Nidularia balachowskii Bodenheimer

Spodek, Malkie & Ben-Dov, Yair, 2014, A taxonomic revision of the Kermesidae (Hemiptera: Coccoidea) in Israel, with a description of a new species, Zootaxa 3781 (1), pp. 1-99 : 79-93

Nidularia balachowskii Bodenheimer

( Figs. 41–48; Plate 6 , figs. a–j)

Nidularia balachowskii Bodenheimer, 1941: 78 –80.

Material examined. Type material studied. Lectotype adult ♀ ( ICVI), and paralectotype 1 adult ♀ (MNHN- 14802-4), 21 km at road from Mardin to Diyarbakir, Turkey, on branches and twigs of Quercus sp., 13.ii.1939, F.S. Bodenheimer. As Bodenheimer (1941) did not select a holotype, we regard the above-mentioned specimens as the original material studied by him as indicated on the slide labels. Other material studied. All non-type material collected off Q. ithaburensis in Israel by M. Spodek, unless otherwise indicated. Daphne Oaks (= current name Horshat Tal Nature Reserve), on Quercus sp., 1.v.1939, F.S. Bodenheimer, (C:4805/ 1 adult ♀). This was the first record of this species from Israel. Alonei Abba Nature Reserve, 11.1.2011, (MC:385/ 4 adult ♀, BMNH); Horshat Tal Nature Reserve, 30.v.2010 (MC:228/ 1 adult ♀), 14.ii.2012 (MC:614/ 5 adult ♀, MNHN), 27.ii.2011 (MC:430/ 2 adult ♀), 13.iii. 2012, Y. Ben-Dov (C:5131/ 1 adult ♀); Yehudiya Nature Reserve, 6.iii.2010 (MC:140/ 1 adult ♀), 10.x.2010, Y. Ben-Dov (C:4891/ 2 adult ♀), 10.x.2010 (MC:308/ 2 adult ♀, MC:309/ 3 adult ♀, MC:310/ 1 adult ♀, MC:311/ 1 adult ♀, MC: 315 adult ♀, MC:317/ 1 adult ♀), 7.xi.2010, Y. Ben-Dov (C:4912/ 10 adult ♀), 7.xi.2010 (MC:355/ 4 adult ♀), 11.i.2011, Y. Ben-Dov (C:4945/ 1 adult ♀), 11.i.2011 (MC:384/ 9 adult ♀), 6.ii.2011, Y. Ben- Dov (C:4970/ 1 adult ♀), 16.x.2011 (MC:587/ 6 adult ♀), 6.xi.2011, Y. Ben-Dov (C:4970/ 1 adult ♀), 3.vi.2012 (MC:690/ 1 adult ♀), 20.ix.2012 (MC:737/ 1 adult ♀), 20.v.2013 (MC:877/ 6 adult ♀); Bet Dagan-Volcani Center, 20.v.2012 (MC:662/4 third-instar ♀); Horshat Tal Nature Reserve, 27.ii.2011 (MC:431/1 third-instar ♀); Yehudiya Nature Reserve, 5.v.2013 (MC:871/3 third-instar ♀), 20.v.2013 (MC:876/9 third-instar ♀); Alonei Abba Nature Reserve, 22.iv.2013 (MC:852/5 second-instar ♀), 2.v.2013 (MC:863/2 second-instar ♀); Horshat Tal Nature Reserve, 27.ii.2011 (MC:431/5 second-instar ♀); Yehudiya Nature Reserve, 3.vi.2012 (MC:690/2 second-instar ♀), 5.viii.2012 (MC:725/1 second-instar ♀), 2.ix.2012 (MC:730/1 second-instar ♀, MC:731/1 second-instar ♀), 5.v.2013 (MC:871/1 second-instar ♀); Bet Dagan-Volcani Center, 13.iii.2012 (MC:635/12 first-instars, USNM), 15.iii.2012 (MC:636/40 first-instars, MNHN); Yehudiya Nature Reserve, 6.iii.2010 (MC:140/15 first-instars, BMNH), 20.iii.2011 (MC:460/68 first-instars); Bet Dagan-Volcani Center, 8.v.2013 (MC:881/ 5 adult ♂); Yehudiya Nature Reserve, 10.v.2013 (MC:870/ 5 adult ♂); Bet Dagan-Volcani Center, 6.vi.2012 (MC:686/ 1 pupa); Yehudiya Nature Reserve, 4.vi.2012 (MC:685/ 1 pupa), 8.v.2013 (MC:869/ 4 pupae), 10.v.2013 (MC:883/ 4 pupae); Alonei Abba Nature Reserve, 22.iv.2013 (MC:854/2 prepupae); Yehudiya Nature Reserve, 26.vi.2012 (MC:708/1 prepupa), 5.v.2013 (MC:868/2 prepupae), 8.v.2013 (MC:884/5 prepupae); Alonei Abba Nature Reserve, 22.iv.2013 (MC:853/1 second-instar ♂); Bet Dagan-Volcani, 6.vi.2012 (MC:683/2 second-instar ♂); Yehudiya Nature Reserve, 4.vi.2012 (MC:684/1 second-instar ♂), 2.ix.2012 (MC:726/1 second-instar ♂), 22.iv.2013 (MC:882/2 second-instar ♂), 2.v.2013 (MC:867/3 second-instar ♂). Additional non-type material of adult females from Turkey collected by M.B. Kaydan on Quercus sp. and deposited at the Yuzuncu Yil University: Van-Koçet Road (alt. 1625 m), 19.vii.2005, (#2056/ 1 adult ♀); Hakkari -Üzümcü Road (alt. 956 m) 15.ix.2005, (#2343/ 1 adult ♀); Van-Hakkari Road (alt. 1266 m), 16.ix.2005, (#2370/ 1 adult ♀); Hakkari-Doğan (alt. 1032 m), 22.v.2005, (#2688/ 1 adult ♀); Bitlis River (alt. 797 m), 23.vi.2006, (#3036/ 1 adult ♀); Bitlis-Kavakbaşı (alt. 1365), 30.v.2007, (#3419/ 1 adult ♀).

