Widespread polytypic species or complexes of local species? Revising bumblebees of the subgenus Melanobombus world-wide (Hymenoptera, Apidae, Bombus) Author Williams, Paul H. 38A45E0C-02A8-407E-8E89-5162D454E9FE Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. paw@nhm.ac.uk Author Altanchimeg, Dorjsuren FB68F32E-4F6D-40C2-A921-20FBAD676D50 Institute of General and Experimental Biology, Peace Avenue 54 b, Ulaanbaatar 13330, Mongolia. altanchimegd@mas.ac.mn Author Byvaltsev, Alexandr B57BAD3E-9E42-4446-994E-4A45A738D404 Novosibirsk State University, ul. Pirogova 2, Novosibirsk, 630090 Russia. byvam@yandex.ru Author Jonghe, Roland De FC98CAB7-B2FF-4BEB-94FF-26F53D33CD04 Langstraat 105, B- 2260 Westerlo, Belgium. roland.de.jonghe@telenet.be Author Jaffar, Saleem 77F70375-0A19-4D0E-A05A-987BB46543C0 South China Agricultural University, Guangzhou 510642, China. saleemjaffar@stu.scau.edu.cn Author Japoshvili, George CCC82B7C-A1E4-4D58-90A3-623116CBAE96 Agricultural University of Georgia, 240 Agmashenebli Alley, Tbilisi, Georgia. g.japoshvili@agruni.edu.ge Author Kahono, Sih F8513496-B409-434C-A182-4146232C89FA Indonesian Institute of Sciences (LIPI), Jakarta, Indonesia. sihkahono@gmail.com Author Liang, Huan A99867E0-C686-4608-8DF7-0EDE8D2D57EC Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China. lianghuan@mail.kib.ac.cn Author Mei, Maurizio 82F344C7-B98A-462C-81E0-D6F3F02348D4 Università di Roma ‘ Sapienza’, Piazzale Valerio Massimo 6, Roma 00162, Italy. maurizio.mei@uniroma1.it Author Monfared, Alireza 48CA77BA-8CF4-4812-89B1-696A11FEDE2D Yasouj University, Zirtol, Yasouj, Iran. alirezamonfared1@yahoo.com Author Nidup, Tshering BE588EE1-5E2C-46CC-8907-CD344D88C869 Sherubtse College, Royal University of Bhutan, Trashigang, Bhutan. tsheringnidup@sherubtse.edu.bt Author Raina, Rifat 48E5AE7A-D5DC-4549-94B7-FD8489D1EF9E Zoological Survey of India, Pali Road, Jodhpur 342005, Rajasthan, India. rifat72001@rediffmail.com Author Ren, Zongxin 27B9DD39-62A8-44D3-9D6A-E6C20D8AAA27 Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China. renzongxin@mail.kib.ac.cn Author Thanoosing, Chawatat 6F4C150C-BC03-4F75-91A6-2A8AF6B5905C Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. c.thanoosing@nhm.ac.uk Author Zhao, Yanhui 299C8EEA-699E-4B15-9BCD-9806E0E7EE63 Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China. zhaoyanhui@mail.kib.ac.cn Author Orr, Michael C. 1E7F46C3-870E-460C-A611-BA1042ED99FB Institute of Zoology (Chinese Academy of Sciences), 1 Beichen West Road, Chaoyang, Beijing 100101, China. michael.christopher.orr@gmail.com text European Journal of Taxonomy 2020 2020-10-02 719 1 120 journal article 10.5852/ejt.2020.719.1107 7ca72f76-4fae-4305-8601-4662f4cd2b96 2118-9773 4064324 A4500016-C219-4353-B81C-5E0BB520547F Bombus prshewalskyi Morawitz, 1880 stat. rev. Figs 3 , 13 , 64–70 , 187 Bombus Prshewalskyi Morawitz, 1880: 342 . Bombus rufocinctus Morawitz, 1880: 343 (non Cresson, 1863: 106 = B. rufocinctus Cresson ). syn. nov . Bombus chinensis von Dalla Torre, 1890 [Jun. 25]: 139 (non Morawitz, 1890 [April 30]: 352), replacement name for rufocinctus Morawitz, 1880: 343 . syn. nov . Bombus prshewalskii – von Dalla Torre 1896: 548 , incorrect subsequent spelling. Bombus przewalskii – Friese 1918: 84, incorrect subsequent spelling. Kozlovibombus przewilskii – Skorikov 1933a: 245 , incorrect subsequent spelling. Bombus rufofasciatus (part) – Williams 1991: 105 ; 1998: 133 (non Smith, 1852a: 48 ). Bombus prshewalskyi was described by Morawitz (1880) without reference to Smith’s (1852a) closely related taxon rufofasciatus , apparently without knowledge of it. Bombus prshewalskyi is listed as a species (as Kozlovibombus przewalskii ) separate