Chinese psyllids in the genus Cacopsylla (Hemiptera: Sternorrhyncha: Psylloidea) associated with Spiraea (Rosaceae)

ABSTRACT The Chinese Cacopsylla species associated with Spiraea are revised. Three new species (Cacopsylla falcata sp. nov., C. nocturna sp. nov. and C. qilianensis sp. nov.) are described, and two [C. hyalinonemae Li and Yang, 1989 and C. spiraeicola (Li, 2011) comb. nov.] are redescribed. A new combination is also proposed for Cacopsylla tetrotaenialis (Li and Yang, 1989) comb. nov. Among the five species, C. falcata and C. qilianensis are affirmed to develop on Spiraea, as fifth instar immatures were found. The host plants of the other species remain uncertain. Based on morphological grounds, it is suggested that C. falcata and C. nocturna are closely related, but that the five species associated with Spiraea do not constitute a monophyletic group.

short and up-curving female terminalia (unpublished data). However, the species on pear are currently considered polygenous (Burckhardt and Hodkinson 1986;Luo et al. 2012).
Spiraea is a genus belonging to Rosaceae, Spiraeoideae, commonly seen in mountains, but not a common host plant for psyllids. Before the present research, only Cyamophiliopsis spp. are known to take Spiraea as the genuine host plant. As a small genus closely related to Cacopsylla, Cyamophiliopsis Li 2011 is undoubtedly considered monophyletic (Luo et al. 2015). Besides Cyamophiliopsis, the only species associated with Spiraea are Euphaleropsis spiraeicola Li, 2011 and Homotoma spiraeae Yang and Li, 1981. In the collections of the last few years, we discovered four more species of Cacopsylla associated with Spiraea, and discovered that Euphaleropsis spiraeicola is rather a Cacopsylla species. In the present paper, we seek to coordinate all of the species, define them, and discuss their phylogenetic relationships.

Materials and methods
All specimens were acquired from the collections of Entomological Museum, China Agricultural University.
Preparation of slides were made following this protocol: whole insect boiled in potassium hydroxide (KOH) solution for 10 minutes, naturally cooled down after heating stopped, then washed in distilled water, and finally mounted on a slide in glycerine. All drawings, examinations and photos were undertaken with an Olympus ® BX41 microscope. In the drawings of fore wings (Figures 1(f), 3(f), 5(f), 6(f), 8(f)), thin dashed lines represent ranges of surface spinules (on the dorsal surface), while thick dashed lines (except the one marking the claval suture) represent ranges of radular spinules (on the ventral surface).
Measurements (Table 1) were taken with a Keyence ® VHX-1000 digital microscope using its measuring function and are given in millimetres (mm). For adults: BL = total body length measured from anterior margin of head to tip of folded fore wing, HW = head width, AL = antennal length, TW = mesoscutum width, WL = fore wing length, TL = metatibial length. For fifth instar immatures: BL = total body length, HW = head width, FL = fore wing pad length, measured as the distance between transverse tangents of anterior angle and posterior margin.
For nomenclature of plants, we follow Lu and Alexander (2003).

