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 keriensis Morawitz, 1887 Figs 16 , 163–172 , 203 , 209 , 212 Bombus keriensis Morawitz, 1887: 199 . Lapidariobombus separandus subsp. meridialis Skorikov, 1914a: 127 . Bombus trilineatus S.-F. Wang, 1982: 441 . Pyrobombus keriensis subsp. karakorumensis Tkalců, 1989: 57 . The taxonomic limits of the species B. keriensis have been a particular long-standing or persistent taxonomic problem. Vogt (1909: 41 , 62) treated B. keriensis s. str. as a species separate from his taxon separandus . Vogt (1911: 58) characterised and distinguished B. keriensis s. str. from B. separandus by having from a few yellow hairs on the face to having the hair of the face entirely pale. Skorikov (1931) listed B. keriensis s. str. and B. separandus as two species but discussed them together under a joint heading, “ Lapidariobombus keriensis (F. Mor.) 1886 und Lapidariobombus separandus (Vogt) 1909 ”. While he mentioned the difference in face colour, he described B. keriensis s. str. and B. separandus as hardly distinguishable (his Lapidariobombus anargumentosus Skorikov, 1931 , on the same page is recognised here from the pattern of punctures on the ocello-ocular area of the head of the syntypes in the ZIN as conspecific with B. ( Pyrobombus ) biroi Vogt, 1911 , syn. nov. , and is not a species of the subgenus Melanobombus ). Reinig (1935) included the taxon keriensis s. str. as a part of a broad species B. keriensis s. lat. that also included the taxa separandus + incertoides + alagesianus . This interpretation was followed by Williams (1991 , 1998 ). Wang (1982) described her taxon trilineatus without reference to B. keriensis , comparing it instead to the more distantly related “ B. occulatus var. haemorrhous Richards ” [sic] (= B. simillimus ) for colour pattern and details of morphology. Our PTP analysis ( Fig. 10 ) of coalescents in the COI gene within the keriensis -complex supports six species including B. keriensis s. str. , corroborated by differences in morphology. These species are also supported by the absence of a positive divergence-with-distance relationship among them ( Fig. 20 ) (see Divergence and geographical distance, page 12). From morphology, after examining>1000 specimens of the keriensis -group from across Asia and after comparing the distribution of states of several characters as part of this study (PW), Vogt’s character state of at least a few yellow hairs on the face does appear to remain one of the most consistently diagnostic character states for B. keriensis s. str. (by comparison with COI sequences), at least for queens in most parts of the species’ range (but see the comments on B. separandus regarding the Mongolian taxon kozlovi ). Several aspects of the colour pattern vary strongly within the species: (1) B. keriensis s. str. includes both yellow-banded and white-banded colour patterns; (2) the extent of pale hair on the face varies; (3) the extent of the black hair between the wing bases varies; (4) the extent of the pale hair on the side and ventral area (leg bases) of the thorax varies; (5) the extent of the pale hair posteriorly on T3 varies (the ‘ciliation’ described by authors); (6) the contrast of the pale posterior fringes on T4–5 varies. Many of these characters might appear to show a continuum of variation that is also continuous with the variation within B. separandus ( Williams 1991 ) , which might then appear consistent with the two taxa being parts of the same species ( Reinig 1935 ). However, evidence from COI barcodes shows that the two gene-coalescent groups differ most consistently in the presence of pale hairs on the face of B. keriensis s. str. , supporting Vogt’s characterisation of the species, and providing integrated morphological support for its status as a separate species. The overlapping colour variation of B. separandus but only in Mongolia (where B. keriensis s. str. does not occur) does not negate the fact that B. keriensis s. str. is recognisable by morphological characters throughout its geographical range. There are some yellow-banded queens from the Pir Panjal and Great Himalaya ranges (part of B. keriensis s. str. according our COI coalescent results, Fig. 10 ) that have very few pale hairs on the face, although these specimens still have a few yellow hairs on the leg bases and have a few black hairs in the pale bands of the thoracic dorsum. Many of the available specimens have failed repeated attempts at sequencing, so the species’ diagnosis remains