Body relatively flat. Vertex, thoracic dorsum and most of fore wing veins with symmetrical long and thick setae, which possess tiny spinules on the surface (Fig. 63, termed ‘prickly setae’ below). Wings held flat over back. Fore wing lacking pterostigma. Hind wing with a single thick anal vein (A) which may result from the reduction of vein A₁ or A₂ or from the combination of them. Posterior aspect of male proctiger enveloped. Aedeagus uni-segmented. Female subgenital plate simple and situated much more proximal than proctiger. Valvula dorsalis of ovipositor lacking flag lobe. Fifth instar immature with symmetrical sectasetae on body dorsum, lacking tarsal arolium on legs.

Adult. Body flat, with abdomen significantly wider than tall. Body dorsum with symmetrical prickly setae on the surface, situated on bulges or projections, distribution as: 4+4 on vertex, 4+4 on pronotum, 1+1 on mesopraescutum, 4+4 (Fig. 53) or 5+5 (Fig. 52) on mesoscutum, 1+1 on mesoscutellum, 1+1 on tegula, 1+1 on humeral plates, 1+1 on metascutellum. Surface of vertex and thoracic dorsum sculptured with granular microscopic structures.

Head slightly inclined from longitudinal body axis. Vertex lacking median suture; two tubercles present along the median line, each bearing a pair of prickly setae. Base of lateral ocelli moderately bulging, each bearing two prickly setae. Vertex consistent with gena. Plane of torulus about perpendicular to that of the vertex. Frons completely fused with vertex and gena, only moderately raised from the surface. Gena not divided into two lateral parts, but firmly compact as one, with roughly symmetrical simple setae (Fig. 47); parts below torulus sometimes produced. Occiput smoothly connected with vertex, not folded below it. Plane of postocular sclerite about perpendicular to that of vertex, not nearly parallel with it. Antennae 10-segmented, surface sculptured with minute spinules arranged in transverse rows; at least six rhinaria present on apices of segments IV-IX, apex of segment III sometimes also with one; segments IV, VI and VIII sometimes possess extra rhinaria; rhinarium with closely packed minute spinules lining below, usually bearing horn-like projections. Clypeus rather short, with no extra seta except the apical pair of setae. Labium rather short, two-segmented, lacking ‘conical sensoria’ (as termed by Liang et al. 2013) on the tip (Fig. 48).

Preepimeron significantly wider than preepisternum. Notopleural sulcus of prothorax well developed. Mesopraescutum near semicircular, not protruding forward to force pronotum to arch. Pleural sulcus of mesothorax reduced, with pleural apophysis relatively small; posterior margin of mesopleurite directed forward. Mesepisternum rather narrow and bulging (Fig. 57). Trochantinal apodeme shallow, present on anterior margin of mesopleurite (Fig. 57). Anapleural cleft of mesothorax widely split (Fig. 57). 1+1 extra sclerites present behind the base of mesothoracic furca (Fig. 59). Heel of mesepimeron swollen, bearing a small tubercle (Fig. 57). Metathoracic pleural sulcus reduced, pleural apophysis poorly developed; metepisternum and trochantin not completely divided (Fig. 57). Trochantinal apodeme of metathorax shallow, present on the anterior margin of metapleurite (Fig. 57). Katepisternum and trochantin of metathorax possess well developed ventral aspect which are convergent in the middle, forming a large and solid plate ventrally (Fig. 59). 1+1 extra sclerites present behind the base of metathoracic furca (Fig. 59) (in contrast with most other psyllids, e.g. Cacopsylla, Fig. 60).

Legs long and slender. The three sensory pores on femora ventrum arranged in a row. Plane of hind legs almost parallel with that of middle legs. Metacoxa with rather large tubercle above apical opening, and lacking meracanthus (Fig. 56). Metafemur without a cluster of thick setae on the outside of apex. Metatibia without genual spine, with 1-3 rows of thick setae; apical part often with a row of tightly packed setae dorsally; apical spurs relatively long and slender, sclerotized at different extents but never reaching the hard and black status as in most other psyllids, forming an open crown. Metabasitarsus lacking sclerotized spurs on the apex, but with one or two (in other psyllids there is only one) pairs of simple setae. Apical tarsus with a pair of short and tapered apical setae. Claws with rounded or rather narrow pulvilli.

Fore wing narrowest in the base and gradually becoming much wider apically, usually widest at subapex or apical 1/4. Costal break present. Pterostigma absent. Vein Rs reaching anterior margin instead of apical margin. Cell cu₁ rather long and flat. Veins A₁ and A₂ touching in the middle. Anal break adjacent to the apex of vein Cu_1b.

