculate pores on the dorsum, and from the pupa of S. cerinus in lacking spinose setae dorsally on the abdomen, in having 10 loculi in each loculate pore, and in having long pleural setae on abdominal segment VIII.

The pupae of E. melnikensis (and probably all Eriococcidae ) and Pseudococcidae have small wingbuds representing the hindwings or hamulolateres, which have not been noted on the pupae of higher neococcids. They also have many loculate pores; these are rare or absent on the pupae of Coccidae which also lack hindwing buds. The pupae of eriococcids differ from those of pseudococcids in: (i) lacking claws and well-developed setae on all legs (distinct on pseudococcids); (ii) the size of the penial sheath, which is barely demarcated from the abdomen on mealybugs but is large and well defined on eriococcids, and in (iii) lacking ostioles, which are present on the abdomen on pseudococcids. In addition, the structure of the loculate pores is different, that on eriococcids being typical of most higher neococcids (with the loculi well defined) whilst the loculi in the pores on pseudococcids are hard to discern, similar to those on the adult females.

Adult male ( Fig. 10). Described from 5 specimens in fair to good condition.

Instar diagnosis: small, total body length about 1.18–1.32 mm; antennae quite short, about 1/2 total-body length, all segments with fleshy setae (fs) and apical 3–5 segments with long or capitate setae; body with few setae, all hair-like (hs), fleshy setae (fs) apparently absent from derm; length of fs on antennae about half width of antennal segments; fs similar to those on antennae. Wings 4/5th total body length and about 0.43 as wide as long.

Head: approximately round to roundly oval; length about 190–200 µm, width across genae about 220–230 µm. Median crest not demarcated and not reticulated but with a long dorsal mid-cranial ridge (dmcr), extending posteriorly to about level with anterior margin of dorsal simple eyes; postoccipital ridge (por) distinct, with lateral arms extending both anteriorly and posteriorly; dorsal surface with (on each side) about 8–10 hs dorsal head setae (dhs) plus 2–5 pores, each rather variable, some irregular in shape, possibly with loculi, others like simple pores. Ventral mid-cranial ridge (vmcr) short and narrow, extending from lateral arms posteriorly to occipital sclerite; without any reticulation laterally, but with 3 or 4 pairs of ventral midcranial ridge setae. Genae (g) not reticulated; with 0–4 hs genal setae on each side just posterior to postoccular ridge (pocr) + 0 or 1 on dorsal end of pocr. Eyes: with two pairs of round, simple eyes; dorsal eyes (dse) rather posterior to ventral simple eyes (vse); simple eyes large, subequal in size, each 40–45 µm wide. Ocelli (o) large, situated laterally, each 15–20 µm wide, just touching postocular ridge. Ocular sclerite (ocs) partially sclerotised, particularly around and between simple eyes; lightly reticulated around each simple eye. Preocular ridge (procr): ventrally possibly extending medially to anterior to each ventral simple eye; dorsally possibly extending to just past dorsal simple eye dorsally. Postocular ridge (pocr) strongly developed, extending from postero-ventrally to each dorsal simple eye anterolaterally past each ocellus and around posterior to each dse. Interocular ridge absent. Dorsal ocular setae (docs) absent. Ventral head setae: with 4–6 hs vhs on each side just anterior to each vse. Preoral ridge poorly developed. Cranial apophysis (ca) short, 18–26 µm long and about 25–30 µm wide, with a trifid apex.

Antennae: 10-segmented and filiform; 530–590 µm long (ratio of total body length to antennal length 1:0.45). Scape (scp) 45 µm long and 45 µm wide, with 12–14 fs + 4 hs. Pedicel (pdc) arising anteriorly from scape: length 62–66 µm, width 45–50 µm; with a few faint concentric ridges on distal end; with 19–21 fs + 10 hs; campaniform sensillum present. Segments III–X each about 30 µm widest; each fs fairly stout, each 15–20 µm long; lengths of segments (µm): III: 100–110; IV: 65–80; V: 50–65; VI: 55–65; VII: 45–55; VIII: 40–45; IX: 40–45 and X: 40–50; approximate number of setae per segment: III: 21–27 fs + 5–8 hs; IV: 21–26 fs + 4 hs; V: 25 fs, 5–7 hs + 2 long setae; VI: 19–22 fs, 5–7 hs + 2 long setae; VII: 18–22 fs, 5–6 hs + 1 or 2 long seta; VIII: 13–17 fs, 5 hs, 2 long setae + 1 bristle; IX: 13–16 fs, 4 or 5 hs, 2 caps, 1 long setae + 1 bristle (abr), and segment X with about 7–10 fs, 4 or 5 hs (1 on apex), 3 large + 2 smaller bristles (one apical) on apical third + 4 or 5 capitate setae (cap); sensilla basiconica not detected; not constricted apically, although bluntly pointed.