Adult female ( Fig. 41). General appearance. Pre-reproductive adult female ( Plate 6 , fig. c). Body oval, soft and flat, 1.2–1.9 mm long, 0.6–0.9 mm wide; dorsum brownish and venter yellowish-white; dorsal surface covered with 5 longitudinal rows of rectangular wax plates, each plate about 0.25 mm long, 0.3 mm wide; median row with 11 plates, mediolateral rows each with 9–11 plates and marginal rows with 7–9 plates. Wax plates largest medially. Gravid female and post-reproductive female ( Plate 6 , fig. d). Body oval, moderately convex and sclerotized, 2.75–3.75 mm long, 2.0–3.0 mm wide, 0.8–1.8 mm high; with 5 longitudinal rows of dark brown wax plates almost fused; with lighter brown wax in between rows of plates.

Mounted young adult female. Body oval, 0.8–3.0 mm long, 0.5–2.5 mm wide.

Margin. Marginal setae spinose, each 12–13 Μm long, in a single complete row of 30–36 on each side. Stigmatic spines absent.

Dorsum. Microtubular ducts with a circular opening about 1 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; present throughout dorsum. Other pore types and dorsal setae absent.

Venter. Eyes circular, each 5–7 Μm diameter, present anterolaterally to each scape. Antennae 1 segmented, 15–25 Μm long, 12–18 Μm wide; with 2–6 fleshy setae. Legs absent. Tentorium 113–155 Μm long, 113–125 Μm wide. Labium 3 segmented, triangular, 100–125 Μm long, 50–63 Μm wide; labial setae setose, as follows: basal segment with 2 pairs 9–20 Μm long, median segment with 1 pair 11–13 Μm long, apical segment with 4 pairs each 9–12 Μm long. Spiracles subequal in size, each peritreme 42–60 Μm long, 31–50 Μm wide, each sclerosis about 50 µm long. Quinquelocular pores each 5 Μm diameter: with 8–11 between mesothoracic spiracles and submarginal band of tubular ducts, and 10–13 in a broad band medially between metathoracic spiracles, plus 2–4 laterad to each metathoracic spiracle; also in a single, complete submarginal band of about 50–70 pores, band one pore wide. Bilocular pores each about 3 Μm wide, with 103–135 per side, dispersed within submarginal tubular duct band. Multilocular pores, each 7–8 Μm diameter with 9–10 loculi: in groups of 3 or 4 between each metathoracic and mesothoracic spiracles; in transverse bands across abdominal segments as follows; segment II with 12–24, III with 14–28, IV with 15–35, V with 18–25, VI with 12–19, VII with 2–3 on each side of vulva. Tubular ducts of Type 2 only, with outer ductule 12–18 Μm long, 5 Μm wide, a sclerotized cup about 5 Μm wide and inner ductule 22–30 Μm long; in a complete submarginal band 2–3 ducts wide, and sparsely medially in thorax. Ventral setae of two types: (i) setose setae: 3 or 4 pairs present in a group anterior to tentorium, each 7–13 Μm long; 2 setae, each 5–8 Μm long, medially to each spiracle; abdominal segments III–VI with transverse rows of 6–10 anterior to multilocular pores, each 7–8 Μm long. Also with 2 pairs anterior to anal ring, each 25–38 Μm long, and a pair postero–laterally to anal ring, 15–25 Μm long. Posterior margin with a pair of long apical setae, each 65–68 Μm long plus 2 pairs similar to marginal setae, each 15–22 Μm long; and (ii) tack-like setae, each 5–13 Μm long, 8 in a longitudinal submarginal row. Microspines, each 1 Μm long, present in 3–5 transverse rows on median and submedian areas of each abdominal segment; also scattered on thorax. Anal ring ventral, diameter 27–35 Μm, composed of 2 semi–circles, each half with 10–12 pores and 3 spinose setae each 65–75 µm long.

