Kermes greeni Bodenheimer

Spodek, Malkie, Ben-Dov, Yair & Protasov, Alex, 2012, Taxonomy of Kermes greeni Bodenheimer (Hemiptera: Coccoidea: Kermesidae) with a new synonymy, Zootaxa 3545, pp. 67-75 : 68-73

Kermes greeni Bodenheimer

Kermes greeni Bodenheimer, 1931: 241

Kermes palestiniensis Balachowsky, 1953: 186 n. syn.

This species was originally described from an adult female collected from Nahalal forest, Israel, on Quercus coccifera L. The junior synonym was described from first-instar nymphs collected from Ras-El Nakura [=current name Rosh Hanikra], Israel, on Q. coccifera .

Material examined: Kermes greeni : Lectotype adult female of K. greeni (here designated and deposited at ICVI), off Q. coccifera , ISRAEL, Nahalal, 2.ii.1926, F.S. Bodenheimer. Paralectotypes: 7 adult females (6 in ICVI, 1 in BMNH), same data as lectotype. Also: 5 first-instar nymphs in 2 of the paralectotype adult females, ISRAEL: Nahalal, K. greeni , 2.ii.1926, F.S. Bodenheimer. Also: Syntype first-instar nymphs of Kermes palestiniensis Ras-el Nakurah [=current name Rosh Hanikra], Q. coccifera , 12.vii.1950, S. Neumark ( MNHN).

Additional non-type material of adult females and first-instar nymphs, all collected off Q. calliprinos by M. Spodek (unless other noted), all material deposited at ICVI: ISRAEL: Eilon , 13.iii.2011 ( ICVI MC:459); Gesher Alkosh, 16.vii.2001, Y. Ben-Dov ( ICVI C:4765); Hanita, 5.vi.2001, Z. Tamari ( ICVI C:4701); 6.vi.2010, 24.i.2011, 8.v.2011, 17.v.2011, 3.vi.2011, 7.v.2012, 3.vi.2012 ( ICVI MC:247, MC:405, C:4997, C:5011, MC:706, MC:668, MC:520); Timrat, 24.vi.2011, 31.vii.2011, 22.iv.2012, 7.v.2012, 30.v.2012 ( ICVI MC:567, MC:568, C:5096, C:5099, MC:705) .

Also: Paratype adult female of K. bytinskii Sternlicht, Tivon, Q. ithaburensis, iii.1957, M. Sternlicht ( BMNH, 1969-627) .

Adult female ( Fig. 1).

General appearance. Dorsum of young adult females with red and black areas and with a transverse red area medially; dorsum round and smooth. Body 2.5–3.7 mm long, 1.9–3 mm wide ( Plate 1a). Gravid female dark red to black and uniform in colour ( Plate 1b). Post-reproductive female. Orange to tan coloured; body spherical and derm smooth and sclerotized. Body 4.2–5.9 mm long, 2.9–5 mm wide, 3.8–4.3 mm high ( Plate 1d). Mounted adult female. Oval to round, 2.5–3 mm long, 1.8–2.5 wide.

Margin. Marginal setae pointed; each about 5 µm long, in a single row of 15–21 on each margin. Stigmatic setae absent.

Dorsum. Bilocular pores oval, each 3 µm long and about 2 µm wide, abundant throughout dorsum. Tubular ducts sparse, much less frequent than bilocular pores, each inner ductule 7–17 µm long, inner cup 5–8 µ m wide and outer ductule 12–15 µm long. Setae 17–25 um long, restricted to a complete submarginal band from cephalic apex to anal lobe, each seta with a cluster of 4–14 loculate pores, each about 8 µm wide with 3–5 loculi, with 10–13 clusters on each side of body. Anal ring dorsal, closed and circular, without setae or cells; diameter 75–100 µm. With 2 quadrate, sclerotized areas, probably referable to anal lobes, present posterior to anal ring, each 50–75 µm long and 63–75 µ m wide with 13–16 hair-like setae placed on and surrounding each area; each seta 30–43 µ m long.