Hind wing with partially thickened anterior margin. Veins A₁ and A₂ combined or one is lost (probably A₁), leaving a thickened vein A; cell a₁ lost (Fig. 49).

Tergite of abdominal segment 1 better developed, with a median sclerite present (Fig. 50). Spiracles of segments 1 and 2 invisible. Sternites of segments 4-6 each with a pair of wax-secreting pore fields laterally, with shape variable.

Male terminalia: In natural status, proctiger, aedeagus and parameres all oriented caudally instead of upwards. Posterior aspect of proctiger enveloped. Aedeagus uni-segmented and simple, sometimes with tiny spines on dorsum. Sperm pump with only basal end plate, lacking apical end plate (Fig. 51).

Female terminalia: Subgenital plate placed much more proximal than proctiger and simple, lacking tip sometimes. Proctiger lacking rows of long setae on the dorsum. Valvula dorsalis of ovipositor without flag lobe. Median valve slender and placed more terminal, apex touching the subapex of ovipositor.

Fifth instar immature. Body dorsum with symmetrical sectasetae. Antennal 7- or 9-segmented, with three rhinaria. Compound eyes with 1+1 or 2+2 ocular setae. Postocular setae present in 2+2 or more. Fore wing pads simple, without humeral lobe. Legs long and slender, lacking specialized seta. Both tarsal segments differentiated. Tarsal claws with pulvilli and without arolium. Apical setae of tarsus both long and capitate. Abdominal sclerites firm, not broken in the middle. Abdominal apex with a pair of bulges. Circum anal pore field lacking additional rings.

Vein M₃₊₄ of fore wing with 3 prickly setae (Fig. 9). Tarsal pulvilli rounded (Fig. 13). Male subgenital plate without long seta on the dorsal-apical angle (Fig. 19). Female proctiger steeply narrowed in the apical 1/3 (Fig. 29).

Adult coloration. Ground color yellow. Long and thick setae on dorsum black. Compound eyes grey. Ocelli yellow. Antennae yellow, with black apices on segments IV, VI, VIII; segments IX-X entirely black. Pronotum, meso- and metascutum each with one pair of orange markings. Legs yellow. Fore wing hyaline and colorless (Fig. 9). Tergites of abdominal segments 3-5 brown. Male and female terminalia yellow.

Structures: Setae on dorsum of body relatively long (Table 1) and based on prominent projections. Gena flat (Fig. 1). Antennal segments III-IX each with a single rhinarium on apex, segments IV, VI and VIII each with one extra rhinarium; rhinarium on segment IX double-pored and with complex horn-shaped projections; proximally based terminal seta about twice as long as the distally based one (Fig. 5).

Mesoscutum with four pairs of prickly setae. Metatibia with three short rows of thick setae, lacking a tightly packed row of short setae on the dorsum (Fig. 13). Apex of metabasitarsus with only one pair of simple setae (Fig. 13). Pulvilli broadly rounded (Fig. 13). Fore wing cell cu₁ tallest in apical 1/3, with vein Cu_1a abruptly curved at the point; vein M₃₊₄ completely decorated with setae; surface spinules absent; fields of radular spinules relatively large (Fig. 9).

Pore fields on abdominal ventrum small oval; pores loosely packed (Fig. 16).

Male terminalia: Proctiger slightly curved backwards (Fig. 19). Paramere small lamellar; apical half of anterior margin with a thin lobe stretching inwards; anterior margin of basal 1/3 emarginated; two long and thick setae present on inner surface, near the anterior margin; inner surface with a curved ridge decorated with thick setae on apical half (Figs 19–21). Aedeagus with a few tiny spines on the dorsum (Fig. 19). Base of subgenital plate with a small cluster of setae (Fig. 19).

Female terminalia (Fig. 29): Oblong in overall shape. Base of proctiger slight raised, anus partly sunken; dorsal view of proctiger constricted at apical 1/3; apical process with small amounts of tiny setae. Subgenital plate lacking tip, with sparse setae on ventral surface.

