Machaerota planitiae, Distant, 1908

Pramanik, M. E. A. & Liang, Ai-Ping, 2019, Description of the immature and adult stages of the jujube tube spittlebug Machaerota planitiae Distant (Hemiptera: Cercopoidea: Machaerotidae) in Bangladesh, Zootaxa 4624 (1), pp. 71-86 : 72-84

Descriptions of male and female adults of M. planitiae

Materials examined (Fig. 70). Bangladesh: Rajshahi division: Chapainawabgonj, Northwestern district, 24.73 0 N 88.20 0 E, 19m, 30 Sept. 2015, Lac Research Station , Bangladesh Agricultural Research Institute (BARI), Ziziphus sp. ( Rhamnaceae ), 20♂, 25♀, 200 nymphs. Rajshahi, Northwestern district, Fruit Research Station , BARI, 24.40 0 N 88.50 0 E, 18m, 16 Oct. 2015, Ziziphus sp. , 50♂, 40♀, 150 nymphs. Bagura, Northern district , OFRD Research field, BARI, 24.78 0 N 89.35 0 E, 20m, 24 Dec. 2015, Ziziphus sp. , 15♂, 20♀, 150 nymphs. Rangpur division: Rangpur, Northwestern district , OFRD research Field , BARI, 25.44 0 N 89.15 0 E, 34m, 10 Jan. 2016, Ziziphus sp. , 15♂, 10♀, 50 nymphs. Dinajpur, Northwestern district , Regional Wheat Research station field, BARI, 25.63 0 N 88.65 0 E, 37m, 15 Jan. 2016, Ziziphus sp. , 12♂, 6 ♀, 20 nymphs. Barishal division: Barishal, Southern district , 22.80 0 N 90.37 0 E, 2m, 25 Jan 2016, Research field, Ziziphus sp. , 10♂, 6♀, 30 nymphs. Patuakhali, southern district, 22.3542 0 N 90.318 0 E, 2m, 27 Jan. 2016, OFRD Research field, BARI, Ziziphus sp. , 15♂, 10♀, 20 nymphs. Chattagram division: Chat- tagram, Southeastern district , 22.3375 0 N 91.8389 0 E, 351m, 2 Feb. 2016, Regional Agricultural Research Station Field , BARI, Ziziphus sp. , 10♂, 5♀, 13 nymphs. Rangamati, Southeastern district , 22.38 0 N 92.12 0 E, 17m, 10 Feb. 2016, Research field, Ziziphus sp. , 12♂, 8♀, 14 nymphs. Dhaka division: Gazipur, 24.00 0 N 90.43 0 E, 20 Feb. 2016, BARI Research Field , Ziziphus sp. , 8♂, 5♀, 10 nymphs. Khulna division: Khulna, Southwestern district , 22.35 0 N 89.30 0 E, 9m, 26 Feb. 2016, Research field, Ziziphus sp. , 5♂, 5♀, 10 nymphs. Mymensingh division: Mymensingh , Northeastern district , 24.38 0 N 90.164 0 E, 19m, 20 Jan. 2017, Research field, Ziziphus sp. , 15♂, 10♀, 50 nymphs. Sylhet division: Sylhet, Northeastern district , 24.35 0 N 90.35 0 E, 35m, 2 Feb. 2017, OFRD Research field, BARI, Ziziphus sp., 12♂, 10♀, 40 nymphs.

Distribution. Bangladesh (Rajshahi, Rangpur, Barishal, Chattagram, Dhaka, Khulna, Mymensingh and Sylhet) and India (West Bengal, Bihar, Punjab and Tamil Nadu).

Adult male ( Figs 1 ─3)

Measurements. Body ( Figs1 ─3) medium sized, slender, length (from apex of vertex to tip of forewings) 5.970± 0.465 mm (5.246─ 6.147 mm); average width ( Figs1 ─3) 1.931± 0.646 mm (1.251─ 2.124 mm); length of head process (Figs 9, 10, 12, 14) 1.832± 0.024 mm (1.752─ 1.86 mm); width of head capsule (Figs 9, 10, 12, 14) 1.514 ± 0.055 mm (1.421─ 1.55 mm); pronotum (Figs 3, 8) length 0.721± 0.064 mm (0.597─ 0.796 mm) and width 2.312± 0.051mm (2.114─ 2.457 mm); additional parameters are presented in Table 1 .

Coloration. Body ( Figs 1 ─3) predominantly dark brown. Head (Figs 9, 10, 12, 14): ochraceous to black dor- sally, brown ventrally; postclypeus and tylus together with 9 setaceous slightly yellowish transverse grooves on each side; impressed lines between two transverse grooves black; median line on postclypeus dark black; anteclyp- eus black dorsally, brown ventrally; frons light brown to white; epicranial suture light brown; frontoclypeal suture black; lateral aspect of vertex dark brown to fuscous. Antenna (Figs 11, 13): supra-antennal plate dark brown; scape dark brown; pedicel brown to white; flagellum creamy white; ocellus white; ocular tubercles dark brown, covered with dense white setae; ocellar area light brown; occipital margin bown. Mouthparts (Figs 12, 15─19): gena light brown to fucuous; mandibular plate or lorum ochraceous to black; mentum dark brown; labrum creamy white; max- illary and mandibular stylets light reddish to brown. Pronotum (Figs 3, 8): median carina creamy white to brown; anterolateral margin and posterolateral margin light brown; pronotal collar brown to black; pronotum brown with numerous tiny creamy white and blackish spots. Scutellum ( Figs 1, 3): laterally brownish to black; scutellar process black-yellowish; basilateral depression with white, yellowish, black semicircle; lateral stripe white; rim of sulcus edge white-blackish; edge of postero-inferior margin white-yellowish encircles by black. Abdomen (Figs 20─22): midline of tergite anterior to posterior creamy white; lateral aspect of tergites 1─9 with apical area (4/5) dark brown; midline of sternite dull white and sternites 1─9 with underneath area (4/5) dirty black in lateral view; spiracles dark black, pin hole fuscous. Forewings (Figs 28─30): subhyaline; apex of 4th apical cell and tip of medial 1─2 cells (upper 1/3) dark brown; basal, discal, medial, cubital cells and clavus light white to transparent; costal, subcostal, radial veins and medio-cubital crossvein dark brown, with 12─16 brownish granules; basal veins almost entirely colorless, with 6─8 brownish granules; peripheral membrane transparent; ambient vein colorless. Hindwings (Fig. 31): hyaline; transparent to white; scattered numerous tiny brownish granules over entire wing; veins and crossveins dark brown. Forelegs (Figs 23, 27): dark brown; stramineous near base and margin; posterior part of femur and an- terior portion of tibia slightly yellow; brown to black near apices. Hindlegs (Figs 24, 25): femur black; tibia slightly yellow and striped with brownish black; tarsomeres brown to blackish; tarsal spines apically castaneous. Genitalia ( Figs 32 ─37): pygofer dark black; anal style black; top of anal tube white; aedeagus with middle part white and tip black in ventral view.

