taxonID	type	description	language	source
03B287EEFFEFFF9A39B48152FE3EFAAC.taxon	description	Torulus Li, 1991: 37, 41. Type species: Torulus sinicus Li, by original designation and monotypy. Syn. nov. Diagnosis: Adult. Body large and robust, beset with long pubescence. Head narrower than mesothorax; inclined from longitudinal body axis at almost 90 °. Vertex subrectangular, lacking anterior transverse suture separating it from the genal processes; genal processes conical, subacute, about as long as vertex along midline. Eyes large, hemispherical. Antenna 10 - segmented with a single subapical rhinarium on segments 4, 6, 8 and 9; 1.3 ‒ 1.9 times as long as head width; flagellar segments sparsely beset with long setae; segment 10 with one terminal seta longer and one shorter than segment. Clypeus small, narrowly pyriform; hidden by procoxae and genal processes; rostrum short, in lateral view only tip of apical segment visible. Pronotum antero-medially curved downward and laterally backward; head posteriorly sometimes hiding pronotum medially. Dorsal outline of mesopraescutum, in lateral view, weakly or strongly curved. Mesosternum angularly concave antero-medially; basisternum indented postero-medially, raised laterally to form a tubercle on either side; katepisternum triangular, large; pleurosternal suture well-developed; precoxale forming a right angle. Legs relatively short and robust; metacoxa with spur-shaped meracanthus, not extended antero-medially; metatibia with a blunt tubercule at base, with 1 + 3 apical spurs, each on a slightly raised tubercle. Forewing lanceolate to obovate, usually with dark pattern; vein C + M + Cu strictly trifurcating; veins beset with conspicuous setae. Hindwing about three quarters length of forewing; costal setae ungrouped; vein M + Cu developed. Abdomen with short dorsal and long ventral setae. Male proctiger strongly bulging posteriorly, beset with setae. Paramere lamellar. Proximal segment of aedeagus slender, strongly curved in basal half; distal segment with weakly inflated, curved apical dilation. Female terminalia cuneate, relatively short. Fifth instar immature. Body weakly sclerotised, elongate, 1.5 – 1.9 times as long as wide. Cephalothorax on either side with an area laterally delimited by eye in anterior third and an area in posterior third densely covered in small, very slender sectasetae; anterior area also has a few scattered long simple setae. Antenna 8 - segmented with an apical rhinarium on segments 4 and 6, and two on segment 8. Thoracic and abdominal tergites densely covered in small, slender sectasetae. Legs robust, 4 - segmented; tibiotarsus longer than femur; tarsus has two moderately sized claws; tarsal arolium longer than claws, broadly triangular, weakly concave apico-medially, with petiole and unguitractor. Forewing pad long and slender, lacking humeral lobe; with marginal stripe of dense, small and slender sectasetae and dorsal stripe in apical third of similar sectasetae. Hindwing pad with a few marginal and a stripe of dorsal, densely spaced small, slender sectasetae. Anus ventral. Outer circumanal ring moderately large, irregularly heart-shaped, slightly wavy, consisting of a single row of elongate and narrow pores. Abdominal venter beset with long normal setae. Distribution: Oriental Region. China (Li 2011), India (Mathur 1975), Japan (Miyatake and Matsumoto 2008), Laos (Cho et al. 2017), Philippines (Crawford 