Fifth instar immature. Body dorsum strongly sclerotized, ventrum weakly sclerotized. Dorsum of head, thorax and abdomen with symmetrical truncate sectasetae varying in size, mixed with a few simple setae (Fig. 33); dorsum and margin of wing pads with roughly symmetrical truncate sectasetae (Fig. 35). Antennae 7-segmented, apices of segments 4-6 each with one single rhinarium, segments 3-6 each with one single truncate sectaseta (Fig. 34). Compound eyes with 2+2 ocular truncate sectasetae, postocular truncate sectasetae present in 2+2 (Fig. 33). Fore wing pad with two pores on dorsum (Fig. 35). Tarsal pulvilli broad and rounded (Fig. 36). Abdominal ventrum with four pairs of spiracles surrounded by peritremes partly fused with central sclerites. Abdominal apex produced as a pair of rounded bulges (Fig. 37). Circum anal pore field present in between the bulges, both outer and inner ring consisting of neat single row of oval pores (Fig. 37).

Figure 1–8. 

Head and antennae of Togepsylla spp. 1, 5 Togepsylla glutinosae sp. n. 2, 6 Togepsylla matsumurana 3, 7 Togepsylla takahashii 4, 8 Togepsylla tibetana 1–4 Head 5–8 Antenna. Scale bar: 0.2 mm.

Figure 9–12. 

Fore wing of Togepsylla spp. 9 Togepsylla glutinosae sp. n. 10 Togepsylla matsumurana 11 Togepsylla takahashii 12 Togepsylla tibetana. Scale bar: 0.5 mm.

Figure 13–18. 

Hind legs and wax-secreting fields of Togepsylla spp. 13, 16 Togepsylla glutinosae sp. n.; 14, 17 Togepsylla matsumurana 15, 18 Togepsylla takahashii 13–15 Metafemora and tibia 16–18 Wax-secreting fields on lateral aspect of sternites of abdominal segments IV–VI. Scale bar: 0.2 mm.

Holotype: ♂, CHINA: Hainan, Danzhou, Nada, 131 m, 19°30.878'N, 109°31.085'E, ex Litsea glutinosa, 12.iv.2016, Xinyu Luo (CAU). Paratypes: 10 ♂, 12 ♀, 15 immatures, same data as holotype (CAU).

Litsea glutinosa (Lour.) C. B. Rob. (Lauraceae)

China: Hainan.

Named after the scientific name of the host plant.

Based on a brief observation in the field, this species was found free living, both immatures and adults are sparsely scattered across the abaxial surface of leaves (no preference for young leaves or shoots is displayed). The immatures do not induce any form of gall or leaf rolling, and from the sectasetae on body margin they produce wax threads of varying lengths, of which the ones from the terminal bulges of abdomen are longest (Fig. 65).

Dorsum of head and thorax brown with large areas of brown patterns. Antennal segments VI and VIII each with two additional rhinaria (Fig. 6). Paramere with a small area of netlike grains on inner surface (Fig. 23). Apical 1/3 of female proctiger strongly curved upwards (Fig. 30).

Adult coloration. Head yellow, vertex with brown patterns. Long and thick setae on dorsum black. Compound eyes light brown. Ocelli yellow. Antennae yellow, segments I-II light brown, apices of segments III, IV, VI, VIII black, segments IX-X entirely black. Thoracic dorsum brown, except for bases of setae which are yellow. Thoracic pleurites light brown. Legs yellow, with apical half of femora light brown, apex of tibiae brown. Fore wing membrane hyaline and colorless; R₁, apices of Rs and M₁₊₂ black (Fig. 10). Abdominal tergites of segments 1-5 black, sternites brown. Male proctiger brown. Female terminalia yellow.

Structures: Setae on dorsum of body relatively short (Table 1) and based on smooth projections. Vertex with a pair of small foveae between median-posterior tubercle and lateral ocelli (Fig. 2). A pair of small tubercles present above toruli (Fig. 2). Genal tubercles strongly protruding (Fig. 2). Antennal segments IV-IX each with a single rhinarium on apex, segment IV with one, segments VI and VIII each with two extra rhinaria; rhinaria without horn-shaped projection; proximally based terminal seta slightly longer than the distally based one (Fig. 6).

Mesoscutum with 5 pairs of prickly setae (Fig. 52). Metatibia with one row of thick setae ventrally, and with a tightly packed row of long setae on the dorsum (Fig. 14). Pulvilli narrow (Fig. 14). Fore wing with broad cell r₁, cell cu₁ tallest in the middle; vein M₃₊₄ without seta; surface spinules rather minute, widely spread across a relatively small area in distal cells; fields of radular spinules relatively large (Fig. 10).

Pore fields on abdominal ventrum large oval, with pores loosely packed (Fig. 17).