Structure. Body ( Figs 1 ─3) densely covered with setae; body length to width 59.70: 19.31. Head process (Figs 9, 10, 12, 14): about 2.5× higher than length of pronotum and height more than 2× depth of eye; strongly vertically produced, anteriorly broadly rounded both in dorsal and lateral view; slightly broader than anterior part of prono- tum; postclypeus smooth and shining, laterally with 9 impressed setaceous transverse grooves on each side; median carina high and sharp in uppermost 1/3, gradually widened and weakened towards anteclypeus; anteclypeus ventral aspect, longer than wide, strongly and broadly raised dorsally along median line and laterally finely punctate and descending; tylus distinctly wider than long, anteriorly broadly rounded, gently descending, evenly convex, more raised and nearly 2× as broad as lateral aspect of vertex, transversely depressed, posteriorly reflexed; lateral aspect of vertex distinctly depressed, 1/2× as long as broad, anteriorly with a transverse carina; compound eyes broader than long, posterior margin curved and distinctly angulate near ventral margin which nearly reaches the posterior margin of head; ocelli behind tylus; ocellar area short, centrally elevated, 2/4 as long as interocellar distance; lorum 3/5 as broad as pleurostome and broader than length of gena proper; head process with length to width 18.32: 15.14. Antenna (Figs 11, 13): located at transition zone between fronto-clypeus and compound eyes; inserted vertically on cuticular wall with articulated socket; composed of a short cone-shaped scape, a cylindrical pedicel and a long thread like flagellum; pedicel narrower at base and concave apex; flagellum forms single antennomere with proxi- mal ampulla and distal filament segments; supra-antennal plate concave, well-defined, about 2/3 as long as broad, 2× as long as ocellar area; antenna located near anterior end of compound eye, base hidden by deep antennae tenta- cles tube like structure; relative length of scape, pedicel and width 1.56: 1.32: 7.86. Labrum (Figs 12, 15): anterior portion narrower and thinner than posterior part; conical shaped; short; joined with anterior margin of anteclypeus; slightly less longer than 1 st. labial segment; covered with few triangular spines; relative length of labrum and 1 st labial segment 52.3: 57.8. Labium (Figs 12, 15─19): tubular-shaped; broad; uniform wide base to tip; deep labial groove scattered entire surface; each side of labial groove with symmetrically distributed sensilla; 1 st labial segment shorter than 2nd and 3rd labial segments; cover up anteclypeus; very few sensilla trichodea I; 2nd labial segment, 1.33× as long as 1 st labial segment; sensilla trichodea I slender, little curved and pointed tip arranged lateral and posterior surface near the junction of middle and 3rd labial segments; sensilla basiconica I short, stout and blunt tip distributed on ventral and dorsal of 2nd labial segment; numerous short dentricles scattered on anterior surface; 3rd labial segment tubular; 1.34× as long as 1 st labial segment; uniform width base to apical; widen apex; trichodea I little; several sensilla basiconica I distributed dorsal of 3rd labial segment; basiconica II present on junction of 2nd and 3rd labial segments. Stylet fascicle (Figs 16, 18, 19): needle shaped, outer two mandibular and middle two maxillary stylets. Mandibular stylet (Figs 16, 19): crescent shaped; inner surface smooth; stylet tip 4─5 nodules in dorsal margin; ventral margin serrate; one row numerous tooth-like projection on lateral edge. Maxillary stylet (Figs 16, 18): surface smooth, internal longitudinal groove interlocking and forming food, salivary and dendritic canals. Pronotum (Figs 3, 8) ≈1.5× as broad as head; about 3.2× as wide as long; strictly convex; anterior of pronotum and ante-callus area 3/4 roundish; posterior 4/5 of pronotum strongly punctate-reticulate, posterior portion of reticula- tions narrower and sharper than anterior portion; distinct calli elongate, oblique; lateral angle with oblique carina apically slightly upraised and distinctly produced into angulate spreading lamina; median carina length ˂1/3× as pronotal width; median carina weak from midpoint to posterior margin very strong at anterior margin, definable only at anterior 1/2, almost reaching summit of head process; anterolateral margin convexly curved, larger than posterolateral margin; posterolateral submarginal area broadly, strongly depressed; posterolateral angle pointed and little produced; posterior margin angularly incised at middle in dorso-caudally view; relative length and width of pronotum process 7.21: 23.12.

Scutellum ( Figs 1, 3) as long as high in profile; scutellar process slender originating from near midpoint of postero-inferior margin, basally compressed; this process 1/3 as long as scutellum proper, tightly extended, curved, tip acute, spike like; lateral stripe of scutellum 3/5 as long as lateral, apically slightly curved, far from meeting rim of sulcus, near to lateroventral margin; rim of sulcus edge sharp, almost 2/3 as long as scutellum; sulcus evenly narrow, about 3/4 as long as scutellum, broader than interocellar distance; superior margin in profile gently curved from end of sulcus to process; basilateral depression as wide as long; entering sulcus and in profile, in compliance with summit of scutellum; postero-inferior margin of scutellum subequal to profile depth of scutellar process at base; scutellum proper with length to basal width 41.12: 15.2; relative lengths and breadths of head, pronotum and scutellum 18.3: 7.12: 41.12 and 15.14: 23.12: 15.1 respectively.