Venter. Eyes absent. Antenna each 4–6 segmented; when 4-segmented, segments I, II and III appear fused; in 5-segmented antennae, segments II and III appear fused; each antenna 105–110 µm long; setal distribution as follows: scape with 1 thin hair-like seta; pedicel with no setae; segment III with 1 fleshy seta; segment IV with 2 fleshy setae; segment V with 1 fleshy seta; apical segment with 5 hair-like setae. Legs well-developed, tibia and tarsus occasionally fused, measurements of hind legs (µm): coxae 30–35, trochanter + femur 70–75, tibia 37–50, tarsus 40–48, claw blunt, each 10–18 µm long; total length of leg 212–250; each trochanter with 2 oval, sensory pores on each side, each pore 4–5 µm long and 2–3 µm wide; tarsal digitules each 25–38 µm long, knobbed apically and extending beyond apex of claw; claw digitules knobbed apically, each 22–25 µm long, shorter than tarsal digitules. Clypeolabral shield 250–287 µm long, 237–250 µm wide. Labium 3-segmented, triangular, 162–212 µm long and 125–162 µm wide; labial setae: basal segment with 4 hair-like seta 3–5 µm long, median segment with 2 hair-like seta 7–8 µm long, and apical segment with 8 setae, each 10–13 µm long. Spiracles subequal in size, peritreme 62–87 µm long and 45–55 wide. Multilocular pores widespread of rather variable size: (i) pores each 5–7 µm in diameter with 5–6 loculi, present in a group of 35–65 surrounding each spiracle; (ii) pores each 7–8 µm in diameter with 9–11 loculi, in 6 or 8 clusters of 30–60 pores on abdominal segments III–VII; (iii) 12–24 other pores on each abdominal segment, each 7 µm in diameter with 5 or 6 loculi, not in clusters; (iv) pores in submedial groups of 64–104 posterior to vulva, each 7–8 µm in diameter with 5 or 6 loculi, and (v) a medial group of 52–60 pores, posterior to vulva, each 6 µm diameter with 4–7 loculi. Tubular ducts in a submarginal band about 10 ducts wide; each duct with inner ductule 7–17 µm long, 5 µm wide, and outer ductule 22–30 µm long. Each abdominal segment with 2 setae, 23–25 µ m long, placed submedially, plus 8 or 10 setae, each 10–12 µ m long, placed medially; also with a group of 40–48 setae posterior to vulva, each 10–13 µm long. Microspines, about 2–3 µm long, arranged in groups of 3 or 4 in about 8 rows on each abdominal segment.

Comments

The above details for the body dimensions, form and body colour of the young adult and post-reproductive female are the same as those in the original description (Bodenheimer, 1931). However, although Bodenheimer noted the presence of tubular ducts and pores, he gave no details. He also observed that the legs and antennae were "reduced" but the appearance of these structures on mounted material of the post-reproductive females appear distorted because the derm is highly sclerotized. However, in our observations of young adult females, the antennae are 4–6 segmented and all segments of the legs are present. The best characters for comparing the lectotype and paralectotypes of K. greeni with the fresh topotypic material were the following: (i) size and shape of spiracles; (ii) dorsal submarginal setae-pore clusters; (iii) shape of anal ring, and (iv) size of anal lobes and distribution of associated setae. Because most other Kermes species from Israel are only known from either first-instar nymphs or post-reproductive females, we were only able to compare this material of K. greeni with a single paratype of K. bytinskii Sternlicht (1969) (deposited in the BMNH). Both species have: (i) fully-developed legs; (ii) clusters of multilocular pores on venter of abdomen; (iii) a submarginal band of tubular ducts on venter, and (iv) setae-pore clusters located submarginally on dorsum. In both species, the anal lobes, represented as sclerotized areas, are located on the dorsum. However, the size of the anal lobes and the frequency of setae on the anal lobes are different. Kermes greeni has wide, quadrate-shaped anal lobes, each 50–75 µm long and 63–75 µm wide, whilst those on K. bytinskii are narrower, 38–40 µm long and 20–37 µm wide. Kermes greeni also has 13–16 hair-like setae on and surrounding each lobe, whereas K. bytinskii has about 5 hair-like setae restricted to each lobe only. Kermes bytinskii also has a dorsal, medial row of derm-spots which are absent in K. greeni .

