Kermes spatulatus Balachowsky

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 : 65-78

Kermes spatulatus Balachowsky

( Figs. 33–40; Plate 5 , figs. a–j)

Kermes spatulatus Balachowsky, 1953: 184

This species was originally described from first-instar nymphs collected from Daphna Oaks, Israel, on Q. ithaburensis.

Material examined. Type material studied. Syntypes: 3 first-instar nymphs, Daphne Oaks (=Horshat Tal Nature Reserve), Israel, Q. ithaburensis, 12.v.1952, H. Bytinski-Salz ( ICVI C:3693, MNHN 1058-5).

Other material studied. All non-type material collected off Q. ithaburensis in Israel by M. Spodek, unless otherwise indicated. Bet Dagan-Volcani Center, 24.iii.2012 (MC:641/ 5 adult ♀, BMNH), 10.v.2012 (MC:757/ 8 adult ♀, MNHN); Horshat Tal Nature Reserve, v.1958, Y. Ben-Dov (C:5250/ 1 adult ♀), 10.vi.1964, Y. Ben-Dov (C:5251/ 1 adult ♀), 26.iv.1967, Y. Ben-Dov (C:5252/ 1 adult ♀); M. Sternlicht; 18.iv.2010 (MC:155/ 1 adult ♀), 3.iv.2011 (MC:475/ 9 adult ♀, USMN), 8.iv.2012 (MC:644/ 4 adult ♀); Mt. Hermon Nature Reserve, 20.vi.2012 (MC:738/ 2 adult ♀); Bet Dagan-Volcani Center, 4.iii.2012 (MC:630/1 third-instar ♀), 13.iii.2012 (MC:628/1 thirdinstar ♀); Horshat Tal Nature Reserve, 20.iii.2011 (MC:465/1 third-instar ♀), 3.iv.2011 (MC:475/2 third-instar ♀), 21.iv.2011 (MC:489/3 third-instar ♀), 8.iv.2012 (MC:644/5 third-instar ♀); Mezar, 10.iii.2013 (MC:830/1 thirdinstar ♀); Mt. Hermon Nature Reserve, 7.v.2012 (MC:655/4 third-instar ♀), 24.v.2012 (MC:703/2 third-instar ♀), 20.vi.2012 (MC:739/1 third-instar ♀), 20.v.2013 (MC:878/4 third-instar ♀); Bet Dagan-Volcani Center, 8.ii.2012 (MC:608/3 second-instar ♀), 4.iii.2012 (MC:629/4 second-instar ♀); Horshat Tal Nature Reserve, 11.i.2011 (MC:381/8 second-instar ♀), 6.ii.2011 (MC:413/5 second-instar ♀), 14.ii.2011 (MC:615/8 second-instar ♀), 27.ii.2011 (MC:429/9 second-instar ♀), 20.iii.2011 (MC:464/10 second-instar ♀), 21.iv.2011 (MC:488/3 secondinstar ♀), 8.iv.2012 (MC:648/1 second-instar ♀), 13.i.2013 (MC:785/6 second-instar ♀); Mezar, 10.iii.2013 (MC:829/1 second-instar ♀); Horshat Tal Nature Reserve, 12.v.2000, Z. Tamari (C:4750/1 first-instar); Horshat Tal Nature Reserve, 2. v.2010 (MC:180/15 first-instars, BMNH), 3.vi.2010 (MC:270/12 first-instars, USNM), 13.v.2010 (MC:496/78 first-instars), 29.viii.2010 (MC:301/1 first-instar), 6.v.2011 (MC:722/38 first-instars, MNHN); Mt. Hermon Nature Reserve, Q. look, 21.viii.2011 (MC:572/4 first-instars), 25.vi.2012 (MC:760/20 firstinstars); Horshat Tal Reserve, 3.iv.2012, Y. Ben-Dov (C:5230/ 5 adult ♂), 2.iv.2013 (MC:842/ 5 adult ♂), 27.iii.2012 (MC:817/ 1 adult ♂); Mezar, 27.iii.2013 (MC:826/ 1 adult ♂); Mt. Hermon Nature Reserve, Q. look, 24.v.2012 (MC:824/ 1 adult ♂), 28.v.2012 (MC:828/ 1 adult ♂); Horshat Tal Nature Reserve, 27.ii.2012 (MC:817/ 1 pupa), 19.iii.2013 (MC: 835/ 8 pupae); Mezar, 17.iii.2013 (MC:827/ 1 pupa); Mt. Hermon Nature Reserve, Q. look, 7.v.2012 (MC:272/ 3 pupae), 24.v.2012 (MC:701/ 1 pupa); Bet-Dagan-Volcani Center, 27.2.2012 (MC:627/1 prepupa); Horshat Tal Nature Reserve, 20.iii.2013 (MC:834/3 prepupae); Mezar, 10.iii.2013 (MC:825/2 prepupae); Mt. Hermon Nature Reserve, Q. look, 7.v.2012 (MC:272/9 prepupae); Bet Dagan-Volcani Center, 8.ii.2012 (MC:607/1 second-instar ♂), 27.ii.2012 (MC: 626/3 second-instar ♂), 24.iii.2012 (MC:637/1 second-instar ♂); Horshat Tal Nature Reserve, 12.iii.2012 (MC:631/3 second-instar ♂), 8.iv.2012 (MC:643/2 second-instar ♂), 14.ii.2013 (MC:807/4 second-instar ♂); Mt. Hermon Nature Reserve, Q. look, 21.viii.2011 (MC:580/2 secondinstar ♂), 7.v.2012 (MC:271/6 second-instar ♂), 24.v.2012 (MC:700/1 second-instar ♂).

