New crane fly species of the subgenus Tipula (Vestiplex) Bezzi (Diptera, Tipulidae) from Qinghai-Tibet Plateau, China

183 New crane fly species of the subgenus Tipula (Vestiplex) Bezzi (Diptera, Tipulidae) from Qinghai-Tibet Plateau, China Pavel Starkevich1, Qiu-Lei Men2, Aidas Saldaitis1, Kristina Valavičiūtė-Pocienė1 1 Nature Research Centre, Akademijos 2, LT-08412 Vilnius, Lithuania 2 School of Life Sciences, Provincial Key Laboratory of the Biodiversity Study and Ecology Conservation in Southwest Anhui Province, Anqing Normal University, Anqing, Anhui 246011, China Corresponding authors: Pavel Starkevich ([email protected]); Qiu-Lei Men ([email protected]) Copyright: © Pavel Starkevich et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Four new crane fly species of the subgenus Tipula (Vestiplex) Bezzi, 1924, T. (V.) apteroides sp. nov. (Yunnan), T. (V.) bidentata sp. nov. (Sichuan), T. (V.) spinaventralis sp. nov. (Sichuan) and T. (V.) ventrobasilata sp. nov. (Sichuan) are described and illustrated, providing discussion on separation of the newly described species from other closely related species. The male genitalia of T. (V.) aestiva Savchenko, 1960 is illustrated and redescribed. Key words: Distribution, hypopygium, new species, ovipositor, taxonomy, Tipulinae Introduction The current paper presents a report of descriptions of four new T. (Vestiplex) species from Qinghai-Tibet Plateau (QTP), a region characterized by high Tipulidae diversity (Yang et al. 2022), including the subgenus which is well represented in QTP. Recent studies revealed 26 new T. (Vestiplex) species from the Eastern Palaearctic and Oriental regions (Men et al. 2017, 2021a, 2021b, 2021c, 2023; Starkevich et al. 2019a, 2019b; Ren et al. 2021; Yang et al. 2021; Starkevich and Young 2023). There are 196 recognized species of T. (Vestiplex) distributed throughout the Holarctic and Oriental regions with 104 species recorded in China (Oosterbroek 2024). The subgeneric characteristics are summarized in Alexander (1934, 1935), Savchenko (1960, 1964) and recent works dealing with the revision of T. (Vestiplex) fauna in Mongolia (Starkevich et al. 2020), Taiwan (Starkevich and Young 2023) and the Korean Peninsula (Starkevich et al. 2021). Material and methods Adult crane flies were collected by net or at ultraviolet light. The hypopygium was removed and boiled in 10% NaOH for 5–10 min, and observed in glycerin under an Olympus SZX10 stereomicroscope (Olympus, Japan). Photographs of adult specimens were taken with a Canon EOS 6D Academic editor: Xiao Zhang Received: 19 October 2024 Accepted: 10 March 2025 Published: 19 December 2025 ZooBank: https://zoobank.org/ ACE99CFC-5ECA-4F70-B181942B71F2E7C2 Citation: Starkevich P, Men Q-L, Saldaitis A, Valavičiūtė-Pocienė K (2025) New crane fly species of the subgenus Tipula (Vestiplex) Bezzi (Diptera, Tipulidae) from QinghaiTibet Plateau, China. ZooKeys 1264: 183–206. https://doi.org/10.3897/ zookeys.1264.139740 ZooKeys 1264: 183–206 (2025) DOI: 10.3897/zookeys.1264.139740 184 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China camera and Canon MP-E65 mm macro lens using a MJKZZ automated focus stacking rail set (MJKZZ.de, Vienna, Austria). Pictures of dissected terminalia were taken with: a) an Infinity1 camera mounted on a Nikon Si-L stereomicroscope; b) a Canon EOS 80D camera mounted on an Olympus SZX10 dissecting microscope; c) an EvolutionTMMP camera mounted on an Olympus SZX10 dissecting microscope; d) a Sony SSC-DC 50AP camera mounted on an Olympus BX 40 dissecting microscope; e) Canon EOS 6D camera and Mitutoyo 10× objective using MJKZZ automated focus stacking rail set. The obtained image layers were stacked using the program ZereneStacker v. 1.04 and edited with Photoshop v. 26.5.0. Often particular genital structures have complicated three-dimensional forms. While these structures have a taxonomic and phylogenetic importance, several photos are provided for the same sclerite to uncover shape which differs depending on the angle, e.g., the inner gonostylus of T. (V.) apteroides sp. nov. or epandrium of T. (V.) spinaventralis sp. nov. Descriptive terminology of the adults generally follows that of Cumming and Wood (2017) and de Jong (2017) for wing venation with some additions for particular features of T. (Vestiplex). The term appendage of sternite 9 (A9s) is adopted from Mannheims (1963); the terms ventral lobe and dorsal lobe of