Forewings (Figs 28─30): about 3× as long as broad with reduced venation; apical margin conspicuously oblique; discoidal cell 5× as long as broad; veins M and Cu basally separated, joined by a short mediocubital (m-cu) crossvein that forms basal margin of discoidal cell; claval vein unbranched; basal cell 4× smaller than discal cell; basal cell tends to be rounded; medial cell long, larger than clavus; C pilose fused to R and Sc veins in node and Sc extends to Rs; ambient and peripheral membrane extend apex to anal angle. Hindwings (Fig. 31): subtriangular, wider than medial cell; mediocubital crossvein very short (m-cu); apical cell 2× as wide as discal cell; cubital cell 2× as wide as medial cell; wing peripheral membrane extend apex to claval fold; jugal fold definable only in anterior 1/5, weak up to posterior margin.

Forelegs (Figs 23, 27): elongate, slender, tibia with lateral claws. Midlegs (Fig. 26): tip wider than base; 4 claws joined with conical shaped pretarsus; outer claws about 1.5× as long as inner. Hindlegs (Figs 24, 25): femur about 2.5× as long as broad, length of tibia about 3/4 breadth of head; lateral spine of tibia very strong, arising before midpoint; inner apical lobe of tarsomere 2 slightly longer than outer one, nearly 1.5× as long as outer lobe, slightly longer than claws. Abdomen (Figs 20─22): subcylindric; apex vertically subtruncate; covered with dull white setae; spiracles tuberculiform, prominent, surrounded by fine rugae, sclerotized peritreme, each segment of thorax and abdomen with one spiracle on each side near the pleural membrane; thoracic spiracle 0.61 mm long, 0.52 mm wide; abdominal spiracles I, II and V 0.65 mm long, 0.48 mm wide; III and IV 0.59 mm long, 0.47 mm wide; VI 0.68 mm long, 0.52 mm wide; VII 0.69 mm long, 0.54 mm wide; VIII 0.62 mm long, 0.49 mm wide (Fig. 39).

Male genitalia ( Figs 32 ─37) with pygofer densely pilose, moderately convex in posterodorsal view, longer than wide in lateral view, periandrium with pilose, hemispherical bulged region in lateral view; in ventral aspect, caudal end with deep notch in middle; anal style large, rounded, apex obtuse, slender at base; subgenital plates elongate, rounded, separated from each-other, anterodorsally-directed, base to end pilose; genital styles short subapically, apical part forming a short neck, apex round directed dorsally in lateral view. Aedeagus with shaft elongate, slender, dorsoposteriorly curved, anterodorsally directed, apical part relatively broader than base and middle in lateral view; gonophore apical elongate, slender on dorsal aspect.

Adult female (Figs 4-6)

Measurements. Adult female (Figs 4─6). Body (Figs 4─6) length (from apex of vertex to tip of forewings) 6.870± 0.387 mm (6.465─ 7.124 mm); average width (Figs 4, 5) 2.311± 0.387 mm (2.214─ 2.471mm); length of head process (Figs 6, 14) 1.861± 0.081 mm (1.831─ 1.914 mm); width of the head capsule (Figs 6, 14) 1.610± 0.061 mm (1.532─ 1.693 mm); interocular distance (Fig. 14) 1.005± 0.032 mm (0.854─ 1.091mm); pronotum (Fig. 6) length 0.812± 0.071 mm (0.643─ 0.889 mm) and width 2.579± 0.046 mm (2.197─ 2.674 mm); eye height (Fig.7) 0.876± 0.043 mm (0.799─ 0.987 mm); eye width 0.665± 0.005 mm (0.595─ 0.699 mm); further measurement of body features are included in Table 1 .

Coloration. Similar to male but differs in the following: dominant yellow and reddish brown body; head shining black; gena, lorum and side piece of vertex tinted with brown; pronotum reddish to light dirty brown with a num- ber of small brown dots, calli and punctures at posterior 5/6 dull brown to black; scutellum almost uniformly dull reddish brown; anterior 3/4 of lateral stripe oblong, ivory white, posteriorly attenuated; apical 1/6─1/5 of scutellar process yellow.

Structure. Similar to male but differs in the following points: body ≈2.97× as long as wide; head process ≈1.13× as long as wide; ≈2.25× as long as pronotum; height 3× depth of eye; epicranial suture 2.5× as long as frontoclypeal suture; posterior of postclypeus 2.5× as wide as anterior part; median carina high at middle 2/3; anteclypeus 2.6× as long as wide; side piece of vertex 3/4 as long as broad; compound eyes raised at uppermost 3/4, 2.8× as long as broad; supra-antennal plate 2× as long as anterior arm of epicranial suture; relative length of scape, pedicel and flagellum 1.6:1.39: 7.98; relative length of 1st, 2nd and 3rd labial segments 5.87: 8.24: 7.84; pronotum proper ≈3.17× as wide as long, mesonotum ≈3.9× as wide as long; metanotum ≈4.5× as wide as long; body length to width 68.7: 23.11; head process length to width 18.61: 16.10; pronotum length to width 8.12: 25.79; eye height to width 8.76: 6.65; scutellum proper with length to basal width 45.81: 16.21; relative lengths and breadths of head, prono- tum and scutellum 18.6: 8.12: 45.81 and 16.10: 25.79:16.21 respectively.

Female genitalia (Figs 38─41): dorsal beak extended downward with a developed apical spine in dorsal view; anal style short, posterior end tapering. Genitalia ducts open via genital openings on the underside of segment 8 or 9. Female abdominal segment 9 shortened, wider than long, dorsal midline about equal in length to abdominal tergites 2 and 3; ovipositor sheath extended beyond apex of abdominal segment 9. Two sets of valvulae (gonapophyses) form the shaft of ovipositor connected to base sclerites called valvifers (gonocoxites) that is articulate with abdominal terga; gonocoxites VIII (valvifer I) short, slender, with distal end broad, rounded; gonocoxites IX (valvifer II) anterior end tapering, apex sharp, middle to distal part wide, rectangular, tip obtuse; 9th abdominal segment covered gonocoxites VIII and, IX and gonapophyses X and, IX; gonocoxites IX and gonapophyses X parts fused together to form ovipositor sheath extending to posterior end; gonapophyses X (valvulae II) and IX (valvulae I) posterior end curved, convex and meet with ovipositor sheath; vesical opening (gonopore) broad, rounded ventrally positioned at apex of orifice of spermathecal gland; ovipore small, rounded near to orifice of spermathecal gland.

