Kermes nahalali 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 : 51-65

Kermes nahalali Bodenheimer

( Figs. 25–32; Plate 4 , figs. a–i)

Kermes nahalali Bodenheimer, 1931: 241

Kermes bytinskii Sternlicht, 1969: 253 n. syn.

This species was originally described from adult females collected in Nahalal forest, Israel, on Quercus coccifera L. The junior synonym, K. bytinskii , was described from adult females collected at Tivon, Israel, on Q. ithaburensis.

Material examined. Type material studied. Lectotype: K. nahalali adult ♀ (here designated, ICVI) and 4 adult ♀. Paralectotypes: Israel, Nahalal, off Q. coccifera in v.1920 by F.S. Bodenheimer. Also paratype: K. bytinskii Sternlicht , adult ♀, Tivon, Israel, Q. ithaburensis,. iii.1957, M. Sternlicht ( BMNH, 1969-627); Paratype: K. bytinskii 5 first-instar nymphs, Tivon, Q. ithaburensis, iii.1957, M. Sternlicht ( BMNH, 1969-627/1 slide). Other material studied. All non-type material collected in Israel, off Q. ithaburensis by M. Spodek, unless otherwise indicated. Alon Hagalil, 25.iv.2010 (MC:170/ 2 adult ♀); Alonei Abba Nature Reserve, 25.iv.2010 (MC:171/ 4 adult ♀, BMNH), 27.ii.2011 (MC:422/ 3 adult ♀, MNHN), 13.iii.2011 (MC:423/ 6 adult ♀), 14.ii.2012 (MC:610/ 4 adult ♀, USMN), 4.iii.2012, Y. Ben-Dov (C:5083/ 1 adult ♀), 24.iii.2012 (MC:642/ 4 adult ♀); Bet Dagan-Volcani Center, 19.ii.2012 (MC:625/ 1 adult ♀); Horshat Tal Nature Reserve, 20.ii.1968, Y. Ben-Dov (C:5249/ 1 adult ♀), 6.ii.2011 (MC:414/ 1 adult ♀), 21.ii.2010 (MC:119/ 1 adult ♀), 27.ii.2011 (MC:428/ 5 adult ♀), 14.ii.2012 (MC:609/ 2 adult ♀), 29.ii.2012, Y. Ben-Dov (C:5081/ 1 adult ♀); Tivon, 15.vi.1955, M. Sternlicht (C:5049/ 1 adult ♀); Alonei Abba Nature Reserve, 11.i.2012 (MC:597/2 third-instar ♀, MC:599/3 third-instar ♀), 14.ii.2012 (MC:609/4 third-instar ♀, MC:610/2 third-instar ♀), 3.i.2013 (MC:805/5 third-instar ♀), 13.i.2013 (MC:784/2 third-instar ♀); Horshat Tal Nature Reserve, 11.i.2011 (MC:386/6 third-instar ♀, MC:387/2 third-instar ♀), 6.ii.2011 (MC:414/3 third-instar ♀), 14.ii.2011 (MC:609/4 third-instar ♀), 27.ii.2011 (MC:430/6 third-instar ♀, MC:428/4 third-instar ♀); Alonei Abba Nature Reserve, 14.ii.2012 (MC:634/1 second-instar ♀), 14.10.2012 (MC:735/10 second-instar ♀), 18.xi.2012 (MC:751/4 second-instar ♀); Bet Dagan-Volcani Center, 15.i.2012 (MC:605/2 second-instar ♀); Horshat Tal Nature Reserve, 11.i.2011 (MC:386/1 second-instar ♀, MC:387/2 second-instar ♀), 14.ii.2012 (MC:623/7 second-instar ♀); Yehudiya Nature Reserve, 6.ii.2011 (MC:412/2 second-instar ♀); Alon Hagalil, 8.iv.2010 (MC:150/12 first-instars, BMNH), 23.iv.2010 (MC:190/20 first-instars, MNHN); Alonei Abba Nature Reserve, 1.iv.2010 (MC:148/25 first-instars, USMN), 25.iv.2010 (MC:171/2 first-instars), 17.x.2010 (MC:320/2 first-instars, MC:322/8 first-instars, MC:323/1 first-instar, MC:325/2 first-instars, MC:333/1 first-instar), 7.xi.2010 (MC:349/1 first-instar), 21.iv.2011 (MC:494/83 first-instars), 16.iv.2012 (MC:649/20 first-instars); Horshat Tal Nature Reserve, 12.v.2000, Z. Tamari (MC:3411/3 first-instar), 6.vi.2010 (MC:294/5 first-instars), 29.viii.2010 (MC:300/4 first-instars), 10.x.2010 (MC:326/6 first-instars, MC:327/7 first-instar, MC:328/1 first-instar, MC:329/ 3 first-instars, MC:331/5 first-instars), 7.xi.2010 (MC:357/3 first-instars, MC:361/4 first-instars); Neve Yaar, 16.v.2001, Y. Ben-Dov (C:4755/8 first-instars); Timrat, 25.iv.2012 (MC:650/13 first-instars); Yehudiya Nature Reserve, 10.x.2012 (MC:313/10 first-instars); Alonei Abba Nature Reserve, 10.ii.2013 (MC:823/ 2 adult ♂), 17.ii.2013 (MC:839/ 7 adult ♂); Horshat Tal Reserve, 17.ii.2013 (MC:862/ 1 adult ♂); Bet Dagan-Volcani Center, 15.i.2012 (MC:603/ 5 pupae); Hanita, 1.v.2012 (MC:816/ 5 pupae); Alonei Abba Nature Reserve, 3.i.2013 (MC:805/5 prepupae); Bet Dagan-Volcani Center, 15.i.2012 (MC:603/2 prepupae), 17.ii.2012 (MC:616/3 prepupae); Alonei Abba Nature Reserve, 11.i.2012 (MC:599/1 second-instar ♂), 18.xi.2012 (MC:751/2 secondinstar ♂); Bet Dagan-Volcani Center, 15.i.2012 (MC:604/4 second-instar ♂), 19.i.2012 (MC:604/1 second-instar ♂), 17.ii.2012 (MC:616/7 second-instar ♂); Horshat Tal Nature Reserve, 11.i.2011 (MC:386/1 second-instar ♂), 14.ii.2012 (MC:623/2 second-instar ♂); Yehudiya Nature Reserve, 3.vi.2012 (MC:707/2 second-instar ♂).