Adult female ( Fig. 33). General appearance. Pre-reproductive adult female ( Plate 5 , fig. c). Body oval to round, 2.0–4.0 mm long, 1.4–3.4 mm wide, 1.4–2.5 mm high; dorsum and venter dark grey to black, with 13–15 dark spots on dorsum in 8 longitudinal rows, extending from anal opening to anterior apex of body, each about 0.1 mm in diameter. Gravid and post-reproductive female ( Plate 5 , fig. d). Body spherical and sclerotized, 5.2–6.0 mm long, 4.9–5.6 mm wide and 3.0–5.0 mm high; body black. Some specimens from Mt. Hermon Nature Reserve with a deep, medial longitudinal groove and 4 transverse grooves on each half body, resulting in a bumpy derm ( Plate 5 , fig. e), whilst dorsum of those from Horshat Tal Nature Reserve are smooth and flat, with 1 deep medial longitudinal groove and 4 longitudinal rows of 13–15 depressions as on pre-reproductive specimens; specimens from both localities occasionally with entire body covered in a thin layer of white wax.

Mounted young adult female. Body oval to round; 2.5–4.0 mm long, 2.0– 3.4 mm wide.

Margin. Marginal setae short and spinose, each 15 Μm long, in a single complete row of 40 on each side.

Dorsum. Dorsal reticulated areas somewhat oval, in about 8 longitudinal rows, each with 8–15 areas 25–60 Μm long and 20–40 Μm wide. Bilocular pores oval, each 8 Μm long and 5 Μm wide, abundant throughout dorsum. Microtubular ducts with circular opening, each about 3 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, scattered over entire dorsum. Seta-pore clusters in 20–30 clusters on each side, extending in a single complete submarginal row, each cluster with 3–7 quinquelocular pores, 5 Μm wide seta and a seta about 20 um long. Anal ring dorsal, circular with anterior gap, without setae or pores, 100 Μm diameter. Anal lobes possibly represented by 2 sclerotized areas, each 50–87 Μm long, 25–40 Μm wide, posterior to anal ring, each with 4 setose setae: 3 setae 50–62 Μm and 1 shorter seta 20–30 Μm long; and with about 18 setae posterior to each anal lobe, each about 25 Μm long.

Venter. Eyes absent. Antennae 5 or 6 segmented; when 5 segmented, segment III and IV appearing fused; 150–175 Μm long; 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 fleshy seta and 1 setose seta; apical segment with 2 fleshy setae and 4 setose setae. Legs well-developed; tibia and tarsus occasionally fused; measurements (in µm) of metathoracic legs: coxa 50–75, trochanter + femur 70–75, tibia 40–50, tarsus 75–80, claw 15–17 µm long; total length 212–250 µm; each trochanter with 2 oval, sensory pores on each side; tarsal digitules narrow, each 40–45 µm long, knobbed apically and extending beyond apex of claw; claw digitules knobbed apically, each 20–25 µm long, shorter than tarsal digitules. Claw denticle absent. Tentorium 350–375 Μm long, 250–275 Μm wide. Labium 3 segmented, triangular, 200–250 Μm long, 125–170 Μm wide; labial setae setose: basal segment with 2 pairs about 7 Μm long; median segment with 1 pair about 15 Μm long, and apical segment with 4 pairs about 10–17 Μm long. Mesothoracic spiracles smaller than metathoracic spiracles; each mesothoracic peritreme 112–130 Μm long, 62– 75 Μm wide; each metathoracic peritreme 150–180 Μm long, 75–80 Μm wide. Tubular ducts of two types: Type 1 with inner ductule 13–15 µm long, cup 10 µm wide and outer ductule 20–25 µm long, in a complete submarginal band 15–20 ducts wide. Type 2 with inner ductule 10–15 µm long, cup 8 µm wide and outer ductule 10–12 µm long, present on median areas of abdominal segments. Quinquelocular pores, each 8 Μm diameter, with about 50 surrounding both anterior and posterior spiracles; also 12–14 pores across each abdominal segment, and 50–100 pores posterior to vulva. Trilocular pores, each 5 Μm diameter, with about 5 surrounding each anterior spiracle. Multilocular pores, each 8–10 Μm diameter with 9–10 loculi, in a group of 20–25 pores mesad to each spiracle; also in 3 or 4 clusters of 15–30 pores on abdominal segments II–VII. Ventral setae spinose, each 10 Μm long, in groups of 7–10 posterior to each leg; with 6–8 setae in transverse rows across each abdominal segment, each 20 Μm long; and in a group of 35–50 setae posterior to vulva, each 23 Μm long. Microspines, each 2–3 Μm long, arranged in groups of 2–5 in 10–12 rows on each abdominal segment and dispersed on thorax.

