Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li

© 2024 Academia Sinica, Taiwan Open Access Jumping Plant Lice of the Genus Petalolyma Scott (Hemiptera, Triozidae) Inducing Galls on Ilex Species (Aquifoliaceae) in Taiwan, with Comments on Torulus Li Yi-Chang Liao1,2,3,*, Daniel Burckhardt4, and Man-Miao Yang3,* 1Department of Entomology, University of California, Riverside, USA. *Correspondence: E-mail: [email protected] (Liao) 2Back to the Wild Insect Ecosurvey CO., LTD., Taichung, Taiwan 3Department of Entomology, National Chung Hsing University, 145 Xingda Rd., Taichung 402, Taiwan. *Correspondence: E-mail: mmy[email protected] (Yang) 4Naturhistorisches Museum, Augustinergasse 2, 4001 Basel, Switzerland. E-mail: [email protected] (Burckhardt) urn:lsid.org:pub:7F0CEA63-F266-4FE7-B323-E84F7C52545E Received 16 December 2023 / Accepted 26 June 2024 / Published 19 December 2024 Communicated by Shen-Horn Yen The triozid genus Petalolyma Scott, 1882 is reviewed, and the monotypic genus Torulus Li, 1991, syn. nov., is considered a junior subjective synonym. Torulus sinicus is transferred to Petalolyma as P. sinica (Li, 1991), comb. nov., producing a secondary homonym of P. sinica Yang & Li, 1984, for which the replacement name Petalolyma lii Liao & Burckhardt, nomen nov. is proposed here. Adults and fifth instar immatures of Petalolyma vittata Liao & Burckhardt, sp. nov. are described and illustrated. The new species induces marginal fold galls on leaves of Ilex ficoidea (Aquifoliaceae). The previously unknown male and female of P. lii and P. nigra, respectively, are described and illustrated, and Petalolyma castanopsis Li & Yang, 1991 and P. fujianensis Li & Yang, 1999, syn. nov., are synonymised. Petalolyma hyalina (Kuwayama, 1910) is considered a nomen dubium as the original description is not diagnostic and the type material cannot be found. Hence, Petalolyma currently includes 13 valid species whose host plant and biogeographic patterns are briefly discussed. Identification keys for the adults and fifth instar immatures are provided for the Taiwanese species. Key words: Sternorrhyncha, Psylloidea, New species, New synonymies, Taxonomy, Marginal fold gall BACKGROUND Jumping plant lice or psyllids (Psylloidea) are characterised within Sternorrhyncha by their generally narrow host ranges, both at species and higher taxonomic levels (Hollis 2004; Hodkinson 2009; Burckhardt et al. 2014; Ouvrard et al. 2015). Some psyllids are known to induce galls on their host plants (Hodkinson 1984; Burckhardt 2005; Yang and Raman 2007). Gall inducers are particularly abundant in the Calophyinae, Phacopteroninae and Triozidae (Burckhardt 2005; Malenovský et al. 2007; Yang et al. 2013), and they seem more frequent in the tropics than in temperate regions. According to Docters van Leeuwen and Docters van Leeuwen-Reijnvaan (1914), in Java (Indonesia) about 5% of all galls are induced by Psylloidea. In Taiwan, more than half of the psyllid species are gallicolous with the highest numbers known to develop on Lauraceae and Myrtaceae (Yang et al. 2006). The number of gall-inducing compared to freeliving psyllid species is probably underestimated as the former are often univoltine, and the period with adult presence is comparatively short (Yang et al. 2006; Carneiro et al. 2013). The effort needed to find adults Citation: Liao YC, Burckhardt D, Yang MM. 2024. Jumping plant lice of the genus Petalolyma Scott (Hemiptera, Triozidae) inducing galls on Ilex species (Aquifoliaceae) in Taiwan, with comments on Torulus Li. Zool Stud 63:28. doi:10.6620/ZS.2024.63-28. Zoological Studies 63:28 (2024) doi:10.6620/ZS.2024.63-28 1 © 2024 Academia Sinica, Taiwan of some species and their presence on non-hosts, due to their capacity to fly, makes the association of adults with the respective galls difficult. A good example of the patchy knowledge