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 eximius Smith, 1852 Figs 1 , 12 , 25–34 , 181 Bombus eximius Smith, 1852a: 47 . Bombus latissimus Friese, 1910: 404 . Bombus rufipes var. [subsp.] tonkinensis Friese, 1914: 11 . Bombus latissimus var. [subsp.] detritus Friese, 1916: 109 . Bombus latissimus var. [subsp.] segmentarius Friese, 1916: 109 . Bombus latissimus var. [subsp.] tricoloratus Friese, 1916: 109 . Bombus rufipes var. [subsp.] lutescens Friese, 1918: 517 (non Pérez, 1890: 154 = B. flavidus Eversmann ). Bombus discrepans Pendlebury, 1923: 64 . Bombus geei subsp. nigribasis Cockerell, 1931: 5 . Bombus eximius var. [subsp.] fulvobrunneus Chiu, 1948: 65 . Bombus eximius var. [subsp.] pallidus Chiu, 1948: 66 (non Cresson, 1863: 92 = B. pensylvanicus (DeGeer)) . Bombus rufipes var. tonkininsis – Friese 1918: 527, incorrect subsequent spelling. Bombus latiszimus – Friese 1931: 304, incorrect subsequent spelling. Figs 25–63. Simplified diagrams for the colour patterns of the hair on the dorsum for the species from the integrative analysis. The dorsum is divided into regions, each of which shows only the predominant or most apparent colour for that region using a simplified colour palette, with olive indicating a mixture of black and yellow hair, and grey indicating a mixture of black and white hair. The rufipes -group . 25 . Queen, China-Taiwan. 26 . Queen, China-Guangdong. 27 . Worker, China-Sichuan. 28 . Worker, Thailand. 29 . Queen, Nepal. 30 . Male, China-Sichuan. 31 . Male, China-Sichuan. 32 . Male, Thailand. 33 . Male, Nepal. 34 . Male, Nepal. 35 . Queen, Indonesia-Sumatra. 36 . Queen, Indonesia-Java. 37 . Male, Indonesia-Java. The festivus -group . 38 . Queen, Nepal. 39 . Queen, China-Sichuan. 40 . Worker, ChinaSichuan. 41 . Worker, China-Xizang. 42 . Worker, China-Yunnan. 43 . Male, Nepal. 44 . Male, ChinaYunnan. 45 . Male, China-Yunnan. The rufofasciatus -group. 46 . Queen, India-Kashmir. 47 . Worker, India-Kashmir. 48 . Male, India-Kashmir. 49 . Male, India-Kashmir. 50 . Queen, India-Arunachal Pradesh. 51 . Queen, Bhutan. 52 . Worker, Bhutan. 53 . Worker, Bhutan. 54 . Male, Bhutan. 55 . Queen, India-Kashmir. 56 . Queen, India-Kashmir. 57 . Worker, India-Kashmir. 58 . Worker, India-Kashmir. 59 . Worker, IndiaKashmir. 60 . Worker, Pakistan. 61 . Male, India-Kashmir. 62 . Male, India-Kashmir. 63 . Male, Nepal. Figs 64–102. Simplified diagrams for the colour patterns of the hair on the dorsum for the species from the integrative analysis. The dorsum is divided into regions, each of which shows only the predominant or most apparent colour for that region using a simplified colour palette, with olive indicating a mixture of black and yellow hair, and grey indicating a mixture of black and white hair. The rufofasciatus -group. 64 . Queen, China-Yunnan. 65 . Queen, China-Sichuan. 66 . Worker, China-Xizang. 67 . Worker, ChinaXizang. 68 . Worker, China-Gansu. 69 . Male, China-Yunnan. 70 . Male, China-Xizang. 71 . Queen, IndiaKashmir. 72 . Queen, India-Kashmir. 73 . Queen, Pakistan. 74 . Worker, India-Kashmir. 75 . Worker, IndiaKashmir. 