Paratachardina pseudolobata Kondo & Gullan

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 : 20-27

Paratachardina pseudolobata Kondo & Gullan , sp. nov.

( Figs 1 N, 2B & C, 7–9)

Type material studied. Holotype. Adult female. U.S.A., Florida: Broward Co., Davie, University of Florida Fort Lauderdale Research Education Center, 20.xi.2002, coll. T. Kondo, ex leaf of Mangifera indica (mango) 1(1) (BME); dimensions for holotype as follows: 1.6 mm long, 1.1 mm wide anteriorly, and 1.5 mm wide posteriorly.

Paratypes. U.S.A., Florida: same label data as holotype, 3(3) (BME), 2(2) ( USNM); same label data as holotype, except on stem of Clusia sp. 2(4) (BME), 1(2) ( MNHN), 1(2) ( USNM); Big Cypress Seminole Indian Reservation, 26°22'N, 81°01'W, coll. 8.vi.2006, S. Schroer & R.W. Pemberton, FL5, ex Myrica cerifera , 2(4) (BME); Everglades National Park, Shark Valley, 25°03.6'N, 80°29.2'W, coll. 8.vi.2006, S. Schroer & R.W. Pemberton, FL2, ex Chrysobalanus icaco , 1(1) (BME); Broward Co., Fort Lauderdale, Secret Woods, 26°05.3'N, 80°10.7'W, 14.vi.2006, coll. S. Schroer & R.W. Pemberton, FL9, ex Psychotria nervosa , 1(2) ( USNM), 2(2 adult females + 10 second-instar nymphs) (BME); Broward Co., Broward, Fern Forest, 26°13.8'N, 80°11.1'W, 14.vi.2006, coll. S. Schroer & R.W. Pemberton, FL10, ex Psychotria nervosa , 4(6) (BME), 2(3) ( USNM); Dade Co., US 27, Tamiami Trail crossing, 26°03.6'N, 80°26.0'W, 8.vi.2006, coll. S. Schroer & R.W. Pemberton, FL1, ex Melaleuca vinifera 5(13: 6 adult females + 1 pharate nymph + 6 secondinstar nymphs) (BME), 2(4) ( USNM).

Paratypes. AUSTRALIA: Christmas Island: Winifred Beach track, 10°30'S, 105°34'E, 20.vi.2006, coll. R. Reeves, ex stem of Celtis timorensis 8(8) (ANIC), 2(2) (USNM), 2(2) (BME); Indian Ocean, WCR, 10.v.2002, coll. K.L. Abbott, ex stem of Syzygium nervosum , tended by Anoplolepis gracilipes , KLAcanb.007–009, 3(3) (ANIC); Indian Ocean, 12.iv.2003, coll. Kirsti Abbott, ex stem of Combretum acuminatum , 6(6 adult females + 16 first-instar nymphs) (ANIC); Dolly Beach track, 10°30'S, 105°40'E, 15.vi.2006, coll. K. Retallick, ex stem of Terminalia catappa 6(5 adult females + 1 first-instar nymph) (ANIC), 1(1) (MNHN); Daniel Roux Road, 10°26'S, 105°40'E, 15.vi.2006, coll. Kent Retallik, ex stem of Celtis timorensis , 9(7 adult females + 1 first-instar nymph) (ANIC), 3(3) (BME).

Paratypes. BAHAMAS: Andros: 24°49'N, 77°49'W, viii.2006, Coll. S. Schroer & R.W. Pemberton, Bh30, ex stem of unidentified host, 1(1) (BME); 24°50'N, 77°53'W, viii.2006, Coll. S. Schroer & R.W. Pemberton, Bh31, ex Chrysobalanus icaco , 1(1) (BME); Grand Bahamas: 26°34'N, 78°51'W, viii.2006, Coll. S. Schroer & R.W. Pemberton, Bh26, ex Eugenia confusa , 9(6 adult females + 39 first-instar nymphs) (BME), 3(3) (FLDA), 8(6 adult females + 14 first-instar nymphs) (USNM); Rand Nature Center, 26°32'N, 78°40'W, viii.2006, Coll. S. Schroer & R.W. Pemberton, Bh12, ex stem of Tetrazygia bicolor , 1(1) (BME); New Providence: Nassau, 25°03'N, 77°27'W, viii.2006, Coll. S. Schroer & R.W. Pemberton, Bh1, ex Exothea paniculata , 1(1) (BME); Nassau, 25°04'N, 77°19'W, viii.2006, Coll. S. Schroer & R.W. Pemberton, Bh2, ex unidentified plant, 1(1) (BME).