Comments. Bodenheimer’s (1941) original description of N. balachowskii is incomplete, lacking such details as presence of: (i) dorsal microtubular ducts; (ii) 8–11 quinquelocular pores between mesothoracic spiracles and submarginal band of tubular ducts, and 10–13 quinquelocular pores between metathoracic spiracles; (iii) bilocular pores interspersed between tubular ducts on venter; (iv) microspines on abdominal segments on venter; and (v) presence of eyes.

The adult female of N. balachowskii shares with N. japonica and the type species, N. pulvinata , the following characters: (i) one-segmented antennae; (ii) legs absent; (iii) seta-pore clusters absent; (iv) anal ring ventral; and (v) anal ring setose, with pores. Nidularia balachowskii can be easily separated from N. pulvinata by the presence of quinquelocular pores lateral to each anterior spiracle and medial to each posterior one in N. balachowskii but lying within each peritreme in N. pulvinata ). Nidularia balachowskii differs from N. japonica in the absence of tubular ducts dorsally (present on thorax and abdominal segments on venter of N. japonica ). The adult female of N. balachowskii differs from K. greeni , K. nahalali and K. spatulatus (characters of the latter in brackets): (i) onesegmented antennae (six-segmented antennae); (ii) legs absent (legs fully-developed); (iii) seta-pore clusters on venter absent (present); (iv) tubular ducts on both venter and dorsum of Type 2 (Type 1 and Type 2); (v) anal ring ventral (dorsal); and (vi) anal ring setose, with pores (absent).

However, along with N. balachowskii , two Israeli species, K. echinatus and K. hermonensis , and two Palaearctic species, K. quercus ( Podsiadlo, 2012) and K. vermilio ( Pellizzari et al., 2012) , share the following traits: (i) one-segmented antennae or with indistinct segmentation; (ii) seta-pore clusters absent; (iii) legs absent or reduced to three-segments; (iv) only Type 2 tubular ducts present on venter; (v) anal ring ventral; and (vi) anal ring setose and with pores.

Third-instar female ( Fig. 42; Plate 6 , fig. b). General appearance. Body oval, soft and flat, 0.5–0.8 mm long, 0.4–0.5 mm wide; dorsum brownish and venter yellowish-white; dorsal surface covered with 5 longitudinal rows of rectangular wax plates, median row with 11 plates, mediolateral rows with 9–11 plates and marginal rows with 7–9 plates. Wax plates largest medially.

Mounted specimen. Body oval; 0.7–0.8 mm long, 0.4–0.5 mm wide.

Margin. Marginal setae spinose, each 10 µm long, 5 µm wide at base, in a complete single line of 21–30 on each side.

Dorsum. Setae and tubular ducts absent. Microtubular ducts with a circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; in 4 longitudinal rows, each with 9 pores.

Venter. Eyes circular, each 5 µm diameter, lateral to each scape. Antennae 1 segmented, 25–30 µm long; with 2 fleshy setae on apical tip. Legs reduced to stumps that appear to be 1 segmented, each 15–20 µm long; occasionally with a round sclerotized area on apex. Tentorium 120–125 µm long, 87–95 µm wide. Labium 3 segmented, triangular, 62–80 µm long, 45–50 µm wide; labial setae setose: basal segment with 1 pair about 5 µm long, median segment with 1 pair 17–20 µm long and with 3 pairs of apical setae, each 8–13 µm long. Spiracles subequal in size, each peritreme 13 Μm long, 8 Μm wide; crescent-shaped sclerosis 28–35 µm long; each spiracle with quinquelocular pores anterolaterally, each 3 µm diameter, each anterior spiracle with 2 pores and each posterior spiracle with 1 pore. Tubular ducts of Type 2 only, with inner ductule 5–8 µm long, inner cup about 5 µm wide and outer ductule 8–13 µm long, arranged in a complete submarginal band 2–3 ducts wide; ducts also scattered medially on head and thorax. Ventral setae of two types: (i) setose setae: with 3 pairs anterior to tentorium, each 8 µm long; abdominal segments II–VII with medial and submedial setae, each 10 µm. Also with a pair just anterior to anal ring, about 7 µm long, plus a pair posterior to anal ring 12 µm long; (ii) tack-like setae, each 3 µm long, with 2 medial to each coxa, and also submarginally in abdominal segments II–VII. Microspines, each 2 µm long, in groups of 1–8 over entire body but in 2–5 transverse rows on abdominal segments. Anal ring ventral, diameter 25 µm, circular with a posterior gap, each half with 3 spinose setae, each 25 µm long, and 5–8 pores. Anal lobes barely developed, each with 2 spinose setae on apex, one 10 µm long and other 12 µm long, each similar in shape to marginal setae, plus a long setose apical setae 48–50 µm long.

