Widespread polytypic species or complexes of local species? Revising bumblebees of the subgenus Melanobombus world-wide (Hymenoptera, Apidae, Bombus) Author Williams, Paul H. Natural History Museum, Cromwell Road, London SW 7 5 BD, UK. Institute of General and Experimental Biology, Peace Avenue 54 b, Ulaanbaatar 13330, Mongolia. Novosibirsk State University, ul. Pirogova 2, Novosibirsk, 630090 Russia. Langstraat 105, B- 2260 Westerlo, Belgium. South China Agricultural University, Guangzhou 510642, China. Agricultural University of Georgia, 240 Agmashenebli Alley, Tbilisi, Georgia. Indonesian Institute of Sciences (LIPI), Jakarta, Indonesia. Kunming Institute of Botany (Chinese Academy of Sciences), 132 Lanhei Road, Kunming, Yunnan 650201, China. Università di Roma ‘ Sapienza’, Piazzale Valerio Massimo 6, Roma 00162, Italy. Yasouj University, Zirtol, Yasouj, Iran. Sherubtse College, Royal University of Bhutan, Trashigang, Bhutan. Zoological Survey of India, Pali Road, Jodhpur 342005, Rajasthan, India. Institute of Zoology (Chinese Academy of Sciences), 1 Beichen West Road, Chaoyang, Beijing 100101, China. paw@nhm.ac.uk Author Altanchimeg, Dorjsuren altanchimegd@mas.ac.mn Author Byvaltsev, Alexandr byvam@yandex.ru Author Jonghe, Roland De roland.de.jonghe@telenet.be Author Jaffar, Saleem saleemjaffar@stu.scau.edu.cn Author Japoshvili, George g.japoshvili@agruni.edu.ge Author Kahono, Sih sihkahono@gmail.com Author Liang, Huan lianghuan@mail.kib.ac.cn Author Mei, Maurizio Author Monfared, Alireza Author Nidup, Tshering Author Raina, Rifat Author Ren, Zongxin Author Thanoosing, Chawatat Author Zhao, Yanhui Author Orr, Michael C. text European Journal of Taxonomy 2020 2020-10-02 719 1 120 journal article 9775 10.5852/ejt.2020.719.1107 7ca72f76-4fae-4305-8601-4662f4cd2b96 4064324 A4500016-C219-4353-B81C-5E0BB520547F Bombus rufofasciatus Smith, 1852 Figs 13 , 71–77 , 188 Bombus rufo-fasciatus Smith, 1852a: 48 . Bombus rufofasciatus var. [subsp.] championi Richards, 1928a: 107 . Bombus waterstoni Richards, 1934: 88 . Bombus rufofasciatus s. str. is revised here as a western species separate from the eastern B. prshewalskyi because of their unique and strongly divergent species coalescents in the COI gene ( Fig. 10 ), corroborated by differences in morphology. The morphological differences are subtle, but do appear to corroborate that the two 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 sister candidate species within a rufofasciatus -complex: 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 (even though it also includes faster-evolving intron regions) shows no diagnostic nucleotide changes. From morphology, B . rufofasciatus s. str. has a slightly lighter colour pattern of the hair than B . prshewalskyi for the females (larger workers and queens), with the white bands often appearing bright white. T2 of workers (and sometimes in the far west, also of queens) is often predominantly yellow. Current restrictions on collecting and sequencing in the areas where these 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 these candidate species rufofasciatus s. str. and prshewalskyi and it remains possible with more sampling from the Himalaya that the two taxa could still prove to be parts of a single 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 vary from having T2 black with a few yellow hairs (in the east) to having T2 predominantly yellow (in the far west in Pakistan ) and T4 is white; whereas the smaller workers have T2 extensively yellow and T4 red. Males have a similar colour pattern to the workers. That the castes and sexes are conspecific has been confirmed from examining variation within one colony from Kashmir ( Williams 1991 ). Diagnosis Females Queens medium-sized body length 20–23 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. prshewalskyi but tending to have T2 with more yellow hair and the scutellum with less black hair. Males Body length 14–16 mm . Can be distinguished reliably at present only by their COI sequence, but tending to have T2 with more yellow hair and the scutellum with less black hair (cf. B. prshewalskyi ). Genitalia ( Fig. 188 ) 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 pronounced adjacent acute teeth, the proximal tooth often slightly shorter than the distal tooth (cf. other rufofasciatus -group species); 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 Lectotype INDIA • ♀ (queen), lectotype of Bombus rufofasciatus Smith, 1852 by designation of Tkalců (1974a) ; “N.[orth]”; NHMUK (examined PW ). Material sequenced ( 6 specimens ) PAKISTAN • 1 ♂; upper Memosh ; 34.7093° N , 76.1208° E ; 5 Sep. 2017 ; S. Jaffar leg.; BOLD seq: 1555E08; UAP : ML256 • 1 ♂; same collection data as for preceding; BOLD seq: 1555E09; UAP : ML257 • 1 ♂; same collection data as for preceding; BOLD seq: 6877H05; UAP : ML421 . INDIA • 1 ♀ (worker); Uttarakhand , Badrinath Valley of Flowers ; 30.7291° N , 79.6193° E , 18 Aug. 1990 ; S. Cameron leg.; BOLD seq: 6877E07; PW : ML389 . NEPAL • ♀ (queen); Karnali , Churta ; 29.4667° N , 82.1° E ; 2 Jun. 2007 ; F. Creutzburg leg.; BOLD seq: 1551H08; NME : ML213 • ♀ (queen); Karnali , Gothichaur ; 29.1972° N , 82.3083° E ; 7 Jun. 2007 ; F. Creutzburg leg.; BOLD seq: 1551H07; NME ML214 . Global distribution (West Himalayan mountain species) Himalaya : PAKISTAN , INDIA : Kashmir, Himachal Pradesh , Uttaranchal; NEPAL . (NHMUK, NME, NMS, PW, SEHU, UAP.) The species is widely distributed within its range and often abundant. Behaviour A colony of this species has been found underground (under large rocks) in a subalpine meadow at 3000 m a.s.l. in Kashmir ( Williams 1991 ). Food-plant generalists and the male mate-searching behaviour is truly territorial with collisions between males in flight and replacement in perch occupancy ( Williams 1991 ).