Overview of the Bee Genus Trachusa Panzer, 1804 (Hymenoptera: Apoidea: Megachilidae: Anthidiini) from China with Description of Three New Species

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Overview of the bee genus Trachusa Panzer, 1804 (Hymenoptera: Apoidea: Megachilidae: Anthidiini) from China with description of three new species

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Overview of the bee genus Trachusa Panzer, 1804 (Hymenoptera: Apoidea: Megachilidae: Anthidiini) from China with description of three new species

ZE-QING NIU1, JOHN S. ASCHER2, MICHAEL C. ORR1, TERRY GRISWOLD3 & CHAO-DONG ZHU1,4,5,6 1Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1 Beichen West Road, Chaoyang District, Beijing, 100101, P. R. China. E-mail: [email protected]; E-mail: [email protected] 2Department of Biological Sciences National University of Singapore, 14 Science Drive 4, Singapore 117543, Singapore. E-mail: [email protected] 3USDA-ARS Pollinating Insects Research Unit, Utah State University, Logan, Utah 84322-5310, USA. E-mail: [email protected] 4State Key Laboratory of Integrated Pest Management, Institute of Zoology, Chinese Academy of Sciences, 1 Beichen West Road, Chaoyang District, Beijing, 100101, P. R. China. 5College of Life Sciences, University of Chinese Academy of Sciences, No.19(A) Yuquan Road, Shijingshan District, Beijing, P. R. China 100049 6Corresponding author. E-mail: [email protected]

Abstract

The Chinese bees of the genus Trachusa Panzer, 1804 are reviewed. Nine species are confirmed to occur in China. Three new species are described and illustrated: Trachusa (Paraanthidium) pingdaensis Niu, sp. nov., T. (P.) staabi Niu, sp. nov. and T. (P.) wuae Niu, sp. nov. The distribution of each species is given. An illustrated key to the Chinese species is provided.

Key words: Apiformes, review, fauna, illustration, taxonomy

Introduction

The genus Trachusa was erected by Georg Wolfgang Franz Panzer in 1804 for the type species Trachusa serratulae Panzer, 1804. Pasteels (1969, 1972) classified the Old World Trachusa in seven subgenera. Protanthidium Cockerell et Cockerell, 1911 was regarded as a synonym of Paraanthidium Friese, 1898 (Mavromoustakis, 1937; Sandhouse, 1943). Later, Michener & Griswold (1994) assigned the genus Archianthidium Mavromoustakis, 1939 as another subgenus of Trachusa and subdivided Trachusa into 11 subgenera (Michener 2000, 2007). Kasparek (2017) gave a key to seven subgenera in the Old World and attributed 49 species to the genus of Old World and New World. Kasparek (2018) published a new species from southern Africa and regarded the subgenus Massanthidium Pasteels as a junior synonym of Paraanthidium, thus reducing the number of subgenera to 10 worldwide (see the subgeneric list in the systematics). At present, only two subgenera (Orthanthidium and Paraanthidium) are recorded in China. Wu (2006) recorded 15 species from China, one under the subgenus Orthanthidium and 14 under the subgenus Paraanthidium, provid- ing morphological characters to distinguish them and a key to the Chinese species of Paraanthidium. There are few clear morphological features which characterize the genus Trachusa unambiguously (Kasparek 2017). Many species assigned to Trachusa by Wu (2006) should be transferred to other genera in the tribe Anthidiini (Niu et al. 2016; Kasparek 2017). Subsequently, only five species of them remain in the genus Trachusa. This study leverages the underused collection of the Institute of Zoology, Chinese Academy of Sciences, Bei- jing, China (IZCAS) to review the Chinese species of this difficult group of bees. We confirm nine species of Trachusa occurring in China (Table 1), including three new species, verify and expand upon the information of previously described Chinese species, and provide illustrations.

Accepted by J. Gibbs: 21 Jun. 2019; published: 24 Jul. 2019 251 TABLE 1. Summary of species of Trachusa currently recognized from China with information on the known sexes and distributions. Sex: ♂ = male; ♀ = female. * represents a new distributional record. Species Name in Wu (2006) Sexes Distribution known Trachusa (Orthanthidium) Trachusa (Paraanthidium) ♀, ♂ China (Beijing, Tianjin*, cornopes Wu, 2004 cornopes Wu, 2004 Shanxi*, Shaanxi, Hubei) Trachusa (Orthanthidium) Trachusa (Orthanthidium) ♀, ♂ China (Guizhou, Fujian, formosana (Friese, 1917) formosanum (Friese, 1917) Guangdong*, Taiwan) Trachusa (Paraanthidium) Trachusa (Paraanthidium) ♀, ♂ India, Nepal, China longicornis (Friese, 1902) longicorne (Friese, 1902) (Yunnan) Trachusa (Paraanthidium) Trachusa (Paraanthidium) ♀, ♂ China (Fujian, Jiangxi, maai (Mavromoustakis, 1953) maai (Mavromoustakis, 1953) Guangxi, Guizhou) Trachusa (Paraanthidium) Trachusa (Paraanthidium) ♀, ♂ China (Zhejiang*, Fujian, muiri (Mavromoustakis, 1937) muiri (Mavromoustakis, 1937) Guangdong, Yunnan) Trachusa (Paraanthidium) pingdaen- ♀ China (Yunnan)* sis Niu, sp. nov. Trachusa (Paraanthidium) staabi ♀ China (Zhejiang*, Fujian*) Niu, sp. nov. Trachusa (Paraanthidium) wuae Niu, ♂ China (Yunnan)* sp. nov. Trachusa (Paraanthidium) xylocopi- ♂ China (Fujian, formis (Mavromoustakis, 1954) Guangdong*)

