Kermes echinatus 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 : 9-24

Kermes echinatus Balachowsky

(Figs. 1–8; Plate 1 , figs. a–i)

Kermes echinatus Balachowsky, 1953: 181 This species was originally described from first-instar nymphs collected at Nahalal forest, Israel, on Quercus coccifera L.

Material examined. Type material studied. Syntypes: 2 first-instar nymphs, Nahalal Forest, Israel, Q. coccifera 10.v.1950, Bytinski-Salz ( ICVI C:3691, MNHN 1065-8). Other material studied. All non-type material collected off Q. calliprinos in Israel by M. Spodek, unless otherwise indicated. Alonei Abba Nature Reserve, 19.vi.2011 (MC:530/ 2 adult ♀, USNM), 26.vi.2011 (C:4999/ 1 adult ♀, MC:549/ 3 adult ♀, MNHN), 3.vi.2012 (MC:711/ 3 adult ♀, BMNH); Eilon, 19.vi.2011 (MC:532/ 3 adult ♀, MC:533/ 4 adult ♀), 22.vi.2011 (MC:541/ 1 adult ♀, MC:542/ 2 adult ♀), 26.vi.2011 (C:4998/ 1 adult ♀, MC:548/ 4 adult ♀), 3.vi.2012 (MC:692/ 2 adult ♀); Nahal Dolev Nature Reserve, 17.vi.2010 (MC:261/ 1 adult ♀), 15.vi.2012 (MC:699/ 2 adult ♀), 17.vi.2011 (MC:528/ 1 adult ♀), 8.vi.2012 (MC:695/ 1 adult ♀), 22.vi.2012 (MC:709/ 4 adult ♀); Hanita, 6.vi.2010 (MC:227/ 1 adult ♀); Nebi Hazuri, 6.vii.2011 (MC:556/ 3 adult ♀); Eilon, 23.v.2011 (MC:508/2 third-instar ♀, MC:510/3 thirdinstar ♀), 12.vi.2011 (MC:531/1 third-instar ♀), 19.vi.2011 (MC:533/1 third-instar ♀, MC:544/4 third-instar ♀), 6.vii.2011 (MC:554/1 third-instar ♀), 3.vi.2012 (MC:691/3 third-instar ♀); Masada, 4.vii.2010 (MC:283/1 thirdinstar ♀); Nahal Dolev Nature Reserve, 17.vi.2010 (MC:257/1 third-instar ♀), 3.vi.2011 (MC:519/5 third-instar ♀), 10.vi.2011 (MC:521/7 third-instar ♀), 17.vi.2011 (MC:540/2 third-instar ♀), 1.vi.2012 (MC:675/3 third-instar ♀), 8.vi.2012 (MC:694/3 third-instar ♀); Nebi Hazuri, 6.vii.2011 (MC:555/7 third-instar ♀, MC:556/1 third-instar ♀); Eilon, 6.vi.2010 (MC:252/1 second-instar ♀), 17.v.2011 (MC:498/3 second-instar ♀), 23.v.2011 (MC:508/4 second-instar ♀), 1.vi.2011 (MC:512/4 second-instar ♀), 17.vi.2011 (MC:499/1 second-instar ♀), Nahal Dolev Nature Reserve, 17.vi.2010 (MC:258/1 second-instar ♀), 15.v.2011 (MC:498/2 second-instar ♀), 27.v.2011 (MC:518/2 second-instar ♀); Nebi Hazuri, 12.vi.2011 (MC:522/2 second-instar ♀), 19.vi.2011 (MC:537/3 secondinstar ♀), 26.vi.2011 (MC:545/1 second-instar ♀); Alonei Abba Nature Reserve, 15.vii.2010 (MC:289/6 firstinstars, USNM), 26.vi.2011 (MC:559/1 first-instar); Eilon, 26.ix.2010 (MC:306/8 first-instars, MNHN), 21.iv.2011 (MC:486/14 first-instars, BMNH), 17.v.2011 (MC:499/2 first-instars), 1.vii.2011 (MC:550/15 first-instars), 22.vii.2012 (MC:718/13 first-instars); Hanita, 6.vi.2010 (MC:247/4 first-instars), 13.iii.2011 (MC:457/7 firstinstars); Nahal Dolev Nature Reserve, 22.viii.2010 (MC:293/6 first-instars), 8.viii.2011 (MC:562/23 first-instars), 1.vii.2012 (MC:717/9 first-instars); Nebi Hazuri, 17.viii.2000, Y. Ben-Dov (C:3409/1 first-instar), 4.vii.2010 (C:4818/1 first-instar), 6.vii.2011 (C:5003/1 first-instar), 17.vii.2011 (MC:561/13 first-instars); Neve Zuf, 10.vii.2000 (C:4752/1 first-instar), 18.vii.2003 (C:4751/1 first-instar), Y. Ben-Dov; Ramah, 12.vii.2001 (MC:56/6 first-instars); Timrat, 21.vii.2011 (MC:563/12 first-instars), 25.iii.2012 (MC:651/3 first-instars); Alonei Abba Nature Reserve, 21.vi.2012 (C:5169/ 1 adult ♂, MC:733/ 4 adult ♂); Nahal Dolev, 1.vi.2012 (MC: 676/ 1 adult ♂), 9.vi.2012 (MC:732/ 7 adult ♂); Nebi Hazuri, 6.vii.2012 (MC:734/ 1 adult ♂); Alonei Abba Nature Reserve, 19.vi.2011 (MC:535/ 1 pupa), 3.vi.2012 (MC:673/ 7 pupae); Eilon, 23.v.2011 (MC:505/ 1 pupa), 1.vi.2011 (MC:513/ 2 pupae), 19.vi.2011 (MC:538/ 3 pupae); Nahal Dolev Nature Reserve, 10.vi.2011 (MC: 521/ 1 pupa), Nebi Hazuri, 19.vi.2011 (MC:536/ 1 pupa), 6.vii.2011 (MC:557/ 2 pupae). Alonei Abba Nature Reserve, 1.vi.2011 (MC:511/1 prepupa), 12.vi.2011 (MC:524/2 prepupae), 19.vi.2011 (MC:535/1 prepupa), Eilon, 23.v.2011 (MC:505/2 prepupae), 1.vi.2011 (MC:513/6 prepupae), 12.vi.2011 (MC:523/1 prepupa), 19.vi.2011 (MC:538/1 prepupa); Hanita, 24.v.2012 (MC:660/1 prepupa); Nahal Dolev Nature Reserve, 10.vi.2011 (MC: 521/1 prepupa); Nebi Hazuri, 19.vi.2011 (MC:536/1 prepupa); Alonei Abba Nature Reserve, 23.v.2011 (MC:507/3 second-instar ♂), 1.vi.2011 (MC:511/10 second-instar ♂), 3.vi.2012 (MC:671/ 1 second-instar ♂); Eilon, 23.v.2011 (MC:505/ 2 second-instar ♂), (MC:508/16 second-instar ♂), 1.vi.2011 (MC:512/9 second-instar ♂), (MC:513/6 second-instar ♂), 3.vi.2012 (MC:689/2 second-instar ♂); Hanita, 7.v.2012 (MC:277/3 second-instar ♂), 24.v.2012 (MC:659/2 second-instar ♂); Nahal Dolev Nature Reserve, 27.v.2011 (MC:509/14 second-instar ♂), 3.vi.2011 (MC:516/7 second-instar ♂), 10.vi.2011 (MC:521/5 second-instar ♂), 17.vi.2011 (MC:539/2 second-instar ♂), 1.vi.2012 (MC:677/6 second-instar ♂), 8.vi.2012 (MC:688/5 second-instar ♂); Nebi Hazuri, 19.vi.2011 (MC:536/4 secondinstar ♂); Timrat, 24.v.2012 (MC:656/9 second-instar).