Male terminalia: Proctiger slightly curved backwards apically (Fig. 22). Paramere broad lamellar, with rather slender base; anterior margin of apical half emarginated and thin; posterior margin with a basal ridge on outer surface; middle of inner surface with a small area of netlike grains; anterior angle with a few short and thick setae on inner surface; posterior margin with a band of inner-curved short setae on apical 2/3 (Figs 22, 23). Tip of aedeagus forming an acute small hook, dorsum of aedeagus lacking tiny spines (Fig. 24). Subgenital plate with moderately produced dorsal-apical angle, and with a few setae on the base (Fig. 22).

Female terminalia (Fig. 30): Short and broad in overall shape. Apical 1/3 of proctiger strongly curved upwards; apical half of proctiger with nearly evenly spaced setae, and with a row of setae along ventral margin of apical process. Subgenital plate with blunt and retracted apex, ventral surface with sparse setae.

Fifth instar immature. Body dorsum firmly sclerotized, with sclerites of thorax and abdomen almost unseparated; body ventrum weakly sclerotized (Fig. 43). Dorsum of head, thorax, and abdomen with symmetrical truncate sectasetae varying in size (Fig. 43). 1+1 projections present before fore wing pads, sheathing the 2+2 long setae on lateral margins of adult pronotum (Fig. 43). Antennae 7-segmented, apices of segments 4-6 each with one single rhinarium; segment 3-6 with truncate sectasetae on dorsum (Fig. 44). Compound eyes with 1+1 ocular truncate sectasetae, postocular truncate sectasetae present in 6+6 (Fig. 43). Fore wing pad with three pores on dorsum, and with outer margin completely decorated with truncate sectasetae (Fig. 43). Tarsal pulvilli narrow (Fig. 45). Abdominal ventrum with five pairs of spiracles (Fig. 43). Abdominal apex emarginated (Fig. 46). Circum anal pore field with both outer and inner ring consisting of neat single row of oval pores (Fig. 46).

Figure 19–28. 

Male terminalia of Togepsylla spp. 19–21 Togepsylla glutinosae sp. n. 22–24 Togepsylla matsumurana 25, 26 Togepsylla takahashii 27, 28 Togepsylla tibetana 19, 22, 25, 27 Male terminalia, in profile 20, 23, 26, 28 Paramere, inner surface 21 Paramere, posterior view 24. Apical half of aedeagus. Scale bar: 0.2 mm.

Figure 29–32. 

Female terminalia of Togepsylla spp. A Togepsylla glutinosae sp. n. B Togepsylla matsumurana C Togepsylla takahashii D Togepsylla tibetana. Scale bar: 0.2 mm.

Figure 33–42. 

Fifth instar immature of Togepsylla spp. 33–37 Togepsylla glutinosae sp. n. 38–42 Togepsylla takahashii 33, 38 Half of head, dorsal view 34, 39 Antenna, dorsal view 35, 40 Wing pads, dorsal view 36 Claws, showing pulvilli and apical setae of tarsus 37 Circum anal ring, ventral view 41 Claws, showing pulvilli 42 Circum anal ring, dorsal view on the left half, ventral view on the right half. Scale bar: 0.2 mm (33–35, 37, 38–40, 42), 0.05 mm (36, 41).

CHINA: 2 ♀, Zhejiang, Qingyuan, Baishanzu, 1300-1500 m, ex Litsea cubeba, 24.ix.1993, Hong Wu (CAU) (type series of Togepsylla zheana); 1 ♂, 2 ♀, Yunnan, Yiliang, Xiaobanchang, 1883 m, 27°48.227'N, 104°21.155'E, 27.iv.2014, Xinyu Luo (CAU); 2 ♂, 3 ♀, Guangxi, Wuming, Mt. Daming, 1341 m, 23°30.421'N, 108°26.084'E, 12.v.2014.v.12, Xinyu Luo (CAU). JAPAN: 2 ♂, 2 ♀, 5 fifth instar immatures, Ibaraki Prefecture, Tsuchiura City, Shishisuka, 15 m, 36°4.8'N, 140°9.54'E, ex Neolitsea sericea, 29.iv.2004, Hiromitsu Inoue (HIC); 3 ♂, 3 ♀, Tochigi Prefecture, Kaminokawa, Kamigo, 65 m, 36°26.4'N, 139°55.98'E, ex Neolitsea sericea, 24.iii.2015, Rikio Sonobe (HIC); 1 ♂, 1 ♀, Fukuoka Prefecture, Mt. Hiko, 26.iv.2001, Hiromitsu Inoue (HIC).

