Paratachardina silvestri (Mahdihassan)

Kondo, Takumasa & Gullan, Penny J., 2007, Taxonomic review of the lac insect genus Paratachardina Balachowsky (Hemiptera: Coccoidea: Kerriidae), with a revised key to genera of Kerriidae and description of two new species, Zootaxa 1617, pp. 1-41 : 27-31

Paratachardina silvestri (Mahdihassan)

( Figs 1 O, 2D, 10)

Tachardia silvestri Mahdihassan, 1923a: 76 ; 1923b: 660.

Tachardina silvestrii (Mahdihassan) ; Chamberlin, 1925: 41. Incorrect subsequent spelling of species name.

Paratachardina silvestrii (Mahdihassan) ; Varshney, 1968: 489.

Tachardina lobata Chamberlin, 1923: 208 . Syn. nov.

Tachardia lobata Ramakrishna Ayyar, 1919: 47 ; 1921: 340. Nomen nudum, discovered by Varshney, 1967: 77.

Tachardia minuta Morrison ; sensu Green, 1922: 414. Misidentification of P. lobata ; discovered by Chamberlin, 1923: 208.

Tachardina lobata (Green) ; Chamberlin, 1925: 41; Kapur, 1958: 37. Incorrect author.

Tachardina lobata var. walczuchii Mahdihassan, 1946: 136 . Probable misidentification [See 'Notes' below for discussion.]

Tachardina lobata var. schmidtii Mahdihassan, 1946: 136 . Probable misidentification [See 'Notes' below for discussion.]

Paratachardina lobata var. schmidti (Mahdihassan) ; Varshney 1984: 370.

Paratachardina lobata var. walczuchae (Mahdihassan) ; Varshney 1984: 370.

Paratachardina lobata schmidti ; Ben-Dov, 2006: 279. Unjustified elevation of rank [See 'Notes' below for discussion.]

Paratachardina lobata walczuchae ; Ben-Dov, 2006: 280. Unjustified elevation of rank [See 'Notes' below for discussion.]

Paratachardina lobata (Green) ; Varshney, 1977: 56; 1984: 370. Incorrect author.

Type material studied. Tachardia silvestri Mahdihassan. Lectotype. Adult female, by subsequent designation ( Varshney 1977). INDIA: Bangalore, Feb.1922, coll. T. Silvestri , ex Ixora parviflora 1(4: lectotype clearly indicated on slide + 3 paralectotypes) ( USNM). Paralectotypes. Same slide as lectotype ( USNM).

Tachardina lobata Chamberlin. Lectotype, hereby designated. Adult female, on slide labelled as " Holotype ", CEYLON (now SRI LANKA): Peradeniya, ex Fluggea leucopyrus [misspelt on label and now Securinega leucopyrus ], from E.E. Green, 1(4: lectotype, clearly marked on label + 3 paralectotypes) (BME). Paralectotypes. Same label data as lectotype, 6(15, including 3 adult females on same slide as lectotype) (BME); 5 boxes of dry material, same label data as lectotype except some labels also with: "Coll. E.E.G., Dec. 1899 " (BME); slide mounted by PJG 2006 from BME dry material labelled as: " Tachard. lobata /Green m.s./ from Fluggea leucopyrus [now Securinega leucopyrus ]/Peradeniya, Ceylon ", 2(2) (BME).

Other material studied. INDIA: Karnataka, Bangalore, coll. S. Mahdihassan, slide mounted by PJG 2006 from BME dry material labelled as: " Tachardia lobata / Chamberlin / On Ixora parviflora / Bangalore, India / Coll. S. Madihassan[sic] / det. E.E.G.", 2(2) (BME); Bangalore, xi.1922, coll. T. Silvestri , ex Ixora coccinea , 1(4) ( USNM); Bangalore, ii. 1923, coll. F. Silvestri , ex Guazuma tomentosa, Let. S. Mahdihassan , 1(8) ( USNM); Bangalore, coll. xi. 1922, T. Silvestri , ex sandalwood tree [ Santalum album ], slide-mounted by PJG 2006 from USNM dry material, 6(6) ( USNM); Bangalore, Malleshwaram Circle, 12°58'N, 77°38'E, 1000 m, "Q", coll. 1–5.v.2006, R.W. Pemberton & S. Schroer, to USDA quarantine Florida, ex Pongamia pinnata , 8(12 adult females + 7 first-instar nymphs) (BME). Tamil Nadu, Coimbator, Onapalayan, 1100 ft [ca. 335 m], 11°01'N, 76°52'E, coll. i.2006, R.W. Pemberton & S. Schroer, ex Pongamia pinnata , M, 4(3 adult females + 10 first-instar nymphs) (BME); same data except, coll. 1–5.v.2006, 2(7: 2 adult females + 5 first-instar nymphs) (BME); same data except, coll. vi.2006, M6, 1(1) (BME);