Adult female (Fig. 1). General appearance. Pre-reproductive adult female ( Plate 1 , fig. c). Body oval to round and slightly convex; 2.5–3.2 mm long, 2.0–3.0 mm wide; dorsum brownish-grey with 4 or 5 black longitudinal lines and 6–9 black transverse lines formed of dots and lines; venter red. Gravid female ( Plate 1 , fig. d). Body highly convex; 2.9–4.4 mm long, 2.7–5.1 mm wide, 3.2–4.8 mm high; dorsum and venter brownishreddish-grey; dorsum with black, longitudinal and transverse lines. Post-reproductive female ( Plate 1 , fig. e). Body hemispherical and moderately convex, 3.0– 4.5 mm long, 3.0– 5.1 mm wide, 2.0–3.0 mm high; body sclerotized; dorsum red with 6–9 transverse black lines represented as reticulated folds.

Mounted young adult female. Body oval to circular; 2.0–3.0 mm long, 2.0– 2.8 mm wide. Margin. Marginal setae conical, each 12–13 µm long, 10–11 µm wide at base; arranged in a single row of 30–38 on each side, extending from cephalic apex to posterior end of body. Dorsal submarginal setae short and spinose; each 7–9 µm long, in a complete line of 28–33 on each side.

Dorsum. Setae conical, each 10–13 µm long, 7–10 wide at base, similar in shape and size to marginal setae, with 14–22 setae randomly placed on dorsum. Bilocular pores oval, each 3 µm long, 2 µm wide; present throughout. Tubular ducts of Type 2, each with outer ductule 12–17 µm long, inner ductule 10–15 µm long and with a sclerotized cup 5 µm diameter; present throughout dorsum.

FIGURE 1.

Venter. Eyes circular, each 20–25 µm diameter, anterolateral to each scape, near body margin. Legs absent. Antennae 1 segmented, 26–35 µm long, 20–31 µm wide, each bearing 5–8 fleshy setae; each scape surrounded by a group of 40–45 multilocular pores, each pore 7–8 µm diameter with 10 loculi. Tentorium 235–250 µm long, 212–225 µm wide. Labium 3 segmented, triangular, 160–175 µm long, 110–135 µm wide; labial setae as follows: basal segment with 2 pairs of setose setae 5–8 µm long; medial segment with a pair of setose setae 12–20 µm long, and apical segment with 4 pairs of setose setae 10–12 µm long. Spiracles subequal in size; each peritreme 50–68 µm long, 30–37 µm wide, in a group of 15–22 loculate pores laterad to each spiracle, each 8 µm diameter with 10 loculi; also with 2 pores with 6 loculi laterad to each anterior peritreme, each 6 µm diameter. Tubular ducts of Type 2, each with outer ductule 10–16 µm long, sclerotized cup 4–5 µm diameter and inner ductule 11–15 µm long, present in a complete, submarginal band 8–11 ducts wide and also sparsely distributed throughout rest of venter. Multilocular pores, each 10 µm diameter with 10–12 loculi, in transverse bands 2 to 3 pores wide on each abdominal segment, totaling 114–120 pores on each segment; also with a group of 52–56 pores just posterior to vulva. Bilocular pores, each 3 µm long and 2 µm wide, interspersed between tubular ducts in submarginal band. Microtubular ducts with circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; interspersed between multilocular pores on abdomen. Ventral setae of three types: (i) setose setae: with about 12 just anterior to tentorium between antennae, each 7–12 µm long; about 8 in median and submedian areas of thorax; and 6-8 setae present in a row on each abdominal segment, each 20–25 µm long; also 1 pair present just anterior to anal ring, each 10–12 µm long; 1 pair present posteriorly to anal ring, each 10–12 µm long; and a pair of apical setae, 33–35 µm long; (ii) conical setae: a pair present on venter slightly above posterior margin, each 10–12 µm long and wide; and (iii) tack-like setae, each 5 µm long, with about 11 on each side mesad to each complete submarginal band of tubular ducts. Microspines, each 1 or 2 µm long, in groups of 3–5 in 3–8 rows on each abdominal segment. Anal ring ventral, forming a complete sclerotized circle, 42–60 µm diameter; pores absent, but with 6 setae, each 25–40 µm long.