76 . Male, Pakistan. 77 . Male, India-Kashmir. 78 . Queen, Burma. 79 . Worker, Burma. 80 . Worker, China-Xizang. 81 . Worker, China-Xizang. 82 . Male, Burma. 83 . Queen, China-Sichuan. 84 . Worker, China-Sichuan. 85 . Worker, China-Sichuan. 86 . Worker, China-Sichuan. 87 . Worker, China-Sichuan. 88 . Worker, China-Xizang. 89 . Male, China-Sichuan. 90 . Male, China-Sichuan. 91 . Queen, China-Beijing. 92 . Queen, China-Beijing. 93 . Worker, China-Shanxi. 94 . Worker, China-Beijing. 95 . Worker, ChinaBeijing. 96 . Male, China-Beijing. 97 . Male, China-Beijing. 98 . Queen, China-Taiwan. 99 . Worker, ChinaTaiwan. 100 . Worker, China-Taiwan. 101 . Worker, China-Taiwan. 102 . Male, China-Taiwan. Figs 103–138. Simplified diagrams for the colour patterns of the hair on the dorsum for the species from the integrative analysis. The dorsum is divided into regions, each of which shows only the predominant or most apparent colour for that region using a simplified colour palette, with olive indicating a mixture of black and yellow hair, and grey indicating a mixture of black and white hair. The tanguticus -group . 103 . Worker, China-Qinghai. 104 . Worker, China-Qinghai. 105 . Queen, China-Qinghai. 106 . Queen, China-Xizang. 107 . Queen, India-Kashmir. The lapidarius -group . 108 . Queen, Russia-North Ossetia. 109 . Worker, Azerbaijan. 110 . Queen, Georgia. 111 . Male, Russia-North Ossetia. 112 . Male, Russia-North Ossetia. 113 . Male, Russia-North Ossetia. 114 . Male, Turkey. 115 . Queen, Morocco. 116 . Queen, Spain. 117 . Queen, Spain. 118 . Queen, UK. 119 . Male, Spain. 120 . Male, Spain. 121 . Male, UK. 122 . Male, UK. The sichelii -group . 123 . Queen, Iran. 124 . Male, Iran. 125 . Queen, India-Kashmir. 126 . Queen, IndiaKashmir. 127 . Male, India-Kashmir. 128 . Male, India-Kashmir. 129 . Queen, Iran. 130 . Queen, RussiaSakha. 131 . Queen, China-Sichuan. 132 . Worker, China-Sichuan. 133 . Queen, Mongolia. 134 . Worker, Mongolia. 135 . Queen, Spain. 136 . Worker, Austria. 137 . Male, Turkey. 138 . Male, Mongolia. Figs 139–180. Simplified diagrams for the colour patterns of the hair on the dorsum for the species from the integrative analysis. The dorsum is divided into regions, each of which shows only the predominant or most apparent colour for that region using a simplified colour palette, with olive indicating a mixture of black and yellow hair, and grey indicating a mixture of black and white hair. The keriensis -group . 139 . Queen, China-Sichuan. 140 . Worker, China-Sichuan. 141 . Queen, China-Sichuan. 142 . Queen, China-Sichuan. 143 . Worker, China-Xizang. 144 . Queen, India-Kashmir. 145 . Male, China-Qinghai. 146 . Male, India-Kashmir. 147 . Male, China-Gansu. 148 . Queen, Armenia. 149 . Queen, Iran. 150 . Male, Georgia. 151 . Worker, China-Qinghai. 152 . Male, China-Qinghai. 153 . Queen, Mongolia. 154 . Worker, Mongolia. 155 . Male, Mongolia. 156 . Male, Mongolia. 157 . Queen, China-Sichuan. 158 . Worker, China-Qinghai. 159 . Worker, China-Qinghai. 160 . Male, China-Qinghai. 161 . Male, China-Qinghai. 162 . Male, China-Qinghai. 163 . Queen, Pakistan. 