appendage of sternite 9 are adopted from Gelhaus (2005); and the term genital bridge is adopted from Dobrotworsky (1968) which is equal to sclerites sp1 and sp2 (Neumann 1958). Abbreviations for institutional collections used herein: ANSP the Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania, USA; NHMUK Natural History Museum, London, United Kingdom; NRC Nature Research Centre, Vilnius, Lithuania; SMOC Silesian Museum, Opava, Czech Republic; USNM United States National Museum of Natural History, Washington, D.C., USA; ZIN Zoological Museum of the Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia. Results Taxonomy Class Insecta Linnaeus, 1758 Order Diptera Linnaeus, 1758 Family Tipulidae Latreille, 1802 Subfamily Tipulinae Latreille, 1802 Genus Tipula Linnaeus, 1758 Subgenus Vestiplex Bezzi, 1924 185 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Tipula (Vestiplex) apteroides Starkevich, Saldaitis & Men, sp. nov. https://zoobank.org/FCADCA7F-AECD-480E-91D7-6BFAD20F747F Figs 1–3 Type material. Holotype: China • ♂ (pinned); Yunnan; Xue Shan nr. Zhongdian; 27°49N, 99°34E; alt. 4000–4100 m; 23 June 1996; Farkač, P. Kábatek and A. Smetana leg.; Mus. Silesiae Opava; Inv.č.: d 005 7-2004; SMOC. Paratypes: China • 2 ♂ (pinned); same data as holotype; SMOC. Comparative material examined. Tipula (Vestiplex) aptera Savchenko, 1955: China • Lectotype ♂; Qinghai; Sanka village, Den-chu River, Kam, Yangtze Basin; 17.VI.1901; Kozlov leg.; ZIN • Paralectotype 2♂; same data as holotype; ZIN. Also, material listed in Starkevich et al. (2019a). Diagnosis. Tipula (V.) apteroides sp. nov. can be easily recognized by dark brown body coloration and greatly reduced wings. Male tergite 9 with lateral corners produced into toothed posterolateral lobes, dorsal surface with dorsomedian anteriorly extended sclerotised area, laterally terminating in obtuse teeth. Dorsomedian area provided with additional median elevated ridge. Inner gonostylus with apical margin terminating in three teeth. Description. Male. Body length 12.0–12.3 mm, wing length 0.8–0.9 mm (N = 3). General body coloration dark brown (Fig. 1). Head. Dark brown, vertex and occiput dusted with grey brown and provided with median dark brown line. Rostrum dark brown with short nasus. Palpus blackish brown. Scape and pedicel dark brown, flagellum broken in all three specimens. Thorax (Fig. 1B). Overall brown, dusted by light greyish brown, medially with dark line. Prescutum and presutural scutum heavily dusted by light brown, with central and lateral stripes ill-defined, recognizable by darker borders. Postsutural scutum brown, dusted with light greyish brown and dark median spot; scutellum dusted with light greyish with median line present in two specimens; mediotergite brown, dusted with light greyish brown with dark median line or spot. Pleura brown, heavily dusted by yellowish grey. Legs with coxae brown, heavily dusted Figure 1. Habitus of Tipula (Vestiplex) apteroides sp. nov. A male, lateral view, holotype B male thorax, dorsal view, paratype. Scale bars: 5 mm (A); 1 mm (B). 186 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Figure 2. Male terminalia of Tipula (Vestiplex) apteroides sp. nov., paratype A hypopygium, dorsal view B hypopygium, lateral view. Abbreviations: gcx, gonocoxite; ig, inner gonostylus; og, outer gonostylus; pll, posterolateral lobe; t9, tergite 9. Scale bar: 0.5 mm. by yellowish grey. Trochanters and femora dark reddish brown, femora slightly darkened at tips; tibiae dark reddish brown with tips darkened; tarsal segments dark brown. Claw without tooth. Wing greatly reduced. Halter dark brown. Abdomen. Abdominal tergites 1–5 dorsally dark reddish brown with darker median stripe. Lateral parts of abdominal tergites brown, with margins broadly pale. Remaining tergites brown. Sternites 1–6 dark reddish brown, remaining dark brown. Hypopygium (Figs 2–4). Tergite 9 in the shape of shallowly emarginated, saucer-shaped plate (Figs 2, 3A, B). Posterior margin blackened and microscopically roughened, broadly emarginated, medially with narrow incision. Posterolateral margins rounded. Lateral corners produced into toothed posterolateral lobes (Fig. 2B). Tergal surface with dorsomedian sclerotised area, extended anteriorly, its lateral margins produced into obtuse teeth. Dorsomedian area provided with additional median elevated ridge, anteriorly and posteriorly terminating in obtuse denticles, so that