Kermes greeni 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 : 24-50

Kermes greeni Bodenheimer

( Figs. 9–16, Plate 2 , figs. a–i)

Kermes greeni Bodenheimer, 1931: 241

Kermes palestiniensis Balachowsky, 1953: 186 ; synonymized by Spodek et al., 2012: 67–75 This species was originally described from adult females collected at Nahalal forest, Israel, on Quercus coccifera L. The junior synonym, K. palestiniensis , was described from first-instar nymphs collected at Ras-El Nakurah [=current name Rosh Hanikra], Israel, on Q. coccifera.

Material examined. Type material studied. Lectotype adult ♀ of K. greeni , off Q. coccifera, Israel, Nahalal, 2.ii.1926, F.S. Bodenheimer ( ICVI). Paralectotypes 7 adult ♀ (6 slides in ICVI, 1 slide in BMNH), same data as lectotype. Also: 5 first-instar nymphs in 2 of the paralectotype adult ♀, Israel: Nahalal, K. greeni , 2.ii.1926, F.S. Bodenheimer. Also: Syntype 3 first-instar nymphs K. palestiniensis Ras-el Nakurah [=current name Rosh Hanikra], Q. coccifera, 12.vii.1950, S. Neumark ( MNHN /1 slide). Other material studied. All non-type material collected in Israel, off Q. calliprinos by M. Spodek, unless otherwise indicated. Bet Dagan-Volcani Center, 6.v.2012 (MC:652/ 1 adult ♀); Eilon, 13.iii.2011 (MC:459/ 1 adult ♀), Y. Ben-Dov (C:4765/ 1 adult ♀); Hanita, 5.vi.2001, Z. Tamari (C:4701/ 1 adult ♀), 6.vi.2010 (MC:247/ 1 adult ♀), 8.v.2011, Y. Ben-Dov (C:4997/ 2 adult ♀, BMNH), 8.v.2011 (MC:493/ 1 adult ♀), 17.v.2011, Y. Ben-Dov (C:5011/ 2 adult ♀, MNHN), 7.v.2012 (MC:668/ 9 adult ♀); Timrat, 24.vi.2011 (MC:567/ 8 adult ♀), 31.vii.2011 (MC:568/ 2 adult ♀, USNM), 22.iv.2012, Y. Ben-Dov (C:5096/ 2 adult ♀), 7.v.2012, Y. Ben-Dov (C:5099/ 2 adult ♀), 7.v.2012 (MC:759/ 11 adult ♀); Eilon, 11.iv.2011 (MC:483/2 third-instar ♀), 21.iv.2011 (MC:491/1 third-instar ♀); Hanita, 21.iv.2011 (MC:485/6 third-instar♀), 8.v.2011 (MC:492/6 third-instar♀, MC:493/7 third-instar ♀), 7.v.2012 (MC:667/4 third-instar ♀); Timrat, 25.ii.2012 (MC:645/7 third-instar ♀), 22.iv.2012 (MC:647/1 third-instar ♀), 7.v.2012 (MC:653/4 third-instar ♀, MC:761/1 third-instar ♀), 18.iv.2013 (MC:841/3 third-instar ♀); Eilon, 11.iv.2011 (MC:482/10 second-instar ♀); Hanita, 13.iii.2011 (MC:442/1 second-instar ♀), 21.iv.2011 (MC:484/13 second-instar ♀), 24.v.2012 (MC:278/1 second-instar ♀); Timrat, 25.iii.2012 (MC:645/1 second-instar ♀), 22.iv.2012 (MC:647/9 second-instar ♀), 18.iv.2013 (MC:841/2 second-instar ♀); Eilon, 13.iii.2011 (MC:459/1 first-instar); Gesher Alkosh, 16.vii.2001 (MC:269/1 first-instar); Hanita, 9.vi.2010 (MC:247/1 first-instar), 24.i.2011 (MC:405/4 first-instars, BMNH), 3.vi.2011 (MC:520/175 first-instars), 3.vi.2012 (MC:706/38 first-instars); Timrat, 31.vii.2011 (MC:568/3 firstinstars, MNHN), 30.v.2012 (MC:705/3 first-instars, USNM), 10.xii.2012 (MC:766/3 first-instars); Eilon, 23.iv.2013 (MC:859/ 1 adult ♂); Hanita, 4.v.2012 (MC:838/ 2 adult ♂), 26.iv.2013 (MC:860/ 4 adult ♂); Timrat, 28.iv.2013 (MC:861/ 3 adult ♂); Eilon, 22.iv.2013 (MC:856/ 2 pupae); Hanita, 24.v.2012 (MC:279/ 3 pupae), 1.v.2012 (MC:816/ 3 pupae), 25.iv.2013 (MC:866/ 2 pupae); Eilon, 15.iv.2013 (MC:857/6 prepupae); Hanita, 16.iv.2013 (MC:858/4 prepupae); Eilon, 11.iv.2011 (MC:482/2 second-instar ♂), 10.iv.2013 (MC:855/2 secondinstar ♂); Hanita, 24.v.2012 (MC:278/2 second-instar ♂), 10.iv.2013 (MC:845/4 second-instar ♂).

Adult female ( Fig. 9). General appearance. Pre-reproductive adult female ( Plate 2 , fig. c). Body oval to round, 2.5–3.7 mm long, 1.9–3.0 mm wide; dorsum with red and black areas and with a transverse red area medially; dorsum convex and smooth. Gravid female ( Plate 2 , fig. d). Body spherical, 4.2–5.9 mm long, 2.9–5.0 mm wide, 3.8–4.3 mm high; body dark red to black and uniform in colour. Post-reproductive female ( Plate 2 , fig. e). Body spherical and derm smooth and sclerotized; body 4.2–5.9 mm long, 2.9–5.0 mm wide, 3.8–4.3 mm high, and orange to tan coloured.

Mounted young adult female. Oval to circular, 2.5–3.0 mm long, 1.8–2.5 wide. Margin. Marginal setae short and spinose, each about 5 Μm long, arranged in a single row of 15–22 on each side. Stigmatic setae not differentiated.

Dorsum. Bilocular pores oval, each about 3 Μm long, 2 Μm wide, abundant throughout dorsum. Tubular ducts of Type 2, each with inner ductule 7–17 Μm long, inner cup 5–8 Μm wide and outer ductule 12–15 Μm long, sparse, much less frequent than bilocular pores. Seta-pore clusters: setae 17–25 um long, each with a cluster of 4– 14 pores, each 8 Μm wide with 3–5 loculi, in 10–13 clusters on each side, extending in a single complete submarginal row. Anal ring dorsal, circular, diameter 75–100 Μm, setae and pores absent. Anal lobes probably represented by two quadrate, sclerotized areas posterior to anal ring, each 50–75 Μm long, 63–75 Μm wide, each with 13–16 setose setae 30–43 Μm long.

Venter. Eyes absent. Antennae 4, 5 or 6 segmented; when 4-segmented, segments I–III appear fused; in 5- segmented antennae, segments II and III appear fused; each antenna 105–110 Μm long; setal distribution as follows: scape with 2 setose setae; pedicel with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 2 fleshy setae; apical segment with 4 setose setae and 1 fleshy seta. Legs well-developed; tibia and tarsus occasionally fused, measurements (in Μm) of metathoracic legs: coxa 30–35, trochanter + femur 70–75, tibia 37– 50, tarsus 40–48, claw blunt, 10–18 Μm long; total length 212–250; each trochanter with 2 oval, sensory pores on each side; tarsal digitules 25–38 Μm long, knobbed apically and extending beyond apex of claw; claw digitules knobbed apically, each 22–25 Μm long, shorter than tarsal digitules. Denticle absent. Tentorium 250–287 Μm long, 237–250 Μm wide. Labium 3 segmented, triangular, 162–212 Μm long, 125–162 Μm wide; labial setae all setose: basal segment with 2 pairs 3–5 Μm long, median segment with 1 pair 7–8 Μm long, and apical segment with 4 pairs 10–13 Μm long. Spiracles subequal in size, each peritreme 62–87 Μm long and 45–55 Μm wide. Multilocular pores of variable sizes distributed as follows: (i) a group of 35–65 surrounding each spiracle, each 5–7 Μm in diameter with 5–6 loculi; (ii) in 6–8 clusters of 30–60 pores on abdominal segments III–VII, each 7–8 Μm in diameter with 9–11 loculi; (iii) with 12–24 on each abdominal segment, not in clusters, each 7 Μm in diameter with 5 or 6 loculi,; (iv) in submedial groups of 64–104 posterior to vulva, each 7–8 Μm in diameter with 5 or 6 loculi; and (v) a medial group of 52–60 pores, posterior to vulva, 6 Μm diameter with 4–7 loculi. Tubular ducts of Type 1, with inner ductule 7–17 Μm long, cup 5 Μm wide, and outer ductule 22–30 Μm long, in a dense complete submarginal band about 10 ducts wide. Setae setose: each abdominal segment with a line of about 8–10 setae, submedially, each 23–25 Μm long, plus 8–10 medially, each 10–12 Μm long; also with a group of 40–48 setae posterior to vulva, each 10–13 Μm long. Microspines, each about 2 or 3 Μm long, arranged in groups of 3 or 4 in about 8 rows on each abdominal segment.