Comments. T he adult female of K. spatulatus shares the following characters with K. greeni , K. nahalali and the Oriental species K. flavus ( Liu & Shi, 1995) , K. miyasakii ( Liu & Shi, 1995) , K. orientalis ( Liu & Shi, 1995) and K. taishanensis ( Hu, 1986) : (i) legs fully developed; (ii) six-segmented antennae; (iii) seta-pore clusters located submarginally on dorsum; (iv) anal ring dorsal; (v) anal ring without setae or pores; (vi) sclerotized anal lobes on dorsum; and (vii) two types of tubular ducts on venter. However, the segmentation of the legs varies: legs 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) ; and legs fully developed in the Nearctic K. nudus and K. rimarum ( Bullington & Kosztarab, 1985) . In addition, the antennae range from four to six segments, mostly with indistinct segmentation (six segmented in K. spatulatus ). Adult female K. spatulatus can be distinguished from those of K. nahalali by the absence of the dorsal median spots; and from K. greeni in having narrow anal lobes with up to five setae on each lobe while K. greeni has wide, quadrate anal lobes with 10–15 setae on each lobe.

Third-instar female ( Fig. 34; Plate 5 , fig. b). General appearance. Body round and convex, 1.7–2.7 mm long, 1.0– 2.5 mm wide, 1.3–1.5 mm high; dorsum and venter brown to black.

Mounted specimen. Body broadly oval, 1.8–3.0 mm long, 1.0– 2.5 mm wide.

Margin. Marginal setae short and spinose, each 10 µm long, 5 µm wide, in a complete, single line of 32–38 on each side.

Dorsum. Seta-pore clusters in 10–12 clusters in a submarginal band on anterior half of body; each cluster with 2–5 quinquelocular pores, each 5 Μm wide and a spinose seta 20–22 um long; posteriorly, submarginal band of setae continues from mid-thorax to anal lobe but without loculate pores. Other setae all setose, each 20 µm long, in 3 broad diagonal bands, each band between diagonal lines of dark-rimmed areas, 1 anteriorly, 1 medially approximately on thorax and another anteriorly on abdomen, each band with 16–20 setae. Bilocular pores oval to circular, each 3 µm long and 2 µm wide, abundant on dorsum. Dark-rimmed areas in 4 diagonal lines; each line with 5–10 areas 30–40 Μm long and 15–20 Μm wide. Anal ring dorsal, 45–50 µm diameter, circular with anterior gap; setae and pores absent. Anal lobes represented by sclerotized rectangular areas, each about 50 µm long, 30 µm wide, sometimes absent; without setae. Each lobe with about 7 setose setae surrounding it, 20 µm long; also with a longer setose setae posterior to each anal lobe, 50 µm long.

Venter. Eyes circular, each 20 µm diameter, laterally to each scape. Antennae 6 segmented, 125–150 µm long; segments II and III appearing fused; scape and pedicel, each with 2 setose setae, segment III with 1 setose seta; IV with 1 fleshy seta and VI with 4 setose setae and 2 fleshy setae. Legs well developed; measurements (in µm) of metathoracic leg: coxa 50–53, trochanter + femur 60–65, tibia 37–50, tarsus 67–75, claw 8–12; total length 210– 250 Μm; trochanter with 2 oval sensory pores on each side, each 5 Μm long and 3 Μm wide; tarsal digitules 21–25 Μm long, knobbed apically, extending slightly beyond claw apex; claw digitules knobbed apically, each 10–18 Μm long, shorter than tarsal digitules; claw denticle absent. Tentorium 165–200 µm long, 163–175 µm wide. Labium 3 segmented, triangular, 137–145 µm long, 90–115 µm wide; labial setae setose: basal segment with 2 pairs 10 Μm long; medial segment with a pair 15 µm long and apical segment with 4 pairs 13 µm long. Spiracular peritreme of mesothoracic spiracles smaller than those of metathorax; each mesothoracic peritreme 25–30 Μm long, 20–25 Μm wide, sclerosis 40–50 µm long; each metathoracic 35–40 Μm long, 25–30 Μm wide, sclerosis 40–55 µm long.

Tubular ducts of two types: Type 1 with inner ductule 12–15 µm long, inner cup 5 µm wide and outer ductule 20– 23 µm long, arranged in a complete, broad submarginal band 6–8 ducts wide; and Type 2, with inner ductule 10–13 µm long, inner cup 5 µm wide and outer ductule 15–18 µm long, arranged in a submedial band 1–2 ducts wide. Quinquelocular pores, each 5 µm diameter; with 2 or 3 pores anterolateral to each mesothoracic spiracle and 8–10 pores anterolateral to each metathoracic spiracle; and with about 6 pores in a line across abdominal segments II– VII and in a group posteriorly. Ventral setae of three types: (i) setose setae in groups of 3–5 posterior to each leg, each 5–10 µm long; and in transverse rows of 6–8 on abdominal segments II–VI, each 10–13 µm long; (ii) spatulate setae: with a total of 10 anterior to scape, each 30 µm long, 8 µm wide; (iii) tack-like setae, each 5 µm long, with 1 on each abdominal segment II–VII. Microspines, each 2 µm long, in groups of 1–4 over entire body; in 10 transverse rows on abdominal segments.