Remarks. M. planitiae is similar to M. coomani Lallemand and M. elegans Maa in regard to head process, pronotum and scutellum, but differs from the latter species in body color. M. coomani Lallemand is light yellowish to deep reddish with reddish subhyaline wings. M. elegans is deep brown to black, with a light reddish scutellum process and subhyaline wings.

Immature stages

Fifth Instar ( Figs 42, 43; 56, 57)

Length 4.770± 0.865 mm; average width 1.231± 0.546 mm; length of head process 1.325± 0.095 mm; width of head capsule 1.166± 0.069 mm; interocular distance 0.695± 0.065 mm; pronotum length 0.517± 0.026 mm; pronotum width 1.879± 0.036 mm; measurements of other body characters are provided in Table 2 .

Head process light yellowish to ochraceous dorsally, dark brown to black laterally; tylus central line light yel- low; postclypeus black; anteclypeus ochraceous; frons yellowish; anterior of epicranial suture 2/3 brown, posterior dull yellow; anterior arm of epicranial suture light brown; frontoclypeal suture black; side piece of vertex yellow dorsally; supra-antennal plate light brown, base of pedicle 1/3 brown, anterior part white; scape junction to flagel- lum light brown; flagellum white transparent; compound eyes black; ocellar area ochraceous, occipital margin yellow; gena brown; mandibular plate black; labrum light brown; maxillary and mandibular stylets light reddish to dull brown; pronotum, median carina creamy white; anterolateral posterolateral margin light yellow; numerous tiny brownish spots scattered in paranotum and pronotum; mesonotum yellow to light brown dorsally; metanotum light reddish to brown; abdominal tergites 1─9 with apical area yellow to light brownish in lateral and dorsal view; forewings with anterior 4/5 black, base light brown; hindwing with anterior 1/3 light reddish and posterior 2/3 dark brown; forelegs and hindlegs creamy white to light brown; dark brown near base and margin; apex of femur and base of tibia creamy white; apex of tibia and tarsomeres light brown ( Figs 56 , 57); genitalia yellowish to creamy white.

Head process ≈1.14 as long as wide; ≈2.6× as long as pronotum; height 1.5× depth of eye; anteriorly rounded in dorsal and lateral view; broader than anterior part of pronotum; epicranial suture 1.5× as long as frontoclypeal su- ture; postclypeus smooth, laterally light transverse groove on each side, posterior 2× as wide as anterior part; median carina high at middle 1/2, gradually weaking toward anteclypeus; anteclypeus 2× as long as wide; broadly raised at median line and descending; vertex depressed laterally, 2/3× as long as broad; compound eyes raised at dorsal 2/3, 3× as long as broad; ocellar area elevated, 1/3× as long as occipital margin; occipital margin reflexed near middle; mandibular plate broader than gena; head process length to width 13.25:11.66; antenna with a short cone-shaped scape, a cylindrical pedicel and a short flagellum; supra-antennal plate convex, longer than broad, 1.5× as long as anterior arm of epicranial suture; relative length of scape, pedicel and flagellum 1.45: 1.25: 7.68; labrum conical shaped, anterior part thinner than posterior, attached to anterior margin of anteclypeus; labium tubular shaped; 3- segments; 2nd segment larger than other 2 segments; relative length of 1st, 2nd and 3rd labial segments 5.68: 7.81: 7.61; pronotum ≈2.5× as wide as long, anterior of pronotum 2/3 smooth; posterior margin of pronotum merged with mesonotum; median carina undefinable; lateral angle with oblique ental carina apically upraised and produced into angulate spreading lamina; mesonotum 3.5× as wide as long; mesonotal lobate wingpad length ≈2× as long as pronotum extending to apex of 3rd dorsal abdomen, with posterolaterally directed thick carina in lateral 1/3; meta- notum ≈4× as wide as long; lobate wingpad length ≈0.7× that of mesonotal wingpad extends apex of 2nd dorsal ab- domen from metanotum; pro- and mesocoxae 2× as long as broad, subquadrate, posteromedially directed, in dorsal view; metacoxae 1.2× as broad as long, fused to anterior margin of sternum, in ventral view; metatrochanter short, subcylindrical; femora length ≈3× that of coxae; slightly flattened, setaceous planter surface, anterior and posterior margin slender than middle, pro-, meso- and metafemora without teeth; tibia length ≈1.3× that of femora, slender, cylindrical, with longitudinal row of numerous setae; tibiae with transverse apical row of eight to ten darkbrowntipped spines on planter surface; pro-, meso and metatarsi each with three tarsomeres; tarsomere 1 subcylindrical, short, without spines; tarsomere 2 cylindrical, setaceous, length ≈4.5× that of tarsomere 1, without spines; tarsomere 3 short, cylindrical, curved, length ≈2× that of tarsomere 1, with pretarsus two dark-brown curved claws; relative length and width of pronotum, mesonotum and metanotum 7.95: 6.17: 4.96 and 21.02: 21.45: 21.01 respectively; abdomen dorsoventrally rounded, dorsally convex, ventrally concave, abdominal segments V─VII diameter 1.7× as wide as abdominal segments I─IV; spiracles tuberculiform, prominent, thoracic spiracle 0.30 mm long, 0.23 mm wide; abdominal spiracles I─V 0.32 mm long, 0.24 mm wide; VI and VIII 0.35 mm long, 0.27 mm wide; VII 0.34 mm long, 0.25 mm wide; female genitalia in abdominal segments VIII─IX and male genitalia in abdominal seg- ment IX; male genitalia with pygofer wider ventrally than dorsally, anterior margin concave but posterior margin straight in lateral view; basal and processes comparatively short with fine spines on apical portion; subgenital plates distinctly separated, short and apically bilobed; genital styles strongly sclerotized, anterior edge with many fine spines in lateral view.

