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 friseanus Skorikov, 1933 Figs 13 , 83–90 , 190 Bombus friseanus Skorikov, 1933b: 62 . Lapidariobombus pyrrhosoma subsp. ĥnei Bischoff, 1936: 10 . Bombus pyrosoma (part) – Williams 1991: 101 (non Morawitz, 1890: 349 ). Our PTP analysis ( Fig. 10 ) of coalescents in the COI gene within the rufofasciatus -group supports nine species including B. friseanus , corroborated by differences in morphology (see the Diagnosis). Females show size-dependent dimorphism in the colour pattern of the hair: large queens have T2 predominantly black; whereas workers (which are smaller) have T2 yellow or light chocolate brown. Males have the thoracic dorsum and T1–2 extensively yellow and T3–5 extensively orange-red. This species is widespread and abundant, but despite variation in COI barcodes shows little geographical variation in colour pattern. Individuals from near Qianggang in Xizang ( 29.998° N , 91.939° E ) have less black hair. One worker (ML5) from Maoshan in Sichuan ( 31.685° N , 103.899° E ) with white bands has been confirmed as this species from a COI sequence. The males have the eye unenlarged relative to the female eye ( Williams et al . 2009 ). Diagnosis Females Queens medium-sized body length 19–22 mm , workers 10–16 mm . Can usually be distinguished in the Hengduan region by their combination of the hair of the thoracic dorsum with yellow bands, side of the thorax yellow, and T3–5 red (cf. B. tibeticus , B. qilianensis , B. miniatus , B. pyrosoma , B. tanguticus ). From other often similar co-occurring species they can be distinguished by their combination of the hind tibia with the distal hairs entirely black (cf. B. ( Pyrobombus ) lepidus Skorikov, 1912 ) with the mid basitarsus with the distal posterior corner broadly rounded (cf. B. ( Thoracobombus ) impetuosus Smith, 1871 ). Figs 205–206. Maps of all available global records with all of the specimens examined and confirmed here. 205 . Bombus richardsiellus ( Tkalců, 1968 ) . 206 . B. tanguticus Morawitz, 1887 . Relief map with hill shading for the region centred on Tibet with (grey lines) borders to national administration as in UN maps. Image created in ArcGIS using World_Shaded_Relief basemap © 2014 Esri. Males Body length 13–15 mm . Can usually be distinguished in the Hengduan region by their combination of the hair of the thoracic dorsum with yellow bands, side of the thorax yellow, and T3–7 red. Genitalia ( Fig. 190 ) 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 possible exceptions of B. prshewalskyi , 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); eye unenlarged relative to female eye. Material examined Holotype CHINA • ♀ (queen), holotype of Bombus friseanus Skorikov, 1933 by monotypy ( Williams 1991 ); Sichuan , Songpan ; [18]94; M. Berezovsky leg.; ZIN (examined PW ). Material sequenced ( 65 specimens ) CHINA – Sichuan Province • 1 ♀ (worker); Maoxian ; 31.6853° N , 103.8994° E ; 10Aug. 2002 ; P.Williams leg.; BOLD seq: 1550E09; PW : ML5 • 1 ♀ (worker); same