Adult female

Unmounted material ( Fig. 2 D). Resin of test often fused; orange, or wine red to dark reddish brown, with tinges of orange. Test with 4 marginal lobes, each anterior lobe with 1–3 (usually 1) ridges, posterior lobes each with three ridges: anterior lobes smaller than posterior lobes; first-instar test incorporated into adult test on mid-dorsum, with a circular opening on an elevated area just posterior to first-instar test. Dimensions of adult female test: 0.9–1.7 mm long, 0.5–1.1 mm wide at anterior lobes, 0.8–1.5 mm wide at posterior lobes, 0.4–0.8 mm high. Lac texture very hard, brittle, shiny.

Mounted material (Fig. 10). Body outline 4-lobed, anterior pair of lobes smaller than posterior pair. Body 1.0– 1.7 mm long, 0.5–1.1 mm wide anteriorly, 0.9–1.8 mm wide posteriorly at widest point (n = 37).

Dorsum. Brachia short, 25–30 µm long, membranous, becoming slightly sclerotized at maturity. Brachial plates subcircular to oblong, often subquadrate, each 75–105 µm long, 55–68 µm wide; brachial crater absent; with a group of 15–28 pseudospines on narrowing side of plate, each pseudospine 5.0–7.5 µm long, with 1 seta on each side of group of pseudospines, each seta 5–6 µm long, setae often absent on one side. Brachial pores each ca. 4.0 µm wide, with 5 (rarely 4) loculi, totalling 2–8 pores per plate, usually present on area just anterior to pseudospines, often 1–3 pores found within pseudospine group around its margin. Anterior spiracles each 50–60 m long, peritremes 25–28 m wide, surrounded by a sclerotized area 80–105 µm long, 50–70 µm wide, bearing 3–6 spiracular pores; canellae represented by a group of 14–18 spiracular pores immediately outside spiracular sclerotization; canellar and spiracular pores similar in shape and size, each 4.0–5.0 µm wide with 4 or 5 (mostly 5) loculi. Dorsal spine well developed, 80–108 µm long, 65–78 µm at base, with a slit-like opening at apex; membranous pedicel about as long as dorsal spine, slightly wider than base of dorsal spine. Anal tubercle well developed, tapering, highly sclerotized; pre-anal plate 55–95 µm long, 153–230 µm wide, slightly less sclerotized than supra-anal plate, each with a fibrous texture, supra-anal plate 100–118 µm long, 120–148 µm wide, with a granulose texture on mid area. Pygidial apodemes slightly to moderately developed, extending from base of each anal tubercle towards body apex. Anal fringe entire, composed of 3 plates, each anal fringe plate 15–38 µm long, 10–20 µm wide, serrated or at least with some teeth. Anal ring entire, 37–40 µm wide, tip of setae surpassing anal fringe. Microducts scarce, present marginally and submarginally, and with several (7–12) present on each antero-anal lobe; diameter of duct rim ca. 3 µm. Spermatoid ducts hard to detect, 0–2 associated with each small-sized microduct and with medium-sized and large-sized microducts in marginal duct clusters, but not associated with microducts on ventral duct clusters. Body setae each 6–10 µm long marginally or submarginally, with longer setae, each 15–20 µm long, in a line running Paratachardina silvestri (Mahdihassan) , adult female.