Fourth Instar (Figs 44, 45; 58, 59)

Length 3.775± 0.397 mm; average width 0.833± 0.345 mm; length of head process 0.787± 0.019 mm; width of the head capsule 0.712± 0.071 mm; interocular distance 0.498± 0.043 mm; pronotum length 0.378± 0.021 mm; pronotum width 1.346± 0.031 mm; measurements of other body characters provided in Table 2 .

Similar to fifth instar, differing in the following points: head shining reddish brown to dark brown; tylus red- dish with brown; occipital margin vertically and laterally whitish; pronotum reddish to slightly brown; midpoint to posterior end of median carina 1/3 creamy white; mesonotum anterior portion and laterally reddish brown; median carina of mesonotum anterior to posterior yellowish; metanotum laterally reddish brown; abdomen dorsally and laterally light yellowish to ochraceous.

Head process ≈1.10× as long as wide; ≈1.31× as long as pronotum; height ≈1.7× depth of eye; epicranial su- ture ≈2× as long as frontoclypeal suture; compound eyes ≈2.5× as long as broad; relative length of scape, pedicel an flagellum 1.32: 1.15: 6.80; relative length of 1st, 2nd and 3rd labial segments 4.87: 6.54: 6.45; midpoint to posterior end of pronotal median carina 1/3 rectangular area differ from other instars; mesonotum anterior and posterior constriction dorsally and laterally prominent; pronotum ≈3× as wide as long; mesonotum 3× as wide as long; mesonotal lobate wingpad length ≈1.5× as long as pronotum extending to apex of 2nd dorsal abdomen, with posterolaterally directed thick carina in lateral 1/4; metanotum ≈3.4× as wide as long; lobate wingpad length ≈0.6× that of mesonotal wingpad extends apex of 1 st dorsal abdomen from metanotum; tibiae with transverse apical row of seven to eight darkbrown-tipped spines on planter surface; relative length and width of pronotum, mesonotum and metanotum 5.98: 4.26: 3.53 and 18.21: 18.64: 17.56 respectively; thoracic spiracle 0.24 mm long, 0.18 mm wide; abdominal spiracles I─V 0.27 mm long, 0.21 mm wide; VI and VIII 0.31 mm long, 0.22 mm wide; VII 0.33 mm long, 0.23 mm wide.

Third Instar (Figs 46, 47; 60, 61)

Length 2.894± 0.324 mm; average width 0.682± 0.234 mm; length of head process 0.685± 0.081 mm; width of the head capsule 0.606± 0.035 mm; interocular distance 0.387± 0.044 mm; pronotum length 0.264± 0.018 mm; pronotum width 1.012± 0.029 mm; measurements of other body characters provided in Table 2 .

Similar to fourth instar, differing in the following: head yellowish to brown; tylus anteriorly ochraceous; prono- tum light reddish to slightly brown; posterior margin of mesonotum with transverse creamy white ridges; anterior to posterior of metanotum 1/2 creamy white, laterally light brown; legs ochraceous to blackish near base and margin, brown near apices; femora and tibiae striped with stramineous; hind tibia and tarsal spines castaneous apically; ab- dominal tergites mostly creamy, yellowish, ochraceous longitudinal fuscous-brown stripe in dorsal and lateral view; wings-pads deep brown to black, margins creamy white.

Head process ≈1.13× as long as wide; ≈1.41× as long as pronotum; height ≈1.6× depth of eye; epicranial suture ≈1.3× as long as frontoclypeal suture; compound eyes ≈2× as long as broad; relative length of scape, pedicel an flagellum 1.25: 1.10: 6.54; labrum trilobed, with central lobe triangular; pilifer well developed, slender; mandibu- lar stylet vestigial but moderately large; maxillary stylet elongate; apical segment with minute sub-apical process; labial palpus well developed, 3-segmented, vestiture moderately smooth dorsally, rough ventrally, fuscous bristles arising latero-ventrally; postclypeus base semirectangular larger than anteclypeus, protruding forward, longitudi- nally depressed; relative length of 1st, 2nd and 3rd labial segments 4.58: 5.68: 5.54; pronotum ≈2.72× as wide as long; mesonotum 2.5× as wide as long; mesonotum lobate wingpad length ≈1.1× as long as pronotum extending to apex of 1 st dorsal abdomen, with posterolaterally directed thick carina in lateral 1/5; metanotum ≈3× as wide as long; lobate wingpad length ≈0.5× that of mesonotum wingpad extends posterior of metanotum; pro-, meso and metatibiae transverse apical row of six to seven brown-tipped spines on planter surface; relative length and width of pronotum, mesonotum and metanotum 4.85: 3.14: 2.65 and 13.21: 16.71: 15.64 respectively; spiracles of abdominal segments slightly increasing in size toward posterior segments; male genitalia nearly narrower, slightly expanded towards rounded apex; sclerotized dorsal half of segment IX, or tegumen, position of the anus near base of tergite.

Second Instar (Figs 48, 49; 62, 63)

Length 1.989± 0.246 mm; average width 0.478± 0.154 mm; length of head process 0.563± 0.067 mm; width of the head capsule 0.510± 0.045 mm; interocular distance 0.235± 0.023 mm; pronotum length 0.198± 0.016 mm; pronotum width 0.598± 0.028 mm; measurements of other body characters provided in Table 2 .

Similar to third instar, differing in the following: head process ≈1.50× as long as pronotum; height ≈1.8× depth of eye; epicranial suture ≈1.5× as long as frontoclypeal suture; compound eyes ≈1.8× as long as broad; relative length of scape, pedicel an flagellum 1.15: 0.98: 5.67; relative length of 1st, 2nd and 3rd labial segments 3.67: 4.56: 4.32; pronotum ≈1.73× as wide as long; mesonotum 2× as wide as long; mesonotal wingpad length ≈0.8× as long as pronotum extending to apex of metanotal wingpad, with posterolaterally directed thick carina in lateral 1/6; metanotum ≈2.5× as wide as long; lobate wingpad length ≈0.3× that of mesonotal wingpad; metanotum ≈1.5× as wide as long; pro- and mesotarsi each with two tarsomeres; tarsomeres 1 and 2 subcylindrical and curved; metatarsi each with three tarsomeres; tarsomere 2 ≈2× length of tarsomere 1; tibiae with transverse apical row of five to six light brown-tipped spines on planter surface; relative length and width of pronotum, mesonotum and metanotum 2.35: 2.65: 1.98 and 6.48: 7.54: 7.21 respectively; spiracles of abdominal segments similar in size.

