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 sichelii Radoszkowski, 1860 Figs 15 , 129–138 , 197 , 211 Bombus SICHELII Radoszkowski, 1860: 481 . Bombus alticola Kriechbaumer, 1873: 339 . Bombus lapidarius subsp. nigritulus Friese, 1905: 518 . Bombus lapidarius subsp. albidulus Friese, 1905: 518 . Bombus sicheli Form [subsp.] uniens Vogt, 1909: 62 . Bombus noster Skorikov, 1910b: 578 . Bombus sicheli var. geogr. [subsp.] czerskiae Vogt, 1911: 59 . Bombus sicheli var. geogr. [subsp.] rehbinderianus Vogt, 1911: 60 . Bombus sicheli var. geogr. [subsp.] cazurroi Vogt, 1911: 60 . Lapidariobombus verticiflavus Skorikov, 1931: 225 . Bombus sicheli f.g. [subsp.] daghestanicus Reinig, 1935: 345 (non Radoszkowski, 1877 : vii = B. sylvarum (Linnaeus)) . Bombus jeholensis Yasumatsu, 1935: 45 . Bombus sicheli subsp. chinganicus Reinig, 1936: 6 . Pyrobombus sicheli subsp. flavissimus Tkalců, 1975: 177 , replacement name for latofasciatus Vogt, 1909: 49 . Pyrobombus erzurumensis Özbek, 1990: 209 . Bombus Sicheli – Radoszkowski 1877: 213 , incorrect subsequent spelling. Since Vogt (1909) , B. sichelii s. lat. has generally been considered as a species separate from B. lapidarius s. lat. Vogt (1909: 46) stated that his “Formen” are subpopulations, which could be considered equivalent to subspecies. Reinig (1935: 333) stated explicitly that his “f.g.” (forma geographica) are equivalent to subspecies. These taxa are considered here to be parts of a sichelii -complex. The taxon erzurumensis was described more recently as a separate species from north-eastern Turkey , although we have seen it also from Iran . Within the sichelii -complex it has an especially narrow band of black hair posteriorly on T2. It was first recognised as likely to be conspecific with B. sichelii s. lat. by Williams (1998) . Our PTP analysis ( Fig. 10 ) supports two coalescents in the COI gene, for a west Asian B . incertus , which is strongly supported, and a widespread Eurasian B. sichelii s. lat. Although the support for a COI coalescent for B. sichelii s. lat. is relatively weak, it is the best supported in this part of the tree, so no alternative interpretation of the member taxa is better supported as species. Corroboration from a positive divergence-with-distance relationship both within ( Fig. 17 ) and among ( Fig. 18 ) the component lineages (Fig. 9) is accepted as additional support for B. sichelii s. lat. as a single broadly distributed species in its long accepted sense. From morphology, B. sichelii s. lat. has a colour pattern of the hair of T2 with black hairs posteriorly that is consistently distinct from B . incertus . Diagnosis Females Queens medium-sized body length 17–20 mm , workers 9–13 mm , Can be distinguished by usually having at least some pale hair on the face at the base of the antenna (cf. B. separandus , B. incertus ), T2 posteriorly usually has at least a narrow line of black hair (cf. B. keriensis , B. separandus , B. alagesianus , B. incertus etc.), and the hindleg corbicular fringes have some extensively orange hairs. Some individuals from eastern Turkey and Iran (the taxon erzurumensis ) resemble B. incertus closely because they have very few black hairs posteriorly on T2 and few or no pale hair on the face, but the queens of B. sichelii are smaller than B. incertus , the red of T4–5 is more orange-red, and the hindleg corbicular fringes have some extensively orange hairs. Males Body length 11–15 mm . Can be distinguished in Europe and West Asia by their combination of yellow or white hair posteriorly on the thoracic dorsum, T2 with yellow or white but posteriorly with some black, and T4–7 a pale red. Genitalia ( Fig. 197 ) with the gonostylus as long as broad, reduced as a rounded flat scale with the inner basal process reduced to a tooth (cf. rufipes- group, festivus- group, rufofasciatus -group); volsella with the inner distal corner broadly produced but without a narrow hook (cf. rufipes- group, festivus- group, rufofasciatus -group); eye unenlarged relative to female eye. Material examined Lectotype RUSSIA • ♀ (queen), lectotype of Bombus sichelii Radoszkowski, 1860 by designation of Tkalců (1974b) ; Kamchatka ; Dittmar leg.; ZMHB (examined PW ). Material sequenced ( 46 specimens , includes some sequences from Lecocq et al. 2015 ) SPAIN • 1 ♂; Catalonia , Cerbi ; 42.646° N , 1.152° E ; 16 Sep. 1993 ; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM02963; ZSM : ML327 • 1 ♂; same collection data as for preceding; BOLD seq: BCZSMHYM02964; ZSM : ML328 . SWITZERLAND • 1 ♀ (worker); 46.2210° N , 7.6203° E ; 22 Jul. 1999 ; P. Williams leg.; BOLD seq: 1555H11; PW : ML313 • 1 ♀ (worker); Ticino , Lago Sambuco ; 46.466° N , 8.648° E ; 28 Jul. 1993 ; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM02961; ZSM : ML326 . AUSTRIA • 1 spec. ; M̹hl; 47.5014° N , 10.7361° E ; D. Michez leg.; GenBank seq: KM458067 ; UMONS : ML285 . FRANCE • 1 spec. ; Col d’Allos; 44.1733° N , 6.3544° E ; O. Poncheau leg.; GenBank seq: KM458070 ; UMONS : ML288 • 1 spec. ; Gavarnie; 42.7119° N , 0.0075° W ; D. Michez leg.; GenBank seq: KM458072 ; UMONS : ML289 . RUSSIA • 1 ♀ (worker); Chuvashiya , Severniy ; 56.2899° N , 47.1931° E ; 24 Jun. 2010 ; A. Lashtukhin leg.; BOLD seq: SHMEL-E11; SDM ML331 • 1 spec. ; Buryatia , Mondy ; 51.4103° N , 100.5936° E ; T. DeMeulemeester leg.; GenBank seq: KC915894 ; UMONS : ML290 • 1 spec. ; same collection data as for preceding; GenBank seq: KC915896 ; UMONS : ML291 • 1 ♀ (worker); Tuva , 31 km Erzin ; 50.0366° N , 95.4348° E ; 18 Jul. 2014 ; A. Lelej leg.; BOLD seq: 1555F11; NSUN : ML268 • 1 ♀ (worker); Sakhalin , south Schmidta Peninsula ; 53.8874° N , 142.7429° E ; 13 Aug. 2003 ; J. Jobe leg.; BOLD seq: 1552F06; SEMC : ML189 • 1 ♀ (queen); Kamchatka ; 53.7887° N , 159.3172° E ; BOLD seq: 6879D08; FSCV : ML217 . TURKEY • 1 ♀ (worker); 41.2008° N , 42.5283° E ; BOLD seq: 6879A04; PW : ML188 • 1 spec. ; Zigana Geçidi ; 40.6353° N , 39.4044° E ; P. Rasmont leg.; GenBank seq: KM458068 ; UMONS : ML286 • 1 spec. ; same collection data as for preceding; GenBank seq: KM458069 ; UMONS : ML287 . IRAN • 1 ♀ (queen); 38.2974° N , 48.0360° E ; 26 May 2006 ; NHMUK seq: IRANPW022; PW : ML15 • 1 ♀ (worker); 38.2724° N , 47.6886° E ; 29 May 2015 ; NHMUK seq: IRANPW020; PW : ML16 • 1 ♀ (worker); 38.2717° N , 47.8515° E ; 31 May 2014 ; NHMUK seq: IRANPW023; PW : ML17 . KYRGYZSTAN • 1 ♀ (queen); Santash ; 42.7389° N , 79° E ; 28 Jun. 2016 ; R. DeJonghe leg.; BOLD seq: 1555F03; RDJ : ML260 . MONGOLIA • 1 ♀ (worker); Ovorhangay , Karakorum ; 47.1667° N , 102.8333° E ; 13–16 Aug. 1990 ; C. Cockell leg.; BOLD seq: 6874D04; PW : ML18 • 1 ♀ (worker); Hovsgol , Tsagaan Nur ; 50.601° N , 101.523° E ; 16 Jul. 2005 ; J. Gelhaus leg.; BOLD seq: 1550G02; SEMC : ML215 • 1 ♀ (worker); Dalbay ; 51.025° N , 100.766° E ; 20 Jul. 2007 ; D. Song leg.; BOLD seq: 9808C12; PCYU : ML332 • 1 ♀ (worker); Dalbay ; 51.0279° N , 100.7667° E ; 4 Aug. 2007 ; D. Song leg.; BOLD seq: 9808E01; PW : ML218 • 1 ♀ (worker); Khovsgol Nuur ; 51.0628° N , 100.7322° E ; 21 Jul. 2004 ; D. Sheppard leg.; NHMUK seq: PW28 ; PW : ML324 . CHINA – Neimenggu Province • 1 ♀ (worker); Moerdaga ; 51.3349° N , 120.6784° E ; 18 Aug. 2009 ; P. Williams leg.; NHMUK seq: CT465; PW : ML465 • 1 ♀ (worker); nr. Huanggangliang ; 43.5423° N , 117.6233° E ; 24 Aug. 2010 ; P. Williams leg.; BOLD seq: 1555B12; PW : ML231 . – Shanxi Province • 1 ♀ (worker); Yingxian ; 39.4008° N , 113.4167° E ; 8 Sep. 2007 ; P. Williams leg.; BOLD seq: 1555C01; PW : ML232 . – Gansu Province • 1 ♀ (worker); Pinnan ; 34.3281° N , 105.6278° E ; 30 Aug. 2009 ; P. Williams leg.; BOLD seq: 1555H01; PW : ML303 . – Sichuan Province • 1 ♀ (worker); Hongyuan ; 32.6403° N , 102.3286° E ; 4 Aug. 2002 ; P. Williams leg.; BOLD seq: 1555H02; PW : ML304 • 16 specs; Hongyuan ; 32.3282° N , 102.4543° E ; Y. Dong leg.; YD seq: DYX53.1, DYX53.2, DYY17, DYX61.1, DYX1.1, DYX18.1, DYX30.2, DYX34.1, DYX11.2, DYZX2, DYX64, DYX1.3, DYX13.1, DYX27.1, DYX13.2, DYX69.1; YD : ML443 to ML458 . Global distribution (Widespread Palaearctic species, in steppes in the north and in mountains in the south) Europe : SPAIN , FRANCE , SWITZERLAND , AUSTRIA . – West Asia : TURKEY , IRAN . – North Asia : KYRGYZSTAN , MONGOLIA , RUSSIA : European Russia, North Caucasus, Ural, West Siberia, Altai Mountains, East Siberia, Far East. – East Asia : CHINA : Xizang , Sichuan , Gansu , Ningxia , Shanxi , Hebei , Neimenggu , Liaoning , Jilin , Heilongjiang ; N KOREA , S. KOREA . (FSCV, IAR, IOZ, NHMUK, PW, SC, SEMC, YUY, ZMHB.) The species is widespread ( Fig. 210 ) and often abundant. Behaviour Food-plant generalists ( Rasmont 1988 ; Neumayer & Paulus 1999 ; Williams et al. 2009 ; An et al. 2014 ). The male mate-searching behaviour is expected to resemble the patrolling of B. lapidarius .