Comments. The adult female of K. echinatus , K. vermilio , a Mediterranean species not present in Israel (Spodek & Ben-Dov, 2012), and K. hermonensis sp. n. described below, all share the following features: (i) legs reduced or absent; (ii) Type 2 tubular ducts present on both venter and dorsum; (iii) dorsum without seta-pore clusters; (iv) anal ring ventral; (v) conical setae on margin and dorsum; and (vi) multilocular pores surrounding each antennal 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 and K. vermilio has pores without setae. They also differ in the number of antennal and leg segments: Kermes hermonensis has three, four, five or six-segmented antennae (segmentation obscure in other two species) and legs reduced to three segments, whereas the two other species have one-segmented antennae and lack legs.

Third-instar female ( Fig. 2; Plate 1 , fig. b). General appearance. Body broadly oval to round, slightly convex, 1.3–1.5 mm long, 1.0– 1.5 mm wide; dorsum orange-red-brown, venter red; dorsum covered with a thin layer of wax, composed of 5 longitudinal rows of rectangular wax plates, median row with 10 plates, mediolateral rows with 9–11 plates and marginal rows with 7–12 plates. Wax plates largest medially.

Mounted specimen. Body oval, 1.0– 1.5 mm long, 0.8–1.4 mm wide. Margin. Marginal setae conical, 10 µm long, 10 µm wide at base, in a complete single line of 30–34 on each side. Dorsal submarginal seta short and spinose, each 5 µm long, in a complete single line of 29–33 on each side.

Dorsum. Setae conical, each about 8 µm long, 5 µm wide at base, distinctly thinner than marginal conical setae; in 4 longitudinal medial and submedial rows, each row with 8 setae. Microtubular ducts with a circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; scattered on entire dorsum. Tubular ducts of Type 2, with inner ductule 13–15 µm long, inner cup about 5 µm wide and outer ductule 13–15 µm long, scattered on dorsum.

Venter. Eyes circular, 10 µm diameter, placed laterally to each scape, near body margin. Antennae 1 segmented, 25 µm long; with 7 fleshy setae on apical tip. Legs reduced to stumps, each 15–28 µm long; prothoracic pair always present, sometimes with 2 setae; other legs sometimes absent. Tentorium 112–180 µm long, 90–150 µm wide. Labium 3 segmented, triangular, 130–142 µm long, 70–98 µm wide; basal segment with 2 pairs of setose setae 5–8 µm long; medial segment with a pair of setose setae 12–20 µm long and apical segment with 4 pairs of setose setae 10–12 µm long. Spiracles subequal in size, each peritreme about 25 Μm long and about 18 Μm wide, each sclerosis about 50 µm long; each mesothoracic spiracle with 2 quinquelocular pores, and each metathoracic spiracle with 1 pore, each 3 µm diameter, placed anterolaterally, Tubular ducts of Type 2, with inner ductule 13–15 µm long, inner cup about 5 µm wide and outer ductule 13–15 µm long, arranged in a complete submarginal band about 5 ducts wide. Bilocular pores oval to circular, each about 3 Μm wide; dispersed among submarginal tubular ducts. Ventral setae of three types: (i) setose setae: with 6–8 setae anterior to tentorium, each about 18 µm long; 6–8 setae in a transverse row on each abdominal segment, each about 10 µm long; 4 setae posterior to anal ring, each 10 µm long, and 4 setae just anterior to anal ring: 2 setae about 13 µm long and 2 setae about 8 µm long; (ii) tack-like setae, each 5 µm long, present medial to each spiracle; also about 10 in a complete submarginal row; and (iii) conical setae, each 8 µm long and 8 µm wide at base, with 2 near margin on posterior apex of body. Microspines each about 2 µm long, present over entire body in groups of 1–8, but arranged in 2–7 transverse rows on abdominal segments. Anal ring ventral, about 35 µm in diameter, circular with posterior gap, each half with 3 spinose setae about 28 µm long; without pores.

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

Second-instar female ( Fig. 3; Plate 1 , fig. a). General appearance. Body oval, tapering posteriorly, 0.7–1.45 mm long, 0.4–0.8 mm wide; dorsum and venter orange-red; dorsum covered with a thin layer of wax, composed of 5 longitudinal rows of rectangular wax plates, each plate about 0.2 mm long and 0.1 mm wide; median row with 10 plates, mediolateral rows with 9–11 plates and marginal rows with 7–12 plates. Wax plates largest medially.