on gallinducers are the psyllids associated with Ilex species (Aquifoliaceae). Ilex, comprising 560 described species worldwide, has been independently colonised by psyllids several times. Two genera with gall inducing species are associated with Ilex: 1. Gyropsylla Brèthes (Aphalaridae, Aphalarinae) with five described species in the Americas, two of which lack host data, and one species in Asia also lacking host data; and 2. Petalolyma Scott (Triozidae) with 13 described species in the Oriental region, of which seven species lack host data. In addition to the two genera, one species each of Cecidotrioza Kieffer and Psylla Geoffroy, and six species of Trioza Foerster have been reported from Ilex (Ouvrard 2023) though some of these host records are doubtful and it is unknown whether any of these species induces galls. Petalolyma is morphologically well diagnosed by adult and immature characters (see diagnosis below). Li (1991) erected the monotypic genus Torulus for T. sinicus Li, known from a single female only, collected on Vernicia fordii (Euphorbiaceae) in Fujian, China. It “differs from Petalolyma in R1 longer than R and mesothorax with two groups of long hairs” (Li 1991). Recently, two males of T. sinicus and an undescribed species, morphologically intermediate between Petalolyma and Torulus, were discovered in Taiwan, the latter inducing marginal fold galls on the leaves of Ilex ficoidea. This new material challenges the validity of Petalolyma and Torulus as separate, monophyletic units, necessitating a review of the two genera. Here we formally describe the new species, including the gall, and provide information of the previously unknown male and female of T. sinicus and P. nigra, respectively. We examine the phylogenetic relationships between Petalolyma and Torulus, review the Taiwanese species, for which we provide identification keys for adults and immatures, and discuss some other Oriental congeners. MATERIALS AND METHODS Specimens were collected by sweeping or by direct search. Galls were bagged in the field and checked monthly for emerged adults. Material is dry mounted or preserved in 70% or 99% ethanol. Some specimens were cleared in 15% potassium hydroxide and orange oil for subsequent microscopical examination in glycerol. Specimens are deposited in or reported from the National Chung Hsing University, Taichung, Taiwan (NCHU), the National Museum of Natural Science, Taichung, Taiwan (NMNS), the Naturhistorisches Museum Basel, Switzerland (NHMB), the Systematic Entomology, Hokkaido University, Japan (SEHU) and the Taiwan Agricultural Research Institute, Ministry of Agriculture, Taichung City, Taiwan (TARI). Photographs of morphological characters were taken with a Leica DM 750 microscope equipped with a digital Canon EOS 600D camera, except for those of the wings, which were taken with a Leica MZ 125 microscope equipped with a digital Olympus EP-1 camera. The photographs were montaged using focus stacking software (Helicon Focus, Helicon Soft). The morphological terminology follows Bastin et al. (2023) and the nomenclature of plants POWO (2023). TAXONOMY Family Triozidae Löw, 1879 Genus Petalolyma Scott, 1882 Petalolyma Scott, 1882: 459. Type species: Psylla basalis Walker, 1858, by original designation and monotypy. 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 anteromedially; 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 page 2 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan 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. Petalolyma includes in the new concept 14 previously described nominal and one new species, described here. Petalolyma hyalina (Kuwayama) is known only from the original description (Kuwayama 1910), and the type material cannot be traced (H. Inoue, pers. comm.; see also below). Based on the forewing shape and venation, the remaining species can