Description
Adult. Colouration (Figure 10(a-b)). Body orange in overall view. Vertex mostly orange, with inner-posterior angles white. Genal processes ochre. Compound eyes purple, ocelli orange. Occiput black. Antenna yellow, with black apices on segments III-XIII, and segments IX-X entirely black. Thoracic dorsum yellow in ground, with the vast majority covered by orange stripes and patterns; anterior half of pronotum irregularly tainted orange; the two triangular patches in mesopraescutum almost covering the whole ground. Metapostnotum with two black patches bilaterally. Ventrum of mesothorax   black except for a transverse narrow band in the middle. Legs yellow, with outer wall of metacoxa dark brown. Fore wing membrane (Figure 1(f)) hyaline, with a dark brown patch covering subapex of cell a 1 , expanding around as a light brown patch near anal break; veins brown. Abdominal terga yellow, with anterior margin of each segment blackish; abdominal sterna yellow in ground, with anterior half of segment III black, other segments with an indistinctly demarcated light brown patch in anterior margin near lateral margin, all segments with a small brown patch in posterior half near lateral margin. Male terminalia yellow, with ventral surface of subgenital plate lightly brownish. Female terminalia yellow, with apex brownish.
Structures. Head (Figure 1(a)) inclined from longitudinal body axis by 80-90°, slightly narrower than mesoscutum transversely. Lateral parts of vertex rather short longitudinally; anterior margin and base of lateral ocelli strongly convex, appearing distinctly contrasted from the inner-posterior angles; boundary between vertex and gena clear. Surface of vertex finely sculptured with scaly microstructures and microscopic setae that gradually grow longer in the antero-inner angles, exceeding the length of the setae around antennal insertion. Gena processes long, gradually growing divergent, with apices subacute; outer margin relatively smoothly curved; genal whip setae relatively widely spaced. Antenna longer than HW; the more distally situated terminal seta only slightly shorter than the more proximally situated terminal seta (Figure 1(g)).
Mesopraescutum strongly produced forward, pressing pronotum to be strongly arched. Metatibia with well developed genual spine, apical spurs arranged in 1 + 1 + 2 + 1. Fore wing (Figure 1(f)) oblong oval, strongly widening until apical 1/3; posterior margin nearly straight; height of cell cu 1 distinctly longer than length of vein Cu 1b ; fields of surface spinules relatively small, slightly contracted at margins of cells; fields of radular spinules as shown in Figure 1(f). Male terminalia: Proctiger (Figure 1(b)) without posterior lobe, gently arched, with anterior surface waved near apex; apical 4/5 of anterior surface setosed. Paramere (Figures 1(b-c)) slightly curved forward, with apex curved backwards; apical tooth curved inwards, with tip subacute and pointed forward; anterior margin expanded into a small rounded lobe at subapex; inner surface with large numbers of long setae curving downwards, posterior margin also with large numbers of long setae. Distal segment of aedeagus (Figure 1(d)) relatively long and slender; apical dilatation sickleshaped; sclerotised end tube of ductus ejaculatorius projected backwards, and gently curved upwards. Subgenital plate (Figure 1(b)) near square in profile, except for the finely rounded ventral surface; weakly sclerotised anterior lobe present.
Female terminalia (Figure 1(e)): Relatively long and simple. Anal ring covering about 2/5 of the total length of proctiger; proctiger slightly curved upwards; longitudinal row of rather long setae in dorsum of apical process formed of: four longest in the base, forming a strongly curved line, one short seta in apical 1/3, and one shortest in apical 1/ 5; fields of peg setae large, almost touching in the middle. Subgenital plate relatively long and narrow in profile, field of peg setae in subgenital plate covering apical 1/2. Valvulae dorsalis and ventralis gently curved upwards, stretching into the cavity beneath proctiger; valvulae lateralis rather narrow.