Venter. Antennae 88–125 µm long, 2 segmented, segmentation poorly defined, with a sclerotized area near base, with 2 longer setae and 2 or 3 shorter setae on sclerotized area at apex of terminal segment. Clypeolabral shield 138–163 µm long, 83–115 µm wide. Labium apparently 1 segmented, 42–58 µm long, 63–75 µm wide. Pre-oral lobes elongate, present along margins of clypeolabral shield on each side; post-oral lobes each 43–50 µm wide, dome shaped, with microtrichia. Legs completely absent. Posterior spiracles much smaller than anterior spiracles, each 40–48 µm long (spiracular apodeme + peritreme), spiracular peritreme 18–23 µm wide; with 6–12 spiracular pores present around each spiracle, each ca. 4.0 µm wide. Marginal duct clusters distinct, oval to elongate oval, 8 pairs in total; each composed of 2 types of microducts: (i) medium-sized microducts with elongate oval rim, each 3.5–4.0 µm wide, most abundant type in each marginal duct cluster, and (ii) large-sized microducts with subcircular rim, each 4.5–5.0 µm wide, present on outer rim of cluster closest to body margin and on inner side of each cluster. Formula for marginal duct clusters as follows: mdc-i: 30– 65 /6–12/10–17; mdc-ii: 10– 15 /2–5/2–6; mdc-iii: 8– 12 /3–4/2–5; mdc-iv: 6– 10 /2–4/3–5; mdc-v: 20– 31 /4– 9/7–10; mdc-vi: 20– 35 /4–8/8–10; mdc-vii: 15– 32 /4–8/4–11, and mdc-viii: 14– 20 /3–6/4–8. Ventral duct clusters subcircular or irregular in shape, all composed of medium-sized microducts, 5 to 6 (rarely 7) pairs in total; pair just anterior to mouthparts (vdc-1) largest with each microduct cluster of pair touching or almost touching ( Fig. 1 O), vdc-1: 10–20 microducts (combined 24–40); a small cluster (vdc-2) present lateral to labium on each side with 4–10 microducts; a small cluster (vdc-3) present just anterior to each metathoracic spiracle with 0–4 microducts, 3 pairs of well-developed clusters on abdominal region (vdc-4: 3–7 microducts; vdc-5: 5–8; vdc-6: 5–9), often with a vestigial cluster (vdc-7) with 0–2 microducts; ventral duct clusters often appearing as a continuous linear group of microducts. Microducts outside ventral and marginal duct clusters smallest, each with rim ca. 3.0 µm wide, present marginally and submarginally, abundant particularly around marginal duct clusters, and also present in 2 linear groups extending from area ventral to dorsal tubercle towards body margin, rest of ventral derm devoid of microducts. Spermatoid ducts hard to detect, similar to those on dorsum, present around body margin, appearing most numerous within each marginal duct cluster (distribution not illustrated). Ventral setae each 7–10 µm long, about 3 pairs anterior to mouthparts, a pair anterolateral to each pre-oral lobe, 1 or 2 setae next to each posterior spiracle, a group of about 5 setae behind each posterior spiracle, a pair on each of last 3 abdominal segments anterior to vulva, 2 pairs on segment posterior to vulva, and a few longer setae, each 15–20 µm long, on submargin of posterior apex, setae absent elsewhere.

Diagnosis. Paratachardina silvestri is similar to P. pseudolobata , but it can be separated from this species by the character states given in the key. It can be diagnosed by the following combination of features: (i) ventral duct clusters totalling 5 to 6 pairs (rarely an additional vestigial seventh pair, each with 0–2 microducts) [4 pairs in P. pseudolobata ], with clusters of most anterior pair touching or almost touching ( Fig. 1 O), each cluster with 10–20 (total of both clusters 24–40) microducts [anterior pair well separated and each cluster with 20–45 microducts in P. pseudolobata ]; and (ii) test of adult females four-lobed with 1–3 ridges on each lobe, resin of test often fused to adjacent tests; orange, wine red to dark reddish brown in colour [test rather smooth and purplish red to dark reddish brown, never orange, in P. pseudolobata ]. For comparison of P. s i l v e s t r i with P. mahdihassani , refer to the 'Diagnosis' of the latter.

Notes. Chamberlin (1925) criticised the publication of Mahdihassan (1923a) vehemently and ended his introduction by stating: "All further comments, changes and additions are made in their appropriate position in the following systematic section." ( Chamberlin 1925: 32). In his section on Tachardina, Chamberlin used the name " Tachardina silvestrii " without explanation, although presumably he changed it deliberately. Mahdihassan did not indicate whether " silvestri " was a noun or an adjective and thus according to Article 31.2.2 of the International Code of Zoological Nomenclature (ICZN 1999) " silvestri " should be treated as a noun in apposition and the original spelling retained. Chamberlin's " silvestrii " becomes an incorrect subsequent spelling and we use Mahdihassan's original spelling for the name of this species. Furthermore, the transfer of this species to Paratachardina was by Varshney (1968), not Varshney & Teotia (1968) as listed in Ben-Dov (2006).