be assigned to six informal, not necessarily monophyletic, species groups (Table 1). While species identification keys with illustrated descriptions are available for the Chinese and Japanese species (Miyatake and Matsumoto 2008; Li 2011), and Mathur (1975) provided an adequate description for the Indian P. basalis, available information on the Taiwanese species is quite incomplete (Yang 1984; Fang and Yang 1986; Yang et al. 2013). Here, identification keys are provided and the taxonomy of the Taiwanese species is reviewed. Keys to Petalolyma species from Taiwan Adults (not including Petalolyma hyalina) 1. Forewing (Fig. 1A, B) with veins Rs weakly and M strongly curved, the two veins thus strongly diverging to wing apex; cell m1 and cu1 small: length ratio M/M1+2 > 3.0, Cu/Cu1b > 4.0 ....... 2 - Forewing (Fig. 1C–E) with veins Rs hardly or not, and M weakly curved, the two veins thus not strongly diverging to wing apex; cell m1 and cu1 large: length ratio M/M1+2 < 2.0, Cu/Cu1b < 2.0 .... ..................................................................................................... 3 2. Forewing (Fig. 1B) with a dark brown patch along anal margin covering cells m1 and cu1, respectively; broadly, irregularly rounded apically. Paramere (Fig. 2H) shorter than male proctiger ............................................... P. lii Liao & Burckhardt, nom. nov. - Forewing (Fig. 1A) with wide black band along anal margin distal of anal break; indistinctly angular apically. Paramere (Fig. 2G) longer than male proctiger. On Ilex ficoidea .......................... ........................................... P. vittata Liao & Burckhardt, sp. nov. page 3 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan 3. Forewing (Fig. 1C, D) rounded apically; wing apex in the middle of cell r2 margin. Dorsal outline of mesopraescutum, in lateral view, strongly curved (Fig. 1J). Paramere (Fig. 2I, L), in lateral view, irregularly lanceolate with forward pointing apical hook. Female proctiger dorsally distinctly curved, blunt apically .......... ......................................................................... P. formosana Yang - Forewing (Fig. 1E) pointed apically; wing apex at distal end of vein M1+2. Dorsal outline of mesopraescutum, in lateral view, weakly curved. Paramere, in lateral view, lamellar with forward pointing apical hook. Female proctiger relatively straight dorsally, subacute apically .......................................................... 4 4. Vein M about 1.2 times as long as M1+2. Paramere blunt apically. On Ilex asprella .................................... P. castanopsis Li & Yang - Vein M1+2 longer than M. Paramere obliquely truncate apically ... .................................................................................. P. nigra Yang Fifth instar immature (not including Petalolyma formosana, P. hyalina, P. lii, P. nigra) 1. On Ilex asprella .................................... P. castanopsis Li & Yang - On Ilex ficoidea ................ P. vittata Liao & Burckhardt, sp. nov. Petalolyma castanopsis Li & Yang, 1991 Petalolyma castanopsis Li & Yang, 1991: 16. Petalolyma fujianensis Li & Yang, 1999: 480; syn. nov. Trichochermes bicolor sensu Fang & Yang, 1986: 157, nec Kuwayama, 1910: 54; Yang & Li 1984: 129, 130. Material examined: Taiwan: 6 ♀, Nantou, Jihyeutan (Sun Moon Lake), 10-III-1985 (S. J. Fang) (NCHU, dry mounted); 4 ♂, 3 ♀, Pingtung, Shizi, Neiwen Village, South Link Highway, N22°14'36.327'', E120°49'56.988'', 480 m, 1-XII-2023 (D. Burckhardt) #23-33(-) (NHMB, 70% ethanol); 1 ♂, 2 ♀, same but Neiwen Village, Dongyuan Road, N22°14'36.575'', E120°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 Table 1. Forewing characters defining species groups of Petalolyma. M/M1+2 = length ratio of veins M and M1+2; Cu/ Cu1b = length ratio of veins Cu and Cu1b; m1/r2 = width ratio of cells