Structures. Dorsum completely covered with minute to long capitate setae, including ocular and postocular setae (Figure 2(a)). Antenna (Figure 2(a)) 7-segmented, with one rhinarium on apex of segments 3 and 5 each, and two rhinaria on segment 7. Wing pads (Figure 2(a)) large; fore wing pad oval, with about 10 long capitate setae on outer margin, anterior half with two pores; hind wing pad knife-shaped, with two long capitate setae on apex, anterior angle with one pore. Dorsum of fore tibialtarsus with one long capitate seta, dorsum of mid and hind tibialtarsus with two long capitate setae (Figure 2(a)). Tarsal arolium (Figure 2(b)) wide fishtailshaped, with expanded areas rather coarse, with relatively short pedicel gradually growing narrow basally, and with well-developed unguitractor. Third pair of sclerites with spiracle in abdominal ventrum completely fused with anal plate (Figure 2(a)). Abdominal ventrum (Figure 2(a)) with long or short capitate setae near the margin, other parts with long or short simple setae. Abdominal margin (Figure 2(a)) with 7 + 7 rather long capitate setae, gradually growing shorter medially, and with 3 + 3 setae close to the lanceolate type (sensu White and Hodkinson 1985). Circum anal pore field (Figure 2(c)) ventral and small, composed of a complete outer ring of single row of slit-shaped pores, and a complete inner ring of jagged single row of oval pores. 1 + 1 setae near the lanceolate type present anterior to anal pore field.
Colouration ). Body light brown in overall view. Vertex pale yellow in ground, with two longitudinal ochreous stripes consistent with those on thoracic dorsum, along with two narrow ochreous stripes beside. Antennal base ochreous except for margin of antennal insertion. Dorsal aspect of genal processes pale yellow in ground, with two longitudinal red stripes; ventral aspect of genal processes brown. Compound eyes red, ocelli orange. Antenna pale yellow, with black apices on segments III-VIII, and segments IX-X entirely black. Thoracic dorsum pale yellow in ground, with ochreous stripes; two longitudinal stripes present through pronotum and mesopraescutum, consistent with the stripes on vertex; lateral stripes on mesoscutum with conspicuous dark brown outlines. Thoracic pleurites mostly brown, dorsal bulge of mesopleuron pale yellow. Legs yellow, coxae black, femora more or less darkened. Fore wing membrane ( Figure 3(f)) golden yellow and hyaline, leaving colourless bands along veins; margin brown, gradually lightening basally; pterostigma dark brown; veins mostly colourless, marginal vein yellow; vein A 1 with several black sections varying in length. Hind wing membrane (Figure 3(g)) hyaline, outer and caudal margin, and part of cell cu 2 black. Abdomen dark brown, tergum of segment III ochreous. Male terminalia ochreous, apical tube of proctiger dark brown, subgenital plate more or less blackish. Female proctiger ochreous, subgenital plate dark brown.
Structures. Head (Figure 3(a)) inclined from longitudinal body axis by about 45°, slightly wider than mesoscutum transversely. Antero-outer and antero-inner angles of vertex protruding as round tubercles; boundary between vertex and gena clear. Surface of vertex finely sculptured with scaly microstructures and microscopic setae. Genal processes long and rather robust, slightly tilted upwards, gradually growing divergent, apices rounded. Demarcation between antennal base and genal process rather distinct as a crease. Antenna relatively slender, more distally situated terminal seta about 2/3 as long as the more proximally situated terminal seta (Figure 3(h)). Mesopraescutum moderately produced forward, pressing pronotum to be gently arched. Metatibia with well developed genual spine, apical spurs arranged in 1 + 3 + 1. Fore wing (Figure 3(f)) oblong oval, strongly widening until apical 1/4; posterior margin nearly straight; height of cell cu 1 distinctly longer than length of vein Cu 1b ; vein A 1 with narrow flag expanded outwards; fields of surface spinules relatively large, contracted at apices of cells; fields of radular spinules as shown in Figure 3(f). Male terminalia: Proctiger (Figure 3(b)) without posterior lobe, gently arched, with nearly evenly spaced setae; posterior margin moderately produced. Paramere (Figure 3 (b-c)) slightly curved forward, outer surface with two ridges basally; apical tooth curved inwards, with tip subacute and pointed forward; inner surface with lots of long setae pointing downwards, posterior margin also with lots of long setae. Distal segment of aedeagus (Figure 3(d)) relatively long, nearly straight; apical dilatation short and stout, slightly hooked apically; sclerotised end tube of ductus ejaculatorius projected backwards, and gently curved upwards. Subgenital plate (Figure 3(b)) nearly round in profile.
Female terminalia (Figure 3(e)): Relatively short. Anal ring covering about 1/3 of the total length of proctiger; longitudinal row of rather long setae in dorsum of apical process not clearly recognisable; setae in bilateral sides of apical process of proctiger gradually turn from short setae basally into peg setae apically. Subgenital plate relatively long and narrow in profile, peg setae present only near apex. Valvulae dorsalis and ventralis gently curved upwards.