three caudally pointed teeth are visible in lateral view (Figs 2B, 3B). Gonocoxite separated from sternite 9 by suture, apically with rounded lobe (Fig. 3C). Outer gonostylus nearly finger-shaped, slightly curved at base (Fig. 3D). Inner gonostylus in the shape of strongly curved plate (Fig. 3E–H). Beak extended and blackened, lower beak broadly flattened with blackened margin. Apical margin of inner gonostylus terminating in three teeth. Dorsal margin terminating in blackened dorsoapical and apical median teeth; median surface apically with dorsomedian ridge which terminates in broad tooth. Genital bridge with sclerite sp2 narrow, fused with both margins of gonocoxite (Fig. 3C). Sclerites sp1 fused into flattened base, distal part narrowed with preapical triangular lobe (Fig. 3J). Adminiculum in the shape of a flattened plate, posterolaterally produced into rounded projection; anterior parts extended (Fig. 3L). Sternite 9 with ventral lobe of A9s narrowing at tip (Fig. 3I). Sperm pump with central vesicle spherical (Fig. 4A). Compressor apodeme flattened, with shallow median incision (Fig. 4B). Posterior immovable apodeme 187 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China relatively short, shorter than compressor apodeme; anterior immovable apodeme narrow. Aedeagus about 1.8 × as long as sperm pump, brownish yellow (Fig. 4A). Female. Unknown. Etymology. The specific epithet is based on the name T. (V.) aptera, a closely related species characterized by a reduced wing. Distribution. China, Yunnan. Remarks. Tipula (V.) apteroides sp. nov. belongs to the erectiloba species group which was proposed by Savchenko (1960), with the following discusFigure 3. Male terminalia of Tipula (Vestiplex) apteroides sp. nov., paratype A tergite 9, dorsal view B tergite 9, anterior view C left gonocoxite and sp2 of genital bridge, lateral view D left outer gonostylus, lateral view E left inner gonostylus, outer lateral view with dorsal part partly visible F left inner gonostylus, mesal view G left inner gonostylus, dorsal view from apex H left inner gonostylus, outer view I adminiculum, ventral view J genital bridge, sclerites sp1, dorsal view. Abbreviations: amt, apical median tooth; b, beak; dat, dorsoapical tooth; dmr, distal median ridge; lb, lower beak; mr, median ridge of tergite 9; sp2, sclerite sp2 of genital bridge; vl, ventral lobe of A9s. Scale bars: 0.5 mm a (A–D, F–J); 0.5 mm b (E). 188 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China sion in Starkevich et al. (2019a). The new species is similar to T. (V.) aptera (China, Qinghai) in having greatly reduced wings and dark brown body coloration. Still the new species can be easily separated by tergite 9 posterolaterally having toothed lobes and median elevated ridge which are absent in T. (V.) aptera (Savchenko 1955: fig. 1). The elevated anterior border of tergite 9 is much broader in T. (V.) apteroides sp. nov. with lateral margins produced into obtuse teeth, while in T. (V.) aptera the border is smaller laterally without teeth. Both species can also be separated by the inner gonostylus, which bears a dorsomedian ridge in T. (V.) apteroides sp. nov., the latter is absent in T. (V.) aptera. Tipula (Vestiplex) bidentata Starkevich, Saldaitis & Men, sp. nov. https://zoobank.org/BF43977D-4FB4-4B1E-BA17-3FC05C55B844 Figs 5–8 Type material. Holotype: China • ♂ (in alcohol); Sichuan, 20 km N. Maoxian; 31°46.310'N, 103°42.898'E; alt. 1820 m; 20 May 2017; A. Saldaitis leg.; NRC; Specimen number NRCE000100; genitalia dissected. Diagnosis. Tipula (V.) bidentata sp. nov. can be recognized by yellow body, elongated antenna reaching middle of abdomen when bent backward and bicolored flagellum. Gonocoxite large, apically bearing longer ventral tooth, dorsal tooth split into two denticles. Description. Male. Body length 11.1 mm, wing length 9.5 mm (N = 1). General body coloration yellow, thorax brownish yellow (Fig. 5A). Head. Brown. Rostrum yellowish brown, nasus small. Palpus brown. Antenna 13-segmented, elongated, if bent backward almost reaching the middle of abdomen. Scape and pedicel brownish yellow, first flagellar segment yellow, segments 3–4 yellow, remaining segments getting yellowish brown to brown. Each flagellomere except the first with darkened basal enlargement, making the bicolor appearance of flagellum; two incisions, the Figure 4. Male terminalia of Tipula (Vestiplex) apteroides sp. nov., paratype