tentative. Particularly influential in the interpretation of these individuals is one sequenced yellow-banded queen with very few pale hairs on the face, on the underside of the thorax, and on T3, from the Pir Panjal range (PW: ML405, Figs 170 , 209 ) that groups with B. keriensis s. str. ( Fig. 10 ) rather than with B. separandus (diagnostic base positions 267T, 286T, 301T, 306C, 339C, 363T, 423C, 498T, 529A, 540C, 541T, 556T). This is one of the specimens that most closely resembles some yellow-banded B. separandus also from Kashmir ( Figs 177 , 210 ) (the difference in the breadth of the black band between the wings is not diagnostic for these species). Diagnosis Females Queens medium-sized body length 17–21 mm , workers 9–14 mm . Can be distinguished in Central Asia by the combination of hair of the face usually with some pale hairs, the leg bases usually with some pale hairs, T2 posteriorly entirely pale without black hairs, with T3 laterally usually with some pale hairs (cf. B. sichelii , B. separandus ). Males Body length 11–15 mm . Can be distinguished reliably at present only by their COI sequence, although in the western Himalaya they may be distinguished by yellow hair between the wing bases and on T3 posterio-laterally. Genitalia ( Fig. 203 ) with the gonostylus shorter than broad, its inner basal projection reduced to a short stub (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 designation CHINA • ♀ (queen), lectotype of Bombus keriensis Morawitz, 1887 ; Keria Mts ; 9000 ft a.s.l. ; Przewalsky leg.; ZIN . Morawitz’s original description of several females of B. keriensis gives the type locality as “Keria” [= Keriya/Yutian, east of Hotan /Hetian / Khotan], presumably not in the low-lying town but higher in the neighbouring Karakax Shan (of the western Kunlun Shan [shan = mountains], southern Xinjiang , and probably near the village of Polu / Pulu/ Pulucun [cun = village], 36.1850° N , 81.4827° E . This village was visited by N. Prshewalsky while searching for a pass southwards into Ladakh during his fourth (and final) Central Asian expedition in the summer of 1885 ( Rayfield 1976 ). If the “9000” on the syntype specimen label (below) were an elevation in feet then this is substantially higher than the elevation of Keriya (ca 4500 feet ), so a collecting site near Pulu is highly likely. These mountains are very arid, so likely sites with a reliable water supply and food plants every summer sufficient to support bumblebee colonies are restricted ( Williams et al. 2015a ). The ZIN collection contains a queen that agrees with the original description and carries seven labels: (1)gold disc;(2)handwritten white label“Keria / Gebirg./ 9000”reverse“2/10/[illegible]”;(3)handwritten white label “Przewalsky”; (4) handwritten white label “ Bombus / keriensis ♀ . / F. Morawitz.”; (5) printed white label in Cyrillic “[K. F. Morawitz ]”; (6) red label partly printed in black “ Lectotypus Bombus / keriensis F. Morawitz / design. Podbolotsk.” (unpublished); (7) red label printed in black “ LECTOTYPE ♀ / Bombus / keriensis / Morawitz, 1887 / det. PH Williams 2017”. This specimen, which is complete, is recognised as one of Morawitz’s syntypes and is designated here as the lectotype in order to reduce uncertainty in the identity and application of the name. Material sequenced ( 7 specimens ) INDIA – Kashmir • 1 ♀ (queen); Gulmarg , Mt Apharwat ; 34.0548° N , 74.3856° E ; Aug. 1985 ; P. Williams leg.; BOLD seq: 6877G02; PW : ML405 • 1 ♀ (queen); Gulmarg , Mt Apharwat ; 34.0212° N , 74.3208° E ; 21 Jul. 2009 ; BOLD seq: 1552D08; RR : ML204 • 1 ♀ (queen); Zanskar , Padum ; 33.4746° N , 76.8871° E ; 30 Jul. 2007 ; BOLD seq: 1552D11; RR : ML203 • 1 ♀ (worker); same collection data as for preceding; 31 Jul. 2011 ; R. Raina leg.; BOLD seq: 1555C12; RR : ML243 • 1 ♀ (worker); Razdaan pass; 34.5390° N , 74.6377° E ; 23 Jul. 2012 ; R. Raina leg.; BOLD seq: 1555D01; RR : ML244 • 1 ♀ (worker); Dawar ; 34.6464° N , 74.7171° E ; 24 Jul. 2012 ; R. Raina leg.; BOLD seq: 1555H03; RR : ML305 . — Himachal Pradesh • 1 ♀ (worker); Saichu Nulla ; 31.6° N , 78.267° E ; 8 Sep. 1986 ; A. Hutchings leg.; BOLD seq: 6877B05; PW : ML159 . Global distribution (West Qinghai-Tibetan-Plateau species) Himalaya : AFGHANISTAN , PAKISTAN , INDIA : Kashmir, Himachal Pradesh . – East Asia : CHINA : Xizang . (IOZ, NHMUK, PW, RR, ZIN, ZSM.) The species is usually not common ( Fig. 212 ). Behaviour Food-plant generalists ( Williams 1991 ). Male mate-searching patrolling behaviour ( Williams 1991 ).