Comments. Of the Israeli species, third-instar female K. spatulatus are very similar to those of K. greeni and K. nahalali . All three species share (character-states for K. echinatus , K. hermonensis and N. balachowskii in brackets): (i) marginal setae short and spinose (conical); (ii) legs fully developed (leg segmentation reduced); (iii) anal ring dorsal (ventral), and (iv) sclerotized anal lobes present on dorsum (absent). The third-instar female of K. spatulatus is distinguished from those of K. greeni and K. nahalali by the absence of setae on the anal lobes ( K. greeni 10–15 setae and K. nahalali with three setae on each lobe). Kermes spatulatus also has an incomplete line of seta-pore clusters, restricted to anterior half of body, whereas the ring is complete on K. nahalali . Of the non-Israeli species, K. spatulatus differs from K. vermilio (character states for latter in brackets) in having six-segmented antennae (one-segmented antennae) and fully-developed legs (one-segmented). Among the Nearctic Kermes , only the third-instar female of K. sylvestris has been described and illustrated ( Bullington & Kosztarab, 1985) and it has a ventral submarginal band of tubular ducts, and the quinquelocular pores are sparse on venter, as on K. spatulatus , but the latter has seta-pore clusters on the dorsum and sclerotized anal lobes on the dorsum, both absent from K. sylvestris .

Second-instar female ( Fig. 35; Plate 5 , fig. a). General appearance. Body oval, 0.5–1.2 mm long, 0.3–0.6 mm wide; dorsum and venter orange; body with marginal fringe of whitish wax filaments.

Mounted specimen. Body oval, 0.9–1.4 mm long, 0.4–0.6 mm wide.

Margin. Marginal setae spatulate, each about 38 µm long, in a single complete line of 30–34 on each side; also with about 5 shorter spatulate setae on each side, each about 23 µm long, from anterior margin to level of labium; tack-like setae, about 5 µm long, present submedially, with about 10 on each side submedially from level of labium to level anterior to anal ring.

Dorsum. Microtubular ducts, with a circular opening, each about 3 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, in longitudinal rows of about 26 pores. Setae setose, each about 5 µm long, in a submedial line of about 10 on each side on thorax and abdominal segments.

Venter. Eyes circular, each 15 µm diameter, lateral to each scape. Antennae 6 segmented, length 150 µm; segments III and VI longest; scape and pedicel each with 2 setose setae, segment III with a setose seta, IV with a fleshy seta, V with a fleshy seta and a setose seta, VI with 4 setose setae and 2 fleshy setae. Legs well developed; measurements (in Μm) of metathoracic legs: coxa 37–50, trochanter + femur 100–105 Μm, tibia 50–58 Μm, tarsus 67–87, claw, 25–28 Μm; total length 280–328 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules each 30–42 Μm long, knobbed apically, extending beyond claw apex; claw digitules knobbed apically, each 20–25 Μm long, shorter than tarsal digitules; claw denticle present. Tentorium 137–145 µm long, 95–112 µm wide. Labium 3 segmented, triangular, 100–107 µm long, 57–70 µm wide; labial setae setose, as follows: basal segment with 2 pairs 3 µm long, median segment with 1 pair 5 µm long and apical segment with 4 pairs about 20 µm long. Spiracles subequal in size, each peritreme about 8–13 Μm wide, crescent-shaped sclerosis 20–25 µm long. Tubular ducts of Type 2 only, with inner ductule 6–10 µm long, cup 5 µm wide and outer ductule 9–17 µm long, arranged in a complete submarginal band about 2 ducts wide. Quinquelocular pores, each 5 µm diameter: with 5 pores anterior to tentorium; 5 or 6 pores anterior to each metathoracic spiracle; 1 pore anterior to each mesothoracic spiracle; and abdominal segments II–VI each with 4 pores. Ventral setae of three types: (i) spatulate setae, similar to marginal setae, anterior to each scape, of two lengths: 6 setae 38 µm long, and 6 setae 10 µm long; (ii) setose setae: with 3 pairs medially anterior to tentorium, each 33 µm long; and segmentally arranged in medial and submedial rows abdominal segments II–VI, medial row setae each 18 µm long, and submedial setae each 10 µm long. Also with a pair just anterior to anal ring about 12 µm long, and a pair posterior to anal ring 38 µm long; (iii) tack-like setae, in a group of 3–5 setae between anterior and posterior spiracles, each 8 µm long, and also segmentally in a submarginal row on metathorax and abdominal segments II–VII. Microspines, each about 2 µm long, in groups of 4 or 5 in 5 or 6 transverse rows on each abdominal segment and scattered on thorax. Anal ring ventral, diameter 37–40 µm, composed of 2 half circles, each half with 3 pointed setae, each 45–50 µm long, and 3 pores. Anal lobes well developed, membranous and somewhat rounded, each with a spatulate-like seta similar to marginal setae, 18 µm long, on inner margin, plus a long flagellate seta on apex, 125–150 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 K. spatulatus is distinct with its spatulate marginal setae and absence of membranous frontal lobes (as in K. vermilio ). Tubular ducts are present on both the venter and dorsum in K. echinatus and K. hermonensis , but are only found on the venter in K. spatulatus . Kermes spatulatus has sixsegmented antennae and well-developed legs, as on K. nahalali and K. greeni and several Nearctic species, such as K. cockerelli , K. concinnulus , N. iselini and O. secunda ( Baer & Kosztarab, 1985) , Kermes echinatus , K. hermonensis and N. balachowskii have reduced antennal and leg segmentation.