Mounted specimen. Body oval, 0.2–0.9 mm long, 0.4–0.7 mm wide. Margin. Marginal setae conical, each 10–13 µm long, 5–8 µm wide at base, in a complete single row of 30–33 on each side. Dorsal submarginal setae absent.

Dorsum. Setae conical, each about 8 µm long and about 5 µm wide at base, shorter than marginal setae, placed 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 complete, with about 7 pores, and each median row with about 6 pores extending from level of labium to level of anal ring.

Venter. Eyes circular, 15 µm wide, placed laterally to each scape near body margin. Antennae 1 segmented, 30–50 µm long, 10–20 µm wide; with 3 setae on basal part of each antenna, each 5 µm long, plus 6 setae on apex, each 10–13 µm long. Legs reduced to 1 segment, 25–40 µm long; with 3 short setae on basal part of each leg. Tentorium 125–130 µm long, 112–125 µm wide. Labium 3 segmented, triangular, 100 µm long, 50–85 µm wide; labial setae: basal segment with 2 pairs of setose setae about 5 µm long; median segment with a pair of setose setae 17–35 µm long, and apical segment with 4 pairs of setose setae 8 µm long. Spiracles subequal in size, each peritreme 7–13 Μm wide and sclerosis 28–32 µm long; each spiracle with 1 or 2 quinquelocular pores, 3 µm diameter, placed anterolaterally. Tubular ducts of Type 2, each with an inner ductule 10–15 µm long, inner cup 5 µm wide and outer ductule 10–15 µm long; arranged in a complete marginal band about 3 ducts wide and also sparsely over entire venter. Bilocular pores oval, each 7 µm long, 5 µm wide, in a line among the submarginal band of tubular ducts. Setae of three types: (i) conical setae, each 7 µm long, 5 µm wide at base, similar in shape to marginal setae, with 2 submarginally on anterior apex of body; (ii) setose setae: with 6 medially, just anterior to level of tentorium, each 38 µm long; also in medial and submedial longitudinal rows of 6 setae on abdominal segments, setae in medial rows about 15 µm long and those in submedial rows about 7 µm long; also with 4 setae placed anterior to anal ring: 2 about 15 µm long and 2 about 45 µm long; also 2 setae posterior to anal ring, each about 17 µm long; and posterior to anal ring: 2 setose setae about 50 µm long, and 2 conical setae, similar to marginal setae, each about 10 µm long, 5 µm wide at base; and (iii) tack-like setae, each 10 µm long, with about 15 setae on each side in a complete submarginal row. 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 27–30 µm, composed of 2 semi-circles, each half with 3 spinose setae 20–28 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) . Of the Israeli species, K. echinatus most closely resembles K. hermonensis but is also similar to K. vermilio (not found in Israel) in having conical marginal setae and reduced antennae and leg segments. However, K. echinatus has one-segmented antennae and one-segmented legs whereas K. hermonensis has 5- or 6-segmented antennae and 3-segmented legs; and K. echinatus lacks the membranous frontal lobes of K. vermilio . In addition, tubular ducts are present on both the venter and dorsum in K. echinatus and K. hermonensis , whereas they are only found on the venter in K. vermilio ; and K. echinatus has two longitudinal rows of conical setae on the dorsum while K. vermilio has one transverse row of setae on dorsum. Other Kermes spp. in Israel have different marginal setae: K. nahalali and K. greeni have robust setose setae and K. spatulatus have spatulate setae. These three species, K. greeni , K. nahalali and K. spatulatus , also have six-segmented antennae and well-developed legs, as in the Nearctic species K. cockerelli , K. concinnulus , Nanokermes iselini and Olliffiella secunda ( Baer & Kosztarab, 1985) .

First-instar nymph ( Fig. 4). General appearance. Body oval, tapering posteriorly, 0.37–0.44 mm long, 0.14–0.2 mm wide; dorsum and venter red; settled first-instar nymph with a fringe of curly, white-wax filaments.

Mounted specimen. Body oval, 0.45–0.49 mm long, 0.2–0.25 mm wide. Margin. Marginal setae conical and slightly curved apically, in 1 to 2 rows; 1 complete row of larger setae, each seta 10–13 Μm long, 5 Μm wide at base, with 17–22 on each side; plus second row of smaller setae, each lying between larger setae, extending from mesothorax to anal lobe, each seta 5–8 Μm long, 3–5 Μm wide at base, with 13–16 on each side.

Dorsum. Derm membranous, inter-segmental lines observable. Dorsal setae setose, with 8 setae in submedian longitudinal rows on thorax, each 6–8 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 7 in each submarginal row on abdomen.