First Instar (Figs 50, 51; 52, 53; 64, 65)

Length 1.084± 0.051mm; average width 0.303± 0.025mm; length of head process 0.354± 0.035mm; width of the head capsule 0.310± 0.050 mm; interocular distance 0.136± 0.012 mm; pronotum length 0.145± 0.017 mm; pronotum width 0.346± 0.025 mm; measurements of other body characters provided in Table 2 .

Similar to second instar, differing in the following: head process light reddish-yellow to ochraceous dorsally, dull black laterally; tylus central line light creamy white to yellow; postclypeus and anteclypeus reddish-brown; frons light brown; median carina of pronotum to metanotum yellow; ponotum yellow; mesonotum laterally reddishbrown; metanotum 2/3 reddish-brown and 1/3 yellow.

Head process ≈1.14× as long as wide; ≈1.52× as long as pronotum; height ≈2× depth of eye; epicranial suture ≈1.3× as long as frontoclypeal suture; compound eyes ≈2.7× as long as broad; relative length of scape, pedicel an flagellum 1.02: 0.85: 4.50; relative length of 1st, 2nd and 3rd labial segments 3.12: 4.12: 4.01; median carina prominent and definable from anterior of pronotum to posterior of mesonotum; median carina weak at midpoint of metanotum to posterior margin; anterior to posterior of mesonotum laterally each side of median carina 2/3 red- dish brown, this rectangular area is a diagnostic character that differs from other instars; pronotum ≈1.62× as wide as long; mesonotum 1.5× as wide as long; meso- and metanotal wingpad not developed, preliminary initiated, like dark-brown broad spot; pro-, meso- and metatarsi each with two tarsomeres; subcylindrical and curved; tarsomere 2 ≈1.5× length of tarsomere 1; pro-, meso and metatibiae transverse apical row of four to five light brown-tipped spines on planter surface; relative length and width of pronotum, mesonotum and metanotum 2.32: 1.95: 1.53 and 3.75: 4.65: 4.12 respectively; spiracles of abdominal segments similar in size.

Key to the immature stages (first to fifth instars)

1a Mesonotal lobate wingpad length ≈0.8× as long as pronotum (Figs 48, 49); pro- and mesotarsi each with two tarsomeres; meta- tarsi with three tarsomeres (Figs 62, 63)................................................................... 2

1b Mesonotal lobate wingpad length ≈1.5× as long as pronotum (Figs 44, 45); pro-, meso and metatarsi each with three tarsomeres (Figs 58, 59)......................................................................................... 4

2a Metatibia with transverse apical row of five spines (Figs 64, 65)........................................ First Instar

2b Metatibia with transverse apical row of seven spines (Figs 60, 61)............................................... 3

3a Tibiae with transverse apical row of five to six spines (Figs 62, 63); mesonotal lobate wingpad length ≈0.8× as long as prono- tum and metanotal wingpad length ≈0.3× that of mesonotal wingpad (Figs 48, 49)........................ Second Instar

3b Tibiae with transverse apical row of six to seven spines (Figs 60, 61); mesonotal lobate wingpad length ≈1.1× as long as pro- notum and metanotal wingpad length ≈0.5× that of mesonotal wingpad (Figs 46, 47)........................Third Instar

4a Mesonotal wingpad length ≈1.5× as long as pronotum and metanotal wingpad length ≈0.6× that of mesonotal wingpad (Figs 44, 45); tibiae with transverse apical row of seven to eight spines (Figs 58, 59)............................ Fourth Instar

4b Mesonotal wingpad length ≈2× as long as pronotum and metanotal wingpad length ≈0.7× that of mesonotal wingpad ( Figs 42, 43); tibiae with transverse apical row of eight to ten spines ( Figs 56 , 57).................................. Fifth Instar

Molting, Nymphal duration and Tube structure of M. planitiae

Male and female adults mate before and after oviposition. Within 7 days old virgin females emitted perfumed like odour which attract the male insect for mating. The one week aged caged females began ovipositing and laid about 45 to 60 eggs. Eggs are deposited singly or in pairs under thin flaps of Jujube stem and leaf. In the optimum conditions, egg hatched within about 20 days. Egg hatching start at the last week of May to early June. Nymphs developing through five instars in July. Adults appear in early august and persist through early November. However, more than 50 days required to complete nymphal period. Newly hatched nymph-producing spittle on host plant; depend- ing on temperature, stadium length 8 to 10 days. Nymphs spend most of their time feeding, primarily on the sap of the green succulent tissue (Figs 71, 72). In the fluid waste is formed into a frothy mass (spittle) in which nymph live. This white frothy fluid excreted by nymph deposited around insects body can soon be coated with the body of the bubble, nymphs in this bubble drops began to use the secretion of the end of the belly to build the nest tube; and in each molting before the nymphs also in the nest of the mouth of the secretion of foam, and in this bubble group finished (Figs 66─68). Seasonal abundance noticed in two-peak period, June and August to November.

Tubes long, cylindrical, white in color at early stage but at latter creamy white (Figs 66─68, 71, 72). Tube width vary from apex to base. Tube length vary from 7.5 to 12.5 mm. Tube apex breadth 1.1 to 1.5 mm; middle portion 1.2 to 1.9 mm and base 1.3 to 2.3 mm (Fig. 68). Insect stay at base of the tube. They secret water which is found at the apex of the tube as a water drops.