First-instar nymph ( Fig. 36). General appearance. Body oval, tapering posteriorly, 0.42–0.49 mm long, 0.14–0.19 mm wide; dorsum and venter orange.

Mounted specimen. Body oval, 0.45–0.50 mm long, 0.17–0.20 mm wide.

Margin. Marginal spatulate setae in a single complete row of 20 each side, each 11–12 µm long, 3 µm wide at apex.

Dorsum. Derm membranous with inter-segmental lines. Spatulate setae in 4 incomplete longitudinal rows, each submedian rows with 8 setae on metathorax and abdominal segments II–VII, each 14–15 µm long, 5–6 µm wide at apex; and each median row with about 6 setae on head and thorax, each 15–17 µm long, 4–5 µm wide at apex. Also with a spatulate seta on inner margin of each anal lobe, 12–17 µm long. 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, with 12–14 ducts in each row.

Venter. Eyes present close to margin, each 12 µm wide. Antennae 6 segmented, 100–110 µm long; segment III longest; scape and pedicel each with 2 setose setae; segment III with a setose seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 setose seta, and apical segment with 2 fleshy setae and 4 setose setae. Legs well developed; measurements (in Μm) of metathoracic leg: coxa 22–30, trochanter + femur 60–75, tibia 27–37, tarsus 50–55, claw 20–22; total length 195–209 µm; trochanter with 2 oval sensory pores on each side; tarsal digitules each 27–32 µm long, knobbed apically, extending slightly beyond claw apex; claw digitules knobbed apically, each 17–22 µm long, shorter than tarsal digitules; each claw with a single denticle near the tip. Tentorium 87–92 µm long, 62–80 µm wide. Labium 3 segmented, triangular, 75–82 µm long, 30–45 µm wide; with 14 setose setae: basal segment with 2 pairs, each 5–10 µm long, median segment with 1 pair on dorsal surface 8–9 µm long, and apical segment with 4 pairs, each 10–14 µm long. Spiracles subequal in size, each peritreme about 5 Μm wide sclerosis 15–20 µm long; each spiracle with 1 quinquelocular pore anterolaterally, 5 µm diameter. Also with a pair of slightly larger quinquelocular pores just anterior to tentorium and 1 medially to each coxa, each 7 µm wide; plus a pair of pores submedially in abdominal segments II–VII. Ventral setae of three types: (i) spatulate setae: a pair anterior to each scape, each 15 um long and 3 µm wide at apex, similar to dorsal submarginal setae; (ii) setose setae: with 6 setae medially between scape and tentorium, each 12–20 µm long; a single setae medial to each coxa, each 10–11 µm long; a line of 4 setae across abdominal segments II–VII, setae in medial rows, each 10–15 µm long; setae in submedial rows each 5–10 µm long. Also with a pair anterior to anal ring, 10 µm long, and a pair latero-posterior to anal ring, each 15–25 µm long; (iii) tack-like setae, each 5–6 µm long, in marginal and submarginal rows, each marginal row complete, with 12–14 setae; and each submarginal row restricted to abdominal segments III–VII. Bilocular pores oval, with 1 pore present between margin and each spiracle, each 3 µm long and 2 µm wide. Microspines, each about 1 µm, arranged medially and submedially on each abdominal segment. Anal ring ventral, diameter 17–20 µm, composed of 2 semi-circles, each half with about 6 pores and 3 pointed setae, each 10–11 µm long. Anal lobes well-developed; apical margin of each lobe with 1 spinose seta, 8–10 µm long, and a long apical seta, 165–200 µm long.

Comments. Kermes spatulatus was described by Balachowsky (1953) based on first-instar nymphs collected in Israel. The present redescription includes several features not originally described, such as presence of: (i) dorsal microtubular ducts in four longitudinal rows; (ii) a complete submarginal row of setae on venter; (iii) a single bilocular pore located submarginally level with each spiracle on venter; (iv) a denticle near tip of each claw; (v) one setose seta mesad to each coxa; (vi) anal ring open both posteriorly and anteriorly; and (vii) microspines on abdominal segments on venter.

Balachowsky (1953) introduced K. spatulatus as a new species after comparing its first-instar nymph with those of K. bacciformis , a similar species known from the Mediterranean and European regions. He observed that the post-reproductive female of both species were spherical, with a dorsal longitudinal median depression, and black in colour ( Plate 5 , fig. d and Plate 7 , fig. a). However, minor differences can be seen in the shape and size of the dorsal spatulate setae of the first-instar nymph. Balachowsky (1953) noted that the median spatulate setae of K. bacciformis are about half the length of those on K. spatulatus . Although type material of K. bacciformis was not available for this study, three specimens collected from Italy (see Material examined for details) were seen. We agree with Balachowsky (1953) and conclude that these species are very similar in the first-instar nymph and postreproductive stages. Perhaps additional studies on the morphology of other stages of K. bacciformis will show more obvious differences between the two. Both K. greeni and K. spatulatus possess spatulate setae on the margin and dorsum but their shape, size and distribution separate these two species.