Litsea cubeba (Lour.) Pers. (Yang 1995), Lindera erythrocarpa Makino, Lindera glauca (Zieb. et Zucc.) Bl (Miyatake 1970), Lindera sp. (Miyatake 1981), Neolitsea sericea (Bl.) Koidz (HIC). (Lauraceae)

China: Guangxi, Taiwan, Yunnan, Zhejiang (Yang 1984; Yang 1995); Japan: Ehime, Fukuoka, Gunma, Ibaraki, Kagoshima, Nagasaki, Nara, Oita, Osaka, Saga, Tochigi, Tokyo (Kuwayama 1949; Miyatake 1970; HIC, OMNH); Nepal: Kathmandu Valley, Mt. Phulchowki (Miyatake 1981).

Miyatake (1970) elaborately recorded the biology of the species on Lindera erythrocarpa and L. glauca. The females lay scattering eggs on the adaxial side of spread young leaves, and by oviposition, pit galls which protrude on the abaxial surface are formed, each is occupied by one later molted immature. The species seems bivoltine, and overwinters as adults on ever green trees (at least in temperate areas of Japan).

Fore wing with yellow bands (Fig. 11). Metabasitarsus with a pair of thickened setae on apex (Fig. 15). Paramere with a sclerotized tooth anteriorly (Figs 25, 26). Female proctiger long and smoothly tapering apically (Fig. 31).

Ground color yellow. Compound eyes grey. Long and thick setae on dorsum black. Ocelli yellow. Antennae yellow, with black spices on segments III-VIII; segments IX-X entirely black. Fore wing hyaline, with four obliquely transverse yellow stripes (Fig. 11). Legs yellow. Abdominal tergites brown. Male and female terminalia yellow.

Structures: Setae on dorsum of body relatively long (Table 1) and based on prominent projections. Torulus produced and slightly turned outwards (Fig. 3). Gena flat (Fig. 3). Antennal segments III-IX each with a single rhinarium on the apex, the ones on segments V and VII with small horn-shaped projections; proximally based terminal seta slightly longer than the distally based one (Fig. 7).

Mesoscutum with four pairs of prickly setae (Fig. 53). Metatibia with two rows of thick setae lateral-ventrally, and with a tightly packed row of short setae on the dorsum (Fig. 15). Apex of metabasitarsus with a pair of thick setae (Fig. 15). Pulvilli narrow (Fig. 15). Fore wing with long and narrow cell r₁, vein M₁₊₂ rather close to vein Rs, cell cu₁ tallest in the middle; vein M₃₊₄ lacking seta; surface spinules as tiny thick spines, widely spread across a large area on wing membrane; fields of radular spinules relatively large (Fig. 11).

Pore fields on abdominal ventrum long, narrow and curved; pores tightly packed (Fig. 18).

Male terminalia: Distal 1/3 of proctiger with posterior surface split and replaced with membranous tissue (Fig. 25). Paramere slender and bilobed; apex of anterior lobe developed into a sclerotized tooth; anterior margin of basal 1/3 emarginated; two long and thick setae present on inner surface, near the anterior margin; apical half with a curved vertical row of small peg setae on inner surface, near posterior margin (Fig. 25, 26). Aedeagus curved forward at apical 1/4, dorsum with a short row of spines that gradually turn smaller apically (Fig. 25). Dorsal-apical angle of subgenital plate produced and with a long seta (Fig. 25).

Female terminalia (Fig. 31): Long and straight in overall shape. Base of proctiger slight raised, apical process without tiny setae. Subgenital plate with acute apex, ventral surface with relatively dense and nearly evenly spaced setae.

Fifth instar immature. Body dorsum firmly sclerotized, with sclerites of thorax and abdomen almost unseparated; body ventrum weakly sclerotized. Dorsum of head, thorax, and abdomen with symmetrical acute sectasetae varying in size (Fig. 38); dorsum and margin of wing pads with roughly symmetrical acute sectasetae (Fig. 40). Head with 1+1 bulges, sheathing the central two pairs of long setae of adult head (Fig. 38); 1+1 projections present before fore wing pads, sheathing the 2+2 long setae on lateral margins of adult pronotum (Fig. 40). Antennae 9-segmented, apices of segments 5, 7 and 8 each with one single rhinarium (Fig. 39). Compound eyes with 1+1 ocular acute sectasetae, postocular acute sectasetae present in 2+2 (Fig. 38). Fore wing pad with two pores on dorsum (Fig. 40). Tarsal pulvilli narrow (Fig. 41). Abdominal ventrum with 5 pairs of spiracles surrounded by peritremes fused with central sclerites. Abdominal apex produced as a small pair of rounded bulges (Fig. 42). Anus terminal, circum anal rings present both dorsally and ventrally. Outer circum anal ring composed of oval pores, significantly expanded bilaterally, anterior aspect strongly crooked; inner circum anal ring composed of minute oval pores, expanded bilaterally, single rowed in the middle (Fig. 42).