collection data as for preceding; BOLD seq: 3262F02; PW : ML7 • 1 ♀ (worker); same collection data as for preceding; BOLD seq: 3263F02; PW : ML199 • 1 ♀ (worker); Hongyuan ; 32.6403° N , 102.3286° E ; 4 Aug. 2002 ; P. Williams leg.; BOLD seq: 3262F01; PW : ML8 • 1 ♀ (worker); Haikou ; 27.3271° N , 102.4651° E ; 19 Aug. 2005 ; P. Williams leg.; BOLD seq: 1555G07; PW : ML297 • 10 specs; Hongyuan ; 32.3282° N , 102.4543° E ; 2017; YD seq: DYX48.2, DYX16.1, DYX40.2, DYX7.2, DYX20.1, DYX6.1, DYX7.4, DYX43.1, DYX6.2, DYX7.1; YD : ML428 to ML437 • 1 ♀ (worker); Ganzi county ; 31.62916° N , 100.22675° E ; 5 Aug. 2018 ; Z. Ren leg.; KIB seq: GZM106001; KIB : ML533 • 1 ♀ (worker); Hailuogou ; 29.93515° N , 101.96387° E ; 7 Aug. 2018 ; Z. Ren leg.; KIB seq: GGSL101019; KIB : ML534 • 1 ♀ (worker); Xinduqiao ; 30.06679° N , 101.76088° E ; 7 Aug. 2018 ; Z. Ren leg.; KIB seq: XDQM101028; KIB : ML535 • 1 ♀ (worker); Daofu county ; 30.78966° N , 101.30716° E ; 5 Aug. 2018 ; Z. Ren leg.; KIB seq: DFM101018; KIB : ML537 • 1 ♀ (worker); Ganzi county ; 31.62916° N , 100.22675° E ; 5 Aug. 2018 ; Z. Ren leg.; KIB seq: GZM102003; KIB : ML540 • 1 ♀ (worker); same collection data as for preceding; KIB seq: GZM101002; KIB : ML541 . – Yunnan Province • 1 ♀ (worker); Lijiang ; 27.0069° N , 100.1751° E ; 26 Jul. 2017 ; A. Moss leg.; BOLD seq: 1555C10; PW : ML241 • 1 ♂; same collection data as for preceding; BOLD seq: 1555G06; PW : ML296 • 1 ♀ (queen); Lijiang ; 27.0156° N , 100.1714° E ; 1 Aug. 2016 ; H. Liang leg.; KIB seq: 24; KIB : ML368 • 1 ♂; same collection data as for preceding; 6 Sep. 2016 ; H. Liang leg.; KIB seq: 40; KIB : ML369 • 1 ♀ (worker); same collection data as for preceding; 1 Aug. 2016 ; H. Liang leg.; KIB seq: 52; KIB : ML370 • 1 ♀ (queen); same collection data as for preceding; 14 Jun. 2012 ; Y. Zhao leg.; KIB seq: 304432147; KIB : ML371 • 1 ♀ (worker); same collection data as for preceding; 26 May 2012 ; Y. Zhao leg.; KIB seq: 40241082; KIB : ML372 • 1 ♀ (worker); same collection data as for preceding; 2 Aug. 2016 ; H. Liang leg.; KIB seq:42; KIB ML373 • 1 ♀ (worker); same collection data as for preceding; 26 Jul. 2015 ; Z. Ren leg.; KIB seq: 2018045; KIB : ML374 • 1 ♀ (worker); same collection data as for preceding; 22 Aug. 2016 ; H. Liang leg.; KIB seq: 2018071; KIB : ML375 • 1 ♀ (worker); same collection data as for preceding; 15 Jul. 2016 ; H. Liang leg.; KIB seq: 2018038; KIB : ML376 • 1 ♀ (worker); same collection data as for preceding; 19 Jul. 2016 ; H. Liang leg.; KIB seq: 53; KIB : ML377 • 1 ♂; same collection data as for preceding; 5 Sep. 2016 ; H. Liang leg.; KIB seq: 84; KIB : ML378 • 1 ♀ (worker); same collection data as for preceding; 1 Sep. 2016 ; H. Liang leg.; KIB seq: 107; KIB : ML379 • 1 ♀ (worker); Cangshan ; 25.88991° N , 100.02974° E ; 14 Jul. 2018 ; Z. Ren leg.; KIB seq: CSL201011; KIB : ML514 • 1 ♀ (worker); Cangshan ; 25.88001° N , 100.02691° E ; 14 Jul. 2018 ; Z. Ren leg.; KIB seq: CSL301005; KIB : ML515 • 1 ♀ (worker); same collection data as for preceding; KIB seq: CSL301016; KIB : ML516 • 1 ♀ (worker); Cangshan ; 25.89461° N , 100.02125° E ; 14 Jul. 2018 ; Z. Ren leg.; KIB seq: CSL101008; KIB : ML517 • 1 ♀ (worker); Baima Snow Mountain ; 28.39838° N , 98.99286° E ; 18 Jul. 2018 ; Z. Ren leg.; KIB seq: BMXSM301021; KIB : ML518 • 1 ♀ (worker); same collection data as for preceding; KIB seq: BMXSM301049; KIB : ML519 • 6 ♀♀ ; Baima Snow Mountain ; 28.3373° N , 99.0771° E ; 14–19 Aug. 2019 ; M. Orr leg.; IOZ seq: 14F2, 14F4, 14F19, OR20, OR24, OR27; IOZ : ML584 to ML589 . – Xizang Autonomous Region • 1 ♀ (worker); Milashan ; 29.85301° N , 92.33378° E ; 27 Jul. 2018 ; Z. Ren leg.; KIB seq: MLSM101027; KIB : ML520 • 1 ♀ (worker); same collection data as for preceding; KIB seq: MLSM101002; KIB : ML521 • 1 ♀ (worker); same collection data as for preceding; KIB seq: MLSM101034; KIB : ML522 • 1 ♀ (worker); Ranwu Lake ; 29.4849° N , 96.64252° E ; 23 Jul. 2018 ; Z. Ren leg.; KIB seq: RWM101017; KIB : ML523 • 1 ♀ (worker); same collection data as for preceding; KIB seq: RWM101057; KIB : ML524 • 1 ♀ (worker); same collection data as for preceding; KIB seq: RWM101001; KIB : ML525 • 1 ♀ (worker); Dongdashan ; 29.57184° N , 98.18467° E ; 20 Jul. 2018 ; Z. Ren leg.; KIB seq: DDSM303001; KIB : ML526 • 1 ♀ (worker); Dongdashan ; 29.62259° N , 97.89246° E ; 20 Jul. 2018 ; Z. Ren leg.; KIB seq: DDSM203003; KIB : ML527 • 1 ♀ (worker); Sejilashan ; 29.64141° N , 94.69818° E ; 26 Jul. 2018 ; Z. Ren leg.; KIB seq: SJLSM101009; KIB : ML528 • 1 ♀ (worker); Dongdashan ; 29.69073° N , 98.11641° E ; 20 Jul. 2018 ; Z. Ren leg.; KIB seq: DDSH301056; KIB : ML529 • 1 ♀ (worker); Sejilashan ; 29.77619° N , 94.74278° E ; 26 Jul. 2018 ; Z. Ren leg.; KIB seq: SJLSL301011; KIB : ML530 • 1 ♀ (worker); Galonglashan ; 29.78346° N , 95.69765° E ; 24 Jul. 2018 ; Z. Ren leg.; KIB seq: GLLSM103018; KIB : ML531 • 1 ♀ (worker); Anjiulashan ; 29.9229° N , 96.67203° E ; 23 Jul. 2018 ; Z. Ren leg.; KIB seq: AJLSM101004; KIB : ML532 • 1 ♀ (queen); Yelashan ; 30.16394° N , 97.2860° E ; 22 Jul. 2018 ; Z. Ren leg.; KIB seq: YLSH101083; KIB : ML536 • 1 ♀ (worker); Zhujiaolashan ; 31.11821° N , 97.01749° E ; 2 Aug. 2018 ; Z. Ren leg.; KIB seq: ZJLSM101015; KIB : ML538 • 1 ♀ (queen); Zonglashan ; 31.2922° N , 97.59554° E ; 3 Aug. 2018 ; Z. Ren leg.; KIB seq: ZLSM201011; KIB : ML539 • 1 ♂; Galonglashan ; 29.82189° N , 95.71101° E ; 24 Jul. 2018 ; Z. Ren leg.; KIB seq: GLLSL301020; KIB : ML542 • 1 ♀ (worker); Xizang ; 29.5032° N , 95.8980° E ; 8 Aug. 2015 ; IBK seq: BB12; IOZ : ML582 . Global distribution (Hengduan mountain species) East Asia : CHINA : Xizang , Qinghai , Sichuan , Yunnan , Guizhou . (AMNH, IAR, IOZ, KIB, KIZ, NHMUK, PW, SAC, USNM, YD, ZIN, ZMHB, ZMUM.) The species is widely distributed within the Hengduan region and often one of the most abundant. Behaviour A colony of this species has been found underground in a coniferous forest at 3300 m in Yunnan (ZR unpublished). Food-plant generalists ( Williams et al. 2009 ). Male mate-searching behaviour appears to resemble patrolling of B. eurythorax (PW pers. obs.).