The other Palaearctic species that possess spatulate setae are K. mutsurensis ( Kuwana, 1931) , K. nakagawae ( Kuwana, 1931) , K. nigronotatus ( Hu, 1986) , K. orientalis ( Liu & Shi, 1997) , K. quercus ( Linnaeus, 1758, Podsiadlo, 2005), K. taishanensis ( Hu, 1986) , K. vastus ( Kuwana, 1931) , K. viridis ( Borchsenius, 1960) and K. williamsi ( Sternlicht, 1970) . Kermes spatulatus differs from these species in having one quinquelocular pore associated with each anterior and posterior spiracle, while K. mutsurensis has two loculate pores within the atrium of each anterior spiracle plus two multilocular pores laterad to each anterior spiracle and lacks posterior spiracles pores; K. nakagawae has three multi-locular pores within the spiracle atrium of each anterior spiracle and lacks posterior spiracular pores; K. nigronotatus has one quinquelocular pore laterad to each anterior spiracle and lacks posterior spiracular pores; K. orientalis has two quinquelocular pores laterad to each anterior spiracle and lacks posterior spiracular pores; K. quercus has two pores associated with each anterior spiracle: one with 3–5-loculi and one with seven-loculi plus one 5–7 with loculi pore associated with each posterior spiracle; K. taishanensis has one quinquelocular pore laterad to only anterior spiracle, and lacks posterior spiracular pores; K. vastus has two multilocular pores within the atrium of each anterior spiracle and lacks posterior spiracular pores; K. williamsi has one pore with seven-loculi and sometimes a second quinquelocular pore associated with each anterior spiracle plus one quinquelocular pore associated with each posterior spiracle.

Adult male ( Fig. 37; Plate 5 , fig. j). As for K. echinatus apart from: General appearance. Newly emerged males brown; body 2.0– 2.5 mm; width across mesothorax 0.5–0.6 mm wide; wings 1.2–1.5 mm long, 0.45–0.75 mm wide; caudal filaments 1.0– 1.5 mm long.

Mounted specimen. Body length 2.0– 2.5 mm long; width across mesothorax 0.5–0.7 mm.

Head. Length 250–300 µm; width across genae 275–300 µm. Dorsal simple eyes (dse) and ventral simple eyes (vse) both 45–50 µm diameter; lateral simple eyes (lse) smaller, each 20–30 µm diameter; each ocellus 20–25 µm diameter. Dorsal head setae (dhs): 8–10 hs on either side of median crest. Ventral head setae (vhs) total 8–10 hs. Antennae: length 950–1000 µm (ratio of total body length to antennal length about 1:0.43). Scape (scp) 40–60 µm long, 44–50 µm wide, with 2 or 3 hs. Pedicel (ped) 70–75 µm long, 40–50 µm wide, with 6–10 hs. Flagellar segments III–X each 60–160 µm long, 30–35 µm wide; segment III longest, segments becoming shorter towards apex; approximate number of setae per segment: III–VI 13–20 fs; VII–X 8–15 fs; VIII 1 ab; IX 1 ab; X 4 ab + 4 cs.

Thorax. Prothorax: as for K. echinatus but sternum (stn1) with faint radial striations and 3 hs prosternal setae (stn1s) on each side. Anteprosternal (astn1s) and antemesospiracular setae (am2s) absent.

Mesothorax: prescutum (prsc) 50–75 µm long, 150–175 µm wide Scutum (sct) with median membranous area 60–65 µm long; with 1 hs on each side laterad to membranous area, 12–15 µm long. Scutellum (scl) 100–150 µm long, 200–210 µm wide, with 1 hs 13–15 µm long on each side. Basisternum (stn2) 150–180 µm long, 250–280 µm wide; basisternal setae absent. Tegula (teg) with 5 or 6 hs tegular setae (tegs). Mesothoracic spiracles (sp2): peritreme 30–35 µm wide.

Metathorax: Metathoracic spiracles (sp3): peritreme 30–35 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: 1200–1500 µm long, 450–475 µm wide (ratio of length to width 1:0.39; ratio of total body length to wing length 1:0.52). Hamulohalteres (h) 100–125 µm long, 35–40 µm wide; with a single apical hamulus, 65–85 µm long. Legs: metathoracic legs longest, length 724–847 µm; prothoracic and mesothoracic legs 716–843 µm long. Coxa: I 86 –100; II 100–110; III 100–115 µm long, with 10–14 hs. Trochanter: I 45 –50; II 40 –53; III 50 –55 µm long, with 6–8 hs; each trochanter with 3 pairs of oval sensory pores in a triangle on each surface. Femur: I 200–230; II 200–205; III 200–220 µm long, with 30–35 hs. Tibia: I 265–300; II 275–280; III 275–300 µm long, with 35–47 hs; all tibia with 2 apical spurs (tibs) 23–25 µm long. Tarsi 2-segmented; tarsus 2 (tar2): I 95 –100; II 90 –110; III 100–110 µm long, with 15–25 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 30 µm long.

