taxonID	type	description	language	source
1A438786793C042C9D16F930FA93FC66.taxon	discussion	We associate a total of 83 published formal names with species of the subgenus Alpigenobombus (including misspellings and infrasubspecific names). Applying the ICZN (1999) rules for an assessment of the status of these names under Article 45.6, just 55 names are considered to be available (most unavailable names are not listed here, but are in a database, PW in prep.). Below is a list of the 11 species recognised from the integrative analysis together with the synonyms that occur most frequently in the literature (identified from morphology) with the changes since the last checklist (Williams 1998). There are few novel arrangements of the names here because there have been so many different re-arrangements in the past, although our results show the currently supported interpretation. The subgenus Mastrucatobombus Krueger, 1917 is a synonym of Alpigenobombus. The subgenus Nobilibombus Richards, 1968 has been included within the subgenus Alpigenobombus (Williams et al. 2008), based on evidence from the five genes obtained by Cameron et al. (2007). The name Nobilibombus was published initially without fixation of a type species (Skorikov 1933 b), or as a junior synonym (Bischoff 1936; Milliron 1961), so the first valid publication is by Richards (1968). The species of Nobilibombus are recognised here as the nobilis - group of species.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793C042C9D16F930FA93FC66.taxon	discussion	Key to species for females of the subgenus Alpigenobombus Future identification of species of the subgenus Alpigenobombus will be most reliable for specimens from which COI-barcode-like data are available for comparison of nucleotide differences with the reference data we provide in supplementary data. Keys using morphological shape, surface sculpturing, and hair-colour-pattern characters follow below. Our results imply that in some cases we should be able to assign most reliably the specimens with locality labels to groups or species on the basis of their collection locality alone. All identifications of species from the key or from the figures should be checked against the species’ diagnoses within the accounts for each species. Diagrams showing the major aspects of variation in the colour-patterns of the dorsal hair are presented in Figs 19 ‒ 105. These diagrams summarise only the major differences (Williams 2007) rather than finer details (e. g., Williams 1991: figs 295 ‒ 310). This is a simplification to aid quantitative comparisons and inevitably requires compromises. Colour-pattern variation within species is established with reference to particular individuals identified from COI barcodes for each diagram. Morphology of the male genitalia is illustrated in Figs 106 ‒ 115.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793C042C9D16F930FA93FC66.taxon	discussion	Key to species for males of the subgenus Alpigenobombus	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	description	Figs 19 ‒ 25, 106	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	discussion	NOTE 1. Alpigenobombus wurfleini subsp. uralicus Tkalců, 1969: 891, by indication of Pittioni (1938). Lectotype by designation of Tkalců (1969: 893): ♀ (queen) central Ural Mountains (‘ Sojmonowsk’), Russia (NHMUK), examined. According to A. Byvaltsev (pers. com.), the type queen of the taxon uralicus was collected on the territory of modern Karabash, in Chelyabinsk Oblast (55.4465 ° N, 60.2151 ° E, ca 320 m a. s. l., but with ridges 20 km to the southwest that reach to ca 1100 m a. s. l.). The Soymonovsky goldmine (named after V. Yu. Soymonov, chairman of the Temporary Mining Commission responsible for the development of gold in the Middle Urals) and the small settlement Sak-Elginsky Vyselok were located here. Later copper deposits were discovered in the area and the Soymonovsky copper smelter was founded in 1834. The Sak-Elginsky settlement then began to be called Soymonovsky or simply Soymonovsk [= ‘ Sojmonowsk’]. Species-taxon concept