Materials and methods

All specimens examined (76/83 specimens) in this study are deposited in the Insect Collection of the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IZCAS). In the species accounts only specimens not from IZCAS have the institution indicated by the city where the museum is located. The specimens were examined with a Nikon SMZ 1500 stereomicroscope and images were recorded with a Nikon D7000 digital camera. The morphological terminology follows Michener (2007). Absolute measurements, in millimeters (mm), are used for length of body. For all other structures, relative measurements are used. Abbreviations used in the description are as follows: BL (body length): measured from the base of the antennal socket to the apex of the metasoma; HL (head length): measured from the apicomedial margin of the clypeus to the upper margin of the vertex in frontal view; HW (head width): measured at the widest point of the head across the compound eyes in frontal view; EW (eye width): the greatest width of eye in lateral view; GW (genal width): the greatest width of the gena in lateral view; MsW (mesosomal width): measured between the outer rims of the tegulae in dorsal view; MtW (metasomal width): the greatest width of the metasomal tergum in dorsal view. Abbreviations T1, T2…, S1, S2…and F1, F2…, respectively, denote the first, second, etc., segments of terga, sterna and flagellomeres. New distribution records are marked by an asterisk (*).

Systematics

Trachusa Panzer, 1804

Trachusa (Archianthidium): Archianthidium Mavromoustakis, 1939: 91. Type species: Anthidium laticeps Morawitz, 1874, by original designation. Trachusab (Congotrachusa): Trachusa (Congotrachusa) Pasteels, 1969: 24. Type species: Anthidium schoutedeni Vachal, 1910, by original designation. Trachusa (Heteranthidium): Anthidium (Heteranthidium) Cockerell, 1904: 292. Type species: Anthidium dorsale Lepeletier, 1841, by original designation. Trachusa (Legnanthidium): Trachusa (Legnanthidium) Griswold & Michener, 1988: 27. Type species: Anthidium ridingsii Cresson, 1878, by original designation.

252 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. Trachusa (Metatrachusa): Trachusa (Metatrachusa) Pasteels, 1969: 22. Type species: Anthidium pendleburyi Cockerell, 1927, by original designation. Trachusa (Orthanthidium): Paraanthidium (Orthanthidium) Mavromoustakis, 1953: 837. Type species: Anthidium formosanum Friese, 1917, by original designation. Trachusa (Paranthidium): Anthidium (Paraanthidium) Friese, 1898: 101. Type species: Apis interrupta Fabricius, 1781, by designation of Cockerell, 1909: 269. Trachusa (Trachusa): Trachusa Panzer s. str., 1804: expl. pl. 14–15. Type species: Trachusa serratulae Panzer, 1804 = Apis byssina Panzer, 1798, by designation of Sandhouse, 1943: 605. Trachusa (Trachusomimus): Trachusomimus Popov, 1964: 406. Type species: Trachusa perdita Cockerell, 1904, by original designation. Trachusa (Ulanthidium): Ulanthidium Michener, 1948: 13. Type species: Ulanthidium mitchelli Michener, 1948, by original designation.

Diagnosis: The genus Trachusa Panzer, 1804 consists of moderate-sized to large megachiliform anthidiine bees, the body length of which varies from 8.5 mm to 20 mm. A carina on the pronotal lobe and the omaulus is often absent, but if present may take the form of a lamella (Kasparek, 2017). The posterior part of the head is well-developed, thus the OOD (ocellocular distance) is shorter than the OCD (ocelloccipital distance) except equal in the subgenus Metatrachusa. The anterior of the median ocellus is closer to the antennal bases than to the posterior margin of the vertex (Grisword & Michener, 1988). The middle tibia is broad, usually nearly as broad as the hind tibia, and both the anterior and posterior margins of the middle tibia are convex. Despite these convexities, the middle tibia is sometimes distinctly narrower than the hind tibia (Michener, 2007). Fore- and mid-tibial spines are produced as blunt, obtuse projections that extend along the tibial surface as carinae (Griswold & Michener, 1988). Vein cu-v of the hind wing is oblique and usually nearly half as long as the second abscissa of M+Cu or longer (Kasparek, 2017). Arolia between the claws are usually present but sometimes greatly reduced or absent (Kasparek, 2017). Black combs on the underside of S4 and S5 of the male are present in some species (Kasparek, 2017). The male T7 is small and curved under the abdomen so that the dorsal surface faces ventrally (Michener, 2007). Females typically have four mandibular teeth but sometimes three, or five, six or seven (Michener, 2007). The number of segments in the maxillary palpus is four or reduced to three (Kasparek, 2017).