Abdomen. Sternites (as) sclerotized on segment VI–VIII; tergites and pleural areas unsclerotized. Abdominal dorsal setae (ads) (totals): segments 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 4–5 hs, 10 µm long, VIII 3 hs, one 85–90 µm long and other 25–30 µm long; ventropleural setae (avps) (on each side) II–VIII 2 hs, 15 µm long; abdominal ventral setae (avs) (totals) II–V 12–20 hs, VI–VIII 6–10 hs, 20–25 µm long. Glandular pouch setae (gls), 142–150 µm long; each pouch with 10– 12 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) 40–50 µm long. Aedeagus (aed) 100–125 µm long Genital capsule without setae dorsally but with 6–8 hair-like setae (gts) ventrally.

Comments. The adult male of K. spatulatus is very similar to those of K. greeni and K. nahalali . The adult male is most similar to K. nahalali in life as both are grey. K. spatulatus differs from these two species in having eight dorsopleural setae on abdominal segment I, whereas K. greeni has four and K. nahalali that has one seta. Metasternal setae are present on K. spatulatus as on most other species, the only exceptions being Kermes echinatus and K. hermonensis , and the Nearctic species E. gillettei ( Miller & Miller, 1993) , which lack metasternal setae.

Fourth-instar male (pupa) ( Fig. 38; Plate 5 , fig. i). 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 brown and elongate oval when removed from test, 1.0– 1.5 mm long, 0.5–1.0 mm wide.

Mounted specimen. Body 1.4–1.6 mm long, 0.5–1.0 mm wide; head 235–250 µm long, 325–425 µm wide.

Margin. Wing-buds: fore-wings 580–600 µm long, 150–200 µm wide; hind-wings: length ratio to mesothoracic wings 1:10. Setae fine and blunt, each 25 µm long, arranged as on K. echinatus .

Dorsum. Eyes absent. Ocular sclerite reticulated, 40–60 µm long, 150–160 µm wide. Setae setose, each 5 µm long, in complete submedial rows of about 5 setae. Anal lobes rounded and slightly sclerotized; each with a seta 80–90 Μm long on apex. Median penial lobe triangular, 105–110 Μm long, 105–130 Μm wide; with 3 pairs of setose setae, each 15–20 Μm long.

Venter. Ocular sclerite as on dorsum. Antennae 10 segmented, 450–550 µm long, 40–50 µm wide; apical segment with 4 or 5 fleshy setae, each about 5 µm long. Spiracles subequal in size, each peritreme 25–30 µm long, 10–20 µm wide; each sclerosis 35–40 µm long; without pores. Legs well-developed, subequal in size; measurements (in Μm) of metathoracic leg; coxa 65–75, trochanter + femur 170–200, tibia 140–165, tarsus 95–100; total length 455–505 Μm; each coxa with 2 short setae, each 5 µm long; claw 5 µm long. Ventral setae setose, with 5 pairs of inter-antennal setae, each 5 Μm long; abdominal setae arranged as on K. echinatus . Microspines as on K. echinatus .

Comments. The pupa of K. spatulatus is greyish-brown in life, similar to those of N. balachowskii and K. nahalali ( K. echinatus and K. hermonensis red; K. greeni orange). The colours of the Nearctic pupae A. kingii and E. gillettei have not been recorded. As with most kermesid pupae, those of K. spatulatus lack quinquelocular pores anterior to each spiracle; the exception is K. hermonensis which has two pores. The pupae of both K. greeni and K. spatulatus have sclerotized anal lobes, unlike all other known kermesid pupae which have membranous lobes.

Third-instar male (prepupa) ( Fig. 39; Plate 5 , fig. h). As for K echinatus apart from: General appearance. Body brown and broadly oval when removed from test, 1.0– 1.5 mm long, 0.5–0.8 mm wide.

Mounted specimen. Body 1.2–1.7 mm long, 0.5–0.8 mm wide.

Margin. Anterior wing-buds each 370–425 µm long, 135–150 µm wide. Setae fine and blunt, each 12–20 µm long, arranged as on K. echinatus . Dorsum. Setae setose, each 5–8 µm long, with 7 pairs on head apex; also with complete submedial longitudinal rows of about 10 setae. Anal lobes rounded apically and membranous, each with a long setose seta 50–65 Μm long and a shorter setose seta 13–15 Μm long. Median penial lobe 60–65 Μm long, 110–115 Μm wide, broadly rounded, with 3 pairs of setose setae, each 5–10 Μm long; also, just anterior to median penial lobe, with 2 pairs of setose setae, each 5–10 Μm long.

Venter. Eyes absent. Antennae 7 or 8 segmented, tapering apically, 250–260 µm long, 75–80 µm wide; apical segment with 5 fleshy setae, each about 5 µm long. Spiracles subequal in size, each peritreme 15–25 µm long, 10– 15 µm wide, each sclerosis 30–35 µm long; without pores. Legs length of metathoracic leg, 215–250 Μm long and 65–75 Μm wide. Ventral setae setose, each 8–10 Μm long; with 3 pairs between scapes and abdominal setae arranged as on K. echinatus . Microspines, as on K. echinatus .