Adult female

Unmounted material ( Fig. 2 B & C). Lac test purplish red to dark reddish brown, often appearing brown or black due to sooty mould; tests of adjacent females often fused if crowded; covering infested twigs and branches of host. Test with 4 marginal lobes, each lobe rounded: anterior lobes (al) smaller than posterior lobes (pl); 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: 1.2–2.0 mm long, 0.7–1.5 mm wide at anterior lobes, 1.0–2.0 mm wide at posterior lobes, 1.0– 1.5 mm high. Lac texture very hard, brittle, shiny.

Mounted material ( Fig. 7 ). Body outline 4-lobed, X-letter or bowtie-shaped, anterior pair of lobes smaller than posterior pair. Body 0.8–1.8 mm long, 0.4–1.3 mm wide anteriorly, 0.5–1.7 mm wide posteriorly at widest point (n = 61).

Dorsum. Brachia short, 25–30 µm long, membranous, becoming slightly sclerotized at maturity. Brachial plates subcircular to oblong, often subquadrate, each 80–98 µm long, 50–75 µm wide; brachial crater absent; with a group of 16–30 pseudospines on narrowing side of plate, each pseudospine 6.0–7.5 µm long, with 1 seta on each side of group of pseudospines, each seta 6.0–8.0 µm long, setae often absent on one side. Brachial pores each ca. 4.0 µm wide with 5 (rarely 4) loculi, totalling 2–7 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 63–65 m long, peritremes 26–30 µm wide, surrounded by a sclerotized area 93–98 µm long, 53– 63 µm wide, bearing 7–17 spiracular pores; canellae represented by a group of 12–18 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, 90–110 µm long, 53–65 µm at base, with a slit-like opening at apex; membranous pedicel short, no longer than half length of dorsal spine, slightly wider than base of dorsal spine. Anal tubercle well developed, tapering, highly sclerotized; pre-anal plate 63– 175 µm long, 133–250 µm wide, slightly less sclerotized than supra-anal plate, each plate with a fibrous texture, supra-anal plate 93–114 µm long, 93–130 µm wide, with a granulose texture on mid areas. Pygidial apodemes slightly to moderately developed, extending from base of each anal tubercle towards body apex. Anal fringe entire, composed of 3 plates, each plate 30–35 µm long, 14–20 µm wide, serrated or at least with some teeth. Anal ring 32–33 µm wide, tip of setae surpassing anal fringe. Microducts scarce, present marginally and submarginally, and with 8–12 ducts on each antero-anal lobe; diameter of duct rim ca. 3 µm. Spermatoid ducts: 1–3 associated with each small-sized microducts. Dorsal setae each 5.0–6.0 m long marginally or submarginally, with longer setae, each 10–18 m long, in a single line running from laterad of anal tubercle to body apex on each side.