Comments. The third-instar female of N. balachowskii can be distinguished from those of Kermes species by its narrow submarginal band of ventral tubular ducts (only one to two ducts wide). All other third-instar nymphs of Kermes species that have been described have a wider submarginal band, five to ten ducts wide. Nidularia balachowskii also resembles K. echinatus , K. hermonensis and K. vermilio in having (characters of K. greeni , K. nahalali and K. spatulatus in brackets): (i) leg segmentation reduced (legs fully developed); (ii) anal ring ventral (dorsal); (iii) tubular ducts of Type 2 on venter (Types 1 and Type 2 on venter); and (iv) sclerotized anal lobes absent (present).

Second-instar female ( Fig. 35; Plate 6 , fig. a). General appearance. Body oval, elongate, 0.5–0.8 mm long, 0.3–0.5 mm wide; dorsum greyish-white and venter yellow-brown; dorsal surface covered with a thin layer of wax, composed of 5 longitudinal rows of rectangular wax plates, median row with about 10–11 plates, submedian rows with about 9–11 plates and marginal rows with 9–12 plates. Wax plates largest medially.

Mounted specimen. Body oval, elongate, 0.6–0.9 mm long, 0.2–0.5 mm wide.

Margin. Marginal setae spinose, each about 8 µm long, 5 µm wide at base, in a single complete line of 24–31 on each side.

Dorsum. Setae and tubular ducts absent. Microtubular ducts with a circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; in 4 complete longitudinal rows, each row with about 25 pores.

Venter. Eyes circular, 10 µm diameter, laterally to each scape. Antennae segmentation indistinct, appears to be 3-segmented, each about 55 µm long; with 1 short seta on basal segment and a setose seta and 2 fleshy setae on apex. Legs reduced to 2 or 3 segments, segmentation indistinct; each leg 30–55 µm long; sometimes with claw on tip, about 3 µm long. Tentorium 70–100 µm long, 70–75 µm wide. Labium 3 segmented, triangular, 60–70 µm long, 40–50 µm wide; labial setae setose: basal segment with 2 pairs about 5 µm long, median segment with 1 pair 17–25 µm long and apical segment with 4 pairs 8 µm long. Spiracles subequal in size, each peritreme 20–25 Μm wide, each crescent-shaped sclerosis 30–50 µm long. Tubular ducts of Type 2 only, with an inner ductule 6–10 µm long, inner cup 5 µm wide and outer ductule 8–10 µm long, arranged in a submarginal band 1 or 2 ducts wide from cephalic apex to anal ring. Quinquelocular pores each 4 µm diameter; with 2 pores anterior to each anterior spiracle; 1 pore anterior to each posterior spiracle. Ventral setae of two types: (i) setose setae: 3 pairs of setae anterior to tentorium, each about 15 µm long; abdominal segments II–VII with setae in medial rows about 8 µm long, and in submedial rows about 5 µm long; 1 seta medial to each coxa 8 µm long. Also, 2 pairs just anterior to anal ring; and a pair posterior to anal ring about 10 µm long; (ii) tack-like setae, each 3 µm long, submarginally on abdominal segments II–VII. Microspines, each about 2 µm long, in groups of 1–8 in 5–8 rows on entire body. Anal ring ventral, diameter 35–45 µm, composed of 2 semi-circles, each half with 3 spinose setae about 30 µm long, and 8–12 pores. Anal lobes well-developed and somewhat rounded, with 2 spinose setae about 13 µm long, similar to marginal setae, with 1 on each inner margin and another on apex of each lobe, plus a long thin apical seta 50–65 µm long.

Comments. Among the Palaearctic species, the only other described second-instar females are those of K. bytinskii ( Sternlicht, 1969) , K. williamsi ( Sternlicht, 1970) , K. quercus ( Podsiadlo, 2012) , and K. vermilio ( Pellizzari et al., 2012) . That of N. balachowskii most closely resembles K. echinatus , K. hermonensis , K. quercus and K. vermilio in having reduced antennae and leg segments. However, N. balachowskii has three-segmented antennae and three-segmented legs, whereas K. echinatus has one-segmented antennae and one-segmented legs, K.

hermonensis has five or six-segmented antennae, K. quercus has six-segmented antennae and three-segmented legs and K. vermilio has three-segmented antennae and one-segmented legs. Also, second-instar female N. balachowskii (and K. vermilio ) lack frontal lobes, and, like K. echinatus , K. hermonensis and K. vermilio , N. balachowskii , lacks quinquelocular pores ventrally on the abdomen. Other Israeli species, i.e. K. greeni , K. nahalali and K. spatulatus , have six-segmented antennae and well-developed legs, as also found on several Nearctic species, including, K. cockerelli , K. concinnulus and N. iselini and O. secunda ( Baer & Kosztarab, 1985) .