from B. rufofasciatus by Skorikov (1923) , although the latter was listed by him as “ Bombi incertae sedis ”. The two were synonymised by Richards (1930) . Here, B. prshewalskyi is revised as an eastern species separate from the western B. rufofasciatus s. str. because of their unique and strongly divergent species coalescents in the COI gene ( Fig. 10 ), corroborated by morphology. These morphological differences between the two are subtle, but do appear to support that the two taxa are separate species within a morphologically more distinctive complex of rufofasciatus s. str. + prshewalskyi . Our PTP analysis ( Fig. 10 ) supports relatively strongly two coalescents in the COI gene for the west Himalayan B . rufofasciatus s. str. and B . prshewalskyi of the eastern QTP. The two coalescent groups differ in COI barcode sequences for at least 22 diagnostic nucleotide positions (3.3% of the barcode region, although some sequences are incomplete). These differences are all synonymous, making no difference to the amino acid sequences at translation. In contrast, examination of sequences for the slower-evolving PEPCK gene (although it also includes faster-evolving intron regions) shows no diagnostic nucleotide changes. From morphology, B . prshewalskyi has a slightly darker colour pattern of the hair than B. rufofasciatus s. str. for the females (larger workers and queens), with the white bands appearing dark greyish because of many black hairs intermixed, especially on the scutellum, and with black hair more extensive on the body generally. T2 has yellow hair conspicuously replaced by black posteriorly. Current restrictions on collecting and sequencing in the areas where the two candidate species are most likely to occur in proximity ( Fig. 13 ) make clarifying their status difficult. However, more evidence is needed to increase confidence in any interpretation of the status of the candidate species rufofasciatus s. str. and prshewalskyi and it remains possible that with more sampling from the Himalaya the two taxa could still prove to be parts of a single broader species, B. rufofasciatus s. lat. Speciation between B. rufofasciatus s. str. and B. prshewalskyi may have arisen through vicariance between populations of the western and eastern Himalaya caused by a period of altered climate (cf. B. miniatus / B. eurythorax ). Females show size-dependent dimorphism in the colour pattern of the hair: large queens have T2 predominantly black and T4 white; whereas the workers (which are smaller) have T2 extensively yellow and T4 red. Males have a similar colour pattern to the workers. Diagnosis Females ( Fig. 3 ) Queens medium-sized body length 19–23 mm , workers 10–15 mm . Can be distinguished by their combination of the hair of T5 white and the hair on the side of the thorax white only in the dorsal half (cf. B. ladakhensis ), closely similar to B. rufofasciatus but tending to have T2 with less yellow hair and the scutellum with more black hair. Males Body length 13–17 mm . Can be distinguished reliably at present only by their COI sequence, but tending to have T2 with less yellow hair and the scutellum with more black hair (cf. B. rufofasciatus ). Genitalia ( Fig. 187 ) with the gonostylus much reduced, less than a quarter as long on its outer side as broad, with the distal edge concave and the inner distal corner with two almost equally pronounced adjacent acute teeth (cf. other rufofasciatus -group species with the exceptions of some B. friseanus , B. pyrosoma ); volsella with the inner distal corner produced as a narrow curved hook ( cf. rufipes -group, B. simillimus , lapidarius -group, sichelii -group, keriensis -group); antenna short and eye enlarged