Adult female ( Fig. 25). General appearance. Pre-reproductive adult female ( Plate 4 , fig. c). Body oval to round, 2.0– 3.7 mm long, 2.0–4.0 mm wide, 1.5–3.1 mm high; dorsum and venter orange to brown, with 6–8 medial dark, circular spots. Gravid female and post-reproductive female ( Plate 4 , fig. d). Body 3.5–5.0 mm long 3.5– 5.0 mm wide, and 3.8–5.0 mm high; grey, occasionally orange, spherical, derm sclerotized with a deep black, longitudinal depression dividing body into 2 parts, depression composed of 6–8 circular dark spots, each about 0.3 mm in diameter; also with 6–8 transverse rows of 13–15 small dark spots, each about 0.1 mm in diameter.

Mounted young adult female. Body oval to round, 1.5–3.5 mm long, 1.4–3.0 mm wide. Margin. Marginal setae short and spinose, each 10 Μm long, in a single complete row of 15–21 on each side.

Dorsum. Dorsal median spots circular, each 125–475 Μm diameter, composed of a central circular depression 13–50 Μm diameter, surrounded with polygonal areas, each 10–20 Μm long, 5–15 Μm wide; with 6–8 spots in a longitudinal, medial row. Dark-rimmed areas each 13–25 Μm long and 15–20 Μm wide, on dorsum in about 8 curved rows of 5–15 areas, extending from anal opening to anterior apex of body. Bilocular pores oval, each 5 Μm long, 3 Μm wide, abundant throughout 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, dispersed over entire dorsum. Seta-pore clusters with a cluster of 1–5 quinquelocular pores, each 5 Μm wide, and a seta 15 um long, in 10–15 clusters on each side, extending in a single complete submarginal row. Anal ring dorsal, 75–85 Μm diameter, circular with anterior gap; without setae or pores. Anal lobes with 2 sclerotized areas posterior to anal ring, probably referable to anal lobes, each 38–40 Μm long, 18–37 Μm wide, with 5 setose setae 22–47 Μm long.

Venter. Eyes absent. Antennae 5 or 6 segmented; when 5 segmented, segment II and III appearing fused; 102– 137 Μm long; segment III longest; setal distribution as follows: scape and pedicel each with 2 setose setae; segment III with a setose seta; IV with a fleshy seta; V with 2 fleshy setae; and 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 30–50, trochanter + femur 45–62, tibia 38–62, tarsus 38–62, claw slightly pointed 17–20 µm long; total length 212–250 µm; each trochanter with 2 oval, sensory pores on each side; tarsal digitules 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; denticle absent. Tentorium 360–375 Μm long, 200–270 Μm wide. Labium 3 segmented, triangular, 175–210 Μm long, 125–150 Μm wide; labial setae setose. as follows: basal segment with 2 pairs about 5 Μm long; median segment with a pair about 15 Μm long, and apical segment with 4 pairs about 17 Μm long. Mesothoracic spiracles clearly smaller than metathoracic spiracles; peritreme of mesothoracic spiracles 50– 100 Μm long, 38–50 Μm wide; peritreme of metathoracic spiracles 87–150 Μm long, 75–90 Μm wide. Tubular ducts of 2 types: Type 1 with an inner ductule 12–15 µm long, inner cup 10 µm wide and outer ductule 20–25 µm long, arranged in a complete submarginal band about 20–30 ducts wide; and Type 2 with inner ductule 12 µm long, inner cup 12 µm wide and outer ductule 12 µm long, a few medially between legs and spiracles. Quinquelocular pores, each 5 Μm diameter: with about 20 pores surrounding both spiracles; also with 6–8 on each abdominal segment, plus 50–100 pores posterior to vulva. Multilocular pores, each 10 Μm diameter with 9–11 loculi: with 6– 10 anterior to each metathoracic spiracle; plus 6–8 clusters of 15–30 pores on each abdominal segment anterior to vulva, each pore 7 Μm diameter. Ventral setae of two types: (i) setose setae: with 6–10 setae on each abdominal segment, each 5 Μm long; and 35–50 setae posterior to vulva, each 23 Μm long; and (ii) tack-like setae, each 3 Μm long, in groups of 7–10 setae between each leg. Microspines, each 3 Μm long, arranged in groups of 3 or 4 in about 8 rows on each abdominal segment and dispersed on thorax.