1917), Taiwan (Yang et al. 2013), Nepal and Sri Lanka (unpublished NHMB data). Host plants: Ilex spp. (Aquifoliaceae). The immatures induce marginal fold galls on the leaves. Mathur (1975) reported Petalolyma basalis (Walker) from India “ on leaves of Quercus dilalata ” [= Quercus floribunda Lindl. ex A. Camus] (Fagaceae). He did not mention immatures nor provide a description of them, suggesting that Quercus is a casual plant rather than a host. Castanopsis tibetana Hance (Fagaceae) is also reported as a host of P. castanopsis Li & Yang by Li and Yang (1991) based on two males, representing a casual host. The host of this species is Ilex asprella Champ. ex Benth. (see below). Similarly, we suspect that Vernicia fordii (Hemsl.) Airy-Shaw (Euphorbiaceae) is a casual plant of Torulus sinicus (= Petalolyma lii Liao & Burckhardt, nomen nov.), as it is based on one specimen only (Li 1991). Comments: Petalolyma, as diagnosed here, constitutes a well circumscribed, putatively monophyletic taxon within the Triozidae. Potential synapomorphies constitute the long and slender apical dilation of the distal aedeagal segment and the distribution pattern of the very slender, small sectasetae in the immature. The phylogenetic significance of these characters is, however, difficult to evaluate without a phylogenetic framework. Petalolyma was not included in the molecular phylogenetic analysis of Percy et al. (2018) and comparable morphological studies are also lacking. To keep Torulus as a separate genus would render Petalolyma paraphyletic, as the latter would be defined by symplesiomorphies only. To prevent this, we propose the following synonymy: Petalolyma Scott, 1882 = Torulus Li, 1991, syn. nov. We transfer Torulus sinicus to Petalolyma as Petalolyma sinica (Li, 1991), comb. nov., which constitutes a secondary homonym of Petalolyma sinica Yang & Li, 1984. We propose the replacement name Petalolyma lii Liao & Burckhardt, nomen nov.	en	Liao, Yi-Chang, Burckhardt, Daniel, Yang, Man-Miao (2024): Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li. Zoological Studies 63 (28): 1-12, DOI: 10.6620/ZS.2024.63-28, URL: http://dx.doi.org/10.5281/zenodo.14704401
03B287EEFFEDFF9E3A9781ACFAB3FBD5.taxon	materials_examined	Material examined: Taiwan: 6 ñ, Nantou, Jihyeutan (Sun Moon Lake), 10 - III- 1985 (S. J. Fang) (NCHU, dry mounted); 4 ò, 3 ñ, Pingtung, Shizi, Neiwen Village, South Link Highway, N 22 ° 14 ' 36.327 '', E 120 ° 49 ' 56.988 '', 480 m, 1 - XII- 2023 (D. Burckhardt) # 23 - 33 (-) (NHMB, 70 % ethanol); 1 ò, 2 ñ, same but Neiwen Village, Dongyuan Road, N 22 ° 14 ' 36.575 '', E 120 ° 50 ' 12.992 '', 450 m, # 23 - 34 (-). Description: Adult. Yang (1984); Li (2011); Yang et al. (2013). Fifth instar immature. Fang and Yang (1986). Member of the divisa species group. Distribution: Taiwan (Nantou) (Fang and Yang 1986; Yang et al. 2013), Pingtung (present paper); China (Fujian Province) (Li 2011). Host plant: Ilex asprella Champ. ex Benth. (Aquifoliaceae) (Li 2011). Comments: Li and Yang (1999), when describing P. fujianensis (divisa species group), listed differences in the forewing venation to P. zhejiangana Yang & Li (bicolor species group) but did not mention P. castanopsis (divisa species group). Li (2011) pointed out the misidentification of Trichochermes bicolor from Taiwan by Fang and Yang (1986), referring the Taiwanese species to P. fujianensis, and separated the latter from