Eggs (Figs 54, 55)

Eggs creamy white which turn off-white during the overwintering period; elongate, well-defined shape, com- paratively narrower neck; mostly parallel sided, pointed lower on one end, slightly rounded the other end, 0.965± 0.034 mm long; broader side 0.452 ± 0.051 mm wide, narrower side 0.154 ± 0.051 mm (average of 15 eggs); oper- culum in collar region dark brown, shallow funnel rounded, smooth borders; exochorion uniform, smooth, homoge- neous; tiny, sparse granulations distributed uniformly on tegument, irregularly in funnel; a few eggs had a narrow, straight, prominent, wrinkled faint demarcation; eggs inserted by oviposition under jujube plant tissue of epidermal layer in which one-third of the egg is covered with a secretion but the remaining portion is exposed; a white cottony secretion covered thickened chorion at midline of micropyle but peripheral chorion covered by epidermis; after fertilization, the broader side of the egg becomes deep brown/blackish which indicates that the eggs are ready for hatching.

Remarks. The nymphal instars are similar, differing primarily in morphometric characters, and previously difficult or impossible to separate due to the absence of a taxonomic key. This work attempts to solve this problem by analyzing body features with morphometric comparisons of both adults and immatures ( Tables 1 & 2). The tube spittlebug attacks the jujube crop at the time of flowering (Fig. 69) and affected flowers are dried completely (Figs 71, 72) and heavy infestation can cause complete failure of the crop ( Prodhan et al. 2016).

Figures

FIGURES 1─14. General body features of Machaerota planitiae. 1─3, Adult male, 1, lateral view, 2, ventral view, 3, dorsal view; 4─6, Adult female, 4, lateral view, 5, dorsal view, 6, head with pronotum, dorsal view; 7─14, Adult male, 7, Compound eye and ocellus; dorsal view; 8, Head and pronotum, dorsal view; 9, Head, ventral view; 10, Head, dorsa-frontal view; 11, Fronto-ventral view of the head capsule showing insertion of the antenna and their relative position; 12, Head capsule showing the position of the mouthparts, frontal view; 13, General view of the antenna in the head capsule showing the 3-segments: the scape, the pedicel and the flagellum; 14, Enlarged dorsal view of head capsule with ocellus. Scale bars: 1─6 =1.00 mm; 7 = 0.10 mm; 8 = 0.50 mm; 9, 10 = 0.10 mm; 11=100 µm; 12, 13 = 0.10 mm; 14 =100 µm.

FIGURES 32─55. Adult male and female genitalia and immatures of Machaerota planitiae. 32, Enlarged view of male genitalia, ventral view; 33, Male genitalia, lateral view; 34, Male genitalia, hind view; 35, Apical part of genital plate, ventrolateral view; 36, Paramere, lateral view; 37, Aedeagus and periandrium, lateral view; 38, Female genitalia, lateral view; 39, Female genitalia, gonapophyses of the ninth abdominal segment (outer), and median gonapophyses of the ninth abdominal (central) segments, ventral view; 40, Enlarged ventral view of female genitalia with ovipositor; 41, Female genitalia ventral view ovipositor with orifice of spermathecal gland, adapted from Moulds (2005); 42, Fifth instar, dorsal view; 43, Head of fifth instar, lateral view; 44, Fourth instar, dorsal view; 45, Head of fourth instar, lateral view; 46, Third instar, dorsal view; 47, Head of third instar, frontal view; 48, Second instar, latero-dorsal view; 49, Head of second instar, frontal view; 50, First instar last stage, latero-dor- sal view; 51, Head of first instar last stage, frontal view; 52, First instar early stage, lateral view; 53, Head of first instar early stage, frontal view. 54, Enlarged view of egg, lateral view; 55, Eggs, dorsal view; Scale bars: 32─39 = 0.10 mm; 40 = 100 µm; 41= 0.10 mm; 42 =1.00 mm; 43 = 0.10 mm; 44=1.00 mm; 45 = 0.10 mm; 46 =1.00 mm; 47 = 0.10 mm; 48 =1.00 mm; 49 = 0.10 mm; 50 =1.00 mm; 51= 0.10 mm; 52 =1.00 mm; 53 = 0.10 mm; 54, 55=100µm.

FIGURES 56─72. Nymphal thoracic legs, sample collection sites, fresh and infested inflorescences, live nymphs, and adult of Machaerota planitiae in tubes. 56, Fifth instar thoracic legs; 57, Enlarged view of fifth instar meta thoracic leg with spinations; 58, Fourth instar thoracic legs; 59, Enlarged view of fourth instar metatibia spinations with tarsal claws; 60, Third instar tho- racic legs; 61, Enlarged view of third instar metatarsomeres; 62, Second instar thoracic legs; 63, Enlarged view of second instar metatibia spinations with tarsomeres; 64, First instar thoracic legs; 65, Enlarged view of first instar metathoracic leg; 66, 67, Live nymph and adult above the tubes at infested jujube flowers in field condition under steriomicroscope; 68, Enlarged view of tubes; 69, Fresh inflorescences; 70, Sample collection sites (Bangladesh); 71, 72, Infested inflorescences; Scale bars: 56─64 =50µm; 65 = 0.05 mm; 66─68 =1.00 mm.

Tables

TABLE 1. Measurements (mean ± standard deviation) and comparison of morphometric characters of adult male and female (Figs1─41) of M. planitiae from Bangladesh