Venter. Antennae 95–130 µm long, 2 segmented, segmentation poorly defined, with a sclerotized area near base, with 2 longer setae and 2 shorter setae on sclerotized area at apex of terminal segment. Clypeolabral shield 135–158 µm long, 85–100 µm wide. Labium apparently 1 segmented, 40–43 µm long, 53–55 µm wide. Pre-oral lobes elongate, present along margins of clypeolabral shield on each side; post-oral lobes each 45–65 µm wide, dome shaped, with microtrichia. Legs completely absent. Posterior spiracles each 43–48 µm long (spiracular apodeme + peritreme), spiracular peritreme 18–21 µm wide; with 11–16 spiracular pores, each 4.0–5.0 µm wide, present around each spiracle. 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 4.0–5.5 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– 60 /6–11/6–19; mdc-ii: 4–12/0– 3/0–4; mdc-iii: 8– 11 /2–3/2–3; mdc-iv: 8– 10 /2–3/3–4; mdc-v: 15– 30 /2–4/4–6; mdc-vi: 20– 30 /4–5/5–10; mdcvii: 15– 32 /4–6/4–10, and mdc-viii: 8– 20 /2–4/4–7. Ventral duct clusters subcircular or irregular in shape, all composed of medium-sized microducts, 4 pairs in total; pair just anterior to mouthparts (vdc-1) largest, each cluster of vdc-1 distinctly separated ( Fig. 1 N) and usually composed of 20–45 (rarely as few as 8; both clusters combined usually 41–86, but one female with 20 combined) microducts, with number of microducts often differing between left and right clusters; a small cluster (vdc-2) present lateral to labium on each side, each with 2–11 microducts; 2 pairs of clusters (vdc-3 and vdc-4) on abdominal region each with 4–9 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 two linear groups extending from area ventrad of dorsal tubercle towards body margin, rest of ventral derm devoid of microducts. Spermatoid ducts hard to detect, similar to those on dorsum, detected around body margin, appearing most numerous within each marginal duct cluster (distribution not illustrated) with 0–2 associated with each medium-sized and large-sized microduct in marginal duct clusters, but not associated with mediumsized microducts on ventral duct clusters. Ventral setae each 5–10 µm long, about 3 pairs present anterior to mouthparts, a pair anterolateral to each pre-oral lobe, 1 or 2 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 setae on submargin of posterior apex, setae absent elsewhere.

Diagnosis. Paratachardina pseudolobata can be diagnosed by the following features: (i) ventral duct clusters totalling 4 pairs, with clusters of most anterior pair well separated ( Fig. 1 N) and each usually with 20– 45 (41–86 combined) microducts, and (ii) test of adult female X-letter or bowtie-shaped with each of the four lobes of the test rather smooth, purplish red to dark reddish brown, often black due to sooty mould. [Refer to 'Diagnosis' of P. mahdihassani and of P. s i l v e s t r i for comparisons.]

Morphological variation. No phenotypic variation related to place of collection was observed among the adult female specimens from Florida ( U.S.A.), the Bahamas, and Christmas Island ( Australia). The number of microducts in each vdc-1 can vary among individuals and between sides of the body of one individual, for example, there may be 13 microducts on the left side and 30 microducts on the right side on vdc-1. On one specimen from Christmas Island, there were 8 microducts on one side and 12 on the other side. This asymmetry occurs in some specimens of every population examined.

Second-instar nymph

Unmounted material. Test of second-instar nymph of same colour as that of adult female. Test of firstinstar nymph incorporated into second-instar test on mid-dorsum, orifice absent or obscure posterior to firstinstar test. Dimensions of teneral second-instar female test: ca. 0.6 mm long, 0.2 mm wide; more developed nymphs 0.7–0.8 mm long, ca. 0.7 mm wide at posterior lobes, 0.4 wide at anterior lobes, 0.2 mm high.

Mounted material ( Fig. 8 ). Body elongate oval, posterior end slightly broader than anterior end. Body 490–690 µm (ca. 0.5–0.7 mm) long, 250–400 µm (ca. 0.3–0.4 mm) wide (n=16).

Dorsum. Brachia short, 45–55 µm long, membranous. Brachial plates subcircular to oblong, each 50–58 µm long, 23–35 µm wide; brachial crater absent; with a group of 4–12 pseudospines on inner side of each plate, each pseudospine 6–8 µm long, with 1 seta on each side of group of pseudospines, each seta 3.0–5.0 µm long, seta rarely absent. Brachial pores each 3.0–4.0 µm wide, with 0 or 5 loculi, totalling 2–8 pores per plate, present on area just anterior to pseudospines, or often with 1–3 simple pores of the same size found within pseudospine group around its margin. Dorsal spine poorly developed, represented by a small lobe present anterior to anal tubercle. Anal tubercle well developed, tapering, fairly sclerotized; pre-anal plate 55–65 µm long, 90–135 µm wide, less sclerotized than supra-anal plate; supra-anal plate 50–55 µm long, 60–80 µm wide, incomplete, with a cleft anteriorly. Anal fringe partial, with 3 or 4 anal fringe plates, each plate 40–45 µm long, 8–10 µm wide, serrated or at least with some teeth. Anal ring 23–25 µm wide, with 10 setae, and an irregular row of simple pores, tip of setae within anal fringe. Microducts scarce, diameter of duct rim 2.0–2.5 µm, present submarginally, with a few ducts present on each antero-anal lobe. Spermatoid ducts hard to detect, visible around submargin, with 0–2 closely associated with each microduct (distribution not illustrated). Dorsal setae slender, 3–5 µm long, present marginally, with 2 parallel rows of about 4 setae extending from area lateral to anal tubercle towards body apex.