m1 and r2 measured at wing margin Species Wing apex M/M1+2 Cu/Cu1b m1/r2References basalis species group basalis (Walker, 1858) rounded, middle of cell margin r2< 2.0 1.4–2.9 < 2.0 Mathur (1975) sinica Yang & Li, 1984 rounded, middle of cell margin r2< 2.0 1.4–2.9 < 2.0 Li (2011) bicolor species group bicolor (Kuwayama, 1910) angular, near M1+2 apex < 2.0 3.0–4.0 < 2.0 Miyatake and Matsumoto (2008) zhejiangana Yang & Li, 1984 angular, near M1+2 apex < 2.0 3.0–4.0 < 2.0 Li (2011) divisa species group castanopsis Li & Yang, 1991 = fujianensis Li & Yang, 1999, syn. nov. angular, near M1+2 apex < 2.0 1.4–2.9 < 2.0 Yang (1984); Li (2011); Yang et al. (2013) divisa (Crawford, 1917) angular, near M1+2 apex < 2.0 1.4–2.9 < 2.0 Miyatake and Matsumoto (2008) nigra Yang, 1984 angular, near M1+2 apex < 2.0 1.4–2.9 < 2.0 Yang (1984); Li (2011); Yang et al. (2013) formosana species group indent Yang, 1984 rounded, middle of cell margin r2< 1.3 1.4–2.9 < 2.0 Yang (1984); Li (2011); Yang et al. (2013) lii species group lii Liao & Burckhardt rounded, middle of cell margin r2> 3.0 > 4.1 < 2.0 present paper vittata Liao & Burckhardt angular, near M1+2 apex > 3.0 > 4.1 < 2.0 present paper yunnanana species group shibatai Miyatake & Matsumoto, 2008 rounded, middle of cell margin r2< 2.0 1.4–2.9 > 2.0 Miyatake and Matsumoto (2008) variegata Li & Yang, 1999 rounded, middle of cell margin r2< 2.0 < 1.3 > 2.0 Li (2011) yunnanana Yang & Li, 1984 rounded, middle of cell margin r2< 2.0 1.4–2.9 > 2.0 Li (2011) nomen dubium hyalina (Kuwayama, 1910) page 4 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan Fig. 1. Forewing (A–E) and habitus (F–N) of Petalolyma spp. A, K–N. P. vittata Liao and Burckhardt, sp. nov. B, G, H. P. lii Liao and Burckhardt, nomen nov. C, D, I, J. P. formosana. E, F. P. nigra. Scale bar = 1 mm. page 5 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan to Li’s (2011) key, the species differ in the length ratio of veins Cu1a/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. Fig. 2. Head (A, B), antennae (C, D), hindlegs (E, F), male terminalia (G–I), male paramere (J–L), distal segment of aedeagus (M–O), female terminalia (P, Q) of Petalolyma spp. A, C, E, G, M, J, P. P. vittata Liao and Burckhardt, sp. nov. B, D, F, H, N, K. P. lii Liao and Burckhardt, nomen nov. I, L, O. P. formosana. Q. P. nigra. Scale bar = 0.1 mm. page 6 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan Petalolyma formosana Yang, 1984 (Figs. 1C–D, I–J, 2I, L, O) Petalolyma formosana Yang, 1984: 234. Material examined: Taiwan: Holotype ♂, paratypes 9 ♂, 9 ♀, Taichung, Anmashan, 22-VII-1982 (C. T. Yang) (NCHU, dry); 1 ♂ 1 ♀, same but 6-VIII1987 (C. T. Yang) (NCHU, dry); 2 ♂, Nantou, Meifeng, 2150 m, 2–4-VI-1980 (L. Y. Chou & C. C. Chen) (TARI, dry); 1 ♂ 1 ♀, Nantou, Tsuifeng, 2300 m, 3-VI-1980 (L. Y. Chou & C. C. Chen) (TARI, dry); 1 ♂, 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. 1C) or two patches along the costal margin (Fig. 1D). Fifth instar immature. Unknown. Member of the formosana species group. Distribution: Taiwan (Nantou, Taichung) (Yang 1984; Yang et al. 2013). Host plant: Unknown. Petalolyma hyalina (Kuwayama, 1910), nomen dubium Trichochermes hyalina Kuwayama, 1910: 55. Petalolyma hyalina, Yang and Li (1984): 130, 137. 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. 1B, G, H, 2B, D, F, H, N, K) urn:lsid:zoobank.org:act:5F01DB62-1BFF-4D09-9ECDA16730C90B74 Torulus sinicus Li, 1991: 37, 41. Petalolyma sinica (Li, 1991), comb. nov. = junior secondary homonym of Petalolyma sinica Yang & Li, 1984: 131, 138. Material examined: Taiwan: 1 ♂, Nantou, Meifeng, 24°03'43.8"N, 121°9'55.8"E, 1972 m, 13II-2012, swept from Eriobotrya deflexa (Y. C. Liao) (NCHU, dry); 1 ♂, 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. 