P. castanopsis in couplet 6 of the identification key of Petalolyma species, suggesting that the two are morphologically most similar. According to Li’s (2011) key, the species differ in the length ratio of veins Cu 1 a / Cu: 1.29 in P. castanopsis versus 1.83 in P. fujianensis. These values appear wrong according to our measurements of Li’s (2011) drawings: 1.33 in P. castanopsis versus 1.36 in P. fujianensis. The difference in the posterior outline of the male proctiger listed by Li (2011) reflects minor intraspecific variation. As the two nominal species are morphologically identical, we propose the following synonymy: Petalolyma castanopsis Li & Yang, 1991 = Petalolyma fujianensis Li & Yang, 1999, syn. nov. Petalolyma formosana Yang, 1984 (Figs. 1 C – D, I – J, 2 I, L, O)	en	Liao, Yi-Chang, Burckhardt, Daniel, Yang, Man-Miao (2024): Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li. Zoological Studies 63 (28): 1-12, DOI: 10.6620/ZS.2024.63-28, URL: http://dx.doi.org/10.5281/zenodo.14704401
03B287EEFFEDFF9E3A9781ACFAB3FBD5.taxon	materials_examined	Material examined: Taiwan: Holotype ò, paratypes 9 ò, 9 ñ, Taichung, Anmashan, 22 - VII- 1982 (C. T. Yang) (NCHU, dry); 1 ò 1 ñ, same but 6 - VIII- 1987 (C. T. Yang) (NCHU, dry); 2 ò, Nantou, Meifeng, 2150 m, 2 – 4 - VI- 1980 (L. Y. Chou & C. C. Chen) (TARI, dry); 1 ò 1 ñ, Nantou, Tsuifeng, 2300 m, 3 - VI- 1980 (L. Y. Chou & C. C. Chen) (TARI, dry); 1 ò, same but 25 - 27 - VI- 1981 (K. S. Lin & W. S. Tang) (TARI, dry). Description: Adult. Yang (1984); Yang et al. (2013). In the type series, the dark forewing pattern is restricted to the basal quarter of the wing with an occasional faint narrow light brown band along the anal margin (Yang 1984; Yang et al. 2013). In some recently collected specimens at hand (TARI), the pattern is more expanded with a broad, well defined dark patch along the anal margin (Fig. 1 C) or two patches along the costal margin (Fig. 1 D). Fifth instar immature. Unknown. Member of the formosana species group. Distribution: Taiwan (Nantou, Taichung) (Yang 1984; Yang et al. 2013). Host plant: Unknown.	en	Liao, Yi-Chang, Burckhardt, Daniel, Yang, Man-Miao (2024): Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li. Zoological Studies 63 (28): 1-12, DOI: 10.6620/ZS.2024.63-28, URL: http://dx.doi.org/10.5281/zenodo.14704401
03B287EEFFE8FF9F3A97867DFBA7FECF.taxon	discussion	Comments: Kuwayama (1910) described Trichochermes hyalina based on two males from “ Formosa ”. No further specimens of this species have been reported in the literature. Yang and Li (1984) transferred the species to Petalolyma but it is not clear on what evidence this decision was based. The original description is not diagnostic and the examination of the types is therefore indispensable to uncover the identity of the species. According to H. Inoue (pers. comm.), the two type specimens are missing from Kuwayama’s collection (SEHU) since 2012, making Petalolyma hyalina a nomen dubium. Petalolyma lii Liao & Burckhardt, nomen nov. (Figs. 1 B, G, H, 2 B, D, F, H, N, K) urn: lsid: zoobank. org: act: 5 F 01 DB 62 - 1 BFF- 4 D 09 - 9 ECD- A 16730 C 90 B 74	en	Liao, Yi-Chang, Burckhardt, Daniel, Yang, Man-Miao (2024): Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li. Zoological Studies 63 (28): 1-12, DOI: 10.6620/ZS.2024.63-28, URL: http://dx.doi.org/10.5281/zenodo.14704401