First-instar nymph ( Fig. 44). General appearance. Body oval, tapering posteriorly, 0.38–0.43 mm long, 0.2–0.3 mm wide; dorsum and venter yellowish-grey.

Mounted specimen. Body oval, 0.42–0.53 mm long, 0.2–0.34 mm wide.

Margin. Marginal setae sharply spinose, pointed apically and slightly curved, each 9–15 Μm long, in a complete single row of 26–33 on one side.

Dorsum. Derm membranous with inter-segmental lines. Microtubular ducts with a circular opening about 1 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; in 4 incomplete longitudinal rows: submarginal lines on thorax and most abdominal segments, and medial lines restricted to abdomen Setae setose: with single setae submedially on each thoracic segment, each 5 Μm long.

Venter. Eyes each 12–15 Μm, near margin. Antennae 6 segmented, length 88–125 Μm; setal distribution as follows: scape and pedicel each with 2 setose setae; segment III with 1 long setose seta; IV with 1 fleshy seta; V with 2 fleshy setae and 2 setose setae; apical segment with 2 fleshy setae and 5 setose setae. Legs well-developed; measurements (in Μm) of metathoracic leg: coxa 25–30, trochanter + femur 63–80, tibia 25–38, tarsus 25–60, claw 13–23; total length 158–213; trochanter with 2 oval, sensory pores on each side; tarsal digitules 25–30 Μm long, knobbed apically, extending beyond claw apex; claw digitules knobbed apically, each 14–20 Μm long, shorter than tarsal digitules; each claw with a single denticle. Tentorium well-developed, 75–90 Μm long, 63–75 Μm wide. Labium 3 segmented, triangular, 82–100 Μm long, 35–38 Μm wide; labial setae setose, as follows: basal segment with 2 pairs 10–15 Μm long, median segment with 1 pair on dorsal surface 10–20 Μm long, apical segment with 4 pairs 12–17 Μm long. Spiracles subequal in size, each peritreme 3–5 Μm wide, each crescent-shaped sclerosis 15– 25 Μm long. Bilocular pores oval, each 2 Μm long and 1 Μm wide; with 1 pore present between margin and each spiracle. Quinquelocular pores, each 3 Μm diameter, with 1 pore just anterior to each spiracle and a pore medial to each coxa; plus 2 on each of abdominal segments IV, V, VI. Ventral setae of three types: (i) sharply spinose setae, similar to those on margin: with 1 pair near anterior margin, each 10–15 Μm; a line of 7 or 8 setae submarginally in metathorax and abdominal segments II–VII, similar to marginal setae but shorter, each about 7 Μm long; (ii) setose setae: 3 pairs between scapes, each 20–30 Μm long; 1 seta medial to each coxa 8–15 Μm long; also segmentally arranged medially and submedially on abdominal segments II–VII, each 10–17 Μm long. Plus a pair anterior to anal ring 12–15 Μm long, and another pair postero-lateral to anal ring 15–25 Μm long; and (iii) tack-like setae, each 5–6 Μm long, extending submarginally from metathorax to abdominal segment VII. Microspines, each about 1 Μm long, arranged in 2–4 transverse rows in median and submedian areas of each abdominal segment. Anal ring ventral, diameter 15–25 Μm, composed of 2 semi-circles, each half with 3 spinose setae 20–30 Μm long, and about 17 pores. Anal lobes well-developed; each lobe with 2 spinose setae on inner margin, each distinctly thicker than marginal setae and 12–15 Μm long, plus a long apical seta 77–125 Μm long.

Comments. There is no unique set of characters that distinguishes the first-instar nymphs of Nidularia from the Palaearctic Kermes examined in this study. Those of N. balachowskii can be distinguished from other Nidularia species by the sharply spinose, apically pointed and slightly curved marginal setae, each 9–15 Μm long; N. pulvinata has setose setae ( Koteja, 1980) and N. japonica has setose setae, somewhat conical at their base ( Kuwana, 1918; Liu et al., 1997).

Adult male ( Fig. 45; Plate 6 , fig. i). As for K. echinatus apart from: General appearance. Newly emerged males brown-black; body 0.8–1.0 mm long; width across mesothorax 0.2–0.3 mm wide; fore-wing 0.8–1.0 mm long, 0.35–0.45 mm wide; caudal filaments 0.5–1.0 mm long.