Three subspecies of P. lobata (= P. silvestri ) are listed by Ben-Dov (2006), i.e., Paratachardina lobata schmidti (Mahdihassan) , P. lobata walczuchae (Mahdihassan) , and P. lobata lobata (Chamberlin) . However, no justification is provided for recognizing as subspecies the two varieties mentioned by Mahdihassan (1946), nor for recognizing P. lobata lobata as proposed by Ben-Dov (2006). According to article 45.6.4 of the International Code of Zoological Nomenclature (ICZN 1999: 50), names of varieties described before 1961 should be treated as subspecies unless their author also expressly gave them infrasubspecific rank or the content of the work unambiguously reveals that the names were proposed for infrasubspecific entities. Mahdihassan's two varietal names fall into the infrasubspecific category, as explained below. Mahdihassan (1946) named two morphologically identical varieties, Tachardina lobata var. walczuchii and T. lobata var. schmidtii from the same locality (in Bangalore, India) but on different host plants ( Michelia champaca and Guazuma tomentosa ), and distinguished them from each other on the basis of morphologically different bacterial symbionts. Furthermore, Mahdihassan (1946: 136) commented as follows: "There is no meaning in naming one insect T. lobata and the other T. lobata variety Walczuchii, for one is as much as the other. Mere museum-specimens cannot but be simply called T. lobata , which might mean one or the other variety; cytologically and by blood smears the varieties can be easily distinguished". Kapur (1958, p. 38) recognized only the " walczuchii " variety and regarded the " schmidtii " variety as a synonym of Tachardina lobata . In a review of the Kerriidae, Varshney (1984) recognizes Mahdihassan's two names as mere varieties of P. lobata . We suggest that it is inappropriate to elevate these varietal names to subspecies level, and here we treat Mahdihassan's Tachardina lobata var. schmidtii and T. lobata var. walczuchii as varieties. However, we believe that Mahdihassan may have been referring to bacterial variation in the species that we describe above as P. mahdihassani (see under 'Etymology and notes' for that species) rather than in P. lobata (now P. s i l v e s t r i). Unfortunately Mahdihassan designated no type specimens nor did he give any descriptions of the morphology for these varieties, and he appears not to have deposited any specimens in a museum. Hence, deliberately we have not considered either of these varietal names as possible valid names for our new species P. mahdihassani , but do list them above in the synonymy for P. silvestri .

Figures

FIGURE 1. Lac insect structures: A, simplex type marginal duct cluster of Kerria lacca; B, duplex type marginal duct cluster of Tachardiella; C, triplex type marginal duct cluster of Austrotachardiella; D, brachial plate of Albotachardina sinensis, arrow pointing to accessory projection (after Zhang 1992); E, long pedicel and dorsal spine of Albotachardina sinensis (after Zhang 1992); F, dorsal spine of Austrotachardia melaleuca; G, dorsal spine of A. australis; H, dorsal spine of A. acaciae; I, dorsal spine of A. angulata; J, marginal duct cluster with two associated auxiliary clusters of Afrotachardina brachysetosa; K, anal ring of Tachardina aurantiaca divided into four sections; L, anal ring of a South African Tachardina sp. divided into four sections; M, marginal duct cluster of T. aurantiaca; N – P, relative position of vdc- 1 and mouthparts in: N, P. pseudolobata; O, P. silvestri; P, P. mahdihassani; Q, typical macroduct of Austrotachardiella.

FIGURE 2. Tests of Paratachardina spp. A, P. decorella, adult female, from W. M. Maskell's dry material; B, P. pseudolobata sp. nov., adult female, ex mango, type material; C, P. pseudolobata sp. nov., adult female after removal of test (al, anterior lobe; pl, posterior lobe; for other abbreviations see Glossary); D, P. silvestri, adult female, ex sandalwood tree, type material; E, P. mahdihassani sp. nov., adult female, type material; F, P. minuta, three adult females with three first-instar nymphs near their bases, type material; G, P. t e r n a t a, adult female, type material; H, P. thea e, two semifused adult females and two male pupae below, type material; I, P. t h e a e, adult females from China (note smoother texture of each ridge of test) (Photos: A, Rosa Henderson, B – I, T. Kondo).