Comments. The adult female of K. greeni shares the following characters with those of K. nahalali and K. spatulatus : (i) fully-developed legs; (ii) six-segmented antennae; (iii) seta-pore clusters located submarginally on dorsum; (iv) anal ring dorsal; (v) anal ring without setae or pores; (vi) anal lobes sclerotized on dorsum; and (vii) two types of tubular ducts, one on venter and other on dorsum. The above characters are also present in adult females of the Oriental species K. flavus Liu (Liu & Shi, 1995) , K. miyasakii ( Hu, 1986) , K. orientalis ( Liu & Shi, 1995) and K. taishanensis ( Hu, 1986) . The segmentation of the legs and antennae varies between the non-Israeli species; legs are reduced to three-segments in K. quercus ( Borchsenius, 1960) , K. roboris ( Saakyan-Baranova & Muzafarov, 1972) , K. sylvestris ( Bullington & Kosztarab, 1985) , and K. williamsi ( Sternlicht, 1970) , while K. nudus and K. rimarum have fully-developed legs ( Bullington & Kosztarab, 1985) as in K. greeni . The Nearctic A. kingii also has fully-developed legs ( Hamon et al., 1976). The number of antennal segment ranges from four to six for the above mentioned species, mostly with indistinct segmentation. Adult female K. greeni can be distinguished from other Kermes species by the presence of quadrate-shaped anal lobes on the dorsum, each lobe with more than five setose setae. The anal lobes of K. spatulatus and K. nahalali are narrower compared to K. greeni , with fewer than five setae. The adult females of K. echinatus , K. hermonensis and N. balachowskii do not have anal lobes.

Third-instar female ( Fig. 10; Plate 2 , fig. b). General appearance. Body round, slightly convex, 1.2–1.7 mm long, 1.0– 1.5 mm wide, 0.8–1.2 mm high; dorsum with orange and brown areas, with a medial, transverse orange area and 4 transverse brown rows.

Mounted specimen. Oval to round, 1.0– 1.8 mm long, 0.6–1.6 mm wide.

Margin. Marginal setae short and spinose, each 10 µm long and 5 µm wide, in a single complete line of 20– 26 on each side. Dorsal submarginal setae robust, in two rows, larger and longer than marginal setae: outer row a single complete line of 20–25 setae on each side, each 25 µm long, each seta frequently associated with 1–3 quinquelocular pores, each 5 µm diameter; inner line of 15–18 setae, similar to submarginal setae, but shorter, each about 16 µm long.

Dorsum. Setae: setose setae in submedial longitudinal rows of about 5 setae, each 8 µm long; also spinose setae, similar to submarginal setae but short, in a group of 3–6 near anterior apex of body. Bilocular pores oval to circular, each 3 µm long and 2 µm wide; abundant on dorsum. Anal ring dorsal, horseshoe shaped, with anterior gap; about 40 µm diameter; setae and pores absent. Anal lobes sclerotized circular areas, each 60 µm long and 50 µm wide, with 12–15 setose setae each 60 µm long.

Venter. Eyes circular, each 10 µm wide, lateral to each scape near body margin. Antennae 6 segmented, 125 µm long; segment III longer than other segments; scape with 2 setose setae; pedicel with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta; apical segment with 4 setose setae and 1 fleshy seta. Legs well-developed; measurements (in µm) of metathoracic leg: coxa 37–50, trochanter + femur 65–74, tibia 70–75, tarsus 32–48, claw 13–20; total length 220–250 Μm; trochanter with 2 oval, sensory pores on each side; tarsal digitules 20–37 Μm long, knobbed apically, slightly extending beyond claw apex; claw digitules knobbed apically, each 20–25 Μm long, shorter than tarsal digitules; claws without a claw. Tentorium 150–175 µm long, 163–180 µm wide. Labium 3 segmented, triangular, 110–125 µm long, 83–100 µm wide; with setose setae as follows: basal segment with 2 pairs, each 5–8 µm long; medial segment with 1 pair about 13 µm long and apical segment with 4 pairs 10 µm long. Spiracles: mesothoracic spiracles smaller than metathoracic spiracles; each mesothoracic peritreme 20 Μm long, 13 Μm wide, with sclerosis 30 µm long; each metathoracic peritreme 28 Μm long, 20 Μm wide, with sclerosis 30 µm long. Tubular ducts of two types: Type 1, with inner ductule 10–13 µm long, inner cup about 5 µm wide and outer ductule 10–13 µm long, arranged in a complete dense submarginal band about 5 or 6 ducts wide; Type 2, with inner ductule 10–13 µm long, inner cup about 5 µm wide and outer ductule 15–18 µm long, arranged in a complete submedial band 1 or 2 ducts wide and scattered medially on thorax. Quinquelocular pores, each 5 µm diameter: with 2 pores anterior to tentorium; 2–4 pores anterolateral to each spiracle; 1 pore medial to each coxa; and with 1 medial and 1 submedial pore on each abdominal segment. Ventral setae of two types: (i) setose setae: 12 anterior to tentorium, each 25 µm long; 3 or 4 anterior to each coxa, each 10–23 µm long; and in medial and submedial longitudinal rows on abdominal segments II–VII, each 10–23 µm long; and (ii) tack-like setae, each 8 µm long, in a submarginal row on abdominal segments II–VII. Microspines, each 2 µm long, in groups of 1–5 in transverse rows over entire body.

Comments. The third-instar female of K. greeni is very similar to those of K. nahalali and K. spatulatus . All three species share (characters for K. echinatus , K. hermonensis and N. balachowskii in brackets): (i) short and spinose marginal setae (conical) (ii) fully-developed legs (leg segmentation reduced); (iii) anal ring dorsal (ventral); and (iv) sclerotized anal lobes on dorsum (absent). The third-instar female of K. greeni is distinguished from K. nahalali and K. spatulatus by the absence of seta-pore clusters on the dorsum and the presence of more than three setae on each anal lobe. Kermes vermilio ( Pellizzari et al., 2012) , a Palaearctic species for which the third-instar female has been described, differs from K. greeni in having (characters of K. vermilio in brackets) sixsegmented antennae (one-segmented antennae) and fully-developed legs (one-segmented). Among the Nearctic Kermes , the third-instar of only K. sylvestris has been described and illustrated ( Bullington & Kosztarab, 1985) and appears to be similar to those of K. greeni in having a ventral submarginal band of tubular ducts, and sparse quinquelocular pores on body margin and venter but K. greeni has two types of ventral tubular ducts, whereas K. sylvestris venter has only one type; also K. greeni has fully-developed legs (total length 220–250 Μm) compared to the much shorter legs (124–160 Μm long) on K. sylvestris .

Second-instar female ( Fig. 11; Plate 2 , fig. a). General appearance Body oval and flat, 0.6–1.3 mm long, 0.3–0.9 mm wide; dorsum and venter bright orange, body flat with fringe of narrow whitish marginal wax filaments and dorsum covered in a thin layer of white wax.

Mounted specimen. Body oval, 0.6–1.5 mm long and 0.4–1.0 mm wide.

Margin. Marginal setae robust and spinose of two sizes: larger each about 60 µm long, in a single complete line of 17–24 on each side; and smaller setae, in a line of 11–16 on each side, lying between larger marginal setae, each about 25 µm long.

Dorsum. Setae tack-like, each about 3 µm long, in submedian longitudinal rows of about 10 setae. Microtubular ducts with a circular opening about 3 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, each 18 µm wide, laterad to each scape near body margin. Antennae 6 segmented, length 108–130 µm; segment III longer than other segments; scape with 2 setose setae; pedicel with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta; apical segment with 3 fleshy setae and 3 setose setae. Legs well-developed; measurements (in Μm) of metathoracic legs: coxa 40–45, trochanter + femur 87–100, tibia 45–50, tarsus 70–75, claw, 20–25; total length 263–295 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 32–38 Μm long, knobbed apically, extending beyond claw apex; claw digitules knobbed apically, each 17–23 Μm long, shorter than tarsal digitules; claws with a small denticle. Tentorium 105–115 µm long, 105–110 µm wide. Labium 3 segmented, triangular, 85–98 µm long, 40–63 µm wide; all setae setose as follows: basal segment with 2 pairs about 8 µm long, median segment with 1 pair about 15 µm long and apical segment with 4 pairs each about 18 µm long. Spiracles subequal in size, each peritreme 8–10 Μm wide; each sclerosis 20–28 µm long. Tubular ducts of Type 2, with inner ductule 6–10 µm long, inner cup 5 µm wide and outer ductule 9–17 µm long, arranged in a complete submarginal band about 1 or 2 ducts wide. Quinquelocular pores, with 5 pores anterolateral to each metathoracic spiracle and 1 anterolateral to each mesothoracic spiracle, each 3 µm diameter; also 1 pore medial to each coxa, 5 µm diameter; and 2 pores submedially on each abdominal segment, each 5 µm diameter. Ventral setae of three types: (i) robust spinose setae similar to marginal setae; with 2 pairs anterior to each scape, each about 60 µm long; and 3 shorter setae, similar to submarginal setae, anterior to each scape, each about 25 µm long; (ii) setose setae: with 6 medially just anterior to tentorium, each about 48 µm long; with 2 setae medial to each coxa, one about 20 µm long and other about 10 µm long; and in medial and submedial lines on abdominal segments II–VII, each medial seta about 30 µm long and each submedial seta about 10 µm long. Also, 1 pair just anterior to anal ring about 53 µm long, plus a pair posterior to anal ring about 10 µm long; and (iii) tack-like setae, each about 5 µm long, submarginally on abdominal segments II–VII. Microspines, each about 2 µm long, in groups of 2–5 in 5–6 transverse rows on each abdominal segment and scattered on thorax. Anal ring ventral, diameter 37–40 µm, circular with anterior gap, each half with 3 spinose setae 65–68 µm long, and about 20–25 pores. Anal lobes very well-developed; each inner margin with a spinose seta about 20 µm long and, on apex of each lobe, a spinose seta about 25 µm long and a flagellate seta 145– 162 µm long.