03B287EEFFE8FF9F3A97867DFBA7FECF.taxon	materials_examined	Material examined: Taiwan: 1 ò, Nantou, Meifeng, 24 ° 03 ' 43.8 " N, 121 ° 9 ' 55.8 " E, 1972 m, 13 - II- 2012, swept from Eriobotrya deflexa (Y. C. Liao) (NCHU, dry); 1 ò, same but Mt. Beidongyen, 24 ° 04 ' 25.8 " N, 121 ° 07 ' 37.2 " E, 2095 m, 25 - I- 2017 (Y. C. Liao & C. Y. Chien) (NCHU). Description: Male. Colouration (Fig. 1 B, G, H). Body and head greenish brown. Antenna yellow with apex of segment 8 and totality segments 9 and 10 black. Thorax yellowish brown. Mesopraescutum with two black longitudinal stripes. Mesoscutum with five black longitudinal stripes, the median one shortest. Pro- and mesotibia yellow. Forewing hyaline, with a dark brown patch at the apex of cell c + sc, and on cells m 2 and cu 1. Hindwing hyaline. Abdomen dark brown dorsally and light yellow ventrally. Structure: Member of the lii species group. Body (Figs. 1 G, H) length 5.3 mm. Vertex 1.4 – 1.5 times as wide as long; genal process (Fig. 2 B) 1.0 times as long as vertex along midline, bearing long hairs. Antenna slender, 1.8 times as long as head width, sparsely covered with long setae (Fig. 2 D); relative length of flagellar segments as 1.0: 0.4: 0.4: 0.4: 0.4: 0.3: 0.2: 0.2. Thorax covered with long setae. Legs (Fig. 2 F) robust; metatibia 0.9 times as long as head width; metacoxa bearing blunt meracanthus. Forewing (Fig. 1 B) oval, 2.6 times as long as wide, 5.2 – 5.6 times as long as head width, widest in the middle, broadly, irregularly rounded apically; veins sparsely covered with long setae; relative length of veins M 1 + 2, M 3 + 4, Cu and Cu 1 b as 5.2: 1.6: 7.4: 1.8. Male proctiger (Fig. 2 H) robust, concave anteriorly, strongly bulging posteriorly; subgenital plate (Fig. 2 H) subglobular; paramere (Fig. 2 K), in lateral view, lamellar and thick, shorter than proctiger, truncate apically, incised anteriorly and subapically; distal segment of aedeagus (Fig. 2 N) slender, weakly inflated and curved in apical two fifths. Measurements in mm (2 ò): Head width 0.83 – 0.89; antennal length 1.48 – 1.65; metatibia length 0.75 – 0.83; male proctiger length 0.43; paramere length 0.85; length of distal aedeagal segment 0.46. Female described by Li (1991); Li (2011). Fifth instar immature: Unknown. Distribution: China (Fujian) (Li 2011), Taiwan (Nantou) (new record). Host plant: Unknown. The single female from China was collected on Vernicia fordii (Hemsl.) Airy-Shaw (Euphorbiaceae) (Li 2011) and one male from Taiwan on Eriobotrya deflexa (Hemsl.) Nakai (Rosaceae). Both are unlikely hosts. Comments: The dark forewing pattern of the female appears slightly more expanded, judging from the original description (Li 2011). We consider these differences intraspecific variation also observed in other Petalolyma species (Mathur 1975; Yang and Li 1984; Li 2011).	en	Liao, Yi-Chang, Burckhardt, Daniel, Yang, Man-Miao (2024): Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li. Zoological Studies 63 (28): 1-12, DOI: 10.6620/ZS.2024.63-28, URL: http://dx.doi.org/10.5281/zenodo.14704401