A sperm pump and aedeagus, lateral view B sperm pump, dorsal view. Scale bar: 0.5 mm. 189 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China distinct incision at base and weak distal incision. Apical flagellomere small, distinctly shorter than preceding flagellomere. Long verticils about 1/3 as long as corresponding flagellomeres. Thorax. Yellowish brown to brown, thinly dusted with grey. Pronotum brown with darker median spot. Prescutum and presutural scutum with four brown stripes, indistinctly bordered by lighter brown. Central stripes separated by light brown ground. Interspaces gray. Postsutural scutum brown, dusted with grey, each lobe with two brown sports, anterior spot twice smaller than posterior one. Scutellum brownish yellow, mediotergite brown with darker median line, both sclerites dusted with grey. Pleura yellowish brown, dusted with grey. Legs with coxae yellow, dusted with grey. Trochanters yellow. Femora basally yellow, getting brownish yellow toward brown tips. Tibiae brownish yellow, tips brown. Tarsal segments brown. Claw without tooth. Wing pale brown, tinged with brown (Fig. 5B). Vein m-cu joining dm at fork of M3+4, cell M1 about 1.5 as long as its petiole. Halter pale yellow. Abdomen. Abdominal segments 1–6 yellow, remaining brownish yellow, trivittate. Lateral lines narrow, dark brown, dorsal line pale brown, interrupted. Hypopygium (Figs 6–8). Tergite 9 forming a concave saucer-shaped plate (Figs 6, 7A). The main body of tergal plate yellow, posterior margin broadly emarginated with shallow median U-shaped notch; lateral angles of plate nearly angular and blackened; anterior portion of tergal plate raised into narrow sclerotised border. Gonocoxite separated from sternite 9 by suture, in the shape of large nearly, rectangular plate (Figs 6, 7B–D). Apex of gonocoxite terminating in longer ventral tooth, with dorsal tooth split into two denticles. Outer gonostylus nearly finger shaped (Fig. 7E). Inner gonostylus in the shape of slightly curved plate (Figs 6A, 7F–H), beak broadly blackened with tip obtuse, lower beak absent; dorsal margin apically with short extension, the margin of extension rough. Genital bridge with sclerite sp2 reduced (Fig. 7D), sclerites sp1 fused at base, forming V-shaped structure (Fig. 7K). Adminiculum triangular in distal half with gonapophyses weakly developed (Fig. 7I, J). Apex ventrally produced into short denticle. Proctiger pale with narrow anal plates (Fig. 7L). Sperm pump with central vesicle flattened (Fig. 8A). Compressor apodeme with deep V-shaped median incision (Fig. 8B). Posterior immovable apodeme extended, slightly flattened (Fig. 8C). Anterior immovable apodeme rounded. Aedeagus about 4 × as long as sperm pump, blackish brown except for yellow distal part. Apex with sagital and frontal incisions, trident in shape (Fig. 8A, D). Female. Unknown. Etymology. The name of the new species refers to the gonocoxite apically possessing two teeth. Distribution. China, Sichuan. Remarks. Tipula (V.) bidentata sp. nov. is closely related to T. (V.) aestiva Alexander, 1936 (China, Qinghai and Gansu) based on the shape of tergite 9, gonocoxite, inner gonostylus and adminiculum (Fig. 10). The new species can be separated by the gonocoxite which has dorsal tooth split into two denticles; tergite 9 of T. (V.) bidentata sp. nov. is posterolaterally angular, while this part is rounded in T. (V.) aestiva; and the inner gonostylus bearing dorsoapical rough extension which is smooth in T. (V.) aestiva. Both species can also be separated by the sperm pump: in T. (V.) aestiva the posterior apodeme is medially produced into a specific short lobe which is absent in T. (V.) bidentata sp. nov. 190 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Figure 6. Male terminalia of Tipula (Vestiplex) bidentata sp. nov., holotype (specimen number NRCE000100) A hypopygium, dorsal view B hypopygium, lateral view. Abbreviations: dt, dorsal tooth; gcx, gonocoxite; ig, inner gonostylus; og, outer gonostylus; t9, tergite 9; vt, ventral tooth. Scale bar: 0.5 mm. Figure 5. Habitus of Tipula (Vestiplex) bidentata sp. nov., holotype (specimen number NRCE000100) A male, lateral view B male wing. Scale bar: 5 mm (A, B). 