1B, 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. Proand mesotibia yellow. Forewing hyaline, with a dark brown patch at the apex of cell c+sc, and on cells m2 and cu1. Hindwing hyaline. Abdomen dark brown dorsally and light yellow ventrally. Structure: Member of the lii species group. Body (Figs. 1G, H) length 5.3 mm. Vertex 1.4–1.5 times as wide as long; genal process (Fig. 2B) 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. 2D); 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. 2F) robust; metatibia 0.9 times as long as head width; metacoxa bearing blunt meracanthus. Forewing (Fig. 1B) 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 M1+2, M3+4, Cu and Cu1b as 5.2: 1.6: 7.4: 1.8. Male proctiger (Fig. 2H) robust, concave anteriorly, strongly bulging posteriorly; subgenital plate (Fig. 2H) subglobular; paramere (Fig. 2K), in lateral view, lamellar and thick, shorter than proctiger, truncate apically, incised anteriorly and subapically; distal segment of aedeagus (Fig. 2N) slender, weakly inflated and curved in apical two fifths. Measurements in mm (2 ♂): 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 page 7 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan Petalolyma species (Mathur 1975; Yang and Li 1984; Li 2011). Petalolyma nigra Yang, 1984 (Figs. 1E, F, 2Q) Petalolyma nigra Yang, 1984: 237. Material examined: Taiwan: Holotype ♂, Taichung, Anmashan, 22-VII-1982 (C. T. Yang) (NCHU, slides); 1 ♀, Nantou, Chiehitashan, 14-VI-1987 (I. C. Hsu) (NCHU, dry); 2 ♀, Nantou, Meifeng, 2150 m, 24–26-VI-1981 (K. S. Lin & W. S. Tang) (TARI, dry); 1 ♀, same but 15-VII-1982 (S. C. Lin & C. N. Lin) (TARI, dry); 1 ♂ 2 ♀, Hsinchu, Beidelaman, 30-IV2011 (C. T. Tang) (NCHU, dry, glycerol) Description: Male. Yang (1984), Yang et al. (2013). Member of the divisa species group. Female: Female terminalia (Fig. 2Q) 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. 1A, K–N, 2A, C, E, G, M, J, P, 3) urn:lsid:zoobank.org:act:E3E03DF3-BCC8-47E2-A11743BF9772BB4F Material examined: Holotype ♂, 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 ♂, 7 ♀, same data as holotype but (NCHU, NMNS, NHMB, dry, glycerol, slide); 1 ♀, same but 18-II-2014, Ilex ficoidea (Y. C. Liao) (NCHU, dry); 6 ♂, 3 ♀, 4 immatures, same but 19-I-2017, Ilex ficoidea (Y. C. Liao) (NCHU, 70% ethanol); 6 ♂ 3 ♀, 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) GA050127-4; empty gall, Taichung, Mt. Daxue 20K, 8-IX-2008, Ilex ficoidea (C. T. Tang) GA080908-4; 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-X2018, (Y. C. Liao) (NCHU, 70% ethanol). Description: Adult: Colouration (Figs. 1A, K– N, 3A). 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. Proand 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. 1K–N) length 6.0–7.0 mm. Vertex 1.5–1.6 times as wide as long; genal process (Fig. 2A) 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. 2E). Forewing (Fig. 1A) 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 M1+2, M3+4, Cu and Cu2 as 6.0: 2.0: 7.6: 1.5. Male proctiger (Fig. 2G) 0.4 times as long as head width, thick, in lateral view, weakly sinuate anteriorly, strongly produced posteriorly; subgenital plate (Fig. 2G) subglobular; paramere longer than proctiger, narrowly lamellar, truncate apically, almost straight anteriorly, incised subapically, truncate apically, slightly wavy posteriorly; distal segment of aedeagus (Fig. 2M) straight basally, weakly dilated and curved in apical third. Female proctiger (Fig. 2P) 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 ♂, 7♀): Head width 0.90–1.05; antennal length 1.40–1.65; metatibia length page 8 of 12Zoological Studies 63:28 (2024) © 2024 Academia Sinica, Taiwan Fig. 3. Habitus and immature of P. vittata Liao and Burckhardt, sp. nov. A, male. B, last instar immature in a gall. C, plant galls. D, immature. E, arolium. F, circumanal ring. G, sectasetae on forewing pad. Scale bar = 0.1 mm. page 9 of 12 Zoological Studies 63:28 (2024)