Venter. Eyes circular, close to margin, each 10–12 Μm wide. Antennae 6 segmented; total length 102–110 Μm; segment III longer than other segments; 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 2 setose setae; apical segment with 2 fleshy setae and 5 setose setae. Legs well-developed; measurements (in Μm) of metathoracic leg: coxa 25–30, trochanter + femur 68–70, tibia 33–38, tarsus 45–50, claw 15–20; total leg length 187–200 Μm; trochanter with 2 oval sensory pores on each side; setae present on all leg segments; tarsal digitules knobbed apically, each 25–30 Μm long, extending beyond apex of claw; claw digitules knobbed apically, each 15–20 Μm long; each claw with a single denticle near tip. Tentorium 73–76 Μm long, 63–75 Μm wide. Labium 3 segmented, triangular, 75–83 Μm long, 45–47 Μm wide; labial setae as follows: basal segment with 2 pairs of setose setae 5–8 Μm long; median segment with 1 pair of setose setae on dorsal surface 12–13 Μm long; apical segment with 4 pairs of setose setae 16–20 Μm long. Spiracles subequal in size, each peritreme 3–5 Μm wide; each with apodeme crescent-shaped, 13– 15 Μm long; each spiracle with 1 quinquelocular pore, 5 Μm diameter, placed anterolaterally. Trilocular pores, each about 3 Μm wide, distributed as follows: a pair between scapes just anterior to tentorium; 1 mesad to each coxa; also pairs submedially on abdominal segments V–VII. Bilocular pores oval, each 3 Μm long, 2 Μm wide; with 1 pore present between margin and each spiracle. Ventral setae of two types: (i) conical setae: a pair on anterior apex of head, each 14–16 Μm long and 5 Μm wide at base; and (ii) setose setae: 6 inter-antennal setae in a longitudinal line medially between scapes, each 38–45 Μm long; also 1 seta, 10–11 Μm long, mesad to each coxa and associated with a trilocular pore; abdominal segments with medial, submedial and submarginal lines of setae, with 1 setae per line per segment; setae in medial and submedial rows each 10–15 Μm long, and submarginal setae each 5–6 Μm long. Also with a pair anterior to anal ring 15–18 Μm long, and a pair latero-posteriorly to anal ring 15–20 Μm long. Microspines, each about 3 Μm long, arranged in 3 or 4 groups in 2 transverse rows on each abdominal segment and sparsely on thorax. Anal ring ventral, composed of 2 semi-circles, diameter 20–25 Μm, each half with 4–6 pores and 3 spinose setae, each 13–18 Μm long. Anal lobes slightly developed; apical margin of each lobe with 1 spinose seta, 10–13 Μm long, 2 or 3 Μm wide, and 1 very long, flagellate seta, 220–275 Μm long.

Comments. Kermes echinatus was first described by Balachowsky (1953) based on the first-instar nymph from Israel. Balachowsky suggested that this species was similar to K. vermilio , a Mediterranean and European species, not known from Israel. The first-instar nymphs of both species are bright red and possess conical marginal and dorsal setae. The morphological characters that distinguish them were discussed in Spodek & Ben-Dov (2012). The first-instar nymph of K. hermonensis , described as new in this paper, is also red and possesses dorsal and marginal conical setae similar to those on K. echinatus and K. vermilio . However, K. echinatus can be distinguished from the two other species by the presence of only one quinquelocular pore associated with each spiracle and the absence of dorsal bilocular pores. Kermes hermonensis has two quinquelocular pores associated with the prothoracic spiracles, usually with one pore larger than the other, whilst K. vermilio has usually one, rarely two, locular-pores, each with 5–8 loculi associated with each spiracle and several dorsal bilocular pores ( Balachowsky, 1950; Pellizzari et al., 2012).

Adult male ( Fig. 5 ; Plate 1 , fig. i). General appearance Newly emerged males alate, elongate and widest across mesothorax; body red, 1.5–2.0 mm long from head apex to apical tip of penial sheath; fore-wings 1.2–1.5 mm long, 0.45–0.75 mm wide; radial wing vein red; with 5 pairs of simple eyes; body with rather few setae fleshy setae (fs) few and hard to separate from hs; pores absent apart from in glandular pouches; hamulohalteres present; abdomen slender, glandular pouches present, extruding white, waxy caudal filaments, 1.0– 1.5 mm long; penial sheath longer than broad.

Mounted specimen. Total length 1.5–2.0 mm long, width across mesothorax 0.4–0.5 mm.

Head. Broadly oval in dorsal view; distinctly separated from thorax by a cervical groove or constriction. Length from head apex to pronotal ridge (prnr) 275–300 µm long; width across genae 275–315 µm. Median crest not reticulated. Postoccipital ridge (por) heavily sclerotized, with anterior arm longer than posterior arm. Midcranial ridge (mcr) with well-developed lateral branches to each antenna; dorsal arm absent. Ocular sclerites (ocs) sclerotized, not reticulated, with 5 pairs of simple eyes and one pair of ocelli (o); dorsal (dse) and ventral simple eyes (vse) largest, each 45–50 µm diameter; 3 pairs of lateral simple eyes (lse), each 35 µm in diameter; ocellus 13–18 µm diameter. Dorsal head setae (dhs) in a longitudinal row on either side of median crest, with 4 or 5 hs in each row. Ventral head setae (vhs) in a transverse row anterior to ventral simple eyes, total 8–10 hs. Genae (g) unsclerotized, not reticulated and without setae. Preoral ridge (pror) well-developed. Anterior tentorial pits and head pores absent. Antennae: filiform, 10 segmented, length 1000–1050 µm (ratio of total body length to antennal length about 1:0.58). Scape (scp) 43–50 µm long, 55–63 µm wide, with 2 or 3 hair-like setae (hs). Pedicel (ped) 50–63 µm long, 40–50 µm wide, with 5–8 hs. Flagellar segments III–X 93 –165 µm long, 20–25 µm wide; segment III longest, segments becoming shorter towards the apex; approximate number of setae per segment (note: fs and hs hard to differentiate): III–VII 15–23 fs; VIII 11–15 fs; IX 14–16 fs + 1 antennal bristles (ab); X 10–14 fs + 2–4 ab +4 capitate setae (cs), each cs 30–40 µm long.