03B287EEFFE9FF913B2E8373FF61FCEC.taxon	materials_examined	Material examined: Taiwan: Holotype ò, Taichung, Anmashan, 22 - VII- 1982 (C. T. Yang) (NCHU, slides); 1 ñ, Nantou, Chiehitashan, 14 - VI- 1987 (I. C. Hsu) (NCHU, dry); 2 ñ, Nantou, Meifeng, 2150 m, 24 – 26 - VI- 1981 (K. S. Lin & W. S. Tang) (TARI, dry); 1 ñ, same but 15 - VII- 1982 (S. C. Lin & C. N. Lin) (TARI, dry); 1 ò 2 ñ, Hsinchu, Beidelaman, 30 - IV- 2011 (C. T. Tang) (NCHU, dry, glycerol) Description: Male. Yang (1984), Yang et al. (2013). Member of the divisa species group. Female: Female terminalia (Fig. 2 Q) cuneate. Proctiger 0.9 times as long as head width; dorsal margin, in lateral view, almost straight distal to circumanal ring except for apex which is slightly downcurved and subacute; beset with long setae in basal third laterally, very long setae in median third and short setae in apical third. Circumanal ring cruciform in dorsal view, 0.5 times as long as proctiger; consisting of a single row of elongate pores. Subgenital plate irregularly triangular in lateral view, 0.3 times as long as proctiger; ventral margin weakly concave; apex truncate; beset with long hairs. Dorsal valvula cuneate; dorsal margin has small dorsal nick. Ventral valvula straight with about 10 dorsal teeth. Lateral valvula membranous, irregularly rounded apically. Fifth instar immature: Unknown. Distribution: Taiwan (Nantou) (Yang 1984; Yang et al. 2013). Host plant: Unknown. Petalolyma vittata Liao & Burckhardt, sp. nov. (Figs. 1 A, K – N, 2 A, C, E, G, M, J, P, 3) urn: lsid: zoobank. org: act: E 3 E 03 DF 3 - BCC 8 - 47 E 2 - A 117 - 43 BF 9772 BB 4 F Material examined: Holotype ò, Taiwan: Nantou, Mt. Beidongyen, 24 ° 04 '. 33.2 " N, 121 ° 08 ' 04.8 '' E, 1940 m, 25 - I- 2017, Ilex ficoidea (Y. C. Liao & C. Y. Chien) (NCHU, dry). Paratypes: Taiwan: 7 ò, 7 ñ, same data as holotype but (NCHU, NMNS, NHMB, dry, glycerol, slide); 1 ñ, same but 18 - II- 2014, Ilex ficoidea (Y. C. Liao) (NCHU, dry); 6 ò, 3 ñ, 4 immatures, same but 19 - I- 2017, Ilex ficoidea (Y. C. Liao) (NCHU, 70 % ethanol); 6 ò 3 ñ, 5 immatures, same but 17 - II- 2017, Ilex ficoidea (Y. C. Liao) (NCHU, 70 % ethanol). Material not included in type series: Taiwan: adult (NCHU, 70 % ethanol), New Taipei, Yulu (Fish Road) Historical Trail, 27 - I- 2005, Ilex ficoidea (M. M. Yang, M. F. Lou & C. C. Shen) GA 050127 - 4; empty gall, Taichung, Mt. Daxue 20 K, 8 - IX- 2008, Ilex ficoidea (C. T. Tang) GA 080908 - 4; early instar immatures extracted from galls, Hsinchu, Rubi, 24 ° 39 ' 59.4 " N, 121 ° 16 ' 46.1 " E, 1420 m, 12 - VIII- 2011 (Y. C. Liao) (NCHU, 70 % ethanol); same but Hsinchu, Beidelaman, 24 ° 44 ' 15.7 " N, 121 ° 16 ' 58.8 " E, 1160 m, (Y. C. Liao) (NCHU, 70 % ethanol); same but Taichung, Mt. Daxue: 32 K, 24 ° 13 ' 34.0 " N, 120 ° 58 ' 20.9 " E, 1970 m, 3 - V- 2012 (Y. C. Liao) (NCHU, 70 % ethanol); same but 14 - X- 2018, (Y. C. Liao) (NCHU, 70 % ethanol). Description: Adult: Colouration (Figs. 1 A, K – N, 3 A). Body and head greenish brown. Antenna light yellow with apical two segments black. Thorax yellowish brown. Mesopraescutum with two black markings. Mesoscutum with four black longitudinal stripes. Pro- and mesotibia yellow. Forewing hyaline, with a dark brown patch at the apex of cell c + sc and a broad dark brown band along anal margin stretching from anal break to apex. Hindwing hyaline. Abdomen brown. Structure: A member of the lii species group. Body (Fig. 1 K – N) length 6.0 – 7.0 mm. Vertex 1.5 – 1.6 times as wide as long; genal process (Fig. 2 A) 1.0 – 1.1 times as long as vertex along midline, bearing long hairs. Antenna slender, 1.6 times as long as head width, sparsely covered with long setae; relative length of flagellar segment as 1.0: 0.5: 0.5: 0.5: 0.4: 0.4: 0.2: 0.2. Thorax covered with long setae. Metatibia 0.8 – 1.0 times as long as head width; metacoxa