Venter. Antennae 48–75 µm long, 2 segmented, segmentation poorly defined, segments delineated by a sclerotized bar near mid length of antenna, with 2 longer setae and 2 or 3 shorter setae on sclerotized area at apex of terminal segment. Clypeolabral shield 105–125 µm long, 73–85 µm wide. Labium apparently 1 segmented, 38–40 µm long, 45–50 µm wide. Pre-oral lobes broad, present along margins of clypeolabral shield on each side; post-oral lobes each 43–43 µm wide, dome shaped, with microtrichia. Anterior spiracles with spiracular apodemes not developed, connection of tracheae and spiracular peritremes clearly visible; each spiracular peritreme 12–13 m wide; peritreme opening into a slightly sclerotized spiracular furrow, with 7–9 spiracular pores within furrow extending from area anterior to each peritreme towards body margin; each spiracular pore 4.0 m wide, with 4 or 5 (mostly 5) loculi. Posterior spiracles well developed, each 23–26 µm long, spiracular peritreme 9–10 µm wide; with 3 spiracular pores present around each spiracle within a shallow spiracular pocket (not illustrated), each pore 4.0 µm wide. Legs completely absent. Marginal duct clusters distinct, oval to elongate oval, 8 pairs in total; each composed of 2 types of microducts: (i) medium-sized microducts, each with rim 2.5–3.0 µm wide, most abundant; and (ii) large-sized microducts, each with rim 3.2–4.0 µm wide, present on outer margin of each cluster closest to body margin, and on inner area of each cluster. Ventral duct clusters subcircular or irregular in shape, all composed of medium-sized microducts, 4 pairs in total; pair just anterior to mouthparts largest, each composed of 4–10 microducts; a small cluster present antero-lateral to labium on each side, each composed of 2–4 microducts; and 2 pairs of clusters on abdominal region each with 1–3 microducts. Microducts outside ventral and marginal duct clusters smallest, each with rim 2.0–2.4 µm wide, present marginally and submarginally, abundant particularly around marginal duct clusters, rest of ventral derm devoid of microducts. Spermatoid ducts similar to those on dorsum, detected around body margin, appearing most numerous within each marginal duct cluster (distribution not illustrated). Ventral setae slender, scarce, each 3–5 µm long, 2 pairs present anterior to tentorial bridge, a row of setae present on each side of abdomen from behind posterior spiracle to near body apex, and a few setae on submargin on posterior apex, setae absent elsewhere.

First-instar nymph

Unmounted material. Test elongate oval, appearing dorsally segmented, body tapering towards posterior apex; colour wine red to crimson red, about 0.4 mm long, 0.2 mm wide.

Mounted material ( Fig. 9 ). Body elongate oval, narrowing towards posterior end. Body 410–435 µm (ca. 0.4 mm) long, 150–200 µm (ca. 0.2 mm) wide (n=26).

Dorsum. Derm membranous, dorsal segmentation clearly visible. Dorsal spine absent. Supra-anal plates each irregularly rectangular, with outer angles broadest, each plate 25–30 µm long, 15–22 µm wide; dorsal surface with a short subapical seta and a very long apical seta, with a simple pore mesad to subapical seta. Anal ring 16–17 µm wide, with 6 setae and an irregular row of translucent wax pores. Dorsal microducts each with rim ca. 1.0 µm wide, associated with 1–3 simple pores, each simple pore subequal in size to a microduct, area around each group of microducts and simple pores sclerotized; one group of microducts + pores present on body margin of most segments, with about 10 groups on each side of body. Spermatoid ducts present marginally and on mid-dorsum, difficult to detect. Dorsal setae absent.