191 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Figure 7. Male terminalia of Tipula (Vestiplex) bidentata sp. nov., holotype (specimen number NRCE000100) A tergite 9, dorsal view B right gonocoxite, lateral view C right gonocoxite and sp2 of genital bridge, caudal view D apical part of gonocoxite E right outer gonostylus, lateral view F right inner gonostylus, inner lateral view G right inner gonostylus, outer lateral view H right inner gonostylus, dorsal view I adminiculum, lateral view J adminiculum, ventral view K genital bridge, sclerites sp1, dorsal view L proctiger, dorsal view. Scale bars: 0.5 mm a (A, D, D–L); 0.25 mm b (C). 198 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Figure 14. Habitus of Tipula (Vestiplex) ventrobasilata sp. nov., holotype (specimen number NRCE000101) A male, lateral view B male wing. Scale bars: 5 mm (A, B). Figure 15. Male terminalia of Tipula (Vestiplex) ventrobasilata sp. nov., holotype (specimen number NRCE000101) A hypopygium, dorsal view B hypopygium, lateral view. Abbreviations: gcx, gonocoxite; ig, inner gonostylus; og, outer gonostylus; t9, tergite 9. Scale bar: 0.5 mm. 199 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China spots, its inner margin bordered by darker brown. Scutellum anteriorly brown, posteriorly yellow with dark median vitta. Mediotergite yellowish brown, with dark median vitta. Legs with coxae and trochanters yellow. Femora yellow, tips brown. Tibiae brownish yellow, tips brown. Tarsal segments yellowish brown. Claw with small tooth. Wing yellowish brown (Fig. 14B). Vein m-cu joining dm at fork of M3+4, cell m1 about 4.3 as long as its petiole. Halter pale yellow, knob pale brown. Abdomen. Abdominal segments 1–4 yellow, segment 5 yellowish brown, remaining segments brown. Lateral stripes narrow, pale brown. Hypopygium (Figs 15–17). Male hypopygium with tergite 9 fused with sternite 9 basally (Fig. 15B). Tergite 9 divided along midline by pale membrane, posteriorly with deep U-shaped notch (Figs 15A, 16A, B). Dorsal porFigure 16. Male terminalia of Tipula (Vestiplex) ventrobasilata sp. nov., holotype (specimen number NRCE000101) A tergite 9 and proctiger, dorsal view B tergite 9, dorsal view C left outer gonostylus D right inner gonostylus, dorsal view E right inner gonostylus, outer lateral view F sternite 9, ventral view, tergite 9 and gonostyli removed G adminiculum, lateral view H left dorsal lobe of appendage of sternite 9. Abbreviations: adm, adminiculum; ap, anal plate; dl, dorsal lobe of A9s; dp, dorsal portion of tergite 9; lb, lower beak of inner gonostylus; mf, median fold of inner gonostylus; pr, proctiger; vl, ventral lobe of A9s; vp, ventral portion of tergite 9. Scale bar: 0.5 mm. 200 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Figure 17. Male terminalia of Tipula (Vestiplex) ventrobasilata sp. nov., holotype (specimen number NRCE000101) A sperm pump and aedeagus, lateral view B sperm pump, dorsal view. Scale bar: 0.5 mm. Figure 18. Female terminalia of Tipula (Vestiplex) ventrobasilata sp. nov., paratype (specimen number NRCE000105) A ovipositor, left lateral view B sternite 8 with hypogynial valves, ventral view C sternite 8 with hypogynial valves, the same sclerite gently pressed by cover glass D sternite 9 and furca, dorsal view E spermatheca. Abbreviations: c, cercus; f, furca; h, hypogynial valve; s8, sternite 8; s9, sternite 9; t10, tergite 10. Scale bars: 0.5 mm a (A); 0.5 mm b (B–E). 201 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China tion with posterior margin produced into nearly triangle lobes on either side of midline, posterolateral part slightly extended, rounded. The surface covered with long setae. Ventral portion large, occupying nearly half of tergite area. Each part flattened; posterior margin slightly emarginated. A pair of blackened, serrulated lobes on either side of midline. The lobe broadly expanded at basal half; distal part narrowed. Gonocoxite partly separated from sternite 9 by suture, unarmed (Fig. 15). Outer gonostylus narrowed at base, expanded in distal third (Fig. 16C). Inner gonostylus nearly crescent plate, terminating in a blackened beak (Fig. 16D, E). Lower beak split in sagital projection into small blackened lobe and yellow, flattened, apically narrowed lobe. Crest with blackened lobe feasibly representing the vestige of outer basal lobe. Outer surface in middle forming elongated, blackened fold. Sternite 9 with ventral lobe of A9s oblong, surface covered with setae (Fig. 16F). Dorsal lobe of A9s in the shape of short, apically rounded lobe (Fig. 16F, H). Adminiculum nearly triangular in ventral view (Fig. 16F). Basal part of