Comments. Kermes bytinskii is here synonymized with K. nahalali because, according to Bodenheimer (1931), the post-reproductive female of K. nahalali has "five to six brown spots present in a median, longitudinal furrow...and the body is kidney or heart-shaped" (translated from German). These traits, which Bodenheimer described and illustrated, can be seen on the paratype adult female of K. bytinksii and on fresh material that we recovered from the type locality of both species.

Although Sternlicht (1969) described the third-instar and adult females of K. bytinskii and suggested that these stages had the same structure, his descriptions and illustration of the former are here considered to represent a young adult female. The present redescriptions are based on fresh specimens plus type material of the adult females of K. bytinskii and K. nahalali (see Material examined).

The adult female K. nahalali shares the following characters with K. greeni and K. spatulatus : (i) fullydeveloped 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) dorsal anal lobes sclerotized, and (vii) two types of tubular ducts on venter. These characters are also present on the adult female of several Oriental Kermes species: K. flavus ( Liu & Shi, 1995) , K. miyasakii ( Liu & Shi, 1995) , K. orientalis ( Liu & Shi, 1995) and K. taishanensis ( Hu, 1986) . The adult female of K. nahalali can be distinguished from K. greeni in having (characters of K. greeni in brackets): (i) narrow anal lobes with up to five setae on each lobe (wide, quadrate lobes with 10–15 setae on each lobe); (ii) two types of tubular ducts ventrally and none dorsally (Type 2 on dorsum and Type 1 on venter); (iii) presence of darkrimmed areas on the dorsum (absent); (iv) presence of tack-like setae near each coxa (absent); (v) presence of microducts on dorsum (absent); and (vi) spiracles of different-sizes (spiracles subequal). The adult female of K. nahalali can be distinguished from K. spatulatus in having (characters of K. spatulatus in brackets): (i) dorsal median spots (absent); (ii) dorsal reticulated areas absent (present); and (iii) lack of multilocular pores posterior to each leg (present).

Third-instar female ( Fig. 26; Plate 4 , fig. b). General appearance. Body convex, 1.3–2.8 mm long, 1.3–2.2 mm wide, 0.8–1.5 mm high; dorsum grey to brown, with 5 longitudinal rows of rectangular wax plates, median wax plates largest.

Mounted specimen. Body oval, 1.4–3.0 mm long, 1.2–1.8 mm wide.

Margin. Marginal setae short and spinose, each 13 µm long, 2 µm wide, in a single complete line of 28–30 on each side. Dorsal submarginal setae short and spinose, each 8 µm long, 3 µm wide, in a single complete line of 23–25 on each side.

Dorsum. Seta-pore clusters restricted to a single complete submarginal band of 18–20 clusters on each side, each cluster with 1–5 quinquelocular pores, each pore 5 Μm wide and each seta 12–15 um long. Bilocular pores oval to circular, each 4 µm long and 3 µm wide; abundant over dorsum. Anal ring dorsal, horseshoe-shaped with anterior gap, 40–42 µm wide; setae and pores absent. Anal lobes represented by sclerotized areas, each about 50 µm long, 30 µm wide, each with 3 or 4 setose setae 38–50 µm long. Also with a pair of setose setae, posterior to anal lobes, 10–13 µm long.

Venter. Eyes circular, each 15 µm diameter, lateral to each scape. Antennae 6 segmented, 95–100 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; V with 2 fleshy setae; and apical segment with 4 setose setae and 1 fleshy seta. Legs welldeveloped; measurements (in µm) of metathoracic leg: coxa 30–50, trochanter + femur 60–80, tibia 50–75, tarsus 60–70, claw 15–18; total length 220–290 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 21–25 Μm long, knobbed apically, extending slightly beyond claw apex; claw digitules, knobbed apically, each 10– 15 Μm long, shorter than tarsal digitules; claw denticle absent. Tentorium 225–250 µm long, 180–200 µm wide. Labium 3 segmented, triangular, 137–155 µm long, 115–150 µm wide; labial setae setose, basal segment with 2 pairs 5 Μm long; medial segment with 1 pair about 15 µm long and apical segment with 4 pairs about 19 µm long. Spiracles peritreme of mesothoracic spiracles smaller than those of metathoracic spiracles; mesothoracic peritremes 30–35 Μm long, 20–25 Μm wide; metathoracic peritreme 40–45 Μm long, 25–30 Μm wide; scleroses of both spiracles 40–60 µm long. Tubular ducts of two types: Type 1 with inner ductule 11–14 µm long, inner cup about 9 µm wide and outer ductule 18–20 µm long, arranged in a broad complete submarginal band, 8–10 ducts wide; Type 2 ducts with inner ductule 10–12 µm long, inner cup about 5 µm wide and outer ductule 10–12 µm long, arranged in a submedial band 3–6 ducts wide from cephalic apex to most anterior abdominal segment. Quinquelocular pores, each 5 µm diameter; with 3–8 pores anterolateral to each spiracle; and 4–8 pores medially and submedially on each abdominal segment. Ventral setae of two types: (i) setose setae: in groups of 3–5 posterior to each coxa, each 5–10 µm long; also in a transverse row of 10–12 on each abdominal segment, each 10– 13 µm long; and (ii) tack-like setae, each 3 µm long, with 1 seta submarginally on each abdominal segment II–VII. Microspines, each 2 µm long, in groups of 1–4 in transverse rows over entire body.