Host plant
Probably Spiraea sp., as although there were only a few adults collected on the plant, no adult was collected from the surrounding plants. Li (2011) listed Syrina oblata as the host plant, but it is most probably a shelter plant or food plant.

Remarks
This species is rather unique within the genus, attributed to the longitudinal stripes through the vertex to mesopraescutum, the red stripes on the genal processes, the less inclined head, and the protruding anterior angles of the vertex. Psylla tetrotaenialis Li and Yang, 1989 is another species with such characters; we regard it as a closely related species to C. hyalinonemae, and hereby assign it to Cacopsylla as Cacopsylla tetrotaenialis (Li and Yang, 1989) comb. nov.

Description
Adult. Colouration (Figure 10(e-f)). Body dark brown to black in overall view. Vertex white in ground, mostly covered by brown markings except for antero-inner angles, antero-outer angles and postero-inner angles; discal foveae dark brown. Genal processes ochre, with apical half brown. Compound eyes black; lateral ocelli orange, medial ocellus brown. Occiput black; postocular sclerites black, with margin entirely white. Antenna brown, with black apices on segments III-XIII, and segments IX-X entirely black. Thoracic dorsum mostly orange, with brown to dark brown stripes and patterns; pronotum with three brown markings in the middle; the two triangular patches in mesopraescutum almost covering the whole ground. Metapostnotum entirely black. Thoracic pleurites almost completely black, except for the antero-dorsal bulging of mesopleuron. Legs yellow, with pro-and mesocoxae black, metacoxa irregularly blackish; all femora black. Fore wing (Figure 4(f)) membrane hyaline, with a black marking near anal break; veins yellow, gradually turning brown apically, with C + Sc, base of A 2 , and outer margin before anal break brown. Abdomen black, with a longitudinal white band across the lateral aspect of terga of segments III-VI. Male and female terminalia black.
Structures. Head (Figure 4(a)) inclined from longitudinal body axis by 90°, slightly wider than mesoscutum transversely. Lateral parts of vertex rather short longitudinally; anterior margin and base of lateral ocelli strongly convex, appearing distinctly contrasted from the inner-posterior angles; antero-outer angles distinctly bulging; boundary between vertex and gena clear. Surface of vertex finely sculptured with scaly microstructures and microscopic setae. Gena processes long and slender, gradually growing divergent, with apices subacute; outer margin strongly emarginated; genal whip setae relatively narrowly spaced. Antenna longer than HW; terminal setae rather long, the more distally situated one about 3/4 as long as the more proximally situated one (Figure 4(g)).
Male terminalia: Proctiger (Figure 4(b)) without posterior lobe, gently arched, with nearly evenly spaced short setae. Paramere (Figure 4(b-c)) slightly curved forward, with apex curved backwards; apical tooth relatively small, curved inwards, with tip subacute and pointed forward; base better expanded, near rectangular; inner surface with large amounts of long setae curving downwards, posterior margin also with large numbers of long setae. Distal segment of aedeagus (Figure 4(d)) relatively long, robust basally, smoothly growing slender apically until apical dilatation; apical dilatation rather small, strongly hooked; sclerotised end tube of ductus ejaculatorius projected upwards, and gently curved forward. Subgenital plate (Figure 4(b)) rather small, with weakly sclerotised anterior lobe.
Female terminalia (Figure 4(e)): Relatively long and simple. Anal ring covering less than 1/3 of the total length of proctiger; longitudinal row of rather long setae in dorsum of apical process formed of: four longest in the base, forming a weakly curved line, one short seta in apical 2/5, and one shortest in apical 1/5; fields of peg setae barely touching in the middle. Subgenital plate relatively long and narrow in profile, field of peg setae in subgenital plate covering apical 2/3. Valvulae dorsalis and ventralis gently curved upwards.

Host plant
Probably Spiraea teniana. Before the light trap, we also observed several adults on the plant at dusk.

Etymology
Named after the time the type series was collected, 'nocturnus' = 'night'.