Comments. Among the Palaearctic species, in addition to those described here, second-instar females are known for K. bytinskii ( Sternlicht, 1969) , K. williamsi ( Sternlicht, 1970) , K. quercus ( Podsiadlo, 2012) and K. vermilio ( Pellizzari et al., 2012) . The second-instar female of K. greeni most closely resembles those of K. nahalali in having robust spinose marginal setae and well-defined anal lobes, but differs in that K. greeni has four longitudinal lines of quinquelocular pores ventrally on the abdominal segments, while K. nahalali has only two lines of pores. This instar of K. greeni also lacks the marginal spatulate setae found in K. spatulatus . Also the second-instar female of K. greeni lack the membranous frontal lobes found in K. vermilio . Tubular ducts are present on both the venter and dorsum in K. echinatus and K. hermonensis , but are found only on the venter in K. greeni . Kermes greeni also has six-segmented antennae and well-developed legs, as on the Nearctic species K. cockerelli , K. concinnulus , K. rimarum ( Baer & Kosztarab, 1985). The second-instar female K. greeni differ from these Nearctic species in the number of quinquelocular pores on the venter: two on head; one medial to each coxa, and two on each abdominal segment II–VII on venter ( K. cockerelli has18; four to six; four; K. concinnulus has 30; six; six, and K. rimarum has 14; two to three; four respectively).

First-instar nymph ( Fig. 12). General appearance. Body oval, tapering posteriorly, 0.5–0.52 mm long, 0.21–0.23 mm wide. Dorsum and venter orange.

Mounted specimen. Body oval, 0.5–0.55 mm long, 0.2–0.25 mm wide. Margin. Marginal setae short and spinose, each about 5 Μm long, in a single row of 6–10 on each side, not including anal lobe setae, extending from about prothorax to anal lobes, with anterior-most pair distinctly longest, 7–8 Μm long.

Dorsum. Derm membranous, with inter-segmental lines. Setae of 2 types: (i) spatulate setae, each 7–8 Μm long and 4–5 Μm wide, in submarginal and submedian rows: submarginal rows with 18–20 on each side, extending along entire margin, plus 1 slightly larger seta on each anal lobe, 9–10 Μm long and 4–5 Μm wide; and each submedian row with 6 or 7 setae extending from head to abdominal segment I inclusive; and (ii) setose setae in submedian rows on abdominal segments II–VII, each 2–3 Μm long. Microtubular ducts with a circular opening about 1 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; with about 24 in submarginal longitudinal rows on abdomen.

Venter. Eyes each 10–12 Μm wide, close to margin. Antennae 6 segmented, length 100–105 Μm; segment III longest; scape and pedicel each with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta, and apical segment with 2 fleshy setae and 6 setose setae. Legs well-developed; measurements (in Μm) of metathoracic leg: coxa 17–20; trochanter + femur 57–70; tibia 28–35; tarsus 45–50 and claw 15–20; total length 175–192 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 27–31 Μm long, knobbed apically; claw digitules knobbed apically, each 20–23 Μm long, extending slightly beyond apex of claw; each claw with a denticle. Tentorium 73–76 Μm long, 63–75 Μm wide. Labium 3 segmented, triangular, 77– 80 Μm long, 40–50 Μm wide; basal segment with 2 pairs of setae, each 7–8 Μm long; median segment with 1 pair each 8–10 Μm long, and apical segment with 4 pairs, each 11–18 Μm long. Spiracles subequal in size, each peritreme 2 or 3 Μm wide, each crescent-shaped sclerosis 11–13 Μm long; each with loculate pores anterolaterally: prothoracic spiracle with 2 pores, 1 quinquelocular and other with 6 loculi, both 3–4 Μm diameter; and each mesothoracic spiracle with 1 quinquelocular pore, 3–4 Μm diameter. Also with 2 slightly larger quinquelocular pores just anterior to tentorium, each 5–6 Μm wide, and 1 pore medially to each coxa, 4–5 Μm wide. Trilocular pores, each 4–5 Μm wide, present submedially on abdominal segments III–V. Ventral setae of two types: (i) spatulate setae: 2 setae on anterior apex of body, similar to those on dorsum but smaller, each 3–6 Μm long, 2–3 Μm wide; and (ii) setose setae: with 3 pairs medially between antennae and tentorium, each 15-20 Μm long; 1 seta medial to each coxa, 12–15 Μm long,; and in medial, submedial and submarginal lines on abdominal segments II– VII, 1 seta per line per segment; each medial and submedial seta 5–8 Μm long, and each submarginal setae, 2–3 Μm long. Also a pair anterior to anal ring 10–12 Μm long, plus a pair posterior to anal ring, each 20–23 Μm long. Bilocular pores, oval, each 3 Μm long, 2 Μm wide, with 1 pore between margin and each spiracle. Microspines, each about 1 Μm long, arranged in 2–4 transverse rows medially and submedially on each abdominal segment. Anal ring ventral, diameter 18–20 Μm, composed of 2 semi-circles, each half with 4 or 5 pores and 3 spinose setae 12–15 Μm long. Anal lobes slightly developed; each lobe with 4 setae: a spatulate seta dorsally 9–10 Μm long and 4–5 Μm wide, and apical margin with 2 thick setae, each 8–12 Μm long, plus and 1 long, setose apical seta 130–175 Μm long.

Comments. Kermes greeni was described from post-reproductive females by Bodenheimer (1931) and K. palestiniensis was described based on first-instar nymphs from Israel by Balachowsky (1953). Kermes palestiniensis was recently synonymized with K. greeni Bodenheimer ( Spodek et al., 2012a). The present redescription includes several features not described by Balachowsky (1953), i.e. presence of: (i) dorsal microtubular ducts; (ii) setae located dorso-medially on abdominal segments; (iii) a six-loculate pore associated with each metathoracic spiracle; (iv) a claw denticle; (v) bilocular pores submarginally level with each spiracle, and (vi) anal ring open at both posterior and anterior ends ( Spodek et al., 2012a).

Kermes greeni has spatulate marginal and dorsal setae as on the first-instar nymphs of the Palaearctic K. bacciformis ( Leonardi, 1920) , K. mutsurensis ( Kuwana, 1931) , K. nakagawae ( Kuwana, 1931) , K. nigronotatus ( Hu, 1986) , K. orientalis ( Liu & Shi, 1997) , K. quercus ( Linnaeus, 1758) , K. taishanensis ( Hu, 1986) , K. vastus ( Kuwana, 1931) , K. viridis ( Borchsenius, 1960) and K. williamsi ( Sternlicht, 1970) . Of the species from Israel, K. spatulatus also has spatulate setae, but K. greeni can be distinguished by the presence of two loculate pores associated with each prothoracic spiracle, one quinquelocular and the other with six loculi, whereas K. spatulatus only has one quinquelocular pore. In addition, K. greeni has two longitudinal rows of dorsal microtubular ducts whereas K spatulatus has four rows.

The first-instar nymph of K. quercus ( Balachowsky, 1950; Borchsenius, 1960; Podsiadlo, 2005), a Mediterranean and European species not present in Israel, has similar morphological traits to K. greeni . Both species possess dorsal spatulate setae (termed tubercle-like setae by Podsiadlo, 2005). However, there are several differences (characters of K. quercus in brackets): (i) absence of spinose setae dorsally on head, thorax and abdomen (present); (ii) trilocular pores present ventrally on abdomen segments (no pores); (iii) two pores associated with each prothoracic spiracle, one with five loculi and one with six-loculi (also has two pores, but one with either three, four or five-loculi and the other with seven-loculi) and (iv) quinquelocular pores on venter of head (absent).

Adult male ( Fig. 13; Plate 2 , fig. h). As for K. echinatus except for: General appearance. Newly emerged males orange, body 1.2–1.5 mm long; wing 1.2–1.5 mm long, 0.3–0.5 mm wide; caudal filaments 1.0– 1.5 mm long.

Mounted specimen. Total length 1.2–1.5 mm long, width at mesothorax 0.3–0.5 mm.

Head. Length 225–250 µm; width across genae 225–250 µm. Dorsal (dse) and ventral simple eyes (vse) each 35–40 µm diameter; lateral simple eyes (lse) smaller, each 20–30 µm diameter; ocellus 15–20 µm diameter. Dorsal head setae (dhs) in rows of 8–10 hs setae. Ventral head setae (vhs) in a row totalling 8–12 hs. Antennae: length 745–780 µm (ratio of total body length to antennal length about 2:1). Scape (scp) 50–60 µm long, 45–50 µm wide, with 2 or 3 hs. Pedicel (ped) 55–65 µm long, 35–45 µm wide, with 4–6 hs. Flagellar segments III–X each 45–160 µm long, 20–25 µm wide, segment III longest, with segments becoming shorter towards apex; approximate number of setae per segment: III–VIII: 8–15 fs; IX 7 or 8 fs + 1 ab; X 7 or 8 fs + 4 ab + 4 cs, each cs 50–55 µm long.

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) 60–65 µm long, 150–165 µm wide. Scutum (sct): membranous area medially quadrate, 70–75 µm long; scutum with 2 hs on each side lateral to membranous area, each 12–15 µm long. Prealare (pra) and triangular plate (tp) present. Scutellum (scl) 68–100 µm long, 150–165 µm wide; with 1 pair hs, each 12– 15 µm long. Basisternum (stn2) 158–175 µm long, 230–270 µm wide; basisternal setae absent. Tegula (teg) with 3 hair-like tegular setae (tegs). Metathorax: Metathoracic spiracles (sp3): peritreme 30–35 µm wide; without loculate pores. Metasternum (stn3) with a total of 20–25 hs setae: 7–10 anterior metasternal setae (amss); 10 or 11 posterior metasternal setae (pmss) and 3 or 4 postmetaspiracular (eps3s), each 20–30 µm long.