The four characters shared by K. greeni and K. bytinskii mentioned above are also present in the adult female Kermes species described or redescribed from China by Hu (1986) and Liu & Shi (1995), namely K. flavus Liu , K. miyasakii Kuwana , K. orientalis Liu & Shi and K. taishanensis Hu. In the Mediterranean and European regions, few teneral adult females of Kermes have been described. We can, therefore, only compare adult female K. greeni with those of K. quercus (Linnaeus) 1758 , K. roboris (Fourcroy) 1785 and K. vermilio Planchon 1864 . Kermes quercus and K. roboris have setae-pore clusters on the dorsum similar to those on K. greeni , but this character is absent in K. vermilio . Multilocular pores are located ventrally in transverse rows on the abdominal segments of all of the above mentioned species. The number of segments in the antennae and legs is also a trait that can distinguish K. greeni from other Palaeractic species of Kermesidae . Kermes quercus has reduced legs with unclear segmentation and 1-segmented antennae, K. roboris has 5 or 6-segmented antennae and thick, short legs, K. vermilio has 1-segmented antennae and no legs whereas adult female K. greeni have 4–6 segmented antennae and well-developed legs, but with the tibia and tarsus occasionally fused. In addition, the anal ring of K. greeni has 3 pairs of setae but no cells, whereas K. vermilio has cells and no setae. The anal rings of both K. quercus and K. roboris are reported to have 2 short setae and no cells ( Borchsenius, 1960; Saakyan-Baranova & Muzafarov, 1972; Kosztarab & Kozár, 1988; Pellizzari et al. 2012).

First-instar nymph ( Fig. 2).

General appearance. Dorsum and venter orange; body oval and tapering posteriorly; body length 0.5–0.52 mm, width 0.21–0.23 mm. Mounted specimens. Oval, 0.41–0.45 mm long and 0.19–0.22 mm wide.

Margin. Marginal setae, each about 5 µm long, in a distinct line of 6–10 on each side (not including anal lobe setae), extending from laterad to labium to level with anal ring, with anteriormost pair of setae distinctly longer, 7–8 µm long.

Dorsum. Derm membranous; intersegmental lines observable. Dorsal setae of 2 types, spatulate and setose; spatulate setae, each 7–8 µm long and 4–5 µm wide, in 4 longitudinal rows; a submarginal row of 18–20 setae on each side, extending along entire margin, and 2 submedian rows of 6 or 7 setae; also with a slightly larger spatulateshaped seta on each anal lobe, each 9–10 µm long and 4–5 µm wide. Setose setae small, each 2-3 µm long, in 2 submedian, longitudinal rows on abdominal segments. Simple, circular pores, each about 1 µm diameter, in a complete submarginal row of 10–12 pores on each side.