164 . Queen, Pakistan. 165 . Queen, Pakistan. 166 . Queen, Pakistan. 167 . Queen, Afghanistan. 168 . Queen, Pakistan. 169 . Queen, Pakistan. 170 . Queen, India-Kashmir. 171 . Male, India-Kashmir. 172 . Male, India-Kashmir. 173 . Queen, ChinaXinjiang. 174 . Queen, Kyrgyzstan. 175 . Queen, Kyrgyzstan. 176 . Queen, Mongolia. 177 . Queen, IndiaKashmir. 178 . Male, Kazakhstan. 179 . Male, China-Xinjiang. 180 . Male, Mongolia. Figs 181‒189. Morphology of the male genitalia for species of the subgenus Melanobombus von Dalla Torre, 1880 from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 181 . Bombus eximius Smith, 1852 , China-Guangxi. 182 . B. rufipes Lepeletier, 1835 , IndonesiaJava. 183 . B. festivus Smith, 1861 , Nepal. 184 . B. simillimus Smith, 1852 , India-Himachal Pradesh. 185 . B. miniatus Bingham, 1897 , Bhutan. 186 . B. eurythorax Wang, 1982 stat. rev. , India-Kashmir. 187 . B. prshewalskyi Morawitz, 1880 stat. rev. , China-Sichuan. 188 . B. rufofasciatus Smith, 1852 , IndiaKashmir. 189 . B. richardsiellus ( Tkalců, 1968 ) , Burma (left volsella distal part missing). Scale bars = 1 mm. Figs 190‒198. Morphology of the male genitalia for species of the subgenus Melanobombus von Dalla Torre, 1880 from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 190 . Bombus friseanus Skorikov, 1933 , China-Yunnan. 191 . B pyrosoma Morawitz, 1890 , ChinaBeijing. 192 . B. formosellus ( Frison, 1934 ) , China-Taiwan. 193 . B. eriophorus Klug, 1807 , RussiaNorth Ossetia. 194 . B. lapidarius ( Linnaeus, 1758 ) , UK. 195 . B. incertus Morawitz, 1881 , Turkey. 196 . B. semenoviaus (Skorikov, 1914) , India-Kashmir. 197 . B. sichelii Radoszkowski, 1859 , Austria. 198 . B. ladakhensis Richards, 1928 , China-Sichuan. Scale bars = 1 mm. Bombus eximius and B. rufipes have a distinctive morphology and previously were placed in a separate subgenus, Rufipedibombus ( Skorikov 1923 ) . A close relationship with other species of the subgenus Melanobombus was supported by an analysis of five genes by Cameron et al . (2007) and these subgenera were then synonymised by Williams et al . (2008) . Our PTP analysis ( Fig. 10 ) of coalescents in the COI gene within the rufipes- group supports two species B. eximius and B. rufipes , corroborated by differences in morphology. It supports as conspecific within B. eximius s. lat. the individuals from Taiwan that may have much of the hair of T2–6 orange (taxon latissimus ), the individuals from southern China and Southeast Asia that often have T2–3 predominantly black and T4–6 orange, and the individuals from the Himalaya that have T2–4 black and T5–6 either orange or black (taxon eximius s. str. ). Figs 199‒204. Morphology of the male genitalia for species of the subgenus Melanobombus von Dalla Torre, 1880 from the dorsal aspect, anterior at the bottom of the image, posterior at the top. 199 . Bombus alagesianus Reinig, 1930 stat. rev. , Georgia. 200 . B. tibeticus sp. nov. , China-Qinghai. 201 . B. incertoides Vogt, 1911 stat. rev. , Mongolia. 202 . B. qilianensis sp. nov. , China-Qinghai. 203 . B. keriensis Morawitz, 1887 , India-Kashmir. 