with thumblike meracanthus (Fig. 2 E). Forewing (Fig. 1 A) oval, 2.6 – 2.7 times as long as wide, 5.2 – 6.0 times as long as head width, widest in the middle, subacute subapically; veins sparsely beset with long setae; relative length of veins M 1 + 2, M 3 + 4, Cu and Cu 2 as 6.0: 2.0: 7.6: 1.5. Male proctiger (Fig. 2 G) 0.4 times as long as head width, thick, in lateral view, weakly sinuate anteriorly, strongly produced posteriorly; subgenital plate (Fig. 2 G) subglobular; paramere longer than proctiger, narrowly lamellar, truncate apically, almost straight anteriorly, incised subapically, truncate apically, slightly wavy posteriorly; distal segment of aedeagus (Fig. 2 M) straight basally, weakly dilated and curved in apical third. Female proctiger (Fig. 2 P) 0.9 – 1.0 times as long as head width, in lateral view, longer than subgenital plate, strongly narrowing and curved downward at apical quarter; rhomboidal in dorsal view; anus cruciform in dorsal view, almost half as long as proctiger; subgenital plate cuneate, acute at apex. Measurements in mm (7 ò, 7 ñ): Head width 0.90 – 1.05; antennal length 1.40 – 1.65; metatibia length 0.80 – 0.93; forewing length 5.13 – 6.19; paramere length 0.43; length of distal aedeagal segment 0.43 – 0.45; female proctiger length 0.85 – 0.93; female subgenital plate length 0.50 – 0.53; length of female circumanal ring 0.38. Fifth instar immature (Fig. 3 B – G): Colouration. Head and body pale green. Antenna and wing pads brown. Structure: Body 1.5 times as long as wide. Antenna 0.7 times as long as forewing pad; relative length of antennal segments 3 – 8 as 1.0: 0.8: 0.7: 0.8: 0.7: 1.8, segments 3 – 7 each bear a long, simple dorsal and ventral seta, segment 3 bears two dorsal and one ventral long simple setae. Small and very slender, apically blunt sectasetae on forewing pad covering a relatively broad band along lateral margin. Tibiotarsus longer than femur. Abdominal margin bearing two short sclerotised spurs apically. Measurements in mm (3 immatures): Body length 2.98 – 3.23; body width 2.05 – 2.20; antenna length 0.98 – 1.05; metatibiotarsus length 0.65 – 0.70; forewing pad length 1.55 – 1.58; caudal plate length 0.98 – 1.08; caudal plate width 1.30 – 1.43; circumanal ring width 0.23 – 0.28. Etymology: From Latin vittatus = wearing or carrying a band or ribbon, referring to the dark brown stripe on the forewing. Distribution: Taiwan (Nantou, Taichung, Hsinchu). Host plant: Ilex ficoidea Hemsl. (Aquifoliaceae), a common species in forests at mid-elevation in Taiwan. Biological notes: This species is univoltine and lives in mountain habitats from 1500 to 2000 m. Adults emerge from the end of January to mid-February. The immatures induce a marginal fold producing a closed gall on the leaves. Each gall contains a single immature. When the final instar is ready to hatch, the gall unrolls and the immature leaves the gall and emerges. Comments: Petalolyma vittata Liao & Burckhardt, sp. nov. differs from other congeners in the combination of a dark brown band on the forewing, the pointed forewing and the small forewing cells m 1 and cu 1. It can be distinguished from other Taiwanese species according to the key.	en	Liao, Yi-Chang, Burckhardt, Daniel, Yang, Man-Miao (2024): Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li. Zoological Studies 63 (28): 1-12, DOI: 10.6620/ZS.2024.63-28, URL: http://dx.doi.org/10.5281/zenodo.14704401