Key to the known species of Trachusa of China (The females of T. xylocopiformis (Mavromoustakis) and T. wuae Niu, sp. nov. as well as the males of T. pingdaensis Niu, sp. nov. and T. staabi Niu, sp. nov. are unknown and are not covered by the key.)

1 Flagellum 10-segmented, female ...... 2 - Flagellum 11-segmented, male ...... 8 2 Scutellum truncate posteriorly (subgenus Orthanthidium) (Fig. 1C; Fig. 2C) ...... 3 - Scutellum broadly rounded but medially emarginate (subgenus Paraanthidium) (Fig. 4C; Fig. 5C; Fig. 6C; Fig. 9D; Fig. 10E) ...... 4 3 Middle-sized (13–15 mm) (Fig. 1A); T1-T5 with narrow yellow subapical bands, the bands with shallow notch laterally (Fig. 1D) ...... Trachusa (Orthanthidium) cornopes Wu, 2004 - Large-sized (18–20 mm) (Fig. 2A); T1-T3 with broad subapical yellow bands, and the bands with black marking laterally, T4- T5 with narrow subapical yellow bands (Fig. 2E) ...... Trachusa (Orthanthidium) formosana (Friese, 1917) 4 T3 with yellow maculation (frequently present also on T2); scutum almost always with yellow maculations, at least near tegulae (Fig. 5C; Fig. 6C) ...... 5 - T2–T3 without yellow maculations; scutum entirely black (Fig. 4C; Fig. 9D; Fig. 10E) ...... 6 5 T2 with punctation increasingly sparse from base to rim, broadly impunctate apically and lacking raised rim (Fig. 5E); anterior edge of fore tarsus expanded laterally and sharply angled from outer to inner surface (Fig. 5A); scutellum dark (Fig. 5C) ...... Trachusa (Paraanthidium) maai (Mavromoustakis, 1953) - T2 densely punctate throughout, up to very narrow raised rim (Fig. 6D); anterior edge of fore tarsus unmodified, rounded over evenly from outer to inner surface (Fig. 6A); scutellum with light marking (Fig. 6C) ...... Trachusa (Paraanthidium) muiri (Mavromoustakis, 1937) 6 All terga black (Fig. 10F); T6 with longer appressed gold-orange setae branched and nearly covering surface (Fig. 10G) ...... Trachusa (Paraanthidium) staabi Niu, sp. nov. - At least T5-T6 yellow (Fig. 4D; Fig. 9F); T6, if with gold-orange setae, shorter and less branched, barely obscuring surface if at all ...... 7 7 T1–T4 black, T5–T6 yellow (Fig. 4D); omaulus strongly carinate, surface concave anteriorly; pronotal lobe strongly carinate, obscure medial bump clearly shorter in height than carina (Fig. 4C) ...... Trachusa (Paraanthidium) longicornis (Friese, 1902)

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 253 - T1–T3 black, apical half disc of T4 and T5–T6 yellow (Fig. 9F); omaulus distinctly angled but lacking carina, anterior surface weakly concave if at all; pronotal lobe at most moderately carinate, distinct medial bump of similar height to carina ...... Trachusa (Paraanthidium) pingdaensis Niu, sp. nov. 8 Metasoma black without yellow markings (Fig. 8D, Fig. 8E) ...... Trachusa (Paraanthidium) xylocopiformis (Mavromoustakis, 1954) - Metasoma black with yellow markings (Fig. 1F; Fig. 3D; Fig. 7D) ...... 9 9 T1 with yellow markings (Fig. 1F; Fig. 3D) ...... 10 - T1 black, without light markings (Fig. 4I; Fig. 7D; Fig. 11D) ...... 11 10 T1–T5 black, each tergum with a narrow submarginal yellow band, all bands linear (Fig. 1F); most median part of clypeus yellow, with adjacent black markings submedially (Fig. 1G) ...... Trachusa (Orthanthidium) cornopes Wu, 2004 - T1–T2 with a narrow submarginal yellow band, with black marking laterally, T3 with interrupted linear yellow band, T4–T5 without submarginal yellow band (Fig. 3D); clypeus yellow (Fig. 3B) ...... Trachusa (Orthanthidium) formosana (Friese, 1917) 11 Scutum with short lateral yellow stripe at each side near tegulae ...... Trachusa (Paraanthidium) maai (Mavromoustakis, 1953) - Scutum black (Fig. 4H; Fig. 7C; Fig. 11C) ...... 12 12 Clypeal punctation sparser and coarser (Fig. 4G); T5 integument entirely yellow (Fig. 4I) ...... Trachusa (Paraanthidium) longicornis (Friese, 1902) - Clyeal punctation denser and finer (Fig. 7B, Fig. 11B)); T5 yellow along rim, slightly thicker medially but still largely dark (Fig. 7E) ...... 13 13 Pronotal carina moderately-developed, as tall as or taller than medial bump of pronotum; propodeal enclosure with band of punctures near at top, of even density throughout; scutellum deeply and clearly medially emarginate (Fig. 7C) ...... Trachusa (Paraanthidium) muiri (Mavromoustakis, 1937) - Pronotal carina weak, shorter in height than medial bump of pronotum; propodeal enclosure with band of punctures at top interrupted or at least sparser medially; scutellum only weakly emarginate medially, nearly straight along rim (Fig. 11C) ...... Trachusa (Paraanthidium) wuae Niu, sp. nov.