Comments. Of the Israeli species, the third-instar female of K. nahalali is very similar to those of K. greeni and K. spatulatus , all sharing the following traits (characters for K. echinatus , K. hermonensis and N. balachowskii in brackets): (i) marginal setae short and spinose (obviously conical); (ii) legs fully-developed (leg segmentation reduced); (iii) anal ring dorsal (ventral); and (iv) anal lobes sclerotized on dorsum (absent). The third-instar female of K. nahalali can be distinguished from those of K. greeni and K. spatulatus as follows: K. nahalali has up to three setae on each anal lobe, K. greeni has about fifteen setae on each lobe, and K. spatulatus has no anal lobe setae. K. nahalali also has a complete submarginal line of seta-pores while K. spatulatus has seta-pore clusters only on the head and thorax and they are absent in K. greeni . Of the species found outside Israel, third-instar nymphs have only been described for K. vermilio , from which K. nahalali differs in having (character-states of K. vermilio in brackets): six-segmented antennae (one-segmented antennae), and legs fully-developed (one-segmented). Among the Nearctic Kermes , only the third instar of K. sylvestris has been described and illustrated ( Bullington & Kosztarab, 1985), and is similar to K. nahalali in having a ventral submarginal band of tubular ducts and a submarginal line of seta-pore clusters on the dorsum but differs in having (characters for K. sylvestris in brackets): (i) a sclerotized anal lobe (absent); (ii) fully-developed legs (reduced); (iii) bilocular pores on dorsum (absent), and (iv) two types of tubular ducts on venter (one type).

Second-instar female ( Fig. 27; Plate 4 , fig. a). General appearance. Body oval, tapering posteriorly, 0.6–1.0 mm long, 0.3–0.6 mm wide; dorsum and venter white, 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– 11 plates, submedian row with 9–11 plates and marginal rows with about 9–12 plates. Wax plates largest medially.

Mounted specimen. Body oval, 0.9–1.3 mm long, 0.54–0.75 mm wide.

Margin. Marginal setae robust and spinose, each about 25 µm long, 8 µm wide at base, with 20–26 on each side extending around entire margin; also with a submarginal band of 13–15 shorter robust spinose setae, each about 20 µm long and 5 µm wide at base, on each side.

Dorsum. Setae setose, each 5 µm long, about 10 in submedial lines. Microtubular ducts each with a circular opening about 3 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long, in 8 compete longitudinal rows, each row with about 20 pores. Tubular ducts of Type 2, with inner ductule 15–18 µm long, inner cup 5 µm wide and outer ductule 15–19 µm long, scattered over entire dorsum.

Venter. Eyes circular, each 15 µm wide, lateral to each scape. Antennae 6 segmented, length 150 µm; scape and pedicel each with 2 setose setae, segment III with 1 setose seta, IV with 1 fleshy seta, V with 2 fleshy setae, and VI with 4 fleshy setae and 1 setose seta. Legs well-developed; measurement (in Μm) of metathoracic legs; coxa 30– 45, trochanter + femur 88–12, tibia 45–50, tarsus 50–75, claw 20–25; total length 233–320 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 30–42 Μm long, knobbed apically, extending beyond claw apex; claw digitules knobbed apically, each 28–30 Μm long, shorter than tarsal digitules; each claw with a single denticle near tip. Tentorium 132–150 µm long, 120–125 µm wide. Labium 3 segmented, triangular, 110–125 µm long, 85–110 µm wide; labial setae setose, as follows: basal segment with 2 pairs about 3 µm long, median segment with 1 pair about 25 µm long and apical segment with 4 pairs about 18 µm long. Spiracles: each mesothoracic peritreme 10–15 Μm wide, sclerosis 18–25 µm long; each metathoracic peritreme larger, 15–20 Μm wide, each sclerosis 25– 30 µm long. Tubular ducts of Type 2, with inner ductule 15–18 µm long, inner cup 5 µm wide and outer ductule 15–19 µm long, arranged in a complete submarginal band about 3–5 ducts wide. Quinquelocular pores, each 5 µm diameter; with 6 pores anterior to tentorium; 1 pore anteriorlaterally to each mesothoracic spiracle and 5 or 6 pores anterior to each metathoracic spiracle; and each abdominal segment with a transverse band of 4 pores in medial and submedial lines. Ventral setae of three types: (i) robust spinose setae, similar to marginal setae, with 6 anterior to scape, each about 25 µm long; (ii) setose setae: with 3 pairs medially just anterior to tentorium, each about 35 µm long; single setae medial to each coxa about 10 µm long; and with medial and submedial rows on abdominal segments II–VIII, medial setae each about 12 µm long and submedial setae each about 8 µm long. Also, with a pair of setose setae just anterior to anal ring about 12 µm long; and (iii) tack-like setae; each about 5 µm long, in a submarginal row from metathorax to anal ring, with about 9 setae on each side. Microspines, each about 3 µm long, in groups of 2–5 in 5–8 transverse rows on each abdominal segment and scattered on thorax. Anal ring ventral, diameter 37–40 µm, composed of 2 semi-circles, each half with 3 spinose setae 50–75 µm long, and about 25 pores. Anal lobes very well-developed, each with 1 spinose seta on inner margin, about 25 µm long, plus another on apex, about 20 µm long, and also with a long flagellate seta 68–88 µm long on apex.