Comments. The prepupa of K. spatulatus is greyish-brown in life, similar to those of N. balachowskii and K. nahalali ( K. echinatus and K. hermonensis red; K. greeni orange). The colours of the Nearctic pupae A. kingii and E. gillettei have not been recorded. The prepupa of K. spatulatus lacks quinquelocular pores anterior to each spiracle, as in K. nahalali , K. greeni and the Nearctic species A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) . One or two pores are present near each spiracle in K. echinatus , K. hermonensis and N. balachowskii .

Second-instar male ( Fig. 40; Plate 5 , fig. g). General appearance. Secretes a white, flocculent (wooly) wax test 1.6–1.8 mm long, 0.6–0.8 mm wide. Body brown, broadly oval when removed from test, 1.0– 1.45 mm long, 0.4–0.8 mm wide; derm membranous with inter-segmental lines.

Mounted specimen. Body oval, 1.5–1.8 mm long, 0.7–0.9 mm wide.

Margin. Marginal setae robust and setose, each 50 µm long, in a complete single row of 30–33 on each side.

Dorsum. Setae robust, each about 25 µm long, shorter than marginal setae, in complete submedial longitudinal rows, with 10 setae each row. Microtubular ducts with circular opening about 3 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; numerous, scattered between tubular ducts on entire dorsum. Tubular ducts of Type 2 only, 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 15–20 µm diameter, located laterally to each scape. Antennae 7 segmented; total length 175–200 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 38–50; trochanter + femur 100–125; tibia 50–65; tarsus 80–85 and claw 23–30; total length 290–315 Μm; trochanter with 2 oval sensory pores on each side, each 5 Μm long and 3 Μm wide; tarsal digitules 35– 40 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 100–125 µm long, 110–125 µm wide. Labium 3 segmented, triangular, 100–125 µm long, 85–90 µm wide; labial setae setose: basal segment with 2 pairs about 13 µm long, median segment with 1 pair 13–20 µm long and apical segment with 4 pairs 15–22 µm long. Spiracles subequal in size, each peritreme 10–13 Μm wide, sclerosis 35–40 µm long; each spiracle with 5 quinquelocular pores anterolaterally, each 5 µm diameter. Tubular ducts of Type 2 only, with inner ductule 8–15 µm long, inner cup 5 µm wide and outer ductule 10–15 µm long, dispersed in a complete submarginal band 1–2 ducts wide. Bilocular pores circular, each 2 µm diameter, with 10–12 pores on each side in a complete single submarginal band. Quinquelocular pores each 5 µm wide: with 5 pairs among setose setae anterior to tentorium; 6 pores in a line across each abdominal segment in medial, submedial and submarginal lines; plus a group of 8 pores laterad to anal ring. Ventral setae of two types: (i) setose: with 10 medially just anterior to tentorium, each 60–88 µm long; 3 setae mesad to each coxa, each 25–38 µm long; and with medial and submedial rows on abdominal segments II– VII, medial setae each about 60 µm long, and submedial setae about 35 µm long. Also with 2 pairs of setae anterior to anal ring, each 20–25 µm long, and a pair of setose setae posterior to anal ring, each about 150 µm long; (ii) tack-like setae present in a complete submarginal row of about 10–15 setae on each side, each 10 µm long. Microspines, each about 3 µm long, in groups of 3–5 in 4–6 transverse rows on each abdominal segment and on thorax. Anal ring ventral, diameter 35–40 µm; composed of 2 semi-circles, each half with 3 pointed setae, subequal in size, 20–35 µm long, and 8–12 pores.

Comments. The second-instar male of K. spatulatus is unique in having quinquelocular pores in six longitudinal rows on abdominal segments ventrally and laterad to anal ring. Pores are absent in K. echinatus and the number of longitudinal rows on the venter of abdominal segments varies in other species; N. balachowskii , K. nahalali (two rows) and K. greeni , K. hermonensis and K. vermilio (four rows) ( Pellizzari et al., 2012). The Nearctic second-instar males of K. cockerelli , K. concinnulus and K. rimarum also have setose marginal setae but have quinquelocular pores scattered on both the dorsum and venter ( Baer & Kosztarab, 1985). The second-instar male K. spatulatus , and other Palaearctic species, have quinquelocular pores that are present only on the venter.

Figures

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 7. Some Palaearctic Kermesidae, general appearance of post-reproductive female; a. Kermes bacciformis (MNHN 10646; France: Bouches-du-Rhône, Marseille, Quercus sp., x. 1950, coll. P. Vayssière); b. K. ilicis (MNHN 7413; France: Vaucluse, Sérignan, Quercus sp., 18. vi. 1978, coll. P. Teocchi & D. Matile-Ferrero); c. K. roboris (BMNH 180; Germany, Q. sessiflora, 1914, coll. E. E. Green); d. K. vermilio (ICVI C: 5132; Italy: Bitonto (Bari), Q. ilex, 25. vii. 2012, coll. F. Porcelli).