Thorax. Prothorax: pronotal ridge (prnr) sclerotized but not fused medially. Pronotal sclerites (prn) present; without lateral or medial pronotal setae. Proepisternum + cercival sclerite (pepcv) well-developed and connected anteriorly to the postocular ridge (pocr). Sternum (stn1) membranous with a transverse sternal ridge; median ridge absent. Sternal, prosternal and anteprosternal setae absent.

Mesothorax: prescutum (prsc) rectangular with a median groove, 70–75 µm long, 150–165 µm wide; not reticulated; with lateral prescutal ridges (pscr) and with a posterior prescutal suture. Scutum (sct) sclerotized, median membranous area 60–65 µm long, with lateral sclerotized margins, without setae. Prescutal and scutal setae absent. Prealare (pra) and triangular plate (tp) present. Scutellum (scl) 60–65 µm long, 145–150 µm wide; without a foramen but with strong scutellar ridges (sclr); marginal fold of notum (rd) present posteriorly; with 2 hs, each 12–15 µm long. Basisternum (stn2) 158–200 µm long, 250–270 µm wide; without a median ridge, but marginal ridge (mr) well developed with a strong precoxal ridge (pcr2); basisternal setae absent. Furca (f) well-developed and sclerotized, arms divergent and extending about halfway to marginal ridge (mr) anteriorly. Lateropleurite (lpl) wide, with a strong extension from marginal ridge anteriorly. Mesopostnotum (pn2) not reticulated; with a welldeveloped postnatal apophysis (pna). Mesepisternum (eps2) not reticulated; sub-episternal ridge (ser) well developed. Mesothoracic spiracles (sp2): peritreme 10–15 µm wide, without loculate pores. Tegula (teg) distinct, with 3–5 hs tegular setae (tegs).

Metathorax: metapostnotum, metatergal setae and dorso-spiracular setae not detected. Ventrally with a pair of transverse sclerotized precoxal ridges (pcr3) not connected to pleural ridge (plr3). Sternal apophyses (stn3a) present. Metathoracic spiracles (sp3): peritreme 10–15 µm wide; without loculate pores. Microspines 1–2 µm long, singly or in groups up to three, in about 10 transverse rows on venter. Metapleural ridges (plr3) well developed, extending anteriorly to hamulohalteres. Metepimeron (epm3) sclerotised, but without setae. Metasternal setae absent.

Wings: hyaline, 1200–1500 µm long, 450–475 µm wide (ratio of length to width 1:0.38; ratio of total body length to wing length 1:0.7); alar lobe (al) well-developed; alar setae absent. Hamulohalteres (h) 113–125 µm wide, 28–40 µm long, with a single apical hamulus, 55–65 µm long.

Legs: metathoracic legs longest, 763–836 µm long; prothoracic and mesothoracic legs 768–810 µm long. Coxa: I: 88–100; II: 107–110; III: 110–115 µm long, with 12–15 hs. Trochanter: I: 50–52; II: 50–63; III: 60–63 µm long, with about 8 hs; each trochanter with 3 oval sensory pores in a triangle on each surface. Femur: I: 225–230; II: 205–213; III: 210–218 µm long, with 27–30 hs. Tibia: I: 275–287; II: 275–280; III: 280–300 µm long, with 53– 56 hs; all tibia with 2 apical spurs (tibs) 20–25 µm long. Tarsi 2-segmented; tarsus I ring-like, tarsus 2 (tar2): I: 100– 113; II: 100–110; III: 100–110 µm long, with 20–30 hs; tarsal digitules (tdgt) setose and knobbed apically, not extending beyond claw, each about 40 µm long. Claw (cl) with a denticle: I: 25–28; II: 26–28; III: 25–30 µm long; claw digitules (cdt) knobbed apically, not extending beyond claw, each about 30 µm long.

Abdomen: Tergites and pleural areas not sclerotized; sternites (as) sclerotized on segments IV–VIII; caudal extension of segment VII absent. Loculate pores absent on segments I–VII. Dorsopleural setae (adps) (each side) 20–25 µm long: segments I–VII: 2 hs, segment VIII: 3 hs, one 75–80 µm long and two 25–30 µm long; ventropleural setae (avps) (on each side) 15 µm long, on each side, on segments IV–VII: 1 hs; dorsal setae (ads) (total), 10 µm long, segments I–VII: 2 hs, segment VIII: 4 hs, 40 µm long; and ventral setae (avs), (total), 20–25 µm long, segments II–III: 2 setae, segment IV–VII: 4 setae. Caudal extension on segment VIII rounded. Glandular pouch (gp) present on segment VIII, deep, each pouch with 2 long, robust setae (gls) 150–155 µm long; each pouch with 9–18 pores, each 3 µm diameter with 3–5 loculi. Microspines 1–2 µm long, singly or in groups of up to 3 in about 10 transverse rows on each abdominal segment on venter. Genital segment: penial sheath (ps) sclerotized, 170–180 µm long, greatest width 110–120 µm. Anal opening not detected. Basal rod (bra) sclerotized, 40–50 µm long. Aedeagus (aed) elongate 100–120 µm long, not extending beyond penial sheath. Genital capsule with a total of 8 hs (gts) dorsally and 16–20 hs (gts) ventrally; apex of ps with a cluster of about 10 small, circular sensilla.