Comments. The second-instar female of K. bytinskii , now a junior synonym of K. nahalali , was first described by Sternlicht (1969). No material from this description was available for examination and therefore the present redescription is based on fresh specimens and includes several features not originally described, such as the presence of: (i) dorsal microtubular ducts; (ii) a pair of sensory pores on trochanter; (iii) a claw denticle; (iv) microspines on venter of abdominal segments; (v) longitudinal rows of setose seta on dorsum; (vi) only one type of tubular duct, and not two types of ducts as suggested by Sternlicht, (1969), and (vii) quinquelocular pores at the base of each scape and in peristigmatic areas.

Second-instar female K. nahalali are very similar to those of K. greeni in having spinose marginal setae and well-defined anal lobes. However, K. nahalali has four quinquelocular pores on the venter of each abdominal segment whereas K. greeni has only two. Also, the second-instar female of K. nahalali does not have frontal lobes as in K. vermilio . Tubular ducts are present on both the dorsum and venter in K. nahalali (as in K. echinatus , K. hermonensis and K. quercus (Podsiadlo, 2012)) . Kermes nahalali also has six- segmented antennae and welldeveloped legs, as on several Nearctic species, including K. cockerelli , K. concinnulus and Nanokermes iselini and Olliffiella secunda ( Baer & Kosztarab, 1985) , in contrast to those of K. vermilio , which has one-segmented legs ( Pellizzari et al., 2012) and K. quercus which has three-segmented legs ( Podsiadlo, 2012).

First-instar nymph ( Fig. 28). General appearance. Body oval, tapering posteriorly with distinct anal lobes, 0.38–0.46 mm long, 0.18–0.2 mm wide; dorsum and venter grey.

Mounted specimen. Body oval, 0.44–0.49 mm long, 0.18–0.20 mm wide.

Marginal setae more or less in 2 rows: setae in more marginal row robust spinose, in a complete row of 19–21 setae on each side, each 10–15 µm long; and with smaller, less robust setae in a second incomplete row of 9–11 smaller setae on each side, extending posteriorly from mesothorax, each 5–8 Μm long.

Dorsum. Derm membranous, with inter-segmental lines. Setae setose, each about 11 µm long; in submedial longitudinal rows of 10 setae. Microtubular ducts, with a circular opening about 2 Μm diameter, inner ductule about 1 Μm long and outer ductule 0.5 Μm long; in longitudinal submarginal rows of about 9 pores, and medial rows of about 6 pores, from abdominal segment I to level of anal ring.

Venter. Eyes each 12 µm wide, present close to margin. Antennae 6 segmented; length 100–110 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 2 fleshy setae, and apical segment with 1 fleshy seta and 4 setose setae. Legs well-developed; measurements (in Μm) of metathoracic leg: coxa 25–37, trochanter + femur 75–85, tibia 35–40, tarsus 50–60, claw 15–23 µm long; total length 221–232 µm; trochanter with 2 oval sensory pores on each side; setae present on each leg segment; tarsal digitules 27–33 µm long, knobbed apically, extending slightly beyond apex of claw; claw digitules knobbed apically, each 17–25 µm long, shorter than tarsal digitules; each claw with a single denticle near tip. Tentorium 88–100 µm long, 70–75 µm wide. Labium 3 segmented, triangular, 75–90 µm long, 37–47 µm wide, with 14 setose labial setae: basal segment with 2 pairs 5–10 µm long, median segment with a pair 10–13 µm long, and apical segment with 4 pairs 12–17 µm long. Spiracles subequal in size, each peritreme 3–5 Μm wide, and sclerosis 15–20 µm long; with 3 quinquelocular pores anterolateral to each prothoracic spiracle and 1 pore anterolateral to each mesothoracic spiracle, each 2 Μm wide. Trilocular or quinquelocular pores, each 2 µm diameter; with a pair just anterior to tentorium; 1 pore medially to each coxa; and 2 pores present submedially on abdominal segments II–VI. Ventral setae of three types: (i) robust spinose setae, similar to marginal setae, 3 pairs anterior to scapes, each 12–18 µm long; (ii) setose setae: with 3 pairs medially between scapes, each 22–25 µm long; 1 seta medial to each coxa, 10–11 µm long; and with a seta medially and submedially on each abdominal segment II–VII, each 7–8 Μm long; and (iii) tack-like setae, each 5–6 Μm long, in a submarginal line on abdominal segments II–VII. Bilocular pores oval, each 3 µm long, 2 µm wide; 1 pore present between margin and each spiracle. Microspines, each 1 µm long, arranged in 2 transverse rows medially and submedially on each abdominal segment. Anal ring ventral, total diameter 17–20 µm, composed of 2 semi-circles, each half with about 6 pores and 3 spinose setae, each 35–37 µm long. Anal lobes well-developed, each inner margin with 2 setae, similar to marginal setae, each 15–19 µm long, plus a long flagellate, apical seta 157–200 µm long.

Comments. The first-instar nymph of K. bytinskii was first described by Sternlicht (1969). One paratype slide (see Material examined) was available but the specimens are in poor condition. Therefore, the present redescription is based primarily on fresh specimens and includes several new features, such as the presence of: (i) dorsal microtubular ducts; (ii) microspines on venter of abdominal segments; and (iii) one bilocular pore between each spiracle and body margin.