Species Accounts

Trachusa (Orthanthidium) cornopes Wu, 2004 (Fig. 1: A–E, female; F–J, male)

Trachusa (Paraanthidium) cornopes Wu, 2004: 546, ♀, ♂. Holotype: ♂, China, Hubei, Lichuan, Xingdoushan, 21.VII.1989, Long-Long YANG; paratypes: 1 ♀, China, Shaanxi, Chang’an, 29.VI.1965, Yong-Lin CHENG, 1 ♀, Shaanxi, Qingling, Fuping, 31.VIII.1983, Ding-Xi LIAO; IZCAS; examined. Trachusa (Paraanthidium) cornopes: Wu, 2006: 182, ♀, ♂ (redescription), Fig. 99. Trachusa (Orthanthidium) cornopes: Kasparek, 2017: 85, ♀, ♂ (redescription).

Material examined: China: Hubei, Lichuan, Xingdoushan (30º02′N, 109º08′E), 800 m, 1 ♂ (holotype), 21.VII.1989, leg. Long-Long YANG; Shaanxi, Chang’an (34º04′N, 108º52′E), 1 ♀ (paratype), 29.VI.1965, leg. Yong-Lin CHENG; Shaanxi, Qinling, Fuping (34º52′N, 109º14′E), 1 ♀ (paratype), 31.VIII.1983, leg. Ding-Xi LIAO; Tianjin, Ji Xian, Jiulongshan (40º00′N, 117º18′E), 4 ♀, 17 ♂, 19.VI.2017, leg. Yong WANG; Shanxi, Jiang Xian (35º29′N, 111º38′E), 1 ♀, 22.VII.1980, leg. Lin-Fu LI. General distribution: China (Beijing, Tianjin*, Shanxi*, Shaanxi, Hubei). Remarks: The assignment of Trachusa cornopes to the subgenus Orthanthidium should be regarded as provi- sional (Kasparek, 2017). Further studies are needed to show whether it is justified to accommodate T. cornopes and T. formosana, two quite different species, in the same subgenus (Kasparek, 2017).

Trachusa (Orthanthidium) formosana (Friese, 1917) (Fig. 2: A–E, female; Fig. 3: A–H, male)

Anthidium formosanum Friese, 1917: 59, ♀. Holotype: ♀, “Formosa, bei Taihorinsho in 1 August, leg. Sauter“ [China: Taiwan], Museum für Naturkunde, Berlin, Germany. Paraanthidium (Orthanthidium) formosanum: Mavromoustakis, 1953: 837, ♀ (colour differences). Trachusa (Orthanthidium) formosana: Pasteels, 1972: 85, ♀ (redescription); Kasparek, 2017: 90, ♀ (redescription). Trachusa (Orthanthidium) formosanum: Wu, 2006: 171, ♀ (redescription), ♂ (new description), Fig. 91.

254 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. Material examined: China: Guizhou, Jiangkou Xian, Fanjing Shan (27º36′N, 108º48′E), 500 m, 1 ♂, 14.VIII.1988, leg. Long-Long YANG; Guangdong, Nanling, Dadongshan (24º46′N, 112º41′E), 1 ♂, 24.VI.2009, leg. Ya-Ping WU, 2 ♀, 22.VI.2009, leg. Bo-Wen YU; Formosa Sauter Fuhosho [Wucheng], IV.1909, 1 ♂, 4 ♀ (Budapest, Logan). General distribution: China (Guizhou, Fujian, Guangdong*, Taiwan). Remarks: Kasparek (2017) only redescribed the female because he thought the male was unknown, but Wu (2006) previously described the male. The coloration pattern of the female specimens collected from Guangdong is different in T3–T5 (especially, T3 with one continuous yellow band on disc, not as one narrow yellow stripe each on both sides of the middle line of T3), scutum and scutellum from that of the specimens described and depicted by Kasparek (2017).

Trachusa (Paraanthidium) longicornis (Friese, 1902) (Fig. 4: A–E, female; F–J, male)

Megachile steloides Bingham, 1896 (nom. praeocc., nec Anthidium steloides Spinola, 1851): 198, ♀, type locality: Sikkim (now India); Bingham, 1897: 490, ♀ (redescription). Anthidium steloides: Friese, 1901: 224. Protanthidium steloides: Cockerell & Cockerell, 1901: 49, ♂ (new description). Anthidium longicorne Friese, 1902: 109, replacement name for Megachile steloides Bingham, 1896. Paraanthidium steloides: Mavromoustakis, 1948: 420: ♀ (redescription); 1953: 836, ♀ (redescription). Trachusa (Protanthidium) steloides: Pasteels, 1972: 78, ♀, ♂ (redescription), Figs 13–16. Paraanthidium longicorne: Wu, 1962: 161 (first record for China); Wu et al., 1988: 64, ♀, ♂ (redescription). Trachusa (Paraanthidium) longicorne: Wu, 2006: 175, ♀, ♂ (redescription), Fig. 92. Trachusa (Paraanthidium) longicornis: Michener & Grisword, 1994: 324 (comments on replacement name); Kasparek, 2017: 113, ♀, ♂ (redescription).