Thorax. Prothorax: pronotal ridge (prnr) well-developed and touching dorsally; pronotal sclerite (prn) represented by a small area dorsolaterally, without lateral pronotal setae. Medial pronotal setae absent; posttergite (pt) possibly present, small, without post-tergital and antero-spiracular dorsal setae or pores. Prosternum apparently unsclerotised, without a median ridge but transverse ridge fairly well developed; without prosternal setae or pores. Anteprosternal absent.

Mesothorax: prescutum (prsc) probably oval, 75–95 µm long, 150–160 µm wide; sclerotised but not reticulated, with 2 or 3 pairs of hs prescutal setae; prescutal ridge (pscr) well developed anteriorly but quickly narrowing; prescutal suture narrow but apparently sclerotised. Scutum (sct): distance of prescutum from scutellum 45–55 µm; scutal setae: 0–2 hs on each side; lateral margins sclerotised but without reticulations laterad to prescutum; prealare ridge and triangular plate well developed. Scutellum (scl) 140–160 µm wide and 55–70 µm long; with a distinct scutellar ridge but no foramen; scutellar setae: 1–2 hs on each side + an occasional small pore; postnotal wing process broad and extending posterolaterally. Basisternum (stn 2) 240–260 µm wide and 155–166 µm long; without a median ridge; bounded anteriorly by a strong marginal ridge (mr) and precoxal ridge (pcr 2); without basisternal setae; lateropleurite (lpl) broad, with a distinct extension from marginal ridge along anterior and lateral margins; furca (f) well developed, narrow-waisted, arms very divergent and extending about half-way to marginal ridge anteriorly. Mesopostnotum well developed; postnotal apophysis well developed. Area bounded anteriorly by scutellum and laterally and posteriorly by meso-postnotum not sclerotised. Mesepisternum (eps 2) not reticulated; subepisternal ridge (ser) long and well developed. Postalare without postalare setae. Mesothoracic spiracle (sp 2): width of peritreme 25 µm, without associated loculate pores. Postmesospiracular setae absent. Tegula present, with 2–5 hs tegular setae (tegs).

Metathorax: with 2 hs metatergal setae (mts) medially and 1 or 2 hs laterally on each side; metapostnotal sclerite (pn 3) small and indistinct or absent. Dorsal part of metapleural ridge (plr 3) present, but without a suspensorial sclerite. Posterior part of metapleural ridge well developed; metepisternum not sclerotised and without postmetaspiracular setae; precoxal ridge lightly sclerotised, extending medially about 80–85 µm. Metepimeron (epm 3) sclerotised, but without setae. Antemetaspiracular setae absent. Metathoracic spiracle (sp 3): width of peritreme 26–28 µm, without loculate pores. Dorsospiracular setae absent. Metasternum (stn 3) membranous, metasternal apophysis small; with 0–2 hs anterior metasternal setae (amss) and 0–2 hs posterior metasternal setae (pmss).

Wings: hyaline, 850–975 µm long, 400–450 µm wide (ratio of length to width 1:0.46; ratio of total body length to wing length 1:0.73); alar lobe (al) well developed; each wing with 2 hs alar setae (als) but no circular sensoria. Hamulohalteres (h) each 95–100 µm long and 25–30 µm wide, each with a single hooked hamulus ham), 66 µm long.

Legs: posterior pair longest. Fleshy setae (fs) present on all segments, similar to those on antenna. Coxae (cx): I: 80–95; II: 95–105; III: 85–110 µm long; setae of coxa III: 15–20 fs + 11 hs; long setae on each coxa not differentiated. Trochanter (tr) + femur (fm): I: 205–230; II: 180–210; III: 205–230 µm long; trochanter III with 5 or 6 fs + 5 or 6 hs; each trochanter with 3 pairs of semi-circular sensoria arranged in a curve medially; long trochanter seta 40–45 µm; femur III with 24–30 fs + 10–16 hs. Tibia (ti): I: 220–255; II: 230–275; III: 250–300 µm; tibia III with 30–34 fs + many hs, many becoming spur-like on distal third; with 2 apical spurs (tibs) on all legs; longest 19–23 µm. Tarsi (ta) two segmented, proximal segment narrow and ringlike, length of both segments combined: I: 70–90; II: 85–90; III: 90–100 µm long (ratio of length of tibia III to length of tarsus III 1:0.35); tarsus III with 3–9 fs + about 18 mainly spur-like setae; tarsal spurs undifferentiated but 11–16 µm long; tarsal campaniform sensillum present; tarsal digitules (tdt) subequal to length of claw. Claws (c) subequal to or slightly longer than width of tarsus; length III: 24–26 µm; with a small denticle; claw digitules (cdt) with minute capitate apices and slightly longer than claw.