Wings: 1100–1200 µm long, 450–500 µm wide (ratio of length to width 1:0.4; ratio of total body length to wing length 1:0.9); Hamulohalteres (h) 15–20 µm wide, 100–110 µm long: hamulus 30–45 µm long. Legs: metathoracic legs longest, 724–847 µm long; prothoracic and mesothoracic legs 665–763 µm long. Coxa: I: 75–80; II: 75–80; III: 80–100 µm long, with 8–12. Trochanter: I 45 –50; II 40 –53; III 50 –55 µm long, with 4 or 5 hs; each trochanter with 3 oval sensory pores in a triangle on each side. Femur: I 200–250; II 185–200; III 200–220 µm long, with 20–25 hs. Tibia: I 240–260; II 250–260; III 250–265 µm long, with 32–38 hs; all tibia with 2 apical spurs (tibs) 20–25 µm long. Tarsi 2-segmented; tarsus 2 (tar2): I 70 –75; II 75 –80; III 75 –85 µm long, with 13–15 hs; tarsal digitules (tdgt) about 35 µm long. Claw (cl): I 25–28; II 25–26; III 25–30 µm long, with denticle; claw digitules (cdt) about 20 µm long.

Abdomen. Tergites and pleural areas unsclerotized; sternites (as) I–VII lightly sclerotized, segment VIII strongly sclerotized. Abdominal dorsal setae (ads) (totals): segments I–VII: 2 hs, 10 µm long; VIII: 6 hs, 40 µm long; pleural setae: dorsopleural setae (adps) (on each side): I–VII: 3 hs, 20–25 µm long, VIII: 3 hs: one 75–80 µm long and two 25–30 µm long; ventropleural setae (avps) (on each side) IV–VII: 1 hs, 15 µm long; abdominal ventral setae (avs) (totals) II–IV: 10–16 hs, V–VIII: 6–8 hs, 20–25 µm long. Glandular pouch setae (gls) 142–150 µm long; each pouch with 8–10 pores, each 3 µm diameter with 3–5 loculi. Genital segment: penial sheath (ps) 160–180 µm long, greatest width 90–110 µm. Basal rod (bra) 40–50 µm long. Aedeagus (aed) 100–125 µm long. Genital capsule with 8 hs (gts), on venter; setae absent from dorsum.

Comments. The adult male of K. greeni is very similar to those of K. nahalali and K. spatulatus , but can be distinguished by the presence of four dorsopleural setae on abdominal segment I ( K. spatulatus has eight and K. nahalali has one). These three species and N. balachowskii have metasternal setae whereas K. echinatus , K. hermonensis and the Nearctic species, E. gillettei ( Miller & Miller, 1993) , lack metasternal setae.

Fourth-instar male (pupa) ( Fig. 14; Plate 2 , fig. g). Develops within a white, flocculent (wooly) wax test secreted by second-instar male; very similar to that of K. echinatus but differing as follows. General appearance. Body 1.0– 1.5 mm long, 0.3–0.5 mm wide; dorsum and venter orange.

Mounted specimen. Body 1.3–1.5 mm long, 0.5–0.6 mm wide; head, 225–235 µm long, 425–450 µm wide.

Margin. Wing-buds: fore-wings 500–550 µm long, 175–185 µm wide; hind-wings: length ratio to mesothoracic wings 1:10. Setae as on K echinatus .

Dorsum. Ocular sclerite 50–60 µm long, 150–185 µm wide. Setae fine and blunt, in two rows of 6 on head apex and also in submedial longitudinal rows. Anal lobes with 1 setose seta on apical tip of each lobe, 55–95 Μm long. Median penial lobe triangular, each 90–95 Μm long, 100–105 Μm wide; with 3 pairs of setose setae, each 15– 20 Μm long.

Venter. Ocular sclerite as on dorsum. Antennae length 525–650 µm, 45–55 µm wide; with 4 or 5 fleshy setae on apical segment, each about 5 µm long. Spiracles: each peritreme 20–35 µm long and 15–20 µm wide. Legs measurements (in Μm) of metathoracic leg; coxa 75–90, trochanter + femur 150–1950, tibia 135–160, tarsus 60– 110; total length 465–505 Μm. Ventral setae: 4 pairs of inter-antennal setae; abdominal setae arranged as on K. echinatus . Microspines as on K. echinatus .

Comments. The pupa of K. greeni is orange in life, while K. echinatus and K. hermonensis are red and N. balachowskii , K. nahalali and K. spatulatus are greyish-brown. The pupa of K. greeni lack quinquelocular pores anterior to each spiracle as also found on Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) , but their colour in life has not been recorded. Both K. greeni and K. spatulatus have sclerotized anal lobes, while the other pupae have membranous lobes.

Third-instar male (prepupa) ( Fig. 15; Plate 2 , fig. f). As for K echinatus apart from: General appearance. Body broadly oval, 0.97–1.5 mm long, 0.4–0.7 mm wide; dorsum and venter orange.

Mounted specimen. Body 1.0– 1.7 mm long, 0.4–0.8 mm.

Margin. Anterior wing-buds each 300–450 µm long, 100–150 µm wide. Setae fine and blunt, in marginal and submarginal rows with 14–20 on each side.

Dorsum. Setae fine and blunt, with 4 pairs on head apex; submedial rows as on K. echinatus . Anal lobes rounded and slightly sclerotized; each with 2 setose setae on apex, one seta 60–75 Μm long and other 13–15 Μm long. Median penial lobe 60–65 Μm long, 110–115 Μm wide, with 8 setose setae, each 5–10 Μm long.

Venter. Antennae 7 or 8 segmented, 250–270 µm long, 65–80 µm wide. Spiracles: each peritreme 18–20 µm long, 10–15 µm wide, with sclerosis 20–25 µm long; without quinquelocular pores. Legs: length of metathoracic leg, 225–250 Μm long, 53–60 Μm wide. Ventral setae fine and blunt, 10–20 Μm long, with 4 pairs between scapes; abdominal setae arranged as on K. echinatus . Microspines as on K. echinatus .

Comments. The prepupa of K. greeni is orange in life, unlike K. echinatus and K. hermonensis which are red and N. balachowskii , K. nahalali and K. spatulatus which are greyish-brown. The colour of the two Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) have not been recorded. Like those of K. nahalali , K. spatulatus , A. kingii and E. gillettei , the prepupa of K. greeni lacks quinquelocular pores anterior to each spiracle, whereas one or two pores are present on K. echinatus , K. hermonensis and N. balachowskii .

Second-instar male ( Fig. 16; Plate 2 , fig. e). General appearance. Secretes a white, flocculent (wooly) wax test, 1.5–1.8 mm long, 0.6–0.8 mm wide. Body orange, broadly oval when removed from test, 0.8–1.45 mm long, 0.4–0.8 mm wide; derm membranous with inter-segmental lines on dorsum and venter.

Mounted specimen. Body oval, 0.9–1.5 mm long, 0.5–0.9 mm wide.

Margin. Marginal setae robust and setose, each 50 µm long, in a single complete row of 24–30 on each side.

Dorsum. Microtubular ducts, each with a circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, numerous, scattered between tubular ducts over entire dorsum, less abundant than tubular ducts. Tubular ducts of Type 2, with inner ductule 8–15 µm long, inner cup 5 µm wide and outer ductule 10–15 µm long, abundant throughout dorsum.

Venter. Eyes circular, each 20 µm diameter. Antennae 7 segmented; total length 150–175 µm; segment III 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 3 fleshy setae and 4 setose setae. Frontal lobes absent. Legs well-developed; measurements (in Μm) of metathoracic legs: coxa 35–50; trochanter + femur 88–100; tibia 50–60; tarsus 75–90 and claw 20–30; total leg length 230–315 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, extending beyond apex of claw; each claw with a single denticle. Tentorium 105– 115 µm long, 88–100 µm wide. Labium 3 segmented, triangular, 75–100 µm long and 55–65 µm wide; labial setae all setose, as follows: basal segment with 2 pairs about 15 µm long, median segment with 1 pair 13–20 µm long and apical segment with 4 pairs, each 15–22 µm long. Spiracles subequal in size, each peritreme 10–13 Μm wide and each sclerosis 25–35 µm long; each spiracle with 4 or 5 quinquelocular pores anterolaterally, each 5 µm diameter. Tubular ducts of Type 2, with outer ductule 5–15 Μm long, a sclerotized cup about 5 Μm in diameter and inner ductule 10–15 Μm long, dispersed in a complete submarginal band 1 or 2 ducts wide. Bilocular pores circular, each 2 µm diameter; with 10–12 pores in a single complete submarginal band. Quinquelocular pores, each 5 µm, with 2 pairs among setae anterior to tentorium; and 4 pores in a line across each abdominal segment. Ventral setae of two types: (i) setose setae: with 5 pairs medially just anterior to tentorium, each 60–88 µm long; 2 medial to each coxa, each 25–38 µm long; also a pair medially on abdominal segments II–VII, each about 60 µm long, plus submedial rows of shorter setae, each about 35 µm long; and (ii) tack-like setae, each 10 µm long, in a submarginal row of 10–15 setae extending from antennae to anal ring, and with a pair anterior to anal ring. Microspines, each about 3 µm long, arranged in groups of 3–5 in 4–8 transverse rows on each abdominal segment and on thorax. Anal ring ventral, total diameter 33–40 µm, composed of 2 semi-circles, each half with 3 spinose setae 40–50 µm long, and 8–12 pores. Anal lobes well developed; each inner margin with 2 spinose setae 15–25 µm long; plus 2 flagellate setae on apex, a shorter seta 50–70 µm long, and a longer seta 100–200 µm long.