relative to female eye. Material examined Syntypes CHINA • 1 ♀ (worker, not a queen), 1 ♂ ( Williams 1991:10 ), syntypes of Bombus prshewalskyi Morawitz, 1880 ; “Gan-su” [= Gansu in a much broader sense than currently: the specimen is probably from modern Qinghai ]; N. Przhevalsky leg.; ZIN (not seen but identity not in doubt). Material sequenced ( 57 specimens ) CHINA – Gansu Province • 1 ♀ (worker); Gahai ; 34.1743° N , 102.5577° E ; 28 Aug. 2009 ; P. Williams leg.; BOLD seq: 6876H09; PW : ML1 • 1 ♂; Hezuo ; 34.9027° N , 102.8445° E ; 27 Aug. 2009 ; P. Williams leg.; BOLD seq: 1555E03; PW : ML251 . – Sichuan Province • 4 specs; Hongyuan ; 32.3282° N , 102.4543° E ; 2017; YD seq: DYX9.1, DYX56.2, DYX22.2, DYX32.2; YD : ML424 to ML427 • 1 ♀ (worker); Hailuogou ; 29.89546° N , 102.01324° E ; 7 Aug. 2018 ; Z. Ren leg.; KIB seq: GGSM102001; KIB : ML555 • 1 ♀ (worker); Que’er shan; 31.89688° N , 99.14853° E ; 4Aug. 2018 ; Z. Ren leg.; KIB seq: QESM202003; KIB : ML560 . – Yunnan Province • 1 ♀ (worker); Lijiang ; 27.0156° N , 100.1714° E ; 27 Aug. 2012 ; Y. Zhao leg.; KIB seq: 402646105; KIB : ML366 • 1 ♂; same collection data as for preceding; 21 Sep. 2012 ; Y. Zhao leg.; KIB seq: 402648064; KIB : ML367 • 1 ♀ (worker); Baima Snow Mountain ; 28.34827° N , 99.05771° E ; 17 Jul. 2018 ; Z. Ren leg.; KIB seq: BMXSM102034; KIB : ML548 • 1 ♀ (worker); Baima Snow Mountain ; 28.37252° N , 98.99982° E ; 16 Jul. 2018 ; Z. Ren leg.; KIB seq: BMXSH101052; KIB : ML549 • 26 ♀♀ , 8 ♂♂; Baima Snow Mountain ; 28.3373° N , 99.0771° E ; 14–17 Aug. 2019 ; M. Orr leg.; IOZ seq: 14F2 to 14F7, 14F9, 14F11 to 14F16, 14F18, 14M1 to 14M5, 16F1, 16M1, 16M2, 16M4, 17F1 to 17F4, 17M1, OR4, OR6 to OR8, OR14, OR16, OR19, OR21, OR25; IOZ : ML599 to ML632 . – Xizang Province • 1 ♀ (worker); Anjiulashan ; 29.6509° N , 96.71712° E ; 23 Jul. 2018 ; Z. Ren leg.; KIB seq: AJLSM103002; KIB : ML550 • 1 ♂; Galonglashan ; 29.78346° N , 95.69765° E ; 24 Jul. 2018 ; Z. Ren leg.; KIB seq: GLLSM201092; KIB : ML551 • 1 ♂; same collection data as for preceding; KIB seq: GLLSM201104; KIB : ML552 • 1 ♀ (queen); same collection data as for preceding; KIB seq: GLLSM101006; KIB ML553 • 1 ♀ (queen); Milashan ; 29.85301° N , 92.33378° E ; 27 Jul. 2018 ; Z. Ren leg.; KIB seq: MLSH101029; KIB : ML554 • 1 ♀ (worker); Honglashan ; 29.92411° N , 92.77629° E ; 19 Jul. 2018 ; Z. Ren leg.; KIB seq: HLSM101103; KIB : ML556 • 1 ♀ (worker); Dangxiong County ; 30.21253° N , 90.62821° E ; 30 Jul. 2018 ; Z. Ren leg.; KIB seq: DXH101006; KIB : ML557 • 1 ♀ (worker); Zhujiaolashan ; 31.10878° N , 96.88469° E ; 2 Aug. 2018; Z. Ren leg.; KIB seq: ZJLSM101045; KIB : ML558 ; 1 ♀ (worker); Ailashan ; 31.61421° N , 98.48028° E ; 1 Aug. 2018 , Z. Ren leg.; KIB-ALSM102002 ( KIB ML559 ) • 1 ♀ (worker); Milashan ; 29.8530° N , 92.3338° E ; 27 Jul. 2018 ; Z. Ren leg.; KIB seq: MLSH109001; KIB : ML578 • 1 ♀ (worker); Galongashan ; 29.7669° N , 95.6991° E ; 24 Jul. 2018 ; Z. Ren leg.; KIB seq: GLLSM104005; KIB : ML579 . Global distribution (Qinghai-Tibetan-Plateau and East Himalayan species) Himalaya : INDIA: Sikkim, Arunachal Pradesh. East Asia : CHINA: Xizang , Qinghai , Gansu , Sichuan , Yunnan . – Southeast Asia : BURMA. (AMNH, IAR, IOZ, KIB, KIZ, MNHN, NHMUK, NME, NMS, PW, SEHU, UAP, YD, ZIN.) The species is widely distributed within its range and often abundant. Behaviour A colony of this species has been found underground in a meadow at 3900 m a.s.l. in Yunnan (ZR unpublished). Food-plant generalists ( Williams et al. 2009 ; An et al. 2014 ). Male mate-searching behaviour appears to resemble the true territoriality of B. rufofasciatus with collisions between males in flight and replacement in perch occupancy (PW pers. obs.).