The first-instar nymph of K. nahalali differs from those of the other Israeli species in having well-developed anal lobes (poorly developed on the other Israeli species). The marginal setae in K. nahalali are spinose and slightly curved apically, resembling those found in the first-instar of N. balachowskii . However, K. nahalali has a marginal and submarginal row of setae, whereas N. balachowskii only has a single marginal row. Sternlicht (1969) suggested that the first-instar nymph of K. bytinskii resembled K. ilicis (Linnaeus) , a Mediterranean and European species not found in Israel, although he did note several differences between the two species. We also consider these two species to be very similar. We studied the redescriptions of K. ilicis by Balachowsky (1950) and Borchsenius (1960), and have examined slide-mounted material of K. ilicis (see Material examined) and we found several characters of K. ilicis not previously mentioned, such as presence of: (i) dorsal setose setae; (ii) dorsal microtubular ducts; (iii) microspines on venter of abdominal segments; and (iv) only one bilocular pore between each spiracle and body margin. The first-instar nymphs of these two species also share the same type of marginal setae, the same distribution of ventral tri- and quinquelocular pores and both have well-developed anal lobes. They differ in the number of quinquelocular pores associated with each anterior spiracle: K. nahalali has three compared to two in K. ilicis . Despite this great similarity, apparently they can be separated at the female post-reproductive stage, as this stage in K. ilicis is described as spherical in shape, with a red-brown to brown-black dorsum and sixsegmented antennae and fully-developed legs ( Leonardi, 1920; Borchsenius, 1960) ( Plate 7 , fig. b), whereas the post-reproductive female of K. nahalali is grey, sometimes orange-brown, and the body is heart-shaped, with a medial depression. It too has six-segmented antennae and fully-developed legs ( Sternlicht, 1969; present study) ( Plate 4 , figs. d–e). Thus, the colour and shape of the post-reproductive female may be useful in distinguishing between K. ilicis and K. nahalali . Future studies of K. ilicis may suggest other traits to separate these two species.

Adult male ( Fig. 29; Plate 4 , fig. i). As for K. echinatus except: General appearance. Newly emerged males brown; body 2.0– 2.5 mm long, width across mesothorax 0.5–0.6 mm wide; wings 1.2–1.5 mm long and 0.45–0.75 mm wide; waxy caudal filaments 1.0– 1.5 mm long.

Mounted specimen. Total length 2.0– 2.5 mm long, width across mesothorax 0.5–0.8 mm.

Head. Length 330–340 µm long; width across genae 290–300 µm. Dorsal (dse) and ventral simple eyes (vse), each 38–40 µm diameter; lateral simple eyes smaller (lse), each 22–25 µm diameter; ocellus 16–18 µm diameter. Dorsal head setae (dhs) in rows on either side of median crest, each with 4–6 hs setae. Ventral head setae (vhs) in a transverse band of a total 10–12 hs. Antennae: 783–1000 µm long (ratio of total body length to antennal length about 1:0.4). Scape (scp) 43–50 µm long, 34–40 µm wide, with 2–4 hs. Pedicel (ped) 50–63 µm long, 40–50 µm wide, with 4–6 hs. Flagellar segments III–X each 52–136 µm long and 20–25 µm wide; segment III longest, with segments becoming shorter towards apex; approximate number of setae per segment: III–VII 15–23 fs VIII–X 10– 16 fs; VIII 1 ab and X 2 ab + 4 cs, each cs 35–40 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) 75–80 µm long, 180–200 µm wide. Scutum (sct): median membranous area 60–65 µm long, without setae but with 2 pairs of hair-like setae laterad to membranous area, each 12–15 µm long. Scutellum (scl) 80–90 µm long, 150–200 µm wide, without setae. Basisternum (stn2) 158–200 µm long, 250–270 µm wide; basisternal setae absent. Tegula (teg) with 5 or 6 hs tegular setae (tegs). Mesothoracic spiracles (sp2): peritreme 10–15 µm wide, without loculate pores.

Metathorax: Metathoracic spiracles (sp3): peritreme 10–15 µm wide. Metasternum (stn3) with a total of 34–40 hs setae: 8–12 anterior metasternal setae (amss); 10–18 posterior metasternal setae (pmss) and 3–9 postmetaspiracular setae (eps3s), each 20–25 µm long.

Wings: 1200–1500 µm long, 450–475 µm wide (ratio of length to width 1:0.37; ratio of total body length to wing length 1:0.55). Hamulohalteres (h) 150–152 µm long and 50–55 µm wide; hamulus, 55–65 µm long.

Legs: metathoracic legs longest, 724–847 µm long; prothoracic and mesothoracic legs 768–810 µm long. Coxa: I 60 –65; II 80 –85; III 94 –98 µm long, with 2–13 hs. Trochanter: I 50 –55; II 50 –63; III: 60–63 µm long, with 2–7 hs; each trochanter with 3 oval sensory pores in a triangle on each surface. Femur: I 232–235; II 205–213; III 244–250 µm long, with 22–35 hs. Tibia: I 296–300; II 293–300; III 294–300 µm long, with 29–43 hs; all tibia with 2 apical spurs (tibs) 20–25 µm long. Tarsi 2-segmented; tarsus 2 (tar2): I 100–113; II 105–110; III 100–110 µm long, with 9–18 hs; tarsal digitules (tdgt) not extending beyond claw, each about 40 µm long. Claw (cl) with denticle: I 27–29; II 26–28; III 30–34 µm long; claw digitules (cdt) not extending beyond claw, each about 30 µm long.