Description
Adult. Colouration (Figure 10(g-h)). Body orange in overall view. Vertex orange. Genal processes yellow. Compound eyes brown, ocelli orange. Antenna yellow, with black apex on segment VIII, and segments IX-X entirely black. Thoracic dorsum pale grey, with reddish orange stripes. Thoracic pleurites mostly orange, trochantin of metathorax more or less blackish. Ventral aspect of mesothorax black, except for a transverse narrow band in the middle. Legs yellow; metacoxa blackish; dorsal surface of femora blackish. Fore wing ( Figure 5(f)) membrane translucent, grey, darker in apical margin, gradually turning light basally; veins slightly darker than membrane. Abdomen orange; lateral end of anterior margin of terga of segments III-VII light brown to black, darkest in segment III, gradually lightening by each segment; ventral surface with two longitudinal brown bands. Male and female terminalia orange.
Structures. Head (Figure 5(a)) inclined from longitudinal body axis by 90°, slightly narrower than mesoscutum transversely. Boundary between vertex and gena clear and nearly straight, antero-outer angles nearly rounded; plane of vertex distinctly higher than that of genal processes; discal foveae relatively shallow. Surface of vertex finely sculptured with scaly microstructures and microscopic setae. Gena processes about half as long as median suture of vertex, moderately divergent apically, with apices blunt. Antenna about as long as HW; terminal setae relatively short, the more distally situated one about 1/3 as long as the more proximally situated one (Figure 5(g)).
Mesopraescutum moderately produced forward, pressing pronotum to be moderately arched. Metatibia without genual spine, with five apical spurs (in one case six), midial ones irregularly grouped. Fore wing (Figure 5(f)) oblong oval, widest in apical 1/3; height of cell cu 1 indistinctly longer than length of vein Cu 1b ; surface spinules small and dense, completely covering the whole fore wing membrane; fields of radular spinules as shown in Figure 5(f).
Male terminalia: Proctiger ( Figure 5(b)) relatively short and thick, with produced posterior margin, gently arched, covered with nearly evenly spaced short setae. Paramere ( Figure 5(b-c)) blade-shaped in profile, strongly curved forward, without distinct apical tooth; inner surface with rather few setae; anterior half of the base with relatively dense setae. Distal segment of aedeagus ( Figure 5(d)) relatively short; apical dilatation near oval, slightly inclined downwards, not hooked; sclerotised end tube of ductus ejaculatorius projected obliquely backwards, and gently curved upwards. Subgenital plate (Figure 5(b)) near square in profile, dorsal margin slightly folded inwards; ventral surface with relatively dense setae.
Female terminalia (Figure 5(e)): Rather short. Longitudinal row of rather long setae in dorsum of apical process not clearly recognisable; bilateral sides of apical process of proctiger without peg setae, short and slender setae present instead. Subgenital plate rather short, forming a wide opening with proctiger, without peg setae. Valvulae dorsalis and ventralis stretching obliquely upwards, partly hidden beneath the cavity of proctiger.
Fifth instar immature. Colouration. Membranous parts orange. All sclerites dark brown. Antenna yellow, with apical three segments black. Compound eyes reddish brown.
Structures. Dorsum completely covered with a type of specialised simple setae (awlshaped, termed conical seta below), including ocular and postocular setae (Figure 6(a)). All bases of setae strongly sclerotised (Figure 6(a)), following the colour of normal sclerites. Antenna (Figure 6(a)) 7-segmented, with one rhinarium on apex of segments 3 and 5 each, and two rhinaria on segment 7; segment 7 relatively short, with the two rhinaria relatively close. Wing pads (Figure 6(a)) large; fore wing pad oval, outer margin decorated with dense conical setae varying in length, anterior half with one pore; hind wing pad knife-shaped, with two long conical setae on apex. Dorsum of mid and hind tibialtarsus with two long capitate setae (Figure 6(a)). Tarsal arolium (Figure 6(b)) fishtailshaped, with expanded areas rather coarse, with relatively long pedicel gradually growing narrow basally, and with well-developed unguitractor. Third pair of sclerites with spiracle in abdominal ventrum completely free from anal plate (Figure 6(a)). Abdominal ventrum (Figure 6(a)) with long or short conical setae near the margin, other parts with long or short simple setae. Abdominal margin (Figure 6(a)) with 7 + 7 rather long conical setae, innermost pair relatively short; and with 2 + 2 sectasetae. Circum anal pore field (Figure 6(c)) ventral, composed of a complete outer ring of single row of slit-shaped pores, and a complete inner ring of jagged single row of oval pores.

Etymology
Named after the type locality, the Qilian Mountains.

Redescription
Adult. Colouration (Figure 10(i-j)). Body brown in overall view. Vertex brown, discal foveae moderately darker. Genal processes slightly darker than vertex. Compound eyes black; lateral ocelli brown, medial ocellus orange. Antenna generally brown to dark brown, with basal half of segments III-V lighter. Thoracic dorsum with dark brown stripes and patterns. Thoracic pleurites generally brown, irregularly darkened along pleural sulci. Legs yellowish brown; pro-and mesocoxae dark brown, metacoxa irregularly darkened; trochanters dark brown; femora more or less darkened. Fore wing (Figure 7(f)) membrane hyaline, with brown to dark brown clouding, leaving breaches at apices of cells r 2 , m 1 , m 2 and cu 1 ; most veins brown, costal margin yellow. Hind wing membrane hyaline, more or less brownish along veins; veins light brown to brown. Abdomen and male and female terminalia dark brown to black.
Structures. Head (Figure 7(a)) inclined from longitudinal body axis by 45°, slightly narrower than mesoscutum transversely. Boundary between vertex and gena clear, without outstanding antero-outer angles; discal foveae relatively shallow, secondary foveae slightly concave. Surface of vertex finely sculptured with scaly microstructures and microscopic setae that gradually grow longer on antero-inner angles, exceeding length of setae around antennal insertion. Plane of genal processes inclined from that of vertex by about 45°. Gena processes about half as long as median suture of vertex, widely divergent apically, with apices nearly rounded. Antenna longer than HW, relatively thick; terminal setae rather short, the more distally situated one about 1/3 as long as the more proximally situated one (Figure 7(g)).
Mesopraescutum moderately produced forward, pressing pronotum to be moderately arched. Metatibia without genual spine, with four apical spurs, arranged in 1 + 2 + 1. Fore wing (Figure 7(f)) oblong oval, widest in the middle; veins relatively thick; height of cell cu 1 indistinctly longer than length of vein Cu 1b ; surface spinules (in dorsal surface) and radular spinules (in ventral surface) completely covering the whole fore wing membrane.
Male terminalia: Proctiger (Figure 7(b)) without posterior lobe, gently arched, covered with nearly evenly spaced short setae. Paramere (Figures 7(b-c)) strongly curved forward at basal 1/4, then gently curved backwards; apical tooth finely congruent with the main part, moderately curved inwards, with tip acute and pointed forward; inner surface with dozens of erect short setae. Distal segment of aedeagus (Figure 7(d)) gently curved downwards; apical dilatation near oval, not hooked; ductus ejaculatorius relatively thick, sclerotised end tube projected obliquely backwards, and gently curved upwards. Subgenital plate (Figure 7(b)) with small anterior lobe, ventral surface waved in profile.
Female terminalia (Figure 7(e)): Relatively long and simple. Anal ring rather small; dorsal surface of proctiger smoothly convex near base of apical process; longitudinal row of rather long setae in dorsum of apical process formed of nearly evenly spaced setae. Subgenital plate relatively long and narrow in profile, without relatively long setae near dorsal margin. Valvulae dorsalis and ventralis slightly curved upwards.