Material examined: China: Yunnan, Xiaomengyang (22º00′N, 100º54′E), 850 m, 1 ♀, 1 ♂, 4.V.1957, leg. Shu- Yong WANG, 850 m, 9 ♀, 4.V.1957, leg. Qiu-Zhen LIANG, 950 m, 2 ♀, 4.V.1957, leg. Ling-Chao ZANG; Yunnan, Cheli (23º06′N, 102º10′E), 650 m, 2 ♀, 26.IV.1957, leg. Shu-Yong WANG, idem, 560 m, 7 ♀, 26.IV.1957, leg. Da-Hua LIU; Yunnan, Xishuangbanna, Meng’a (22º06′N, 100º18′E), 1000 m, 1 ♀, 20.X.1958; 4 ♀, 23.X.1958, leg. Zhi-Zi CHENG; idem, 1050–1080 m, 1 ♀, 20.V.1958, leg. Shu-Yong WANG; idem, 700 m, 1 ♀, 1.VI.1958, leg. Chun-Pei HONG; Yunnan, Xishuangbanna, Menglun (22º00′N, 100º48′E), 600 m, 1 ♀, 22.IV.1994, leg. Long-Long YANG. General distribution: India, Nepal, China (Yunnan).

Trachusa (Paraanthidium) maai (Mavromoustakis, 1953) (Fig. 5: A–E, female)

Paraanthidium maai Mavromoustakis, 1953: 834–836, ♀. Holotype: ♀, “S.E. China: Psingshanpu, 600 m., Changtinghsien (Fukien Province), leg. Tsing-chao Maa” [Changdin Xian, Fujian, China], coll. of G. Mavromoustakis; Mavromoustakis, 1954: 250, ♂ (new description). Trachusa (Proanthidium) maai: Pasteels, 1972: 82, ♀ (redescription), Figs 21–24 (assigned to Proanthidium, but listed under Protanthidium); Wu, 2006: 175, ♀ (description); Kasparek, 2017: 118, ♀, ♂ (redescription).

Material examined: China: Guangxi, Longsheng (25º42′N, 110º00′E), 1 ♀, 12.VII.1983, leg. no information on label; Guizhou, Leishan, Taojiang (26º20′N, 108º09′E), 1000 m, 1 ♀, 5.VII.1988, leg. Long-Long YANG; Fujian, Jiangle Xian (26º42′N, 117º24′E), 490 m, 1 ♀, 3.VII.1991, leg. Xing-Jian WANG. General distribution: China (Fujian, Jiangxi, Guangxi, Guizhou). Remarks: Wu (2006) only described the female of the species, and thought the male is unknown. However, the male was described by Mavromoustakis (1954) (see also redescription in Kasparek 2017).

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 255 FIGURE 1. Trachusa (Orthanthidium) cornopes Wu, 2004, A–E, female, F–J, male. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Metasoma in dorsal view; E: Fore wing in frontal view; F: Body in lateral view; G: Head in frontal view; H: Mesosoma in dorsal view; I: T3–T6 in dorsal view; J: Metasoma in ventral view. Scale bars: 1.0 mm.

256 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. FIGURE 2. Trachusa (Orthanthidium) formosana (Friese, 1917), A–E, female. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Fore wing in frontal view; D: Metasoma in dorsal view. Scale bars: 1.0 mm.

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 257 FIGURE 3. Trachusa (Orthanthidium) formosana (Friese, 1917), A–H, male. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: T1–T3 in dorsal view; E: T4–T6 in dorsal view; F: Metasoma in ventral view; G: Genitalia in lateral view; H: Genitalia in ventrolateral view. Scale bars: 1.0 mm.

258 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. FIGURE 4. Trachusa (Paraanthidium) longicornis (Friese, 1902), A–E, female, F–J, male. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Metasoma in dorsal view; E: Fore wing in frontal view; F: Body in lateral view; G: Head in frontal view; H: Mesosoma in dorsal view; I: Metasoma in dorsal view; J: Metasoma in ventral view. Scale bars: 1.0 mm.

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 259 FIGURE 5. Trachusa (Paraanthidium) maai (Mavromoustakis, 1953), A–E, female. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Fore wing in frontal view; E: Metasoma in dorsal view. Scale bars: 1.0 mm.

Paraanthidium muiri Mavromoustakis, 1937: 155–157, ♀. Holotype: ♀, “China: How Lik, 1911, leg. F. Muir” [Dinghushan Biosphere Reserve, Guangdong, China], Natural History Museum, London, UK; Mavromoustakis, 1954: 249, ♂ (new description); Wu, 1962: 162 (list); Wu et al., 1988: 60, ♂ (redescription, misidentification). Trachusa (Paraanthidium) muiri: Wu, 2006: 176, ♂ (redescription, misidentification), Fig. 93; Kasparek, 2017: 121, ♀, ♂ (redescription).