Comments. Of the Israeli species, the adult male of K. echinatus is most similar to those of K. hermonensis — both are red in life, with reddish wing veins, and lack metasteranal setae, which are present in most other species (e.g., Allokermes kingii ( Hamon et al., 1976) , K. greeni , K. nahalali , K. quercus ( Koteja & Zak-Ogaza, 1972) , K. spatulatus and N. balachowskii ). For differences with other species, see key.

Fourth-instar male (pupa) ( Fig. 6; Plate 1 , fig. h). General appearance. Develops within a white, flocculent (wooly) wax test secreted by second-instar male; body red and elongate oval when removed from test; 1.2–1.5 mm long, 0.4–0.6 mm wide; derm membranous with distinct inter-segmental lines on dorsum and venter; anterior wingbuds extending laterally past coxa of metathoracic legs; posterior wing-buds very short; penial lobe triangular.

Mounted specimen. Body 1.3–1.5 mm long. 0.5–0.6 mm wide; head 225–250 µm long, 375–400 µm wide, slightly demarcated from thorax; thorax and abdomen not clearly demarcated. Margin. Wing-buds: fore-wings 485–550 µm long and 165–200 µm wide; hind-wings: length ratio to mesothoracic wings 1:10. Setae fine and blunt, in marginal and submarginal rows, each with 7 setae segmentally arranged, extending from abdominal segments I–VII each 12–15 µm long.

Dorsum. Eyes absent. Ocular sclerite reticulated, 45–50 µm long, 150–158 µm wide, extending transversely from body margin to mid-body; also present on venter. Setae setose, each 5 µm long, distributed as follows: in rows of 6 on head apex and also in complete submedial longitudinal rows, each row with about 6 setae. Anal lobes rounded, membranous, each with 1 setose seta on apex, 55–95 Μm long. Median penial lobe triangular, extending past anal lobes, slightly sclerotized, each 95–105 Μm long, 95–115 Μm wide; with 3 pairs of setose setae, each 15– 20 Μm long.

Venter. Ocular sclerite as 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; pores absent. Legs well-developed; subequal in size, measurements (in Μm) of metathoracic leg; coxa 75–100, trochanter + femur 155–200, tibia 140–175, tarsus 90–110; total length 455–505 Μm; each coxa with 2 short setae, each 5 µm long; distal end of tarsus with short pointed, triangular claw, 5 µm long. Ventral setae setose; interantennal setae in 2 rows of 3, each 5 Μm long; also in medial, submedial and submarginal rows on abdominal segments II–VII, each 15–25 Μm long, each row with 6 setae segmentally arranged. Microspines, each about 3 Μm long, arranged in groups of 3 or 4 in 9 or 10 transverse rows on each abdominal segment and scattered on thorax.

Comments. The pupae of K. echinatus are most similar to those of K. hermonensis and both are red in life, but those of K. echinatus differ in lacking quinquelocular pores anterior to each spiracle. The pupae of other Israeli species are differently coloured, e.g. K. greeni is orange, and K. nahalali and K. spatulatus are greyish-brown. The pupae of the Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) also lack spiracular pores but their colour was not recorded.

Third-instar male (prepupa) ( Fig. 7; Plate 1 , fig. g). General appearance. Develops within a white, waxy test secreted by second-instar male; body red and broadly oval when removed from test, 1–2 mm long, 0.4–0.8 mm wide; derm membranous, with distinct inter-segmental lines on both dorsum and venter; anterior wing-buds well developed; posterior wing-buds absent.

Mounted specimen. Body 1.1–1.38 mm long, 0.5–0.6 mm wide; separation between head, thorax and abdomen not clearly demarcated. Margin. Anterior wing-buds each 250–325 µm long, 140–165 µm wide, extending to about level with metathoracic coxa. Setae fine and blunt, each 12–15 µm long, in a complete marginal row of 15–20 on each side; and in a submarginal row on abdominal segments I–VII, similar in size and shape to marginal setae.

Dorsum. Setae setose, each 5–8 µm long, with 3 pairs on head apex; also with complete submedial longitudinal rows, segmentally arranged, each with about 10 setae. Anal lobes rounded and membranous, each with 2 setae on apex, one 35–40 Μm long and other 16–20 Μm long. Median penial lobe broadly rounded 95–110 Μm long, 50–75 Μm wide; with a total of 8 setose setae, each 5–10 Μm long.

Venter. Eyes absent. Antennae 7 or 8 segmented, 250–290 µm long, 65–80 µm wide; apical segment rounded; setae absent. Spiracles subequal in size; each peritreme 15–20 µm long, 10–15 µm wide, with sclerosis 30–35 µm long; each spiracle with 2 quinquelocular pores anterolaterally, each 5 Μm diameter. Legs present as 3-segmented lobes, subequal in size; total length of metathoracic leg, 175–250 Μm, width 60–75 Μm; claw and setae absent. Ventral setae setose, 8–10 Μm long; with 3 pairs of setae between scapes; abdomen with medial, submedial and submarginal longitudinal lines of setae, each line with 6 setae, segmentally arranged. Microspines, each about 3 Μm long, arranged in groups of 3 or 4 in 9 or 10 transverse rows on each abdominal segment and scattered on thorax.

Comments. The prepupa of K. echinatus is most similar to those of K. hermonensis : both are red in life and both have two quinquelocular pores anterior to each spiracle. Other Israeli prepupae are differently coloured, e.g. K. greeni orange, and K. nahalali and K. spatulatus greyish- brown; and all three species lack spiracular pores. The prepupa of N. balachowskii is brown and has one quinquelocular pore anterior to each spiracle. The prepupae of the Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) lack spiracular pores but their colour in life was not recorded.