adminiculum slightly broadened and indistinctly protruded medially; apex curved, funnel-shaped (Fig. 16G). Genital bridge absent. Proctiger pale brown with anal sclerites small (Fig. 16A). Sperm pump with central vesicle spherical (Fig. 17A). Compressor apodeme with shallow median incision (Fig. 17B). Posterior and anterior immovable apodemes narrowed. Aedeagus about 3 × as long as sperm pump (Fig. 17A). Female. Body length 18.4–28.8 mm, wing length 18.5–21.4 mm. Generally similar to male by body coloration. Female terminalia (Fig. 18). Tergite 10 shiny brown. Cercus brown, nearly straight, slightly shorter than tergite 10 (Fig. 18A); dorsal and ventral margins smooth, without visible serration with. Hypogynial valve extending before base of cercus. Sternite 8 with hypogynial valve yellow, blade-shaped, its outer margin darkened. (Fig. 18B, C). Posterior margin laterally to hypogynial valve with small, rounded extension covered with setae. The lateral angle of sternite 8 rounded. Median incision between hypogynial valves basally provided with setae. Sternite 9 nearly horseshoe-shaped, posterior margin slightly extended with median incision (Fig. 18D). Furca in the shape of narrowed stripe (Fig. 21F). Spermatheca nearly spherical (Fig. 18E). Etymology. The name of the new species refers to the basally expanded ventral lobes of tergite 9. Distribution. China, Sichuan. Remarks. Tipula (V.) ventrobasilata sp. nov. is considered here as a member of the divisotergata species group, which was proposed by Savchenko (1964), with the following discussion in Starkevich et al. (2019a) and Starkevich and Young (2023). The new species is close to T. (V.) paramonovi Men, Sun & Starkevich, 2023 (China, Xizang) in having basally expanded ventral lobes of tergite 9 (Men et al. 2023: figs 33–44) but differs in details of the male genitalia. Tipula (V.) ventrobasilata sp. nov. has ventral lobes relatively short and distinctly narrow in the distal half, while in T. (V.) paramonovi the lobes are relatively stout and longer. The blackened fold of the inner gonostylus is absent in T. (V.) paramonovi. Both species can also be separated by the shape of the appendage of sternite 9, which is broad at the base and narrow distally in T. (V.) paramonovi while T. (V.) ventrobasilata sp. nov. possess a short and apically rounded appendage of sternite 9. 202 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Tipula (Vestiplex) aestiva Savchenko Figs 19–21 Tipula (Vestiplex) aestiva Savchenko, 1960: 191; 1964: 215; Gao et al. 2023. Material examined. China • Holotype ♂; Qinghai, south shore of Kuku-nor Lake; August 1901; Kozlov leg.; ZIN • Paratype ♀; same data as for preceding; ZIN; • Paratype ♂; Qinghai, Sogon-Gomba, I-chu River, up to Yangtze River; July 1900; Kozlov leg.; ZIN; • Paratype ♂; Qinghai; shore of Orin-nor Lake, Huang-he Basin; May–June 1901; Kozlov leg.; ZIN. Additional material examined. China • ♂ (in alcohol); Gansu, near Xiche (Labrang); 35°11.968'N, 102°33.545'E; alt. 2900 m; 23 May 2017; A. Saldaitis leg.; NRC; Specimen number NRCE000090; genitalia dissected. Diagnosis. Tipula (V.) aestiva can be recognized by brownish-yellow body and elongated antenna reaching beyond base of abdomen. Hypopygium (Figs 19–21). Tergite 9 forming a concave saucer-shaped plate (Figs 19, 20A). The main body of tergal plate yellow, posterior margin emarginated with median U-shaped notch; lateral angles of plate rounded and blackened; anterior portion of tergal plate raised into narrow sclerotised border. Gonocoxite separated from sternite 9 by suture, in the shape of a large, nearly rectangular plate (Figs 19; 20B, C). Apex of gonocoxite terminating in dorsal and ventral blackened teeth. Outer gonostylus nearly leaf-shaped, narrow at base, rest of body flattened (Fig. 20D). Inner gonostylus in the shape of slightly curved plate (Fig. 20E–G), beak broadly blackened with tip rounded, lower beak absent; dorsal margin apically with short extension. Genital bridge with sclerite sp2 small (Fig. 20C), sclerites sp1 fused at base forming V-shaped structure (Fig. 20K). Adminiculum triangular in distal half with short wrinkled gonapophysies (Fig. 20H, I). Apex ventrally produced into short denticle. Proctiger pale with small, uniform anal plates (Fig. 20J). Sperm pump with central vesicle flattened (Fig. 21A). Compressor apodeme with rounded median incision (Fig. 21B). Posterior Figure 19. Male terminalia of