S L. Measurement (mm)		Male		Female
No.		Mean ± SD	Range	Mean ± SD	Range
01	Total length	5.970±0.465	5.246─6.147	6.870±0.387	6.465─7.124
02	Average width	1.931±0.646	1.251─2.124	2.311±0.387	2.214─2.471
03	Width of head capsule	1.514 ± 0.055	1.421─1.55	1.610 ± 0.061	1.532─1.693
04	Length of head process	1.832 ± 0.024	1.752─1.86	1.861 ± 0.081	1.831─1.914
05	Interocular distance	0.885±0.087	0.684─0.984	1.005±0.032	0.854─1.091
06	Pronotum length	0.721±0.064	0.597─0.796	0.812±0.071	0.643─0.889
07	Pronotum width	2.312±0.051	2.114─2.457	2.579±0.046	2.197─2.674
08	Eye height	0.845±0.043	0.780─0.976	0.876±0.043	0.799─0.987
09	Eye width	0.643±0.005	0.584─0.695	0.665±0.005	0.595─0.699
10	Eye radius	0.209±0.006	0.187─0.234	0.220±0.006	0.211─0.226
11	Ommatidial length	0.256±0.003	0.236─0.263	0.265±0.007	0.245─0.714
12	Labrum length	0.523±0.003	0.501─0.536	0.5422±0.006	0.521─0.545
13	1 st. labial seg. length	0.578±0.006	0.545─0.581	0.587±0.004	0.561─0.592
14	2nd. labial seg. length	0.798±0.023	0.775─0.802	0.824±0.028	0.795─0.834
15	3rd. labial seg. length	0.774±0.002	0.748─0.782	0.784±0.005	0.764─0.794
16	Stylet fascicle length	0.009±0.0006	0.004─0.011	0.014±0.0003	0.008─0.018
17	Scape length	0.156±0.004	0.132─0.165	0.162±0.004	0.152─0.168
18	Pedicel length	0.132±0.005	0.118─0.137	0.139±0.007	0.125─0.143
19	Flagellum length	0.786±0.031	0.765─0.794	0.798±0.048	0.769─0.809
20	Length of forewing	5.431±0.624	4.644─5.678	5.784±0.512	5.642─5.824
21	Average width of forewing	1.713±0.214	1.564─1.793	1.956±0.348	1.873─2.104
22	Length of hindwing	3.453±0.215	3.324─3.526	3.712±0.316	3.624─3.796
23	Average width of hindwing	1.952±0.342	1.856─1.968	2.153±0.361	1.926─2.241
24	Abdominal length	4.456±0.324	4.125─4.648	5.624±0.461	5.324─5.768
25	Head cap body diameter	1.251±0.156	1.109─1.264	1.2841±0.168	1.250─1.293
26	Head body diameter	1.179 ± 0.067	1.089─1.194	1.198 ± 0.087	1.156─1.206
27	Mid body diameter	2.451±0.196	2.314─2.567	2.469±0.188	2.452─2.841
28	Post mid body diameter	1.112±0.168	0.985─1.118	1.118±0.189	0.994─1.120
29	Anal body diameter	1.002±0.223	0.986─1.091	1.112±1.113	1.012─1.117
30	Hind tibia length	0.589±0.047	0.569─0.596	0.612±0.086	0.598─0.616
31	Scutellum length	4.112±0.234	3.96─4.116	4.581±0.148	4.314─4.653
32	Scutellum width	1.52±0.534	1.25─1.544	1.621±0.136	1.354─1.654

TABLE 2. Measurements (mean ± standard deviation) and comparison of morphometric key characters of five nymphal instars (Figs 42─65) of M. planitiae from Bangladesh

Key features	1st instar	2nd instar	3rd instar	4th instar	5th instar
Total length	1.084±0.051	1.989±0.246	2.894±0.324	3.775±0.397	4.770±0.865
Average width	0.303±0.025	0.478±0.154	0.682±0.234	0.833±0.345	1.231±0.546
Width of head capsule	0.310 ± 0.050	0.510 ± 0.045	0.606 ± 0.035	0.712 ± 0.071	1.166 ± 0.069
Length of head process	0.354 ± 0.035	0.563 ± 0.067	0.685 ± 0.081	0.787 ± 0.019	1.325 ± 0.095
Interocular distance	0.136±0.012	0.235±0.023	0.387±0.044	0.498±0.043	0.695±0.065
Pronotum length	0.232±0.161	0.373±0.065	0.485±0.047	0.598±0.154	0.795±0.216
Pronotum width	0.375±0.025	0.648±0.028	1.321±0.029	1.821±0.031	2.102±0.036
Mesonotum length	0.195±0.017	0.265±0.016	0.314±0.018	0.426±0.021	0.617±0.026
Mesonotum width	0.465±0.112	0.754±0.109	1.671±0.121	1.864±0.134	2.145±0.152
Metanotum length	0.153±0.017	0.198±0.016	0.265±0.018	0.353±0.021	0.496±0.026
Metanotum width	0.412±0.112	0.721±0.109	1.564±0.121	1.756±0.134	2.101±0.152
Eye height	0.234±.012	0.287±0.024	0.374±0.032	0.548±0.038	0.645±0.043
Eye width	0.198±0.001	0.264±0.002	0.352±0.003	0.521±0.004	0.596±0.005
Eye radius	0.100±0.002	0.121±0.003	0.134±0.004	0.165±0.005	0.184±0.006
Ommatidial length	0.151±0.001	0.176±0.002	0.194±0.003	0.211±0.004	0.240±0.005
Labrum length	0.311±0.002	0.345±0.004	0.367±0.005	0.398±0.006	0.412±0.007
1 st. labial seg. length	0.312±0.001	0.367±0.002	0.458±0.003	0.487±0.004	0.568±0.003
2nd. labial seg. length	0.412±0.013	0.456±0.015	0.568±0.016	0.654±0.017	0.781±0.018
3rd. labial seg. length	0.401±0.003	0.432±0.004	0.554±0.005	0.645±0.006	0.761±0.007
Stylet fascicle length	0.003±0.0001	0.004±0.0002	0.005±0.0003	0.006±0.0004	0.007±0.0005
Scape length	0.102±0.001	0.115±0.002	0.125±0.003	0.132±0.004	0.145±0.006
Pedicel length	0.085±0.002	0.098±0.003	0.110±0.004	0.115±0.005	0.125±0.007
Flagellum length	0.450±0.012	0.567±0.016	0.654±0.019	0.680±0.021	0.768±0.028
Abdominal length	0.513±0.124	1.127±0.194	2.134±0.587	2.975±0.754	3.249±0.968
Head cap body diameter	0.265±0.012	0.345±0.098	0.564±0.124	0.612±0.138	1.001±0.146
Head body diameter	0.310 ± 0.050	0.510 ± 0.045	0.606 ± 0.035	0.712 ± 0.071	1.166 ± 0.069
Mid body diameter	0.425±0.035	0.715±0.092	1.243±0.127	1.634±0.146	2.104±0.178
Post mid body diameter	0.301±0.045	0.498±0.096	0.597±0.157	0.615±0.185	0.987±0.199
Anal body diameter	0.213±0.078	0.324±0.121	0.398±0.178	0.594±0.185	0.897±0.199