and variation The taxon concept of the species B. wurflenii is revised here from the widely-cited interpretation since Tkalců (1969), to distinguish B. wurflenii from the separate taxon concept for the species B. mastrucatus (see the comments below on B. mastrucatus), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see below and the keys). The morphological results (see the keys) support the interpretation that the individuals with the divergent colour patterns of the taxa wurflenii s. str. (Figs 20 ‒ 25) and uralicus (lectotype queen Fig. 19) are conspecific, although only a single queen of the taxon uralicus is available. Only two other individuals of the taxon uralicus from the Urals are definitely known at present (A. Byvaltsev pers. com.): a male from 1909 and a female from 1929 (both ZIN). Variation in the colour-pattern diagrams for B. wurflenii in Figs 19 ‒ 25 is arranged approximately from north (Urals) to south (Turkey). Bombus wurflenii, with its white-banded red-tailed female colour pattern in Turkey and the Caucasus, appears to mimic the more abundant B. (Melanobombus) incertus Morawitz, 1881, B. (Ml.) eriophorus Klug, 1807, and similar species (Williams 2007: fig. 5 l).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	materials_examined	Type material Bombus WURFLENii Radoszkowski, 1860: 482. Type not seen, but identity not in doubt.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	diagnosis	Morphological diagnosis Female Wings nearly clear with veins dark brown, hair long, oculo-malar area shorter than broad, clypeus adjacent to the labrum not raised in the centre but flat so that the shallow transverse anterior groove is not interrupted, this central area uniformly with many small punctures and not shining, ocello-ocular area with the punctures anteriorly to the unpunctured area at least as dense as posterior to it (cf. B. mastrucatus); hair of the thoracic dorsum black, with bands anteriorly and usually posteriorly grey-white. Male Wings nearly clear with veins dark brown, hair long, oculo-malar area shorter than broad; genitalia (Fig. 106) with the gonostylus long and distally near its midline axis convexly rounded, length on its outer side about a quarter as long as on its inner side with the two inner corners of the distal lobe rounded, the margin between them nearly straight, penis-valve head strongly recurved, the recurved hook much longer than broad and distally at the end of the hook broadly rounded (cf. B. mastrucatus); hair of the thoracic dorsum between the wing bases either predominantly pale cream-yellow or with many pale cream-yellow hairs.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	materials_examined	Material sequenced in Fig. 12 GEORGIA • 1 ♀ (queen); Lagodekhi; 41.8714 ° N, 46.3645 ° E; 22 Jun. 2015; G. Kirkitadze leg.; BOLD seq: 1555 A 01; IEAUG: AG # 065.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	distribution	Global distribution Western Asia: Turkey, Georgia, Armenia, Azerbaijan, Iran, Russia (Caucasus, Urals): IEAUG, NHM, PW, ZIN. This species is recorded at elevations of 1600 ‒ 2570 m in Turkey in the montane and subalpine coniferous forest zone (Rasmont & Flagothier 1996) and in the alpine zone in the northern Caucasus (Skorikov 1910).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867936042B9E80FB55FDBAFE66.taxon	biology_ecology	Behaviour Male eye not obviously enlarged relative to female eye: males are expected to show ‘ patrolling’ behaviour in search of mates (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	description	Figs 1, 26 ‒ 34, 107	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	discussion	NOTE 1. The earlier name B. helferanus f. [orma] apfelbecki Pittioni, 1939: 87 (= B. humilis Illiger) was given infrasubspecific status (Pittioni 1939: 51), so that the name apfelbecki Reinig, 1988 remains available and a replacement name is unjustified. Species-taxon concept and variation The taxon concept of the species B. mastrucatus is revised here from the widely-cited interpretation (since Tkalců 1969), to distinguish B. mastrucatus