Material examined: China: Zhejiang, Hanzhou (30º15′N, 120º13′E), 1 ♀, 7.VII.1980, leg. no information on label; Guangdong, Shenzhen, Luohu, Fairy Lake Botanical Garden (22º35′N, 114º10′E), 1 ♀, 4 ♂, 6.V.2016, leg. Feng YUAN; Guangdong, Yangchun, Chuncheng, Panlong (23º09′N, 112º56′E), 1 ♂, 17.IV.2018, leg. Feng YUAN. General distribution: China (Fujian, Zhejiang*, Guangdong, Yunnan). Remarks: Wu (2006) thought the female was unknown, but the female had been described by Mavromoustakis (1937). Wu et al. (1988) and Wu (2006) redescribed the male specimens collected from Yunnan and identified them as the male of T. muiri, but the characters show it is misidentified, it should be Trachusa (Paraanthidium) wuae Niu, sp. nov. As to the locality of Paraanthidium muiri Mavromoustakis, 1937, Mavromoustakis (1937) only recorded “China: Kow Lik”, We confirm it is “Dinghushan Biosphere Reserve, Guangdong, China” (Dr. Max Kasparek, pers. comm.).

Trachusa (Paraanthidium) xylocopiformis (Mavromoustakis, 1954) (Fig. 8: A–H, male)

Paraanthidium xylocopiforme Mavromoustakis, 1954: 251–252, Fig. 1, ♂. Holotype: ♂, “S.E. China: Chungan, Upper Kuatun (Fukien Province), 1,800 m., 1. viii. 1945, leg. Tsing-chao Maa” [Fujian, China], coll. of G. Mavromoustakis. Trachusa (Paraanthidium) xylocopiformis: Kasparek, 2017: 132, ♂ (redescription).

Material examined: China: Guangdong, Lian Xian, Dadongshan (24º59′N, 112º41′E), 1 ♂, 10.VII.1998, leg. Bai PING; Fujian, Wuyi Shan, Guadun (27º16′N, 117º38′E), 10.VIII.2018, 1 ♂, leg. Hai-Tian SONG. General distribution: China (Fujian, Guangdong*).

Trachusa (Paraanthidium) pingdaensis Niu, sp. nov. (Fig. 9: A–H, female)

Diagnosis: This new species is only known from the female; the mandible is five-toothed, the second recurrent vein ends distal to second submarginal crossvein, the subantennal suture is slightly outwardly arcuate, and the scutellum is broadly rounded but emarginate medially, the lateral margin of axilla convex, so it belongs to the subgenus Para- anthidium. The new species is very similar to T. (P.) longicornis, but it can be distinguished from T. (P.) longicornis by the following characters: Clypeus without a shining longitudinal slightly elevated middle line, with fine and dense punctures (Fig. 9C); pronotal lobe small and without obvious sharp lamella directed upwards, distinct medial bump of similar height to carina (Fig. 9D); apical half disc of T4 yellow, not black (Fig. 9F). Description: Female, BL=9.0–10.0 mm (Fig. 9A); head broader than long, HW: HL=79: 72 (Fig. 9B); gena narrower than eye, GW: EW=12: 18; MtW: MsW=95: 83 (Fig. 9D, Fig. 9E); ocelloccipital distance longer than ocellocular distance, OCD: OOD=15: 9. Clypeus broader than long, concave with apical margin slightly emarginate in middle, without a shining longitudinal slightly elevated middle line starting from basal to middle of clypeus, punctures fine and dense (Fig. 9C); subantennal suture slightly arcuate outwardly (Fig. 9C); mandible dull, minutely roughened and bearing very short hairs, broadened apically, with five black teeth; vertex polished and shining, with sparse and rounded punctures (Fig. 9B); omaulus distinctly angled but lacking carina, anterior surface weakly concave if at all; pronotal lobe lamellate, but relatively small, without obvious sharp lamella directed upwards (Fig. 9D); scutum polished and shining, with large and dense rounded punctures (Fig. 9D); scutellum broadly rounded, emarginate medially, overhanging metanotum, with dense and rounded punctures (Fig. 9D); lateral margin of axilla convex, with punctures similar to those of scutellum (Fig. 9D); disc of T1–T3 with dense and rounded punctures

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 261 (Fig. 9E); vein cu-v of hind wing oblique, nearly half as long as second abscissa of M+Cu (Fig. 9H), second recurrent vein of fore wing ends distal to second submarginal crossvein (Fig. 9G); arolia present, claw with inner median tooth (Fig. 9F). T1–T3 black, basal half of disc of T4 black, apical half of disc of T4 yellow, marginal zone of T4 yellowish- brown, T5–T6 yellow (Fig. 9F); scopa bright yellow (Fig. 9A); wings fuscated throughout (Fig. 9G, Fig. 9H). Male, unknown. Type material: Holotype, ♀, China, Yunnan, Longling Xian, Pingda (24º21′N, 98º56′E), 1600 m, 19.V.1955, leg. Bing-Rong OU (IZCAS); Paratypes: 3 ♀, same label information as the holotype (IZCAS); paratype, 1 ♀, Fukien, S. China, Changting, Tsingshanpu (25º48′N, 116º18′E), 30.VI.1940, leg. T. Maa (Nicosia). General distribution: China (Yunnan*). Etymology: The type location Pingda (Yunnan, China) is given as the specific name. Remarks: Wu (2006) assigned the four specimens of the new species to T. (P.) longicornis.