Second-instar male ( Fig. 8; Plate 1 , fig. f). General appearance. Secretes a white, waxy test, 1.5–1.8 mm long, 0.6–0.8 mm wide; when removed from test, body red, broadly oval, 0.7–1.45 mm long, 0.4–0.8 mm wide; derm membranous, inter-segmental lines observable on both venter and dorsum.

Mounted specimen. Body oval; 0.9–1.5 mm long, 0.5–0.8 mm wide. Margin. Marginal setae conical, each 10 µm long and 8 µm wide at base, in a complete single row of 30–34 on each side.

Dorsum. Setae of two types: (i) conical setae in a submedial longitudinal row of 4 setae on head and each thoracic segment, each about 7 µm long and about 5 µm wide at base, shorter than marginal setae; and (ii) tack-like setae in a medial longitudinal row of 6–8 setae, extending from mid-thorax to anal ring, each 8 µm long. Microtubular ducts, each with a circular opening about 5 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, 15–20 µm diameter, placed laterally to each scape, near body margin. Antennae 7 segmented, total length 155–185 µm; segment III longer than other segments; scape and pedicel each with 2 thin 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 membranous, one mesad to each scape, each 45–80 Μm long, 35–75 Μm wide, not always visible. Legs well-developed; subequal in size, measurements (in Μm) of metathoracic legs: coxa 50–75; trochanter + femur 110–125; tibia 50–60; tarsus 70–85 and claw 20–25; total length 300–325 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 denticle. Tentorium 113–125 µm long, 100–125 µm wide. Labium 3 segmented, triangular, 90–110 µm long and 85–90 µm wide; labial setae: basal segment with 2 pairs of setose setae about 5 µm long, median segment with 1 pair of setose setae 17–35 µm long and apical segment with 4 pairs of setose setae 8 µm long. Spiracles subequal in size, each peritreme 10–13 Μm wide, with sclerosis 28–35 µm long; with 4 quinquelocular pores extending in a transverse row, from peritreme to margin of body, each 4 µm diameter. Tubular ducts of Type 2, each with outer ductule 10–14 Μm long, a sclerotized cup about 5 Μm in diameter and inner ductule 8–10 Μm long, in a complete submarginal band 1 or 2 ducts wide. Bilocular pores circular, 2 µm diameter, with 10–12 pores on each side of body, in a complete submarginal line. Ventral setae of three types: (i) conical setae, similar in shape and size to marginal setae, with a pair on anterior apex of body, each 10 µm long and 5 µm wide at base; also 2 about 15 µm long and 5 µm wide at base, posterior to anal ring; (ii) setose setae: with 5 pairs medially, just anterior to tentorium, each 42–63 Μm long; with 4 medial to each coxa, with two 13–15 µm long and two 25–40 µm long; also in medial and submedai rows on abdominal segments II–VII, setae in medial rows each about 28 µm long, setae in submedial rows each about 10 µm long; also anterior to anal ring, with 4 setae, two about 25 µm long and two about 13 µm long; plus 2 setae posterior to anal ring, each about 25 µm long; and (iii) tack-like setae, each 5 µm, long in a complete submarginal row of 10–15. Microspines, each about 3 µm long; in groups of 1 or 2 in 6–8 transverse rows on each abdominal segment and scattered on thorax. Anal ring ventral, diameter 30–35 µm, composed of 2 semi-circles, each half with 3 spinose setae, 32–43 µm long, and 9–12 pores.

Comments. Among the Palaearctic Kermes species, second-instar males have only been described for K. bytinskii (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) , plus the four other species from Israel, K. greeni , K. hermonensis , K. spatulatus and N. balachowskii . All bar K. hermonensis and K. vermilio clearly differ from K. echinatus because their marginal setae are robustly setose (conical in K. echinatus and K. vermilio ). The secondinstar males of the Nearctic K. cockerelli , K. concinnulus and K. rimarum have long, setose marginal setae, tubular ducts on the dorsum and quinquelocular pores scattered on both the dorsum and venter ( Baer & Kosztarab, 1985).

The second-instar males of K. echinatus , K. hermonensis and K. vermilio are similar in possessing conical marginal setae and a pair of membranous frontal lobes between the antennal scapes. These lobes were first described by Williams (1985) on several Eriococcidae species and later were recognized by Marotta & Tranfaglia (1999) and Pellizzari et al., (2012) on K. vermilio . These authors also found them in second-instar and prepupal males and in the second- and third-instar females. The function of the frontal lobes has not been established, but it might indicate a close relationship between Eriococcidae and Kermesidae (Marotta & Tranfaglia, 1999) . Apart from these three species, frontal lobes are unknown in the Kermesidae .

The second-instar males of K. echinatus differs from those of K. hermonensis and K. vermilio as follows (characters of K. hermonensis and K. vermilio in brackets): (i) ventral quinquelocular pores absent anterior to tentorium (present); (ii) each prothoracic spiracle with four quinquelocular pores (eight and two); and (iii) ventral abdominal quinquelocular pores absent (present in four longitudinal rows ventrally on abdominal).

Figures

PLATE 1. Kermes echinatus Balachowsky, general appearance; a. second-instar female; b. third-instar female; c. young adult female; d. gravid female; e. post-reproductive female; f. second-instar male; g. prepupa; h. pupa; i. adult male.

FIGURE 5. Kermes echinatus Balachowsky, adult male.