from the separate taxon concept of the species B. wurflenii (see the comments above on B. wurflenii), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). Bombus mastrucatus is a widespread European species that had long been well-known by the use of this name by many authors (e. g., von Dalla Torre 1896; Friese & von Wagner 1912; Skorikov 1923; Reinig 1935; Pittioni 1937; aside from occasional confusion with B. lefebvrei Lepeletier de Saint-Fargeau, [1835] (= B. pomorum (Panzer, 1805 )) by Strand 1901). Then Tkalců (1969) reduced the taxon mastrucatus to the status of a subspecies within Alpigenobombus wurfleini [sic], although without a detailed justification. Subsequently, the name B. wurflenii (in various spellings) has been used in most publications for this bumblebee species in Europe (e. g., in influential keys by Løken 1973; Hagen & Aichhorn 2003; Ornosa & Ortiz-Sanchez 2004; Rasmont et al. 2021). Nonetheless, the name mastrucatus remains familiar to European bee workers from these same publications, even if only as what has often been interpreted as a subspecies. The PTP and morphological results (Fig. 12, keys) both support the interpretation that B. mastrucatus and B. wurflenii are separate species. The available COI-barcode-like sequences may all be low-divergence neonumts (Fig. 11). The PTP and morphological analysis results (Fig. 12, keys) also support the interpretation that the divergent colour patterns of the taxa mastrucatus s. str. (Alps: Figs 28 – 29, 32 – 33) and flavicante (Pyrenees: Figs 26 – 27, 31 – 32), as well as the disjunct population of the taxon brevigena (Scandinavia: Figs 30, 33 – 34) are conspecific (no fresh samples were available from eastern Europe for sequencing). No diagnostic characters published for the other named taxa for this group in Europe provide any reason to consider them as candidates for separate species. Variation in the colour-pattern diagrams of B. mastrucatus in Figs 26 ‒ 34 is arranged approximately from south-west (Pyrenees) to north-east (Norway). Bombus mastrucatus, with its unbanded red-tailed colour pattern in northern Europe and its yellow-banded red-tailed colour pattern especially in south-western Europe, appears to mimic the local colour patterns of the abundant B. (Melanobombus) lapidarius (Linnaeus, 1758) and similar species (Williams 2007: fig. 5 e, j).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	materials_examined	Type material Bombus mastrucatus Gerstaecker, 1869: 326. Type not seen (not found by Tkalců 1969: 892), but identity not in doubt.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	diagnosis	Morphological diagnosis Female Wings nearly clear with veins dark brown, hair long, oculo-malar area shorter than broad, clypeus adjacent to the labrum not raised in the centre but flat so that the shallow transverse anterior groove is not interrupted, this central area uniformly with many small punctures and not shining, ocello-ocular area with the punctures anteriorly to the unpunctured area in the outer half less dense than posterior to it (cf. B. wurflenii); hair of the thoracic dorsum either entirely black or sometimes with bands anteriorly and posteriorly yellow. Male Wings nearly clear with veins dark brown, hair long, oculo-malar area shorter than broad; genitalia (Fig. 107) with the gonostylus long and distally near its midline axis convexly rounded, length on its outer side about a quarter as long as on its inner side with the two inner corners of the distal lobe rounded, the margin between them nearly straight, penis-valve head strongly recurved, the recurved hook much longer than broad and distally at the end of the hook weakly pointed (cf. B. wurflenii); hair of the thoracic dorsum between the wing bases predominantly black, usually with few or no yellow hairs.