Comments. Of the species described in Israel, the second-instar male of K. greeni most closely resembles that of K. nahalali because of the presence of distinct anal lobes and spinose marginal setae. The other species in Israel and two from the Palaearctic (i.e. K. echinatus , K. hermonensis , K. quercus ( Podsiadlo, 2012) , K. spatulatus , K. vermilio ( Pellizzari et al., 2012) and N. balachowskii ) appear to lack anal lobes. Kermes greeni can be distinguished also from the other species in Israel by the presence of quinquelocular pores in four longitudinal lines ventrally on abdominal segments; K. echinatus lacks abdominal pores, K. nahalali and N. balachowskii have only two lines, and K. spatulatus has six lines. Kermes hermonensis also has four lines of pores, similar to K. greeni however the marginal setae are conical in K. hermonensis and robust setose in K. greeni . The second-instar males of the Nearctic K. cockerelli , K. concinnulus and K. rimarum also have robust setose marginal setae but have quinquelocular pores scattered on both the dorsum and venter ( Baer & Kosztarab, 1985), whereas K. greeni has quinquelocular pores only on the venter.

Kermes hermonensis Spodek & Ben-Dov sp. nov. ( Figs. 17–24; Plate 3 , figs. a–j)

The type series of this species is restricted to material collected from Mt. Hermon, Israel, off Quercus look. Etymology. The species epithet is derived from Mount Hermon Nature Reserve from where this species was first collected.

Material examined. Type material studied. Most type material is deposited in ICVI, with 1 paratype adult ♀ and 1 slide of first-instar nymphs (non-type) at BMNH, MNHN and USNM depositories. Holotype adult ♀ 20.vi.2012 (C:5180:5). Paratypes 30 adult ♀, 20.vi.2012 (C:5180/2 on each slide/15 slides), 13 third-instar ♀, 24.v.2012 (C:5309/2 slides), 34 third-instar ♀, 20.v.2013 (C:5310/3 slides), 20 second-instar ♀ (C:5307/2 slides), 3 second-instar ♀, 2.v.2013 (C:5308/1 slide), 11 second-instar ♂, 20.v.2013 (C:5311/2 slides), 6 second-instar ♂, 20.vi.2012 (C:5312/1 slide); 14 prepupae, 20.v.2013 (C:5313/2 slides); 10 pupae, 26.v.2013 (C:5314/1 slide); 3 adult ♂, 20.vi.2012 (C:5315/3 slides); 13 first-instars, 21.viii.2011 (C:5305/1 slide), 32 first-instars, 4.xi.2012 (C:5306/2 slides). Other material studied. All non-type material collected off Q. look in Israel by M. Spodek, unless otherwise indicated. Mt. Hermon Nature Reserve, 20.vi.2012 (C:5107/ 44 adult ♀), 20.vi.2012 (MC:712/ 7 adult ♀, MC:744/ 4 adult ♀), 4.xi.2012 (MC:755/ 1 adult ♀); Mt. Hermon Nature Reserve, 24.v.2012 (MC:681/11 third-instar ♀); Mt. Hermon Nature Reserve, 7.v.2012 (MC:664/11 second-instar ♀), 24.v.2012 (MC:682/2 secondinstar ♀), 20.vi.2012 (MC:742/4 second-instar ♀, MC:743/4 second-instar ♀); Mezar, Q. ithaburensis, 4.xi.2012 (MC:756/27 first-instars), 24.v.2012 (MC:680/8 first-instars), 20.vi.2012 (MC:713/23 first-instars, MC:741/5 firstinstars); Mt. Hermon Nature Reserve, 29.v.2013 (MC:830/ 7 adult ♂); Mt. Hermon Nature Reserve, 20.vi.2012 (MC:836/ 2 pupae); Mt. Hermon Nature Reserve, 20.v.2013 (MC:874/14 prepupae); Mezar, Q. ithaburensis, 10.iii.2013 (MC:821/2 second-instar ♂); Mt. Hermon Nature Reserve, 7.v.2012 (MC:665/1 second-instar ♂), 24.v.2012 (MC:681/4 second-instar ♂), 20.vi.2012 (MC:742/10 second-instar ♂), 2.v.2013 (MC:865/8 secondinstar ♂).

Adult female ( Fig. 17). General appearance. Pre-reproductive adult female ( Plate 3 , fig. c). Body oval, 2.5–3.2 mm long, 2.0–3.0 mm wide; dorsum and venter red with a thick layer of wax covering dorsum; body soft and slightly convex. Gravid female and post-reproductive female ( Plate 3 , fig. f). Body highly convex, 3.0–5.0 mm long, 3.0–5.0 mm wide, 3.2–5.0 mm high; with a thick layer of white wax enclosing entire body; body not sclerotized.

Mounted young adult female. Body oval, 2.0–3.0 mm long, 2.0– 2.8 mm wide.

Margin. Marginal setae conical, each 10–13 µm long, 10–11 µm wide at base, arranged in a single complete row of 27–33 on each side. Dorsal submarginal setae spinose, each 7–9 µm long, in a complete single line of 27– 33 setae on each side.

Dorsum. Setae conical, each 10–13 µm long and 7–10 wide at base, similar in shape to marginal setae, randomly distributed on dorsum, total 14–22 setae. Bilocular pores oval, each 3 µm long, 2 µm wide, present throughout dorsum; more abundant than tubular ducts. Tubular ducts of Type 2, with outer ductule 12–17 µm long, inner ductule 10–15 µm long and with a sclerotized cup 8 µm diameter, sparse throughout dorsum.

Venter. Eyes circular, each 20 µm diameter, anterolateral to each scape. Legs reduced to 3 segments, sometimes appearing to be 1 or 2 segmented; 80–140 µm long; claw 18–23 µm long, usually with a denticle but this not always visible; tarsal digitules 20–25 µm long; claw digitules absent. Antennae 3 to 6 segmented, segments III–V often appearing fused; total length 75–100 µm, scape and pedicel each with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta; apical segment with 4 setose setae and 2 fleshy setae; each scape surrounded by a group of 40–45 multilocular pores, each pore 7–8 µm diameter with 10 loculi. Tentorium 250–275 µm long, 220–250 µm wide. Labium 3 segmented, triangular, 170–180 µm long, 120–135 µm wide; labial setae as follows: all setae setose, basal segment with 2 pairs 8–10 µm long; medial segment with 1 pair 15–20 µm long; apical segment with 4 pairs 10–15 µm long. Spiracles subequal in size; each peritreme 75–90 µm long, 55–60 µm wide, each sclerosis 60–65 µm long; loculate pores with 3, 7 and 10 loculi, each 8 µm wide, in a group of 55–67 surrounding each spiracle. Tubular ducts of Type 2, with outer ductule 10–16 µm long, inner cup 4 or 5 µm wide and inner ductule 11–15 µm long, present in a single, dense submarginal band 15–20 ducts wide and also sparsely throughout rest of venter. Bilocular pores, each 3 µm long and 2 µm wide, sparse, interspersed between tubular ducts in submarginal band. Multilocular pores, each 10 µm diameter with 10–12 loculi, arranged in 2–4 transverse rows of 114–120 pores on each abdominal segment; also with a group of 52–56 pores just posterior to vulva. Microtubular ducts with circular opening about 4 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, interspersed between multilocular pores on abdominal segments; total pores on each segment 110–150. Ventral setae of two types: (i) setose setae: 4 pairs of setae just anterior to tentorium, each 35–50 µm long; about 10 setae in a line from antennae to anal ring mesad to each submarginal band of tubular ducts, each 15–20 µm long; with 6–10 setae in a transverse row on each abdominal segment, each 13–30 µm long. Also with a pair just anterior to anal ring, each 10–12 µm long, and 3–4 pairs posterior to anal ring, each 10–20 µm long; and (ii) conical setae: a pair similar in shape to marginal setae and 10–12 µm long and 10 µm wide, close to posterior margin. Microspines, each 1 or 2 µm long, in groups of 3–5 in 3–8 rows on each abdominal segment and in rows on thorax. Anal ring ventral, forming a complete sclerotized circle 60–65 µm diameter, with 6–8 pores and 3 pairs of setae, each 50–75 µm long.

Comments. Even though the derm of K. hermonensis produces a thick, waxy ovisac that covers its' membranous body during oviposition ( Plate 6 , figs. e–f), we assign K. hermonensis sp. n. to the family Kermesidae because its microscopic characters match other species of this family. The derm of most post-reproductive female Kermesidae is sclerotized and lacks a waxy ovisac but two species of Nidularia , N. japonica and N. pulvinata , produce a waxy ovisac posterior to female's body.

Adult female K. hermonensis share several traits with K. echinatus and K. vermilio (a Mediterranean species, not found in Israel): (i) legs reduced or absent; (ii) same type of tubular duct on both venter and dorsum; (iii) setapore clusters absent from dorsum; (iv) anal ring ventral; (v) conical setae on both margin and dorsum; and (vi) presence of multilocular pores surrounding each scape. These species differ in the structure of the anal ring: K. echinatus has three pairs of setae without pores; K. hermonensis has three pairs of setae with pores, while K. vermilio has pores without setae. Kermes hermonensis also differs from K. echinatus and K. vermilio in the number of antennae and leg segments: K. hermonensis has three, four, five or six-segmented antennae and three-segmented legs, while the two other species have one-segmented antennae and no legs. Of the Nearctic species, K. hermonensis can be distinguished from Allokermes spp. by the absence of pit or tooth-shaped pores on middorsum. Nanokermes species, also from North America, have two types of tubular ducts while K. hermonensis only has one type. Olliffiella species have eight-shaped pores on mid-dorsum ( Bullington & Kosztarab, 1985), absent in K. hermonensis .

Third-instar female ( Fig. 18; Plate 3 , fig. b). General appearance. Body oval to round, slightly convex, 1.3– 1.8 mm long, 1.0– 1.2 mm wide; dorsum red with tufts of white wax and venter red.