Tipula (Vestiplex) aestiva (specimen number NRCE000090) A hypopygium, dorsal view B hypopygium, lateral view. Abbreviations: dt, dorsal tooth; gcx, gonocoxite; ig, inner gonostylus; og, outer gonostylus; t9, tergite 9; vt, ventral tooth. Scale bar: 0.5 mm. 203 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China immovable apodeme extended, slightly flattened with additional short median lobes on inner side (Fig. 21C). Anterior immovable apodeme nearly triangle. Aedeagus about 4.8 × as long as sperm pump, blackish brown except yellow apex. Apex with sagittal and frontal incisions, trident in shape (Fig. 21A, D). Figure 20. Male terminalia of Tipula (Vestiplex) aestiva (specimen number NRCE000090) A tergite 9, dorsal view B left gonocoxite, lateral view C left gonocoxite and sp2 of genital bridge, caudal view D left outer gonostylus E left inner gonostylus, outer lateral view F left inner gonostylus, outer dorsal view G left inner gonostylus, mesal view H adminiculum, lateral view I adminiculum, dorsal view J proctiger, dorsal view K genital bridge, sclerites sp1, dorsal view. Abbreviations: ap, anal plate; gph, gonapophysis; sp2, sclerite sp2 of genital bridge. Scale bar: 0.5 mm. 204 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Remarks. The identification of the specimen collected in Gansu Province is based on comparative analysis with type specimens and morphology of dissected genitalia. The male genital structures of the male from Gansu are identical to that of the dissected paratype (Starkevich 2012, fig. 40; illustrations are also available in Oosterbroek 2024), including details of sperm pump having a specific median lobe on the inner side of the posterior immovable apodeme. Acknowledgements We are grateful to the following Museum curators for their help and assistance in obtaining specimens: Dr D. Sivell, NHMUK, United Kingdom; Dr J. Roháček, SMOC, Czech Republic; Dr J. Gelhaus, ANSP, USA and Dr T. Dikow, USNM, USA. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding The visit of P. Starkevich to ANSP was supported by the European Social Fund (project No 09.3.3-LMT-K-712-14-0146) under grant agreement with the Research Council Figure 21. Male terminalia of Tipula (Vestiplex) aestiva (specimen number NRCE000090) A sperm pump and aedeagus, lateral view B sperm pump, caudal view C sperm pump, dorsal view D apical part of aedeagus. Abbreviations: aia, anterior immovable apodeme; ca, compressor apodeme; ml, median lobe; pia, posterior immovable apodeme. Scale bars: 0.5 mm a (A–C); 0.25 mm b (D). 205 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China of Lithuania (LMTLT). The visit to the BMNH (GB-TAF-5424) was supported by the SYNTHESYS (http://www.synthesys.info/), which was financed by European Community Research Action un-der the FP7 Integrating Activities Programme. Author contributions All authors have contributed equally. Author ORCIDs Pavel Starkevich https://orcid.org/0000-0002-0746-0398 Qiu-Lei Men https://orcid.org/0000-0002-1233-6439 Aidas Saldaitis https://orcid.org/0000-0003-0999-3996 Kristina Valavičiūtė-Pocienė https://orcid.org/0000-0003-4444-1388 Data availability All of the data that support the findings of this study are available in the main text. References Alexander CP (1934) New or little-known Tipulidae from eastern Asia (Diptera). XVII. Philippine Journal of Science 52: 395–442. Alexander CP (1935) New or little-known Tipulidae from eastern Asia (Diptera). XXV. Philippine Journal of Science 57: 81–148. Cumming JM, Wood DM (2017) Adult morphology and terminology. In: Kirk-Spriggs AH, Sinclair BJ (Eds) Manual of Afrotropical Diptera, Volume 1: Introductory Chapters and Keys to Diptera Families. Suricata 4. South African National Biodiversity Institute, Pretoria, 89–133. de Jong H (2017) Limoniidae and Tipulidae (Crane Flies). In: Kirk-Spriggs AH, Sinclair BJ (Eds) Manual of Afrotropical Diptera, Volume 2: Nematocerous Diptera and lower Brachycera. Suricata 5. South African National Biodiversity Institute, Pretoria, 427–477. Dobrotworsky N (1968) The Tipulidae (Diptera) of Australia I. A review of the genera of the subamily Tipulinae. Australian Journal of Zoology 16: 459–494. https://doi. org/10.1071/ZO9680459 Gao Y, Zhang B, Yang D, Li Y (2023) The mitochondrial genome of Tipula (Vestiplex) aestiva Savchenko, 1960 (Diptera: Tipulidae). Mitochondrial DNA. Part B, Resources 8(2): 233–235. https://doi.org/10.1080/23802359.2023.2172975 