204 . B. separandus Vogt, 1909 stat. rev. , Kyrgyzstan. Scale bars = 1 mm. Diagnosis Females ( Fig. 1 ) Queens very large body length 23–30 mm , workers 13–20 mm . Can be distinguished by their combination of wings yellow with the veins orange (cf. B. rufipes ), the hair and integument of mid and hind tibiae and of all barsitarsi orange (cf. some B. rufipes ). The labral lamella has the anterior edge broad and nearly straight (cf. B. ( Pyrobombus ) flavescens Smith, 1852 , B. ( Alpigenobombus ) genalis Friese, 1918 ). Males Body length 16–19 mm . Can be distinguished by their combination of wings yellow with the veins orange (cf. B. rufipes ), similarly coloured to the worker but with the hair of the face black intermixed with grey (cf. B. rufipes ) and the thoracic dorsum black or sometimes intermixed with grey (cf. B. festivus ). Genitalia ( Fig. 181 ) with the gonostylus reduced to a transverse ‘S’-shaped band (cf. non- rufipes -group), the proximal inner projection not broadened distally and simple (cf. B. rufipes ); volsella projecting beyond the gonostylus by ca 6 × its breadth at its midpoint (cf. B. rufipes ), and without an obvious inner distal process or hook (cf. non- rufipes -group); penis valve head with the outer flange greatly expanded as a triangular projection (cf. non- rufipes -group); eye unenlarged relative to female eye. Material examined Holotype BANGLADESH • ♀ (queen), holotype of Bombus eximius Smith, 1852 by monotypy ( Williams et al. 2009 ); “ Silhet ” [= Sylhet ]; NHMUK (examined PW ). Material sequenced ( 7 specimens ) THAILAND • 1 ♀ (worker); Chiang Mai , Doi Inthanon ; 18.553º N , 98.48º E ; 2 Feb. 2007 ; Y. Areeluck leg.; BOLD seq: 3773E10; PCYU : ML180 • 1 ♀ (worker); same collection data as for preceding; 16 Feb. 2007 ; Y. Areeluck leg.; BOLD seq: 3773E11; PCYU : ML195 • 1 ♀ (worker); Chaiyaphum , Tat Tone NP ; 15.988º N , 102.041º E ; 12 Oct. 2006 ; Y. Areeluck leg.; BOLD seq: 3261H06; PCYU : ML196 . CHINA • 1 ♀ (queen); Guangdong ; 22.2933º N , 111.2061º E ; 16 Jan. 2013 ; BOLD seq: 1555A07; PW : ML178 • 1 ♀ (worker); Chongqing ; 30.13º N , 108.1018º E ; Aug. 2010 ; BOLD seq: 1555A08; PW : ML179 . TAIWAN • 1 ♀ (queen); Taichung , Dashuishan road; 24.2561º N , 121.0108º E ; 18 Jun. 2004 ; C. Dietrich leg.; BOLD seq: 1555A06; PW : ML177 . BURMA • 1 ♀ (worker); Chin , Natmataung ; 21.2198º N , 93.9401º E ; 2 Jun. 2010 ; D. Zimmermann leg.; BOLD seq: 6880E04; NHMW : ML502 . Global distribution (Himalayan, south Chinese and southeast Asian mountain species including some of the northern islands) East Asia : CHINA : Xizang , Sichuan , Yunnan , Chongqing , Jiangxi , Guizhou , Fujian , Guangdong , Guangxi , Taiwan. – Himalaya : INDIA : Sikkim , Darjiling Bengal, Arunachal Pradesh , Meghalaya ; NEPAL , BHUTAN . – Southeast Asia : BANGLADESH , BURMA , THAILAND , VIETNAM . ( IAR , IOZ , KIZ , NHMUK , NHMW , NME , PCYU , PW , RMNH , SEHU , SMNS , USNM , ZMHB , ZMUM .) The species is widely distributed but not common in collections. Behaviour Expected to be food-plant generalists ( Kjellsson et al. 1985 ).