Mounted specimen. Body oval, 1.4–2.0 mm long. 1.0– 1.5 mm wide.

Margin. Marginal setae conical, each 15 µm long. 5 µm wide at the base, in a single complete line of 37–40 on each side. Dorsal submarginal setae short and spinose, each 5 µm long, 2 µm wide, in a single complete line of 20–25 on each side.

Dorsum. Setae conical, each 10 µm long, 5 µm wide, distinctly narrower than marginal conical setae, in longitudinal medial and submedial rows, each row with 9–11 setae. Tubular ducts of Type 2, inner ductule 20–25 µm long, inner cup about 5 µm wide and outer ductule 20–25 µm long, scattered on dorsum. Bilocular pores and microducts absent.

Venter. Eyes circular, each 2 µm diameter, lateral to each scape. Antennae 95–100 µm long, segmentation indistinct, probably with six segments; scape pronounced and wide at base; scape and pedicel each with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta; apical segment with 4 setose setae and 4 fleshy setae. Leg segmentation indistinct, probably reduced to 3 segments; total length 110–125 µm; tarsal digitules each 80–85 Μm long, knobbed apically, extending beyond the apex of the claw; claw distinct, 25 µm long; claw denticle not visible; claw digitules, knobbed apically, each 35–40 Μm long, shorter than tarsal digitules. Tentorium 175–200 µm long, 150–175 µm wide. Labium 3 segmented, triangular, 160–175 µm long, 100–110 µm wide; all setae setose, as follows: basal segment with 2 pairs about 3 µm long, median segment with 1 pair about 30 µm long, and apical segment with 4 pairs, each about 15 µm long. Spiracles subequal in size; each peritreme 40 Μm long, 25 Μm wide; each sclerosis 38–50 µm long; each spiracle with 2 quinquelocular pores, 5 µm diameter, placed anterolaterally. Tubular ducts of Type 2, inner ductule 20–25 µm long, inner cup about 5 µm wide and outer ductule 20–25 µm long, arranged in a dense complete submarginal band 8–10 ducts wide; ducts also scattered over entire venter. Bilocular pores oval to circular, each 3 Μm wide, dispersed among the submarginal tubular ducts from head apex to anal ring. Ventral setae of three types: (i) setose setae: with 2 pairs anterior to tentorium, each 13 µm long; and 6–8 setae in transverse row across each abdominal segment, each about 25 µm long; also with 2 pairs just anterior to anal ring, each 13 µm long; (ii) conical setae: with 2 pairs of short setae posterior to anal ring, each 15 µm long, 5 µm wide, plus 2 longer setae, each 95 µm long; also 2 setae on most posterior apex of body, each 10–15 µm long, 10 µm wide; and (iii) tack-like setae, each 5 µm long, with about 10 setae in a complete submarginal row. Microspines, each 2 µm long, in groups of 1–8 present over entire body. Anal ring ventral, circular, 50 µm diameter, with a posterior gap; each half circle with 2 or 3 pores and 3 spinose setae about 40 µm long.

Comments. The third-instar female of K. hermonensis is very similar to those of K. echinatus and K. vermilio (a species not found in Israel) (characters for the other Israeli species, K. greeni , K. nahalali and K. spatulatus , in brackets): (i) marginal setae conical (non-conical) (ii) leg segmentation reduced (legs fully developed); (iii) anal ring ventral (dorsal) and (iv) sclerotized anal lobes absent (present). The third-instar female K. hermonensis are distinguished from K. echinatus in having (characters for K. echinatus in brackets): five or six-segmented antennae (one-segmented); and three-segmented legs with a claw (one segmented, without a claw); it can be distinguished from K. vermilio in having (characters for K. vermilio in brackets): four-segmented antennae (three-segmented) and dorsal tubular ducts present (absent).

Second-instar female ( Fig. 19; Plate 3 , fig. a). General appearance. Body oval, tapering posteriorly, 0.7– 1.45 mm long, 0.4–0.8 mm wide; dorsum and venter pink to reddish; dorsal surface covered with a thin layer of wax, composed of 5 longitudinal rows of rectangular wax plates, each plate about 0.2 mm long, 0.1 mm wide; median row with 10 plates, submedial rows with 9–11 plates, and marginal rows with 7–12 plates. Wax plates largest medially. Dorsum also covered with thin, stringy, wax filaments.

Mounted specimen. Body oval, 0.97–1.5 mm long, 0.5–1.0 mm wide.

Margin. Marginal setae conical, each 10–15 µm long, 5 µm wide at base, in a single complete row of 30–33 on each side.

Dorsum. Derm membranous. Setae conical, each about 8 µm long, about 5 µm wide at base, shorter than marginal setae, in submedial longitudinal rows, extending from cephalic apex to mid-abdomen, with 5 setae in each row. Microtubular ducts with a circular opening about 3 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, in 4 longitudinal rows, each submarginal row with about 7 pores, from head to level of anal ring, and each median row with about 6 pores, from level of labium to level of anal ring. Tubular ducts of Type 2, with inner ductule 10–15 µm long, inner cup 5 µm wide and outer ductule 10–15 µm long, sparse throughout dorsum.

Venter. Eyes circular, each 15 µm wide, lateral to each scape. Antennae 3 to 6 segmented, several segments often appearing fused; total length 75–110 µm; scape and pedicel each with 2 setose setae, segment III with 1 setose seta, IV with 1 fleshy seta, V with 1 fleshy seta and 1 setose seta, and V1 with 2 fleshy setae and 4 setose setae. Legs reduced to 3 segments, coxa and trochanter plus femur, tibia and tarsus fused; legs subequal in size, each 78–100 µm long; claw 10 µm long; claw denticle absent; tarsal digitules about 20 µm long, knobbed apically and extending beyond apex of claw; claw digitules knobbed apically, each 15–20 µm long, shorter than tarsal digitules. Tentorium 125–150 µm long, 113–125 µm wide. Labium 3 segmented, triangular, 125–138 µm long, 63–75 µm wide; labial setae setose: basal segment with 2 pairs about 10 µm long, median segment with 1 pair about 15 µm long, and apical segment with 4 pairs 15 µm long. Spiracles subequal in size, each peritreme 18–20 Μm long, 13–15 Μm wide, each sclerosis 25–33 µm long; each spiracle with 2 quinquelocular pores, 5 µm diameter, anterolaterally. Tubular ducts of Type 2, with inner ductule 10–15 µm long, inner cup 5 µm wide and outer ductule 10–15 µm long, arranged in a complete submarginal band 3 or 4 ducts wide and dispersed singly over entire venter. Bilocular pores circular, each 2 µm long and 2 µm wide, dispersed among tubular ducts in submarginal band. Ventral setae of three types: (i) conical setae, each 15 µm long and 5 µm wide at base, similar to marginal setae, with 2 on anterior apex of body; (ii) setose setae: 3 pairs medially just anterior to tentorium, each 25–38 µm long; also 4 setae on abdominal segments II–VIII, medial setae about 15 µm long, and submedial setae about 10 µm long. Also with 2 pairs anterior to anal ring, each about 5 µm long; 2 setae posterior to anal ring, each about 20 µm long, and 2 long setae, each 30–75 µm long and 2 conical setae, similar to marginal setae, about 13 µm long, 5 µm wide at base, on posterior apex of body; and (iii) tack-like setae, in a submarginal row from abdominal segment III to anal ring, each 5 µm long, totaling 6 setae on each side. Microspines, each about 3 µm long, arranged in groups of 1 or 2 in 4 or 5 transverse rows on each abdominal segment and scattered on thorax. Anal ring ventral, diameter 27–35 µm, composed of 2 semi-circles, each half with 3 spinose setae, each 20–30 µm long, and 4–7 pores.

Comments. Among the Palaearctic species, the only other described second-instar females are of K. bytinskii ( Sternlicht, 1969) , K. williamsi ( Sternlicht, 1970) , K. quercus ( Podsiadlo, 2012) and K. vermilio ( Pellizzari et al., 2012) . Those of K. hermonensis most closely resembles K. echinatus and K. vermilio (not found in Israel) in having conical marginal setae and reduced antennal and leg segments, but can be distinguished from K. echinatus in having (characters for K. echinatus in brackets): five or six-segmented antennae (one-segmented), and threesegmented legs (one-segmented); and from K. vermilio in lacking membranous frontal lobes between the scapes. Also, K. echinatus and K. hermonensis have tubular ducts (Type 2) present on both the venter and dorsum in whereas they are only on the venter in K. vermilio . Kermes echinatus also has two longitudinal rows of dorsal conical setae whereas K. vermilio has a transverse row of dorsal setae. Other Kermes spp. in Israel have differentlyshaped marginal setae: K. nahalali and K. greeni (robust setose), and K. spatulatus (spatulate). These latter three species also have six-segmented antennae and well-developed legs, as on the Nearctic A. galliformis , K. cockerelli , K. concinnulus , K. rimarum ( Baer & Kosztarab, 1985) and E. gillettei ( Miller & Miller, 1993) . The second-instar female of K. hermonensis is distinct from E. gillettei in having a ventral anal ring (dorsal in E. gillettei ) and in having tubular ducts on both dorsum and venter (absent in E. gillettei ); and it differs from A. galliformis , K. cockerelli , K. concinnulus and K. rimarum in lacking quinquelocular pores on abdominal segments (present).

First-instar nymph ( Fig. 20). General appearance. Body oval, tapering posteriorly, 0.5–0.7 mm long, 0.2– 0.3 mm wide; dorsum and venter red.

Mounted specimen. Body oval, 0.6–0.8 mm long, 0.2–0.3 mm wide.