Figure 43–46. 

Fifth instar immature of Togepsylla matsumurana. 43 Overall view, dorsal view on the left half, ventral view on the right half 44 Antenna, dorsal view 45 Claws, showing pulvilli 46 Circum anal ring, ventral view. Scale bar: 0.5 mm (43), 0.2 mm (44, 46), 0.05 mm (45).

Figure 47–53. 

Various parts of Togepsylla spp. 47–51, 53 Togepsylla takahashii 52 Togepsylla matsumurana 47 Head, ventral view 48 Labium 49 Hind wing 50 Tergites of abdominal segments 1–3 51 Sperm pump. Scale bar: 0.2 mm (47, 48, 50, 51), 0.5 mm (49), 0.32 mm (52, 53).

Figure 54–56. 

Comparison of coxa of different psyllid taxa. 54, 55 Cacopsylla sp. 56 Togepsylla takahashii 54 Mesocoxa and trochanter 55 Metacoxa and trochanter; 56. Metacoxa. Scale bar: 0.2 mm. Abbrevations: atc = anterior trochanteral condyle; de = dorsal edge; mr = meron; mrc = meracanthus; ptc = posterior trochanteral condyle; tct = trochanter; td = trochanteral tendon.

CHINA: 18 ♂, 21 ♀, 10 fifth instar immatures, Fujian, Shaxian, ex Lindera communis, 1.ix.1974, Chikun Yang and Fasheng Li (CAU, type series of Togepsylla minana); 35 ♂, 82 ♀, Guangxi, Liuzhou, 8.vi.1984, Fasheng Li (CAU); 40 ♂, 51 ♀, Guangxi, Lingchuan, Longkou, 5.vi.1984, Fasheng Li (CAU); 10 ♂, 5 ♀, Taiwan, New Taipei, Wulai, Fushan Nature Reserve, ex Lindera communis, 8.vi.2013, Xinyu Luo (CAU).

Lindera communis Hemsl., L. megaphylla Hemsl. (= L. oldhamii) (Lauraceae) (Takahashi 1936).

China: Fujian, Guangxi, Taiwan.

Biology: Takahashi (1936) and Li (2011) recorded that the immatures of the species feed on the abaxial surface of young leaves, inducing the edge of leaves to curl downwards, forming leaf-rolling galls that harbor large amounts of the insect. The immatures also secrete wax and honey dew. Severe damages to the host can cause most of the shoots to twist and shrink.

Paramere with large area of netlike grains covering the inner surface of apical half, anterior margin serrated (Figs 27, 28). Female proctiger short, curved upwards only at the tip (Fig. 32).

Adult coloration. Ground color yellow. Long and thick setae on dorsum yellow. Compound eyes grey. Ocelli yellow. Antennae yellow, with black spices on segments IV, VI, VIII; segments IX-X entirely black. Fore wing hyaline and colorless (Fig. 12). Male and female terminalia yellow.

Structures: Setae on dorsum of body relatively short (Table 1) and based on smooth projections. A pair of small tubercles present above toruli (Fig. 4). Gena moderately swollen bilaterally (Fig. 4). Antennal segments IV-IX each with a single rhinarium on the apex, segments IV, VI and VIII each with an extra rhinarium; all rhinaria with horn-shaped projections; proximally based terminal seta about equally long with the distally based one (Fig. 8).

Mesoscutum with five pairs of prickly setae. Metatibia with one row of thick setae ventrally, and with a tightly packed row of long setae on the dorsum. Pulvilli narrow. Fore wing with broad cell r₁, cell cu₁ tallest in the middle; vein M₃₊₄ with one seta on the base; surface spinules rather minute, widely spread across a large area in distal cells; fields of radular spinules unclear (Fig. 12).

Pore fields on abdominal ventrum large oval, with pores loosely packed.

Male terminalia: Proctiger completely sealed, with apex slightly thickened (Fig. 27). Paramere broad lamellar, with rather slender base; anterior margin of apical half emarginated, thin and serated; posterior margin with a basal ridge; apical half of inner surface with netlike grains; anterior angle with a few short and thick setae on inner surface; posterior margin with a row of inner-curved short setae on apical half (Figs 27, 28). Aedeagus curved backwards apically, dorsum lacking tiny spines, tip forming a small acute hook (Fig. 27). Subgenital plate near rectangular in profile, dorsal-apical angle with one long seta, ventral surface with sparse setae (Fig. 27).