Abdomen. Sternites (as) lightly sclerotized on segments V–VIII; tergites and pleural areas unsclerotized; 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–VII 2 hs, 20–25 µm long, VIII 3 hs, one 85–90 µm long plus others 25– 30 µm long; ventropleural setae (avps) (on each side) IV–VII 1 hs, 15 µm long; abdominal ventral setae (avs) (totals) II–III 12–20 hs, IV 8 hs, V–VIII 4 hs 20–25 µm long. Glandular pouch (setae (gls) 142–145 µm long; each pouch with 10–12 pores, each 3 µm diameter with 3–5 loculi. Genital segment: penial sheath (ps) 170–190 µm long, greatest width 110–120 µm. Basal rod (bra) 40–50 µm long. Aedeagus (aed) 100–120 µm long. Genital capsule with a pair of hair-like setae on dorsum and 4 or 5 pairs of hair-like setae (gts) ventrally.

Comments. The adult male of K. bytinskii , now a junior synonym of K. nahalali , was first described by Sternlicht (1969) from Israel. No male type material of K. bytinskii is available and therefore the present redescription is based on fresh specimens and includes several features not originally noted: (i) absence of pores on anterior and posterior metasternal regions (several were recorded by Sternlicht, 1969); (ii) abdominal tergites not sclerotized, only sternites are sclerotized; and (iii) presence of microspines on venter of abdominal segments and metathorax.

The adult male of K. nahalali is very similar to those of K. greeni and K. spatulatus , but can be distinguished by having only one dorsopleural seta on abdominal segment I ( K. spatulatus has eight and K. greeni has four). These three species plus N. balachowskii have metasternal setae, while K. echinatus , K. hermonensis and E. gillettei , a North American species ( Miller & Miller, 1993) do not. Kermes nahalali has more than 25 metasternal setae, and more than two dorso-pleural setae on abdominal segments II–VII. Another Palaearctic species not present in Israel, K. quercus ( Koteja & Zak-Ogaza, 1972) , also has 28–40 metasternal setae but can be distinguished from K. nahalali by the more abundant ventral head setae (28–42) and dorsal head setae (13–23) ( K. nahalali has 10–12 and 4–6 setae respectively).

Fourth-instar male (pupa) ( Fig. 30; Plate 4 , fig. h). 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 1.3–1.5 mm long, 0.5–1.0 mm wide; head 225–135 µm long, 425–450 µm wide.

Margin. Wing-buds: fore-wings 550–600 µm long, 200–210 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. O cular sclerite reticulated, 45–50 µm long, 140–158 µm wide. Setae setose, each 5 µm long, distributed as follows: 4 pairs on head apex, and in submedial longitudinal rows, each row with about 6 setae. Anal lobes slightly developed, rounded and membranous; each with 1 setose seta 65–95 Μm long on apex. Median penial lobe triangular, 95–115 Μm long, 95–115 Μm wide; with 3 pairs of setose setae, each 13–20 Μm long.

Venter. Ocular sclerite reticulated as on dorsum. Antennae 10 segmented, length 475–500 µm, width 30–45 µm; setae absent. Spiracles subequal in size, each peritreme 25–30 µm long and 20–30 µm wide; each sclerosis 35–40 µm long; without associated pores. Legs well-developed, subequal in size; measurements (in Μm) of metathoracic leg: coxa 75–90, trochanter + femur 170–185, tibia 155–165, tarsus 100–110, total length 515–540 Μm; each coxa with 2 short setae, each 5 µm long; claw 5 µm long. Ventral setae setose, with 4 pairs of interantennal setae each 5 Μm long, and abdominal seta arranged as on K. echinatus . Microspines, as on K. echinatus .

Comments. The pupa of K. nahalali is greyish-brown in life, similar to those of N. balachowskii and K. spatulatus (other Israeli kermesids: K. echinatus and K. hermonensis red, and K. greeni orange). As with all other Israeli pupae (bar K. hermonensis ) those of K. nahalali lack quinquelocular pores anterior to each spiracle, and has membranous lobes, similar to other kermesid pupa apart from K. greeni and K. spatulatus , which have sclerotized anal lobes.

Third-instar male (prepupa) ( Fig 31; Plate 4 , fig. g). General appearance. As for K echinatus apart from: General appearance. Body brown and oval when removed from test, 1.0– 1.5 mm long, 0.5–0.8 mm wide.

Mounted specimen. Body 1.25–1.75 mm long, 0.54–0.9 mm wide.

Margin. Anterior wing-buds 355–450 µm long, 135–200 µm wide. Setae fine and blunt, each 12–15 µm long, in 2 rows as on K. echinatus .

Dorsum. Setae setose, each 5–8 µm long, with 4 pairs on head apex, and with submedial longitudinal rows as on K. echinatus . Anal lobes 50–80 Μm long, 90–110 Μm wide, each with 3 pairs of setose setae, each 5–10 Μm long; also with 2 pairs of similar setae just anterior to median penial lobe, each 5–10 Μm long.

Venter. Eyes absent. Antennae 7 or 8 segmented, tapering apically, 250–275 µm long, 70–80 µm wide; with a short fleshy seta on segments V and VI and 5 on apical segment, each about 5 µm long. Spiracles subequal in size, each peritreme 20–25 µm long, 10–15 µm wide, each sclerosis 30–40 µm long; without pores associated with spiracles. Legs: length of metathoracic leg, 190–260 Μm, width 70–75 Μm; claw and setae absent. Ventral setae setose, each 8–10 Μm long; with 4 pairs between scapes; abdominal setae arranged as on K. echinatus . Microspines, as on K. echinatus .