Margin. Marginal setae conical and slightly curved apically, in 1 to 2 rows; 1 row complete, with 21–24 on each side, each setae 10–13 Μm long, 5 Μm wide at base; second row with smaller conical setae, each setae 7–10 Μm long, 3–5 Μm wide at base, with 12–14 on each side, located between larger setae and extending from mesothorax to anal lobe.

Dorsum. Derm membranous with inter-segmental lines. Setae setose, with 4 pairs in each submedian line on head and thorax, each 5–7 Μm long. Microtubular ducts with a circular opening about 1 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; in submarginal longitudinal rows on abdomen and thorax, each row with 8 pores.

Venter. Eyes circular, each 10–12 Μm wide, close to body margin. Antennae 6 segmented, length 113–130 Μm; segments III and VI longest; setal distribution as follows: scape and pedicel each with 2 setose setae; segment III with 1 setose seta; IV with 1 fleshy seta; V with 1 setose seta and 1 fleshy seta; apical segment with 2 fleshy setae and 4 setose setae. Legs well-developed; measurements (in Μm) of metathoracic leg: coxa 25–38, trochanter + femur 80–88, tibia 45–50, tarsus 50–63, claw 22–25; total leg length 190–200 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 40–45 Μm long and knobbed apically, extending beyond apex of claw; claw digitules knobbed apically, each 18–20 Μm long; each claw with a single denticle near tip. Tentorium 83–88 Μm long, 86–88 Μm wide. Labium 3 segmented, triangular, 85–90 Μm long, 50–63 Μm wide; labial setae setose as follows: basal segment with 2 pairs 5–8 Μm long, median segment with 1 pair on dorsal surface, each 12–15 Μm long, and apical segment with 4 pairs, each 16–20 Μm long. Spiracles subequal in size, each peritreme 3–5 Μm wide; each with apodeme crescent shaped, 20–25 Μm long; each prothoracic spiracle with 2 quinquelocular pores anterolaterally, one 3 Μm diameter and one 5 Μm diameter, each mesothoracic spiracle with 1 quinquelocular pore anterolaterally, 3 Μm diameter. Trilocular pores, each about 5 Μm wide, distributed as follows: 1 pore medial to each scape, just anterior to tentorium; 1 mesad to each coxa; and pairs submedially on abdominal segments V–VI. Bilocular pores oval, each 3 Μm long and 2 Μm wide, present near body margin from lateral to prothoracic spiracle to abdominal segment VI, with 5 on each side. Ventral setae of two types: (i) conical setae: a pair on anterior apex of head, each 11–13 Μm long, 5 Μm wide at base; and (ii) setose setae:, with 3 pairs in longitudinal lines medially between scapes, each 25–30 Μm long; also a seta mesad to each coxa, associated with trilocular pores, each 10–13 Μm long; and with 6 setae across each abdominal segment, medial and submedial setae each 10–18 Μm long, and submarginal setae 5–6 Μm long. Also with a pair anterior to anal ring 18–23 Μm long, and a pair latero-posterior to anal ring 20–25 Μm long. Microspines, each about 3 Μm long, arranged in groups of 3 or 4 in 2 transverse rows on each abdominal segment and sparsely on thorax. Anal ring ventral, diameter 20–25 Μm; composed of 2 semicircles, each half with 3 spinose setae, each 13–18 Μm long; pores absent. Anal lobes small; apical margin of each with 1 spinose seta, 10–15 Μm long, 3–5 Μm wide, and 1 very long flagellate seta, 220–275 Μm long.

Comments. The first-instar nymph of K. hermonensis is most similar to those of K. echinatus and K. vermilio . All three species are red in life and possess dorsal and marginal conical setae. However, K. hermonensis has two quinquelocular pores associated with the prothoracic spiracles, usually with one pore larger in diameter than the other, whereas K. echinatus has only one quinquelocular pore associated with each spiracle and K. vermilio has usually one pore, rarely two, but each with 5–8 loculi ( Balachowsky, 1950; Pellizzari et al., 2012). The first-instar of K. hermonensis also lacks the dorsal bilocular pores dorsum, present on K. vermilio .

Adult male ( Fig. 21; Plate 3 , fig. h). As for K echinatus except: General appearance. Body 1.46–1.8 mm long, widest across mesothorax 0.44–0.45; wings about 2/3rds body length; radial wing vein red; white waxy caudal filaments 1.0– 1.3 mm long.

Mounted specimen. Total length 1.5–2.0 mm long, width across mesothorax 0.5–0.6 mm.

Head. Length 225–275 µm long; width across genae 260–275 µm. Dorsal (dse) and ventral simple eyes (vse), each 45–50 µm diameter; lateral simple eyes smaller (lse), each 35 µm diameter; ocellus 18–20 µm diameter. Dorsal head setae (dhs) in a row of 6-8 hs setae. Ventral head setae (vhs) in a transverse row of 6–8 hs. Antennae: 1150–1200 µm long (ratio of total body length to antennal length about 1:0.65). Scape (scp) 50–62 µm long, 55–63 µm wide, with 2 or 3 hs. Pedicel (ped) 50–63 µm long, 40–50 µm wide, with 5–8 hs. Flagellar segments III–X each 125–163 µm long, 20–25 µm wide; segment III longest, segments becoming shorter towards apex; approximate number of setae per segment (note: fs and hs hard to differentiate): III–V 3–8 fs; VI– VIII 12–19 fs; IX 13–15 fs + 1 ab; X 9–11 fs+ 4 ab + 4 cs, each cs 63–65 µm.

Thorax. Prothorax: as for K. echinatus .

Mesothorax: prescutum (prsc) 75–80 µm long, 160–175 µm wide. Scutum (sct): membranous area medially 50–70 µm long. Prescutal and scutal setae absent. Scutellum (scl) 80–85 µm long, 165–175 µm wide; with 2 pairs of hs (scls), each 13–15 µm long. Basisternum (stn2) 175–200 µm long, 250–270 µm wide; basisternal setae absent. Tegula (teg) with 3–5 hs (tegs). Mesothoracic spiracles (sp2): peritreme 15–20 µm wide; without loculate pores.

Metathorax: Metathoracic spiracles (sp3): peritreme 15–20 µm wide; without loculate pores. Metasternal setae absent.

Wings: 1200–1250 µm long, 450–475 µm wide (ratio of length to width 1:0.37; ratio of total body length to wing length 1:1.68). Hamulohalteres (h) 113–125 µm long, 32–40 µm wide; hamulus, 55–65 µm long. Legs: metathoracic legs longest, 851–921 µm long; prothoracic and mesothoracic legs 789–827 µm long. Coxa: I 90 – 113; II 100–107; III 115–125 µm long, with 12–20 hs. Trochanter: I 50 –70; II 50 –65; III 75 –80 µm long, with about 5 or 6 hs; each trochanter with 3 oval sensory pores in a triangle on each surface. Femur: I 235–250; II 220– 225; III 240–250 µm long, with 25–37 hs. Tibia: I 275–287; II 275–285; III 270–300 µm long, with 36–45 hs; all tibia with 2 apical spurs (tibs) 25–28 µm long. Tarsi 2-segmented; tarsus 2 (tar2): I 113–115; II 100–105; III 108– 118 µm long, with 20–25 hs; tarsal digitules (tdgt) each about 50 µm long. Claw (cl): I 18–25; II 25–30; III 30–33 µm long; claw digitules (cdt) each about 35 µm long; claw denticle present.

Abdomen. Tergites and pleural areas unsclerotized; sternites (as) lightly sclerotized on segments III–VIII. Abdominal dorsal setae (ads) (totals): segments I–VII 2 hs, 10 µm long; VIII 2 hs, 40 µm long; pleural setae: dorsopleural setae (adps) (on each side): I 3 hs and II–VII 2 hs, 10 µm long, VIII 3 hs, one 80–85 µm long and two 50–55 µm long; ventropleural setae (avps) (on each side) II–VIII 2 or 3 hs, 15 µm long; abdominal ventral setae (avs) (totals) II–III 2 hs, IV–VII 4 hs 20–25 µm long. Glandular pouch setae (gls), 110–117 µm long; each pouch with 9–18 pores, each 3 µm diameter with 3–5 loculi. Genital segment: penial sheath (ps) 180–200 µm long, greatest width 110–120 µm. Basal rod (bra) 60–75 µm long. Aedeagus (aed) 130–140 µm long. Genital capsule with 4 hs (gts) dorsally and 16 hs (gts) ventrally.

Comments. Of the Israeli species, the adult male of K. hermonensis is most similar to that of K. echinatus — both are red in life with reddish wing veins and both lack metasternal setae. Metasternal setae are present on all other known males ( Hamon et al., 1976; Koteja & Zak-Ogaza, 1972; Hu, 1986; Liu et al., 1997).

Fourth-instar male (pupa) ( Fig. 22; Plate 3 , fig. g). Develops within a white, flocculent (wooly) wax test secreted by second-instar male; very similar to that of K. echinatus but differing as follows. General appearance. Body 1.5–1.7 mm long, 0.6–0.7 mm wide.

Mounted specimen. Body 1.5–1.8 mm long, 0.6–0.8 mm wide; head 225–250 µm long, 400–425 µm wide.

Margin. Wing-buds: fore-wings 485–650 µm long, 200–250 µm wide, length ratio to mesothoracic wings 1:10. Setae as on K echinatus .

Dorsum. Eyes absent. O cular sclerite reticulated, 50–75 µm long, 150–158 µm wide. Setae setose, with 4 pairs on head apex, each 20 µm long; also in submedial longitudinal rows. Anal lobes rounded, membranous, each with a long setose seta 75 Μm long on apex. Median penial lobe triangular, 100–130 Μm long, 100–125 Μm wide, with 3 pairs of setose setae, each 25–30 Μm long.