Female terminalia (Fig. 32): Short and broad in overall shape. Apex of proctiger moderately curved upwards; apical half of proctiger with nearly evenly spaced setae, and with a row of setae along ventral margin of apical process. Subgenital plate with blunt and retracted apex, ventral surface with sparse setae.

Figure 57–58. 

Lateral aspect of thorax. 57 Togepsylla takahashii 58 Cacopsylla sp. Scale bar: 0.2 mm (57), 0.5 mm (58). Abbrevations: aac = anterior axillary cord; abas = anterior basalare sclerite; acl = anapleural cleft; adk = anepimeral disk; aepm = anepimeron; aeps = anepisternum; apwp = anterior pleural wing process; cx = coxa; cxc = coxal condyle; epm = epimeron; eps = episternum; hepm = heel of epimeron; kepm = katepimeron; keps = katepisternum; nt = notum; pa = pleural apophysis; pac = posterior axillary cord; pbas = posterior basalare sclerite; pcb = precoxal bridge; pbr = prealar bridge; pls = pleural suture; pnt = postnotum; ppt = parapteron; ppwp = posterior pleural wing process; psc = praescutum; sc = scutum; scl = scutellum; tems = transepimeral suture; tg = tegula; trn = trochantin.

Figure 59–61. 

Comparison of ventral aspect of thorax. 59 Togepsylla takahashii, ventral aspect of meso- and metathorax 60 Cacopsylla sp., ventral aspect of mesothorax 61 Trialeurodes vaporariorum, ventral aspect of meso- and metathorax. Scale bar: 0.2 mm. Abbrevation: es = extra sclerites.

Figure 62–63. 

SEM photographs of Togepsylla matsumurana. 62 Wax-secreting field on the sternite of abdominal segment 5, black arrow showing empty pore, white arrow showing wax thread secreted 63 Prickly setae on the vertex, showing detailed structure.

Fifth instar immature. Unknown.

CHINA: 49 ♂, 69 ♀, Tibet, Nyingchi, Mafenggou, 3050 m, ex Litsea sericea, 1.vi.1978, Fasheng Li (CAU, type series).

Litsea sericea (Nees.) Hook. f. (Lauraceae)

China: Tibet.

Yang and Li (1981) recorded that the adults gather among the clusters of young leaves by large amount. The record of a habit similar with T. takahashii by Li (2011) seems artificial.

Reassessment of morphology

Hind legs

Psyllids jump powerfully, then cast a mid-air rotation. Such a somersault, however, involves not only the strong muscles supported by the specialized metathoracic furca, enlarged metatrochanteral tendon and expanded meral part of the metacoxa but also a kicking of both hind legs on parallel planes (Burrows 2012), which are also parallel to the longitudinal body axis. This longitudinal placement of hind legs is caused by an inward twist of the metacoxa.

To discuss the formation of the enlarged and twisted metacoxa, one must seek reference from the mesocoxa. Mid and hind legs are both appendages of winged thoracic segments; additionally, in immature psyllids, they are equal in every detail, although differing from the forelegs in some aspects, indicating that hind legs of adults emerged from the model of mid legs. An undescribed Cacopsylla species is used as example:

The mesocoxa (Fig. 54) are relatively small and are connected to the coxal condyle of the mesopleurite by a dorsal-most articulation. Starting from the articulation, a thickened vertical edge runs down the outer surface, facing the lateral aspect, and is termed here as the ‘dorsal edge’ of the coxa. The coxa connects to the trochanter via two ‘trochanteral condyles’, which are longitudinally positioned, thus respectively termed ‘anterior-’ and ‘posterior trochanteral condyle’. Such longitudinal positioning of trochanteral condyles places the mid legs on a transverse plane, a plane nearly perpendicular to the longitudinal body axis. Besides, a normally developed trochanteral tendon originates on the inner-dorsal edge of the mesotrochanter, stretching into the chamber of the mesocoxa, clinging onto the corresponding muscles.

Compared with mesocoxa, the metacoxa (Fig. 55) first experienced an enlargement of the coxal wall, which pivots over the elongation of the dorsal edge and is primarily characterized as the expansion of the prearticular part of the coxal wall and thickening of the meron. Simultaneously, because of the unequal development of the prearticular part and the meron, the entire metacoxa is twisted backwards at approximately 90°, turning the two trochanteral condyles into a transverse position. The plane of hind leg is therefore turned longitudinal (Figs 66, 67). This pair of straightly backwards-reaching hind legs provides a much better concentration of jumping force, thereby driving the powerful jump described above. Additionally, the trochanteral tendon is magnified and possesses a tortuous apex, serving to support the strong jumping muscles.