Mounted specimen. Total length 0.8–1.0 mm long, width across mesothorax 0.2–0.4 mm.

Head. Length 175–200 µm long; width across genae 200–215 µm. Dorsal and ventral simple eyes each 30–45 µm diameter; lateral simple eyes (lse) smaller, 15 µm diameter; ocellus 10–13 µm diameter. Dorsal head setae (dhs) 8–10 hs on either side of median crest. Ventral head setae (vhs) few, total 4–6 hs. Antennae: length 375–400 µm (ratio of total body length to antennal length about 1:0.43). Scape (scp) 30–35 µm long, 35–40 µm wide, with 2 or 3 hs. Pedicel (ped) 50–55 µm long, 35–40 µm wide, with 2 or 3 hs. Flagellar segments III–X each 35–65 µm long and 20–22 µm wide; segment III longest, segments becoming shorter towards apex; approximate number of setae per segment: III–VIII 2–8 fs; IX–X 10–14 fs; X 2 ab + 4 cs.

Thorax. Prothorax: as for K. echinatus but sternum (stn1) with faint radial striations and 1 hs prosternal setae (stn1s) on each side. Anteprosternal (astn1s) and antemesospiracular setae (am2s) absent.

Mesothorax: prescutum (prsc) rectangular, 70–80 µm long, 100–125 µm wide. Scutum (sct): median membranous area 50–60 µm long; with 1 hs on each side, lateral to membranous area, 13–15 µm long. Scutellum (scl) 40–45 µm long, 125–130 µm wide; with 1 hs 13–15 µm long on each side. Basisternum (stn2) 125–130 µm long, 200–210 µm wide; basisternal setae absent. Tegula (teg) distinct, with 3–5 hair-like tegular setae (tegs).

Metathorax: Metathoracic spiracles (sp3): peritreme 15–18 µm wide. Metasternum (stn3) with a total of 20–25 hs setae: 7 or 8 anterior metasternal setae (amss); 15 or 16 posterior metasternal setae (pmss) and 3 or 4 postmetaspiracular setae (eps3s), each 20–30 µm long.

Wings: 870–1000 µm long and 350–400 µm wide (ratio of length to width 1:0.4; ratio of total body length to wing length 1:1). Hamulohalteres (h) 25–35 µm long, 10–15 µm wide; with a single apical hamulus, 15–20 µm long. Legs: subequal in size, length 494–521 µm. Coxa: I 45 –50; II 50 –63; III 50 –60 µm long, with 7 or 8 hs. Trochanter + femur: I 155–162; II 158–163; III 158–164 µm long, with 6–8 hs; each trochanter with 3 oval sensory pores in a triangle on each surface. Tibia: I 140–150; II 140–150; III 120–150 µm long, with 8–10 hs; all tibia with 2 apical spurs (tibs), 20–25 µm long. Tarsi 2-segmented; tarsus 2 (tar2): I 55 –65; II 58 –65; III 60 –75 µm long, with 8–10 hs; tarsal digitules (tdgt) about 38 µm long. Claw (cl) with denticle: I 25–28; II 25–28; III 28–30 µm long; claw digitules (cdt) about 25 µm long.

Abdomen. Tergites (at), sternites (as) and pleural areas unsclerotized. Abdominal dorsal setae (ads) (totals): I– VII 2 hs, 10 µm long; VIII 4 hs, 40 µm long; pleural setae: dorsopleural setae (adps) (on each side): I 8 hs and II– VII 4 or 5 hs, 10 µm long, VIII 3 hs, one 60–65 µm long and others 25–30 µm long; ventropleural setae (avps) (on each side) II–VII 3 or 4 hs, 15 µm long; abdominal ventral setae (avs) (totals) II–VII 10–14 hs, VIII 4 hs, 28–30 µm long. Glandular pouch setae (gls) 88–100 µm long; each pouch with 7–10 pores, each 3 µm diameter with 3–5 loculi. Genital segment: penial sheath (ps) 120–130 µm long, greatest width 70–80 µm. Basal rod (bra) 30–35 µm long. Aedeagus (aed) 63–70 µm long. Genital capsule with 2 pairs of hs (gts) dorsally and 4 or 5 pairs hs (gts) ventrally

Comments. The major feature distinguishing the adult male of N. balachowskii from the other five Israeli and Nearctic species is body size. The adult male of N. balachowskii is only 0.8–1.1 mm long and 0.2–0.3 mm wide at metathorax whereas the other species are 1.5–2.0 mm long and 0.4–0.5 mm wide. Nidularia balachowskii can be distinguished from N. japonica by the absence of setae on the prosternum ( N. japonica has eight setae) and the presence of one pair of dorsal setae on the abdominal segments ( N. japonica has two pairs) ( Liu et al., 1997).