Host plant
Unknown. In April of Liaoning province and Beijing, all psyllids are still in hibernation. Spiraea spp. recorded here are probably shelter plants.

Discussion
Due to their conspicuous diversity in morphology, the five species involved in this article do not constitute a monophyletic group. However, Cacopsylla falcata sp. nov. and Cacopsylla nocturna sp. nov. display great resemblance in: female not conspicuously larger than male in size, vertex rather short longitudinally in lateral aspect and with strongly bulging anterior portion, long and slender genal processes, oval fore wing widest at apical 1/3, brown marking near anal break of fore wing, and rather long setae on apical process of female proctiger arranged in transverse rows. These characters define the two species as close relatives. Taking the immature characters of C. falcata (long capitates setae on body margin, 3 + 3 setae close to the lanceolate type on abdominal margin) into account, they are relatively more typical Cacopsylla members.  Li and Yang, 1989; (e-f) C. nocturna sp. nov.; (g-h) C. qilianensis sp. nov.; (i-j) C. spiraeicola (Li, 2011). a, c, e, g, i. lateral view; b, d, f, h, j. dorsal view. Scale bar = 1 mm.
Although C. spiraeicola lacks genual spine on metatibia, possesses only four apical spurs, and bears rather dense surface spinules and radular spinules covering entire fore wing membrane, these characters are not crucial for Cacopsylla. The patterns on fore wing superficially resemble some Elaeagnus-feeding species such as Cacopsylla gossypinmaculosa Li 2011, but are intrinsically different from the latter. Patterns of C. spiraeicola leave breaches at the middle of cells, whereas that of the Elaeagnus feeders leave breaches around the apex of veins. We provisionally assign the species to Cacopsylla as it still resembles Cacopsylla more than it does Euphaleropsis, as the latter is an Indigofera-feeding genus of Macrocorsinae, mainly distributed in the Oriental Region, characterised as possessing black spots on the veins of the fore wing.
Cacopsylla qilianensis sp. nov. is another unique species that currently cannot be grouped with any other species. It possesses a rather distinctly outlined vertex whose plane is conspicuously above that of the gena and postocular sclerite, rather dense surface spinules covering the entire fore wing membrane, uniform radular spinules covering rather large areas on the fore wing, posteriorly produced male proctiger, paramere lacking conspicuously sclerotised apical tooth, female proctiger lacking peg setae and rows of long setae on apical process, and conical seta on dorsum of immature. These characters cannot be found in any other Cacopsylla species. Further findings are needed to determine the particular systematic position of C. qilianensis sp. nov.
Both Cacopsylla falcata sp. nov. and C. nocturna sp. nov. were collected in the mountainous areas in northern Zhaotong, a city in north-eastern Yunnan province, lying south of the Yangtze River (Figure 9). The physiognomy of forest above 1000 m in this area appears much more temperate than tropical, presenting us many undescribed Rosaceae-feeding Cacopsylla species, most of which follow the similarity of morphology stated above, and develop on host plants never recorded for psyllids. Our expedition in 2014 was very whistle-stop, and some further species around these two in future collections in this area can be expected.