Trachusa (Paraanthidium) staabi Niu, sp. nov. (Fig. 10: A–H, female)

Diagnosis: This new species is only known from the female; it belongs to subgenus Paraanthidium, for the mandible is five-toothed, the second recurrent vein ends distal to second submarginal crossvein, the subantennal suture is slightly outwardly arcuate, and the scutellum is broadly rounded but emarginate medially. The new species is very similar to T. ovata, but in T. ovata the basal half of each wing is strongly fuscate and, the outer half is milky white, the scutum is densely covered with reddish-brown erect pubescence; the new species wings fuscate throughtout, the scutum polished and shining, with large and dense rounded punctures. Description: Female, BL=13.5 mm (Fig. 10A); head broader than long, HW: HL=105: 95 (Fig. 10B); gena slightly narrower than eye, GW: EW=24: 28; MtW: MsW=134: 130 (Fig. 10E, Fig. 10F); ocelloccipital distance longer than ocellocular distance, OCD: OOD=26: 20. Clypeus broader than long, concave with apical margin slightly emarginate in middle, with a shining longitudinal slightly elevated middle line starting from basal to middle of clypeus, punctation irregularly and craggy (Fig. 10C); subantennal suture slightly arcuate outwardly (Fig. 10C); mandible dull, minutely roughened and bearing very short hairs, broadened apically, with five black teeth (Fig. 10, D); vertex polished and shining, with sparse and fine punctures (Fig. 10B); omaular carina present, extending down to middle of mesepisternum (Fig. 10A); pronotal lobe large, lamellate (Fig. 10E); scutum polished and shining, with large and dense rounded punctures (Fig. 10E); scutellum broadly rounded, emarginate medially, overhanging metanotum, punctation irregular and wrinkled (Fig. 10E); lateral margin of axilla convex (Fig. 10E); disc of T1–T2 with sparse and rounded punctures, posterior marginal zone of T1–T2 with dense and shallow small punctures (Fig. 10F), vein cu-v of hind wing oblique, nearly as long as second abscissa of M+Cu, second recurrent vein ends distal to second submarginal crossvein (Fig. 10H); arolia present, claw with inner median tooth (Fig. 10A, Fig. 10G). Body black, without yellow markings, only scutellum and basal middle of T2 dull reddish black (Fig. 10E, Fig. 10F); T5 with sparse fulvous hairs, T6 with dense fulvous hairs (Fig. 10G); scopa fulvous (Fig. 10A); wings fuscate throughout (Fig. 10H). Male, unknown. Type material: Holotype, ♀, China, Zhejiang, Gutianshan (29º14′N, 118º07′E), 1542 m, 4.VI.2013, leg. Mi- chael STAAB (IZCAS). General distribution: China (Zhejiang*, Fujian*). Etymology: The specific name is dedicated to Dr. Michael Staab for his collection of the specimens.

Trachusa (Paraanthidium) wuae Niu, sp. nov. (Fig. 11: A–F, male)

Diagnosis: This new species is only known from the male, and it is very similar to T. muiri and T. rufobalteata. Wu et al. (1988) and Wu (2006) misidentified it as T. muiri. Kasparek (2017) reexamined the allotype (male) of T. muiri in the collection of Mavromoustakis, redescribed the male and provided images. The main differences between the new species and T. muiri are as follows: T2 with a yellow linear stripe on each side (Fig. 11D); the genitalia with

262 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. FIGURE 6. Trachusa (Paraanthidium) muiri (Mavromoustakis, 1937), A–E, female. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Metasoma in dorsal view; E: Fore wing in frontal view. Scale bars: 1.0 mm.

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 263 FIGURE 7. Trachusa (Paraanthidium) muiri (Mavromoustakis, 1937), A–H, male. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Metasoma in dorsal view, showing the T1–T4; E: Metasoma in dorsal view, showing the T5–T7; F: Genitalia in dorsal view; G: S7 in ventral view; H: S8 in ventral view. Scale bars: A–F, 1.0 mm; G–H: 0.5mm.

264 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. FIGURE 8. Trachusa (Paraanthidium) xylocopiformis (Mavromoustakis, 1954), A–H, Male. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Metasoma in dorsal view, showing T1–T3; E: Metasoma in dorsal view, showing T4–T6; F: Metasoma in dorsal view, showing T7; G: Metasoma in ventral view; H: Fore wing in frontal view. Scale bars: 1.0 mm.

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 265 FIGURE 9. Trachusa (Paraanthidium) pingdaensis Niu, sp. nov., A–H, female. A: Body in lateral view; B: Head in frontal view; C: Supraclypeal area in frontal view, showing subantennal suture; D: Mesosoma in dorsal view; E: Metasoma in dorsal view, showing T1–T3; F: Metasoma in dorsal view, showing T4–T6; G: Fore wing in frontal view, showing the position of the second recurrent vein; H: Hind wing in frontal view, showing vein cu–v and f M+Cu. Scale bars: 1.0 mm.