Comments. The prepupa of K. nahalali is greyish-brown in life, similar to those of N. balachowskii and K. spatulatus (other Israeli prepupae: K. echinatus and K. hermonensis red and K. greeni orange). The prepupae of K. nahalali lack quinquelocular pores anterior to each spiracle, as also in K. greeni , K. spatulatus and the Nearctic A. kingii ( Hamon et al., 1976) and E. gillettei ( Miller & Miller, 1993) . Kermesid prepupae that have one or two quinquelocular pores anterior to each spiracle are K. echinatus , K. hermonensis and N. balachowskii .

Second-instar male ( Fig. 32; Plate 4 , fig. f). General appearance. Secretes a white, flocculent (wooly) wax test, 1.5–1.9 mm long, 0.6–0.9 mm wide; body brown, broadly oval when removed from test, 1.0– 1.5 mm long, 0.4–0.8 mm wide; derm membranous, with inter-segmental lines.

Mounted specimen. Body oval, 1.5–2.0 mm long, 0.6–1.0 mm wide.

Margin. Marginal setae robust and setose, more or less in complete double rows, marginal setae in a row of 23–30 on each side, each 25 µm long, and in a submarginal row of 20–25 on each side, each 15 µm long.

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

Venter. Eyes circular, each 25 µm diameter, lateral to each scape. Antennae 7 segmented, length 175–200 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, VI with 1 fleshy seta and 1 setose seta, and apical segment with 4 fleshy setae and 3 setose setae. Frontal lobes absent. Legs well-developed; measurements (in Μm) of metathoracic legs: coxa 40–60; trochanter + femur 125–150; tibia 70–80; tarsus 95–100 and claw 25–30; total length 300–350 Μm; trochanter with 2 oval sensory pores on each side; tarsal digitules 30–35 Μm long, knobbed apically; claw digitules knobbed apically, each 20–30 Μm long, extending beyond apex of claw; each claw with a single denticle. Tentorium 135–200 µm long, 100–125 µm wide. Labium 3 segmented, triangular, 110–125 µm long, 60–85 µm wide; labial setae setose: basal segment with 2 pairs about 10 µm long, median segment with 1 pair 20–28 µm long, and apical segments with 4 pairs 15–22 µm long. Spiracles subequal in size, each peritreme 10–13 Μm wide, each sclerosis 25–30 µm long; each with 6–8 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, in a complete submarginal band 1 or 2 ducts wide. Bilocular pores circular, each 2 µm diameter, with 10–12 pores on each side, in a single complete submarginal band. Quinquelocular pores, each 5 µm wide; with 4 pairs between setae anterior to tentorium, and with a pair submedially on each abdominal segment. Ventral setae of three types: (i) robust setose, similar to marginal setae, with 3 pairs anterior to scapes, each about 30 µm long; (ii) setose setae: with 3 pairs of setae just anterior to tentorium, each about 40 µm long; 3 setae mesad to each coxa, each 25–50 µm long; and medially and submedially on abdominal segments II–VII, each about 45 µm long medially, and about 10 µm long submedially. Also with 2 pairs anterior to anal ring, each 20–25 µm long; (iii) tacklike setae, each 5 µm long, in a complete submarginal row of about 10–15 setae on each side. Microspines, each about 3 µm long, in groups of 3–5 in 9 or 10 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 spinose setae, 43–50 µm long, and 18–20 pores. Anal lobes developed; each inner margin with a spinose seta, 15–20 µm long at base, and another 13–15 µm long on apex; also a long flagellate setae, 100–200 µm long, on apex.

Comments. The second-instar male nymph of K. bytinskii was first described by Sternlicht (1969). No material was available and therefore the present redescription is based on fresh specimens and includes several new features, such as the presence of: (i) antennae seven-segmented; (ii) one type of tubular duct on dorsum and venter (not two types of ducts as suggested by Sternlicht, 1969); (iii) simple dorsal pores; (iv) a pair of sensory pores on trochanter; (v) a denticle on the claw; (vi) microspines ventrally on abdominal segments; (vii) bilocular pores interspersed ventrally among the submarginal band of tubular ducts; and (viii) four pairs of quinquelocular pores at the base of each scape.

Of the species described in Israel, the second-instar male of K. nahalali most closely resembles K. greeni because of the presence of anal lobes and pointed marginal setae. The other Israeli species ( K. spatulatus , K. hermonensis , K. echinatus and N. balachowskii ) lack obvious anal lobes. Of the other Palaearctic species, K. williamsi ( Sternlicht, 1970) , K. quercus ( Podsiadlo, 2012) and K. vermilio ( Pellizzari et al., 2012) also lack anal lobes. Of the Israeli species, Kermes nahalali can be distinguished from all bar N. balachowskii by the presence of quinquelocular pores in two longitudinal rows on venter of abdominal segments, but K. nahalali has eight loculate pores associated with each anterior spiracle and N. balachowskii has three. In addition, K. nahalali has four pairs of quinquelocular pores anterior to the tentorium, whilst these are absent in N. balachowskii . The Nearctic K. cockerelli , K. concinnulus and K. rimarum have long, setose marginal setae, and quinquelocular pores scattered on both the dorsum and venter ( Baer & Kosztarab, 1985), whereas K. nahalali lacks quinquelocular pores on the dorsum.

Figures

PLATE 4. Kermes nahalali Bodenheimer, general appearance; a. second-instar female; b. third-instar female and male test; c. young adult female; d. post-reproductive female, dorsal view; e. post-reproductive female, lateral view; f. second-instar male; g. prepupa; h. pupa; i. 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).