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 rufipes Lepeletier, 1835 Figs 12 , 35–37 , 182 Bombus rufipes Lepeletier de Saint-Fargeau, [1835] : 473. Bombus flavipes Handlirsch, 1888: 225 . Bombus rufipes var. [subsp.] obscuripes Friese, 1914: 10 . Bombus rufipes var. [subsp.] intermissus Friese, 1918: 516. Bremus rufipes var. [subsp.] richardsi Frison, 1930: 6 . Bombus rufipes and B. eximius are now considered parts of the subgenus Melanobombus (see the comments on B. eximius ). 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. From morphology, the black wings, black hair of the male face, and the matching form of the male genital volsella and gonostylus support as conspecific the individuals from Sumatra, which usually (not always) have the mid and hind tibiae orange (taxon obscuripes ), together with the individuals from Java, which more usually have the mid and hind tibiae predominantly black (taxon rufipes s. str. ). The gonostylus interior proximal process is less strongly bifid for the male from Sumatra. For the male labelled Borneo, the black wings, black facial hair, and form of the volsella and gonostylus all match B. rufipes . For this male, the hair of the mid and hind basitarsi is orange, although the hair of the mid tibia is black and the hair of the hind tibia is orange at the base and black at the tips. A few individuals from Sumatra and Peninsular Malaysia have the hair of the scutellum brown or orange (taxon richardsi ). A queen of ‘ostindischen’ origin is described as having T5–6 orange-red (taxon flavipes , not seen). Diagnosis Females Queens large body length 24–26 mm , workers 14–18 mm . Can be distinguished by their combination of the hair of the metasoma entirely black with the wings nearly black with the veins black (cf. B. eximius ), the hair and integument of mid and hind tibiae and of all barsitarsi either orange or black. The labral lamella has the anterior edge broad and nearly straight (cf. B. ( Megabombus ) senex Vollenhoven, 1873 , B. ( Megabombus ) melanopoda Cockerell, 1910 , from Sumatra ). Males Body length 15–18 mm . Can be distinguished by their combination of the hair of the face black with the metasoma entirely black with the wings nearly black with the veins black (cf. B. eximius ), and the thoracic dorsum anteriorly black (cf. B. festivus ). Genitalia ( Fig. 182 ) with the gonostylus reduced to a transverse ‘S’-shaped band (cf. non- rufipes -group), the proximal inner projection broadened distally and bifid (cf. B. eximius ); volsella projecting beyond gonostylus by ca 5 × its breadth at its midpoint (cf. B. eximius ), 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 INDONESIA • ♀ (worker), holotype of Bombus rufipes Lepeletier, 1835 by monotypy; Java ; OUMNH (examined PW from photographs). The original description refers to a single worker with no evidence that the author had additional specimens in front of him, so this worker can be recognised as the holotype by monotypy. Recognising this specimen as the holotype invalidates the neotype designation by Frison (1930: 11) . Material sequenced ( 2 specimens ) INDONESIA • 2 ♂♂ ; East Java , Mt Argopuro ; 7.9756° S , 113.5193° E ; 5 Mar. 2010 ; BOLD seq: 1555E01, 1555E02; SK: ML249, ML250 . Global distribution (Southeast Asian mountain species primarily in the southern islands) Southeast Asia : MALAYSIA : Peninsular Malaysia , INDONESIA : Sumatra , Java ,? Kalimantan ,? Sulawesi . (NHMUK, NHMW, PW, OUMNH, SK.) It was reported previously (by Williams in Starr 1989 ) that there is a male labelled “Bandjar/ Borneo” (= Banjar?) in the NHMUK collection (examined PW). The only other bumblebee species recorded apparently from Borneo is the holotype of B. folsomi ( Frison, 1923 ) , which is labelled “Kina Bala/ N. Borneo” (= Gunung Kinabalu, Sabah). However, the latter specimen appears to be a mislabelled queen of B. ( Pyrobombus ) ephippiatus Say, 1837 , probably originating from Costa Rica or Panama ( Starr 1989 ; Williams 1998 , examined PW). The NHMUK “Bandjar/ Borneo” specimen of B. rufipes reached this collection from B. Pittioni’s collection that was assembled in Austria (the specimen carries a det. label from F. Maidl, who was at Vienna’s NHMW museum, and a label “Pittioni Coll./Turner Bequest/ B.M. 1954–79 ”). The NHMW collection has a further three workers and two males with the same locality data in the same handwriting and with the same identification labels. Of nine localities named ‘Bandjar’ identified in Malaysia and Indonesia from US military gazetteers ( USBGN 1968 , 1970 ), one is in Borneo and six are in Java. Frison (1930) interpreted ‘Bandjar’ as referring to Java for a queen, two workers, and the seven males of B. rufipes loaned to him from the ‘Zoological Museum, Buitenzorg, Java’ (= Bogor) collection. Two further workers of B. rufipes without locality labels are listed by Frison that were loaned to him by Maidl from the NHMW. Handlirsch (1891) wrote that all but one of the B. rufipes then in the NHMW collection were from eastern Java (for the other, see below). The occurrence of B. rufipes on Borneo, perhaps on the higher Maratus mountains of south-eastern Kalimantan near Banjar (across the Java Sea from Java and Sumatra), remains possible, but would need to be confirmed. Two other records appear to extend the known range. One is a queen in the NHMUK from “ Selangor / Bukit Kutu” (Peninsular Malaysia , examined). There is no obvious reason to doubt this record. Another is a queen in the NHMW collection labelled “N. Celebes / 1908” (= northern Sulawesi , examined PW). The occurrence of B. rufipes in northern Sulawesi would need to be confirmed because it is ca 2000 km from the nearest sites with confirmed records for B. rufipes on Java (see the Discussion). The species is widely distributed but not common in collections. Behaviour Some aspects of the behaviour of this species have been described ( Michener & Amir 1977 ; Kato et al. 1992 ).