266 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. FIGURE 10. Trachusa (Paraanthidium) staabi Niu, sp. nov., A–H, female. A: Body in lateral view; B: Head in frontal view; C: Supraclypeal area in frontal view, showing subantennal suture; D: Mandible in frontal view, showing teeth; E: Mesosoma in dorsal view; F: Metasoma in dorsal view, showing T1–T3; G: Metasoma in dorsal view, showing T4–T6; H: Fore wing in frontal view, showing the position of the second recurrent vein. Scale bars: 1.0 mm.

OVERVIEW OF THE BEE GENUS TRACHUSA Zootaxa 4646 (2) © 2019 Magnolia Press · 267 large, forked gonostylus with two equally strong arms (Fig. 11F-c). This new species can be distinguished from T. rufobalteata by the lower half of its supraclypeus being yellow (Fig. 11B), scutellum slightly emarginate medially (Fig. 11C), whereas the latter species has an immaculate supraclypeus, scutellum deeply emarginate medially.

FIGURE 11. Trachusa (Paraanthidium) wuae Niu, sp. nov., A–F, male. A: Body in lateral view; B: Head in frontal view; C: Mesosoma in dorsal view; D: Metasoma in dorsal view; E: Metasoma in ventral view, showing combs on S5; F: Drawing of S7 (a), S8 (b), and genitalia (c), source: Wu (2006). Scale bars: 1.0 mm.

Description: Male, BL=8.5 mm (Fig. 11A); head broader than long, HW: HL=66: 58 (Fig. 11B); gena slightly narrower than eye, GW: EW=12: 15; MtW: MsW=174: 180 (Fig. 11C, Fig. 11D); ocelloccipital distance as long as ocellocular distance, OCD: OOD=10: 10 (Fig. 11B). Antenna long, reaching the apex of the scutellum, second to tenth flagellomere nearly equal in length, eleventh one slightly longer (Fig. 11A). Clypeus broader than long, apical margin slightly emarginate in middle, with fine and sparse punctures (Fig. 11B); subantennal suture obviously arcuate outwardly (Fig. 11B); mandible with three black teeth (Fig. 11B); vertex dull, with dense and fine punctures (Fig. 11B); omaular carina present, extending down to middle of mesepisternum; pronotal carina weak, shorter in height than medial bump of pronotum (Fig. 11C); scutum shining, with dense rounded punctures (Fig. 11C); scutellum shining, broadly rounded and slightly emarginate medially, nearly straight along rim, with dense rounded punctures

268 · Zootaxa 4646 (2) © 2019 Magnolia Press NIU ET AL. (Fig. 11C); propodeal enclosure with band of punctures at top interrupted or at least sparser medially; disc of T1–T3 with dense rounded punctures (Fig. 11D). Head with strong, lemon yellow maculations present on clypeus except for a narrow, brown and translucent apical margin; mandible except the black apex; lower paraocular area entirely up to antennal sockets; and lower half of supraclypeal area (Fig. 11B); scape black (Fig. 11B); scutum, scutellum, axilla, and pronotal lobe black (Fig. 11C); tegula largely black, but center dull reddish black (Fig. 11C); T1 black, T2 largely black but with a lemon yellow linear stripe on each side, T3–T4 largely black but with a transverse and lemon yellow discal stripe widely notched at each side of the center (Fig. 11A, Fig. 11D), T5 yellow but with a black marking on each side (Fig. 11A); underside of S4–S5 with black combs (Fig. 11E); S7 (a), S8 (b) as shown in Fig. 11F-a, Fig. 11F-b respectively; genitalia with large, forked gonostylus with two equally strong arms (Fig. 11F-c). Female, unknown. Type material: Holotype, ♂, China, Yunnan, Cheli (23º06′N, 102º10′E), 560 m, 26.IV.1957, leg. Da-Hua LIU (IZCAS). General distribution: China (Yunnan*). Etymology: The specific name is dedicated to Professor Yan-Ru Wu of the Institute of Zoology, Chinese Acad- emy of Sciences for her contributions to bee taxonomy.

Acknowledgements

The authors would like to express their sincere thanks to Prof. Yan-Ru Wu, Dr. Max Kasparek, Dr. Maxim Pros- chalykin, Dr. Yan-Zhou Zhang, and Dr. Jason Gibbs for their comments and suggestions to streamline and improve the earlier drafts of the manuscript. We thank staff at the Mavromoustakis Collection, Ministry of Agriculture, Nico- sia, CYPRUS; the Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, CANADA; and the Hungarian Natural History Museum, Budapest, HUNGARY, for access to specimens of Trachusa. This work was supported mainly by the grant from the National Science Foundation, China (31625024, 31772487) and the grant (Y229YX5105) from Key Laboratory of Zoological Systematics and Evolution, Chi- nese Academy of Sciences. Michael Orr has been supported by PIFI program from Chinese Academy of Sciences (2018PB0003). Chao-Dong ZHU and John S. Ascher acknowledge supports Singapore-China Joint Research Grant from NSFC (41761144068). Study of museum material by TG was supported by funding from USDA-APHIS under cooperative agreements 16-8130-0736-CA and AP17PPQS&T00C053 and US National Science Foundation grant DEB-0742998.

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