Venter. Eyes present as semi-circles close to margin, each 10–12 µm wide. Antennae each 6-segmented; total length 100–105 µm; segments III and VI longer than other segments; scape with 2 thin hair-like setae; pedicel with 1 thin hair-like seta; segment III with 1 thin, hair-like seta; IV with 1 fleshy seta; V with 1 fleshy seta and 1 hairlike seta, and apical segment with 6 fleshy setae and 3 hair-like setae. Legs well-developed; segment lengths of hind legs (µm): coxae 17–20; trochanter + femur 57–70; tibia 28–35; tarsus 45–50 and claw 15–20; total leg length 175–192 µm; trochanter with 2 oval sensory pores on each side, each 1–2 µm long; tarsal digitules 27–31 µm long, knobbed apically; claw digitules knobbed apically, each 20–23 µm long, extending slightly beyond apex of claw; each claw with a denticle. Clypeolabral shield 73–76 µm long and 63–75 µm wide. Labium 3-segmented, triangular, 77–80 µm long and 40–50 µm wide; with 14 hair-like labial setae: basal segment with 4 setae, each 7–8 µm long; median segment with 2 on dorsal surface, each 8–10 µm long, and apical segment with 8 setae, each 11–18 µm long. Spiracles subequal in size; peritreme about 2–3 µm wide, crescent shaped sclerosis 11–13 µm long; each prothoracic spiracle with 2 loculate pores anterolaterally; 1 quinquelocular and other with 6 loculi, both 3–4 µm diameter; each mesothoracic spiracle with 1 quinquelocular pore, anterolaterally, 3–4 µm diameter. Also with slightly larger quinquelocular pores, each 5–6 µm wide, on head and thorax, with 2 just anterior to clypeus, and 1 medially to each coxa, each 4–5 µm wide. Trilocular pores, each 4–5 µm wide, present submedially on abdominal segments III–V. Ventral setae: 2 spatulate setae, smaller than those on dorsum, each 3–6 µm long, 2–3 µm wide, present on anterior apex of body; also 6 hair-like setae, each 15–20 µm long, located medially between antennae and clypeus; 1 seta, 12–15 µm long, medial to each coxa; 6 longitudinal rows of setae on abdomen: 2 medial rows, each with 6 setae, 7–8 µm long; 2 submedial rows, each with 6 setae, about 5 µm long, and 2 submarginal rows, each with 6 setae, about 2–3 µm long. Single bilocular pores, oval, each 3 µm long and 2 µm wide, present submarginally about level with each spiracle. Microspines present medially and submedially on each abdominal segment, in 2–4 transverse rows; each spine about 1 µm long. Anal ring ventral, composed of 2 semicircles, total diameter 18–20 µm; each semi-circle with 4 or 5 cells and 3 pointed setae, subequal in size, each seta 12–15 µm long. Also, with 2 setae, each 10–12 µm long, placed anterior to anal ring, plus 2 setae, each 20–23 µm long, posterior to anal ring. Anal lobes slightly developed; each lobe with 3 setae: 1 thick seta on lateral margin, 8–10 µm long; 1 thick setae on inner margin 8–12 µm long, and an apical seta 130–175 µm long.

Comments

The main characters used here to compare Balachowsky's syntype, the first-instars within the female paralectotype of K. greeni Bodenheimer and the freshly collected crawlers were: (i) type and distribution of spatulate setae on the dorsum, (ii) presence of quinquelocular pores associated with each spiracle, (iii) number of trilocular pores present submedially on venter of abdominal segments III–V, and (iv) distribution of setae on venter of abdominal segments. These were all found to be identical.

The present description includes several features that were not described by Balachowsky (1953) but were observed by us in the type material. These features are presence of: (i) dorsal simple pores; (ii) dorsal setae located medially on abdominal segments; (iii) one, six-loculate pore associated with each metathoracic spiracle; (iv) a denticle near the tip of each claw; (v) one bilocular pore located submarginally level with each spiracle, and (vi) anal ring open at both posterior and anterior ends.

Some morphological characters of the first-instar nymph of K. greeni are shared with other Palaeractic and Neararctic Kermes species : (i) 6-segmented antennae; (ii) 3-segmented labium; (iii) simple pores forming longitudinal line or lines on the dorsum; (iv) marginal, submarginal and medial setae on the dorsum; (v) anal ring with cells and setae, and (vi) denticle on tip of claw (data taken from Kuwana, 1931; Balachowsky, 1950, 1953; Baer & Kosztarab, 1985; Hu, 1986; Liu & Shi, 1995; Podsiadlo, 2005b). Kermes greeni also possesses spatulate marginal and dorsal setae, thus resembling the following Palaerarctic species: K. bacciformes Leonardi , K. mutsurensis Kuwana , K. nakagawae Kuwana , K. nigronotatus Hu , K. orientalis Liu & Shi , K. quercus (Linnaeus) , K. spatulatus Balachowsky , K. taishanensis Hu , K. vastus Kuwana and K. viridis Borchsenius. Kermes vermilio Planchon , a Palaearctic species that has recently been redescribed ( Pellizzari et al. 2012), differs from K. greeni in several traits: (i) presence of simple pores on the dorsum of K. greeni (absent from K. vermilio ); (ii) spatulate dorsal setae ( K. vermilio has conical marginal and dorsal setae), and (iii) anal ring with setae and cells (only setae on K. vermilio ).