Gelhaus JK (2005) Systematics and biogeography of the desert crane fly subgenus Tipula (Eremotipula) Alexander (Diptera: Tipulidae). Memoirs of the American Entomological Society 46: 1–235. Mannheims B (1963) 15. Tipulidae. In: Lindner E (Ed.) Die Fliegen der palaearktischen Region 3(5)1, Lief, 137–176. Men Q-L, Young CW, Starkevich P, Yu Y-F, Lei X-P (2017) Two new species of Tipula (Vestiplex) from Southern China based on morphological and molecular data, with redescription of Tipula (Vestiplex) bicalcarata (Diptera, Tipulidae, Tipulinae). ZooKeys 658: 63–80. https://doi.org/10.3897/zookeys.658.9738 Men Q-L, Starkevich P, Saldaitis A (2021a) Two new crane fly species of the subgenus Vestiplex Bezzi, 1924 (Diptera, Tipulidae, Tipula) from Yunnan and Sichuan, China, with a key to species in the immota species group. ZooKeys 1040: 33–48. https://doi. org/10.3897/zookeys.1040.64376 206 ZooKeys 1264: 183–206 (2025), DOI: 10.3897/zookeys.1264.139740 Pavel Starkevich et al.: New Vestiplex crane flies from China Men Q-L, Starkevich P, Sun M-W (2021b) Two New Species of the Genus Tipula Linnaeus, 1758 (Diptera: Tipulidae) from China. Annales Zoologici 71: 243–250. https://doi.org /10.3161/00034541ANZ2021.71.2.004 Men Q-L, Starkevich P, He L-F, Shi J-X, Shi M-Y, Zhang Z-X, Hu J-H, Chen A, Zhang Y-X (2021c) Tipula (Vestiplex) from Yunnan and Tibet, China: one new species and redescriptions of five known species (Diptera: Tipulidae). Acta Entomologica Musei Nationalis Pragae 61: 341–362. https://doi.org/10.37520/aemnp.2021.020 Men Q-L, Sun Z, Starkevich P (2023) Four new crane fly species of the subgenus Vestiplex (Diptera: Tipulidae: Tipula) from Tibet and Yunnan, China. European Journal of Taxonomy 856(1): 170–192. https://doi.org/10.5852/ejt.2023.856.2037 Neumann H (1958) Der Bau und die Funktion der männlichen Genitalapparate von Trichocera annulata Meig. und Tipula paludosa Meig. (Dipt. Nematocera). Deutsche Entomologische Zeitschrift 5: 235–298. Oosterbroek P (2024) Catalogue of the Craneflies of the World (CCW). https://ccw.naturalis.nl/index.php [April 10, 2024] Ren J, Li Y, Yang D (2021) Two new species of Tipula (Vestiplex) Bezzi (Diptera, Tipulidae) from Northwest China. Zootaxa 4903. https://doi.org/10.11646/zootaxa.4903.3.4 Savchenko EN (1955) Two new species of crane-flies (Diptera, Tipulidae) with shortwinged males. Entomological Review 34: 287–297. Savchenko EN (1960) A contribution to the taxonomy of crane-flies (Diptera, Tipulidae) of the subgenus Vestiplex Bezzi of the genus Tipula L. Horae Societatis Entomologicae Rossicae Unionis Soveticae 47: 143–216. Savchenko EN (1964) Crane-flies (Diptera, Tipulidae), Subfam. Tipulinae, Genus Tipula L., 2. Fauna USSR, N.S. 89, Insecta Diptera, 1–503 pp. Starkevich P (2012) Taxonomic and phylogenetic review of the subgenus Tipula (Vestiplex) Bezzi, 1924 (Diptera, Tipulidae). Doctoral dissertation. Vilnius University, Vilnius, Lithuania. Starkevich P, Young CW (2023) Taxonomic revision of Tipula (Vestiplex Bezzi) crane flies (Diptera, Tipulidae) in Taiwan with descriptions of six new species. Insects 14: 616. https://doi.org/10.3390/insects14070616 Starkevich P, Men Q-L, Saldaitis A (2019a) Four new crane fly species of subgenus Tipula (Vestiplex) (Diptera, Tipulidae) from China. Zootaxa 4679: 69–86. https://doi. org/10.11646/zootaxa.4679.1.4 Starkevich P, Saldaitis A, Men Q-L (2019b) Tipula (Vestiplex) butvilai sp. nov., a new crane fly (Diptera, Tipulidae) from Yunnan, China. ZooKeys 869: 91–101. https://doi. org/10.3897/zookeys.869.34166 Starkevich P, Podenas S, Gelhaus JK (2020) Taxonomic review of Tipula (Vestiplex Bezzi) crane flies (Diptera: Tipulidae) in Mongolia. Zootaxa 4837: 1–88. https://doi. org/10.11646/zootaxa.4837.1.1 Starkevich P, Podėnas S, Podėnienė V, Park S-J, Kim A-Y (2021) Tipula (Vestiplex) crane flies (Diptera, Tipulidae) of Korea. ZooKeys 1061: 23–55. https://doi.org/10.3897/ zookeys.1061.49999 Yang Q, Li Y, Pan Z, Liu X, Yang D (2021) Six new species of Tipula (Vestiplex) Bezzi (Diptera, Tipulidae) from Tibet. European Journal of Taxonomy 764(1): 152–192. https:// doi.org/10.5852/ejt.2021.764.1477 Yang Q, Chen W, Qian L, Yang D, Liu X, Wang M (2022) The effect of environmental factors on the diversity of crane flies (Tipulidae) in mountainous and non-mountainous regions of the Qinghai-Tibet plateau and surrounding areas. Insects 13: 1054. https:// doi.org/10.3390/insects13111054