Abdomen: segments I–VII: tergites and sternites unsclerotised and without obvious oval membranous areas in inter-segmental membranes. Caudal extension (ce) of segment VII absent. Without loculate pores. Dorsal setae (ads) (totals): segments I–VII: 4–6 hs. Pleural setae: dorsopleural setae (dps) on each side: I–VII: 2–4; ventropleural setae (vps) on each side: I–VII: 1 hs. Ventral setae (avs) (totals): II: 2 hs; III–VII: 4–6 hs.

Segment VIII: tergite (at) sclerotised in a transverse band; sternite (as) less heavily sclerotised; with 2 hs ante-anal setae; sternite with 1 or 2 hs ventral abdominal setae (avs) on each side; caudal extension (ce) rounded, each with 4–5 hs pleural setae, one long, 50–85 µm long. Glandular pouches (gp) present, moderately deep, each with numerous loculate pores more or less restricted to pouches; each glandular pouch with 2 setae (gls) of different lengths and both slightly capitate or obviously blunt, length: long 150–170 µm, short 125–130 µm.

Genital segment: penial sheath (ps) divided into two sections: a broad anterior part (probably representing segment IX) and a short, triangular posterior section; anterior part with anal opening dorsally, 16–20 µm wide; ventrally with a broad, possibly unsclerotised, area around and anterior to basal rod; with 3 or 4 pairs of long setae (each 30–35 µm long) along anterior margin and 2 pairs of similar setae laterally. Triangular (posterior) part of penial sheath with 2 pairs of long setae, each 75–85 µm long, arising just above style. Length of penial sheath (from anterior border to apex: 210–230 µm, width across base 130–155 µm); posterior part 75–90 µm long. Copulatory structures complex: basal rod possibly absent, aedeagus (aed) fairly broad anteriorly, narrowing abruptly about half-way along length and sharply pointed (length about 100–105 µm); with two elongate sclerites ventral to aedeagus, more or less on each side of aedeagus, rather bulbous anteriorly but each with a sharp lateral spine posteriorly (length 80–90 µm), and with a median sclerite, perhaps dorsad to aedeagus, which broadens abruptly into two blade-like spines (width across outer spines 100–105 µm).

Comment. The most striking feature of this male is the copulatory apparatus, which is more complex than on any other male known to the authors and appears to be highly vicious. The homologies of the lateral and medial sclerites are unknown but the former are also shown as being present in males of Eriococcus araucariae Maskell and E. spuria (Modeer) (as Gossyparia spuria ) ( Afifi, 1968). The double scythe-like median sclerite would appear to be unique so far. However, these extra sclerites and the presence of the spines are probably to ensure a successful copulation, and may account for the sclerotisation of the area around the vulva on the adult female.

The macropterous adult males of Eriococcidae differ from those of Pseudococcidae (character-states of latter in brackets) in having: (i) claw digitules apically knobbed (setose); (ii) glandular pouches, when present, in distinct indentations, each with a pair of long setae (not in a distinct indentation and each with 3 or more setae); (iii) wing sensoria absent (present); (iv) postoccipital ridge well developed and laterally forked (weak and unforked); (v) loculate pores rare or absent (generally present), and (vi) ostioles absent (often present). Eriococcid males share many of the above characters with the males of Beesoniidae but differ from the latter family (character-states on Beesoniidae in brackets) in having: (i) fleshy setae, when present, generally entirely restricted to antennae and legs and always rather elongate — exceptions being Pseudotectococcus rolliniae Hodgson & Gonçalves ( Hodgson et al., 2004) and Calycicoccus merwei Brain ( Gullan et al., 2006) which have many fleshy setae on the body (abundant thick fleshy setae throughout body, each often almost as wide as long); (ii) antennae with 8 or more segments, and almost always 10 segmented (antennae with 7 or fewer segments); (iii) simple eyes rarely enlarged (often very much enlarged), and (iv) hamulohalteres present or absent (absent).

Figures

FIGURE 5. Adult female Eriococcus melnikensis Hodgson & Trencheva, spec. nov. For labels, see Fig. 1

FIGURE 6. Second-instar female Eriococcus melnikensis Hodgson & Trencheva, spec. nov. For labels, see Fig. 1.

FIGURE 7. Second-instar male Eriococcus melnikensis Hodgson & Trencheva, spec. nov. For labels, see Fig. 1.

FIGURE 8. First-instar Eriococcus melnikensis Hodgson & Trencheva, spec. nov. For labels, see Fig. 1.

FIGURE 9. Pupa Eriococcus melnikensis Hodgson & Trencheva, spec. nov., where ads = dorsal abdominal setae; an = anus; avs = ventral abdominal setae; ce = caudal extension; dhs = dorsal head setae; dps = dorsal pleural setae; lp = loculate pore; ps = penial sheath; sp2 = anterior spiracle; vps = ventral pleural setae; wb1 = anterior wingbud, and wb2 = posterior wingbud.