Margin. Outline smooth except for indentation near anterior spiracle on each side. Eyes located on margin on area lateral to each antennal scape. Spiracular cleft with 3 pseudospines: 2 on anterior side and 1 on posterior side of cleft, each pseudospine 9–11 µm long. Pseudocerarius present on each side on margin ventral to supra-anal plate, composed of a pair of pseudospines with a sclerotized base, each pseudospine 17–21 (23– 27 if sclerotized bar at base included) µm long, with a simple pore, 1–2 µm wide, present at their base.

Venter. Derm membranous. Antennae 120–125 µm long, each 6 segmented, with 3rd segment longest; setae as follows: 3 hair-like setae on segment I, 2 setae and a campaniform sensillum on segment II, 1 hair-like seta on segment III; 1 single fleshy seta on segment IV; 1 fleshy seta plus 1 setose seta on segment V; and 5 hair-like setae, 4 fleshy setae and with 1 short seta between paired fleshy setae on segment VI. Mouthparts normal; clypeolabral shield 88–98 µm long, 58–60 µm wide; labium apparently 1 segmented, 33–35 µm long, 45–50 µm wide. Legs well developed; trochanter + femur 60–65 µm long, tibia + tarsus 58–73 µm long. Tarsal digitules similar, both knobbed, longer digitule 25–35 µm long, shorter digitule 18–22 µm long. Claws each 14–15 µm long, with a denticle; claw digitules similar, knobbed, each 14–16 µm long, one slightly broader than other, apex of each digitule slightly surpassing apex of claw. Anterior and posterior spiracles similar, each with a well developed muscle plate, width of peritremes each 4.0–5.0 µm. Spiracular disc-pores each pore ca. 3.0 µm wide with 4 or 5 (mostly 5) loculi; pores totalling 6–9 between anterior spiracle and body margin in spiracular furrow, and 1 pore just anterior to each posterior spiracle. Pairs of long submedian abdominal setae present on first 5 abdominal segments, plus a seta on each side on area between fore coxa and mid coxae, and between mid coxa and hind coxa, each seta 20–30 long; with 3 pairs of setae in a longitudinal line extending from interantennal area towards mouthparts, each seta 20–25 µm long. Ventral microducts absent. Submarginal setae slender, of 2 size categories, distributed as follows: (i) short setae, each 6–8 µm long, 6 anteriorly between eyes and, on each side, 1 on mid area between each eye and spiracular cleft, 1 present of each side of spiracular cleft, and (ii) very short setae, each 2–3 µm long, present mostly in segmental pairs between each spiracular cleft and body apex, with the setae nearest to spiracular furrow and pseudocerarii present singly.

Notes. As noted above, adult females of this species display no morphological variation related to the known geographic localities, presumably because the species is exotic to each of its currently known areas. We list all specimens studied as part of the type series because of this lack of geographic variation and because the description of the adult female is based on the full range of specimens examined. Males were not found in the present study, nor in previous studies (in which this species is referred to as Paratachardina lobata lobata (Chamberlin)) ( Howard et al. 2004; Mannion et al. 2005). This species appears to be parthenogenetic. Two winged parasitoid adults and two pupae of an unidentified encyrtid wasp were extracted from adult females of P. pseudolobata collected by T. Kondo on mango in Fort Lauderdale, Florida, U.S.A. and were kindly identified as encyrtids by Dr. Steve Heydon, University of California, Davis. P. pseudolobata is one of several polyphagous scale insect species causing canopy dieback on Christmas Island, although this species is not identified in any of the published papers on the subject (e.g., O'Dowd et al. 2003; Abbott 2005).

As explained in the introduction, this species has been known mistakenly as P. lobata in all previous literature discussing its biology and pest status. We identified this error in early 2006 when we began taxonomic work on this group. In their online identification key for scale insects, Miller et al. (2007) describe and illustrate the adult female of P. pseudolobata under the name " Paratachardina sp. nr. lobata ", and also provide diagnostic notes based on information that we provided.

Etymology. The species epithet ' pseudolobata ' is composed of the Greek word ' pseudo ' meaning superficially resembling the original subject, or false, and ' lobata ' the species with which it has been mistaken.

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).

FIGURE 7. Paratachardina pseudolobata sp. nov., adult female.

FIGURE 8. Paratachardina pseudolobata sp. nov., second-instar nymph female.

FIGURE 9. Paratachardina pseudolobata sp. nov., first-instar nymph female.