Fourth-instar male (pupa) ( Fig. 46; Plate 6 , fig. h). Develops within a white, flocculent (wooly) wax test secreted by second-instar male. Morphologically very similar to that of K. echinatus but differing as follows. General appearance. Body brown, 0.9–1.0 mm long, 0.2–0.36 mm wide.

Mounted specimen. Body 0.92–1.0 mm long, 0.34–4.0 mm wide; head 165–175 µm long, 250–275 µm wide.

Margin. Wing-buds: fore-wings 375–440 µm long, 100–140 µm wide; hind-wings: length ratio to mesothoracic wings 1:11. Setae fine and blunt, each 12–15 µm long, distributed as on K. echinatus .

Dorsum. Eyes absent. Ocular sclerite reticulated, 35–50 µm long, 75–90 µm wide. Setae setose, each 5 µm long, with 4 pairs on head apex; and in submedial longitudinal rows. Anal lobes rounded and membranous, each with a setose seta 55–95 Μm long on apex. Median penial lobe triangular, 95–105 Μm long, 95–115 Μm wide; with 3 pairs of setose setae, each 15–20 Μm long. Venter. Ocular sclerite as on dorsum. Antennae 10 segmented, total length 400–460 µm, width 25–35 µm; setae absent. Spiracles subequal in size, each peritreme 18–20 µm long, 10–15 µm wide, each sclerosis 25–33 µm long; without pores. Legs well-developed, subequal in size, measurements (in Μm) of metathoracic legs: coxa 50–75, trochanter + femur 125–150, tibia 100–110, tarsus 60–75; total length 335–410 Μm; each coxa with 2 short setae, each 5 µm long; incipient claw 5 µm long. Ventral setae setose: with 4 pairs of interantennal setae, each 5 Μm long; abdominal setae arranged as on K. echinatus . Microspines, as on K. echinatus .

Comments. The pupa of N. balachowskii is greyish-brown in life, similar to those of K. nahalali and K. spatulatus but unlike other Israeli kermesids, K. greeni is orange and K. echinatus and K. hermonensis are red. As in all known kermesid pupae bar K. hermonensis , the pupa of N. balachowskii lacks quinquelocular pores anterior to each spiracle. Also N. balachowskii has membranous anal lobes, as in all described kermesid pupae apart from K. greeni and K. spatulatus , which have sclerotized anal lobes.

Third-instar male (prepupa) ( Fig. 47; Plate 6 , fig. g). As for K echinatus apart from: General appearance. Body brown and broadly oval when removed from test, 0.5–1.0 mm long, 0.2–0.3 mm wide.

Mounted specimen. Body 0.5–1.0 mm long, 0.2–0.3 mm wide; separation between head, thorax and abdomen not clearly demarcated. Margin. Anterior wing-buds each 250–300 µm long, 90–110 µm wide, extending to about level with metathoracic coxae. Setae fine and blunt, arranged as on K. echinatus .

Dorsum. Setae setose, each 5–10 µm long, with 4 pairs on head apex; and also with submedial longitudinal rows. Anal lobes rounded apically, with 2 setae on apex of each lobe, 1 long and setose, 50–60 Μm long, and other spinose, 10–15 Μm long. Median penial lobe 35–40 Μm long, 50–65 Μm wide, broadly rounded; with 6 setose setae, each 5–10 Μm long.

Venter. Eyes absent. Antennae 8, 9 or 10 segmented, tapering apically; length 250–290 µm, 65–80 µm wide; with 2 short fleshy setae, about 5 µm long, on scape and 1 on segments VIII, IX and X. Spiracles subequal in size, each peritreme 18–20 µm long, 10–15 µm wide, each sclerosis 20–30 µm long; each spiracle with 1 quinquelocular pore anterolaterally, 3 Μm diameter. Legs 3 segmented lobes, subequal in size; total length of metathoracic leg, 145–175 Μm and 60–75 Μm wide; claw absent; each coxa with 2 short setae about 5 Μm long. Ventral setae setose, each 8–10 Μm long; with 4 pairs between scapes, and abdominal setae arranged as on K. echinatus . Microspines arranged as on K. echinatus .

Comments. The prepupa of N. balachowskii is greyish-brown in life, similar to K. nahalali and K. spatulatus but unlike the other Israeli kermesids, K. greeni is orange and K. echinatus and K. hermonensis is red. The prepupa of N. balachowskii has a single quinquelocular pore anterior to each spiracle, while those of K. echinatus and K. hermonensis have two pores. The other Israeli kermesids ( K. greeni , K. nahalali and K. spatulatus ) and the Nearctic species A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) lack spiracular pores.