Venter. Ocular sclerite as on dorsum. Antennae 10 segmented; total length 575–675 µm, width 50–65 µm; setae absent. Spiracles subequal in size, each peritreme 20–25 µm long, 10–15 µm wide; each sclerosis 30–40 µm long; each spiracle with 2–4 quinquelocular pores anterolaterally, each 5 µm diameter. Legs well-developed, subequal in size; measurements (in Μm) of metathoracic leg: coxa 100–125, trochanter + femur 200–250, tibia 175–200, tarsus 105–115; total length 580–690 Μm. Ventral setae setose and each 10 Μm long, with 4 pairs of inter-antennal setae; abdominal setae arranged as on K. echinatus . Microspines arranged as on K. echinatus .

Comments. The pupa of K. hermonensis and K. echinatus are both red in life. Those of K. hermonensis differ from those of K. echinatus in having two quinquelocular pores anterior to each spiracle (absent on K. echinatus ). The other Israeli species are: K. greeni orange and N. balachowskii , K. nahalali and K. spatulatus greyish brown. In addition, all other Israeli species lack spiracular pores. The pupae of the Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) also lack spiracular pores; their colours in life were not recorded.

Third-instar male (prepupa) ( Fig. 23; Plate 5 , fig. h). As for K echinatus apart from: General appearance. Body 1.5–1.8 mm long, 0.7–0.8 mm wide.

Mounted specimen. Body 1.5–1.9 mm long, 0.7–0.9 mm wide.

Margin. Anterior wing-buds each 380–435 µm long, 185–200 µm wide; posterior wing-buds absent. Marginal setae fine and blunt setae, each 15–20 µm long, in a single complete marginal row of 15–20 on each side; submarginal setae absent.

Dorsum. Setae setose, each 10–15 µm long, with 3 pairs on head apex and in complete submedial longitudinal rows of about 10 setae. Anal lobes rounded apically and membranous, each with 2 setae on apex, a long setose seta 40–50 Μm long, and a shorter spinose seta 20–25 Μm long. Median penial lobe broadly rounded, 75–80 Μm long, 125–135 Μm wide, with 3 pairs of setose setae, each 20–25 Μm long.

Venter. Eyes absent. Antennae 7 or 8 segmented, tapering apically and 250–300 µm long, 75–80 µm wide; apical segment with 5 fleshy setae, each about 5 µm long. Spiracles subequal in size; each peritreme 20–25 µm long, 20–25 µm wide, with sclerosis 35–40 µm long; each spiracle with 2 quinquelocular pores anterolaterally, each 5 µm diameter. Legs 3 segmented lobes, subequal in size; total length of metathoracic leg, 250–300 Μm, width 70–75 Μm; claw and setae absent. Ventral setae setose, each 10–15 Μm long, with 4 pairs between scapes; abdominal setae arranged as on K. echinatus . Microspines, arranged as on K. echinatus .

Comments. The prepupae of both K. hermonensis and K. echinatus are red in life and both have two quinquelocular pores anterior to each spiracle but K. hermonensis has two pairs of dorsal setae on abdominal segment VIII and three pairs of setae on the penial lobe, whereas K. echinatus has no dorsal abdominal setae on segment VIII and four pairs of setae on the penial lobe. Of the other prepupae in Israel, K. greeni is orange, and K. nahalali and K. spatulatus are greyish-brown and all three have spiraclular pores, whilst that of N. balachowskii is brown and has one quinquelocular pore anterior to each spiracle. The prepupa of the Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) lack spiracular pores; their colour in life was not recorded.

Second-instar male ( Fig. 24; Plate 3 , fig. e). General appearance. Secretes a white, flocculent (wooly) wax test, 1.8–2.0 mm long, 0.6–1.0 mm wide; body red and broadly oval when removed from test, 0.7–1.45 mm long, 0.4–0.8 mm wide; derm membranous with inter-segmental lines on both dorsum and venter.

Mounted specimen. Body oval, 0.95–1.5 mm long, 0.5–0.8 mm wide.

Margin. Marginal setae conical, each 15 µm long, 10 µm wide at base, in a single complete row of 30–34 on each side.

Dorsum. Setae of two types: (i) conical setae, in submedial longitudinal lines of 5 setae extending from head apex to abdominal segment I, each seta 10 µm long, 7 µm wide at base, shorter than marginal setae; and (ii) tacklike setae, each 7 µm long, in medial longitudinal rows, extending from prothorax to anal ring, with 6–8 setae each row. Microtubular ducts, each with a circular opening about 3 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, numerous, scattered between tubular ducts over entire dorsum. Tubular ducts of Type 2, with inner ductule 8–10 µm long, inner cup 5 µm wide and outer ductule 10–14 µm long, abundant throughout dorsum.

Venter. Eyes circular, each 20–25 µm diameter, lateral to each scape. Antennae 7 segmented; total length 163–217 µm; segments III and VII longest; scape and pedicel each with 2 setose setae; segment III without setae; IV with 1 setose seta; V with 1 fleshy seta; VI with 1 fleshy seta and 1 setose seta and apical segment with 4 setose setae and 2 fleshy setae. Frontal lobes membranous, each 38–100 Μm long, 43–75 Μm wide; not always visible. Legs well-developed; measurements (in Μm) of metathoracic legs: coxa 60–100; trochanter + femur 135–150; tibia 55–85; tarsus 85–100 and claw 25–30; total length 330–410 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 28–40 Μm long, knobbed apically; claw digitules knobbed apically, each 25–35 Μm long, extending beyond claw apex; each claw with a denticle. Tentorium 113–125 µm long, 100–130 µm wide. Labium 3 segmented, triangular, 100–125 µm long, 68–80 µm wide; labial setae setose as follows: basal segment with 2 pairs about 13 µm long; median segment with 1 pair 20 µm long, and apical segment with 4 pairs, each 8 µm long. Spiracles subequal in size, each peritreme 10–15 Μm wide, with sclerosis 25–38 µm long; each spiracle with 5–8 quinquelocular pores, each 5 µm diameter, in a band from spiracle to margin of body. Tubular ducts of Type 2, with outer ductule 10–14 Μm long, sclerotized cup about 5 Μm in diameter and inner ductule 8–10 Μm long, scattered in a complete submarginal band 1 or 2 ducts wide. Bilocular pores circular, in a complete submarginal band of 10–12 on each side. Quinquelocular pores, with 2 pairs of pores among setose setae anterior to tentorium, each 5 µm wide; also pairs medially and submedially on each abdominal segment, each 3 µm wide. Ventral setae of four types: (i) setose setae: with 6 pairs medially, just anterior to tentorium, each 30–50 µm long; 4 setae mesad to each coxa, two 8–15 µm long and two 25–40 µm long; also abdominal segments II–VII each with a pair of setae 38 µm long medially, and a pair about 13 µm long submedially. Also with 2 pairs anterior to anal ring, each 10 µm long, and a pair posteriorly, each about 33 µm long; (ii) tack-like setae, in a complete submarginal row of 10–15, each 10 µm long; (iii) a pair of spinose setae, about 100 µm long, near posterior margin; and (iv) a pair of conical setae, similar to marginal setae, about 18 µm long, 7 µm wide posterior to anal ring, on body margin. Microspines, each about 3 µm long; arranged in groups of 3–5 in 6–9 transverse rows on each abdominal segment and scattered on thorax. Anal ring ventral, diameter 30–40 µm, composed of 2 semi-circles, each half with 3 spinose setae, each 32–45 µm long, and with 18–20 pores.

Comments. Other known Palaearctic second-instar male Kermes are those of K. bytinskii (here made a junior synonym to K. nahalali and redescribed in this study), K. williamsi ( Sternlicht, 1970) , K. quercus ( Podsiadlo, 2012) and K. vermilio ( Pellizzari et al., 2012) . Those of K. greeni , K. hermonensis , K. spatulatus and N. balachowskii are described here. All but K. echinatus and K. vermilio clearly differ from K. hermonensis in having long robust setose marginal setae, whereas those of K. hermonensis are conical and spinose.

Second-instar male K. echinatus , K. hermonensis and K. vermilio have a pair of membranous, frontal lobes. The second-instar male of K. hermonensis differs from K. echinatus and K. vermilio as follows (characters of K. echinatus and K. vermilio in brackets): (i) presence of quinquelocular pores on venter, anterior to tentorium (absent); (ii) 5–8 quinquelocular pores in each spiracular band (four in K. echinatus and two in K. vermilio ); and (iii) quinquelocular pores present in abdominal segments II–VII (absent).

Of the known Nearctic second-instar males, K. cockerelli , K. concinnulus and K. rimarum have long setose marginal setae, and the tubular ducts and quinquelocular pores are scattered on both the dorsum and venter ( Baer & Kosztarab, 1985); K. hermonensis has conical marginal setae, and, although it does have tubular ducts scattered on the dorsum, they are restricted to a narrow marginal band on the venter; in addition, the quinquelocular pores on K. hermonensis are only present on the venter.

Figures

PLATE 2. Kermes greeni Bodenheimer, general appearance; a. second-instar female; b. third-instar female; c. young adult female; d. gravid female; e. post-reproductive female with crawlers; f. second-instar male; g. prepupa; h. pupa; i. adult male.

PLATE 3. Kermes hermonensis Spodek & Ben-Dov, general appearance; a. second-instar female; b. third-instar female; c. young adult female; d. gravid female; e. female with eggs; f. second-instar male; g. prepupa; h. pupa; i. adult male, dorsal view; j. adult male, ventral view.

PLATE 5. Kermes spatulatus Balachowsky, general appearance; a. second-instar female; b. third-instar female; c. young adult female; d. post-reproductive female; e. post-reproductive female from Mt. Hermon; f. male test; g. second-instar male; h. prepupa; i. pupa; j. adult male.

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.