By contrast, Togepsylla possesses half-modified metacoxae (Fig. 56). The enlargement is almost complete, but the positioning of the two trochanteral condyles is shifted at a limited level. For this reason, the hind legs of Togepsylla retain a posture similar to that of the middle legs, as shown in the habitus photograph (Fig. 64). Additionally, the trochanteral tendon is also half-enlarged: the relative size is much smaller, and the apex, although also expanded, is a simple flat surface instead of tortuous. According to the field observations by Xinyu Luo, adults of Togepsylla glutinosae sp. n. can only leap forward like frogs, at a short distance and without mid-air rotations.

Lateral aspect of thorax

Most psyllids possess an apophysis on meso- and metepisternal complex, termed ‘trochantinal apodeme’ (Ouvrard et al. 2002). This is an autapomorphy of Psylloidea. For mesopleuron, this structure may be on the anterior margin or median portion, depending on the taxon (Ouvrard et al. 2002). However, there are some cases like Togepsylla and Pseudophacopteron in which the trochantinal apodeme is placed on the anterior margin and reduced to an obscure vestige.

According to Ouvrard et al. (2002), the modification of psyllid metapleurite relative to mesopleurite is due to a curving of the pleural sulcus. For the metapleurite of most psyllids, taking Cacopsylla as example, the pleural sulcus turns downwards over the coxal condyle, becoming congruent with the elongated and internally ridged dividing suture of episternum and trochantin (Fig. 58). In Togepsylla, the metapleuron represents a halfway modification. The dividing suture of metepisternum and trochantin is absent, the trochantinal apodeme is shallow and in anterior position, as in mesothorax (Fig. 57).

Figure 64–67. 

Habitus, showing difference in the ways that hind legs are held. 64 Togepsylla glutinosae sp. n., adult 65 Togepsylla glutinosae sp. n., immature 66 Trioza urticae (Linnaeus), adult 67 Cyamophila hexastigma (Horvath), adult.

Wax-secreting fields on abdominal sternites

Togepsylla possesses three pairs of fields of pores on sternites of abdominal segments 4–6, in both sexes. Wax secretions from these pores have been observed on T. matsumurana (Fig. 62). Similar structures, several pairs of wax plates composed of many small wax-secreting pores, is one of the defining characters of adult whiteflies: Whiteflies kick the wax secretions of these glands with the hind legs, and then spread the shattered wax particles over the entire body surface (Byrne and Bellows 1991). In females of Aleyrodinae, two pairs of wax plates are found, on segments 3–4, whereas in Aleurodicinae, four, on segments 3–6; in males of Aleyrodinae, four pairs are present on segments 3–6, whereas three pairs appear on segments 3–5 in Aleurodicinae and Udamoselinae (Gill 1990; Martin 2007).

All the four members of Sternorrhyncha are known to secrete wax through integumental wax gland/pores. In scale insects whose wax glands are studied the most, these structures are highly variable in ultrastructure (shape and number of loculars of each pore) and distribution (all over the body or restricted to a certain region) (Foldi and Pearce 1985; Foldi and Lambdin 1995). Some aphid families/subfamilies possess wax gland plates, which also vary in shape and distribution, on body dorsum (Chen and Qiao 2012). These, however, are not so far known to reveal the same arrangement as Togepsyllinae and whiteflies, nor does the lack of detailed ultrastructural study of Togepsyllinae support their resemblance.

Psyllid immatures possess wax-secreting pores on their caudal plates. These pores are arranged in various patterns, mostly with a basic circum-anal ring (possibly homologous with the circum-anal ring of female adults), and on many occasions with extra pore fields (Brown and Hodlinson 1985). Extra pore fields can sometimes be succeeded by the adults, appearing on their more terminal (usually segments 7 and/or 8) abdominal tergites, e.g. Agonoscena pegani Loginova, 1960 and A. sabulisa Li, 1994 (in Li et al. 1994) (Luo 2016). Although it is not currently possible to accurately decide the homology between abdominal segments between immatures and adults, one can still roughly judge and count the separate segments of immatures by the dorsal and ventral setae rows. So far, the immature of not any species possess wax secreting pores on areas that are possibly homologous with abdominal sternites 4–6.

This is the first time that a psyllid adult is found with such fields of wax-secreting pores. Compared with those of whiteflies, wax pore fields of Togepsylla are strongly constricted, and the segment correspondence is different. It is uncertain whether these structures of Togepsylla and Aleyrodoidea are homologous or not.