Second-instar male ( Fig. 48; Plate 6 , fig. f). General appearance. Secretes a white, wooly wax test, 1.0– 1.5 mm long, 0.4–0.8 mm wide; body yellow-brownish, broadly oval when removed from test, 0.7–1.45 mm long, 0.4– 0.8 mm wide; derm membranous, with inter-segmental lines.

Mounted specimen. Body oval, 0.7–1.8 mm long, 0.3–0.9 mm wide.

Margin. Marginal setae robust and setose, each 18 µm long, in a complete single row of 26–30 on each side.

Dorsum. Microtubular ducts with a circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; numerous, scattered over entire dorsum. Tubular ducts of Type 2 only, with inner ductule 10–15 µm long, cup 5 µm wide and outer ductule 9–12 µm long, abundant throughout dorsum. Setae setose, each 10–15 µm long, in submedial longitudinal rows, each row with 5 setae.

Venter. Eyes circular, each 10 µm diameter, lateral to each scape. Antennae 7 segmented, length 100–125 µm; segments III and VII longest; scape and pedicel each with 2 setose setae; segment III without setae; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta, VI with 1 fleshy seta and 1 setose seta, and apical segment with 4 fleshy setae and 3 setose setae. Frontal lobes absent. Legs well-developed; measurements (in Μm) of metathoracic legs: coxa 35–50; trochanter + femur 100–105; tibia 40–50; tarsus 75–80 and claw 20–25; total length 250–275 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 25–30 Μm long, knobbed apically; claw digitules knobbed apically, each 20–25 Μm long; each claw with a single denticle. Tentorium 90–100 µm long, 75–90 µm wide. Labium 3 segmented, triangular, 80–100 µm long, 45–50 µm wide; labial setae setose: basal segment with 2 pairs about 10 µm long, median segment with 1 pair 10 µm long and apical segment 4 pairs 15 µm long. Spiracles subequal in size, each peritreme 10–13 Μm wide, each sclerosis 25–30 µm long; each spiracle with 3 quinquelocular pores anterolaterally, each 5 µm diameter. Tubular ducts of Type 2 only, with inner ductule 10– 15 µm long, inner cup 5 µm wide and outer ductule 9–12 µm long, in a complete submarginal band 1 or 2 ducts wide. Bilocular pores circular, each 2 µm diameter, with 10–14 pores in a single complete submarginal line. Quinquelocular pores, each 3 µm wide, with 2 pairs between scapes; 1 pore medial to each coxa; also submedially on abdominal segments II–VII; and with 10–12 pores in a single complete submarginal line. Ventral setae of three types: (i) robust setose setae: with a pair similar to marginal setae on anterior margin between scapes, each about 14 µm long; (ii) setose setae: with 3 pairs medially just anterior to tentorium, each about 30 µm long; 3 mesad to each coxa, each 25–35 µm long; abdominal segments II–VII with medial rows, each about 15 µm long, and submedial rows, each about 10 µm long. Also with 2 pairs anterior to anal ring, each 10–20 µm long, and 2 pairs posterior to anal ring, one pair 8–10 µm long and other pair long and flagellate, each 80–85 µm long; (iii) tack-like setae in a submarginal row on abdominal segments II–VII, each 5 µm long. Microspines, each about 3 µm long, in groups of 3–5 in 6 or 7 transverse rows on each abdominal segment and on scattered on thorax. Anal ring ventral; diameter 25–30 µm, composed of 2 semi-circles, each half with 3 spinose setae, each 28–33 µm long, plus 10–16 pores.

Comments. The second-instar males of the other two Nidularia species, N. japonica and N. pulvinata , have not been described. Nidularia balachowskii lacks the anal lobes found on K. greeni and K. nahalali and has no frontal lobes, as found in Kermes echinatus , K. hermonensis and K. vermilio ( Pellizzari et al., 2012) . In addition, what distinguishes N. balachowskii from other Israeli kermesids is the presence of three quinquelocular pores associated with each spiracle. All the other species have more or fewer than three pores: K. echinatus (4) K. greeni (five), K. hermonensis (eight), K. nahalali (eight), K. quercus (4–9) ( Podsiadlo, 2012), K. spatulatus (five) and K. vermilio (two) ( Pellizzari et al., 2012). The Nearctic second-instar males of K. cockerelli , K. concinnulus and K. rimarum differ in having long, setose marginal setae and, unlike N. balachowskii which only has quinquelocular pores on the venter, have them scattered on both the dorsum and venter ( Baer & Kosztarab, 1985).

Figures

PLATE 6. Nidularia balachowskii Bodenheimer, general appearance; a. second-instar female; b. third-instar female; c. young adult female; d. post-reproductive female; e. male test and exuviae (arrow); f. second-instar male; g. prepupa; h. pupa; i. adult male, dorsal view, j. adult male, ventral view.