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	materials_examined	Material sequenced in Fig. 12 AUSTRIA • 2 ♀♀ (queens); Carinthia, Innere Eben; 46.978 ° N, 12.759 ° E; 26 Jul. 2019; W. Wallner leg.; BOLD seq: ABOL 19 - 1511 ABOL 19 - 1512; HNS: AG # 164, AG # 165 • 1 ♀ (worker); Tyrol, Hohe Tauern; 47.023 ° N, 12.238 ° E; 20 Jul. 2020; S. Schoder leg.; BOLD seq: ABOLHT 20 - S 038; SS: AG # 166. FRANCE • 1 ♀ (queen); Roussillon, Prats de Mollo; [42.400 ° N, 2.483 ° E]; 25 Aug. 1993; GenBank seq: AY 181174; AG # 066 • 1 ♀ (queen); Alpes Maritimes, Col Turini; 43.999 ° N, 7.429 ° E; 11 Jul. 2010; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM 07560; SNSB: AG # 169 • 1 ♀ (queen); Alpes Maritimes, Trende; 44.105 ° N, 7.448 ° E; 9 Jul. 2010; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM 07561; SNSB: AG # 170. NORWAY • 1 ♀ (queen); Buskerud, Mörkedalen; [60.9 ° N, 8.3 ° E]; 2 Jul. 1994; GenBank seq: AY 181175; AG # 067. Additional sequences in Fig. 10 and haplotype duplicates FRANCE • 1 ♀ (queen); Alpes Maritimes, Col Turini; 43.999 ° N, 7.429 ° E; 11 Jul. 2010; C. Schmid-Egger leg.; BOLD seq: BCZSMHYM 07559; SNSB: AG # 168. GERMANY • 1 ♀ (worker); Saxony, Carlsfeld; 50.435 ° N, 12.585 ° E; 16 Jul. 2004; F. Burger leg.; BOLD seq: BCZSMHYM 08976; SNSB: AG # 171 • 1 ♁; Thuringia, Orlamünde; 50.775 ° N, 11.541 ° E; 19 Sep. 2001; F. Burger leg.; BOLD seq: BCZSMHYM 08978; SNSB: AG # 172 • 1 ♁; Saxony, Carlsfeld; 50.435 ° N, 12.585 ° E; 16 Jul. 2004; F. Burger leg.; BOLD seq: BCZSMHYM 08979; SNSB: AG # 173.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	distribution	Global distribution Europe: Spain, France, Belgium, Italy, Bosnia and Herzegovina, Montenegro, Serbia, Macedonia, Greece, Germany, Switzerland, Austria, Czechia, Slovakia, Poland, Bulgaria, Romania, Moldova, Ukraine, Norway, Sweden, Russia (Murmansk): HNS, NHMUK, PW, RMNH, SNSB, SS, ZMHB. This species is recorded at elevations of 1740 – 2740 in the Eyne Pyrenees (Iserbyt et al. 2008), at 1000 ‒ 2450 m in Languedoc-Roussillon, France (Rasmont 1988), and at 1500 ‒ 2600 m in the Alps (Pittioni 1937; Neumayer 1998; Neumayer & Paulus 1999). It has been associated in general with the mountain-forest zone (Løken 1973; Rasmont et al. 2021).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786793104159E7AFDC6FD08FBE2.taxon	biology_ecology	Behaviour Male eye not obviously enlarged relative to female eye: males show ‘ patrolling’ behaviour when searching for mates in Austria (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	description	Figs 2 – 3, 35 ‒ 48, 108	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	discussion	NOTE 1. Acceptance of B. kashmirensis as the valid name in preference to B. stramineus follows Skorikov (1923) as the First Reviser (ICZN 1999: Article 24). Species-taxon concept and variation The taxon concept of the species B. kashmirensis here agrees with the recent interpretation (Williams 2022 a) that it is separate from the taxon concept of the species B. rainai (see the comments below on B. rainai), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that the divergent colour patterns of the taxa kashmirensis s. str., stramineus, and meinertzhageni are parts of a single species (Williams 1991). The other named taxa (tetrachromus, pulcherrimus) appear to be re-descriptions of this species. Variation in the colour-pattern diagrams of B. kashmirensis in Figs 35 ‒ 48 is arranged approximately from the Hengduan and outer Himalayan ranges (e. g., Pir Panjal: Figs 35 ‒ 37, 42 ‒ 44) to the trans-Himalaya (e. g., Zanskar: Figs 39 ‒ 41, 46 ‒ 48). Bombus kashmirensis, with the white-banded colour pattern in the south (Himalaya) and east (Hengduan) more mesic parts of its range, appears to mimic the abundant B. (Melanobombus) rufofasciatus Smith, 1852, and B. (Ml.) prshewalskyi Morawitz, 1880 (Williams 2007: fig. 5 g), whereas the yellow-banded colour pattern in the north-west (trans-Himalayan mountains) more arid parts of its range appears to mimic the abundant B. (Ml.) keriensis Morawitz, 1887, and similar species (Williams 2007: fig. 5 j).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	materials_examined	Type material Bombus mastrucatus var. [subsp.] kashmirensis Friese, 1909 [September: Tkalců 1974]: 673. Lectotype by designation of Tkalců 1974: ♀ (queen) Kashmir, India (ZMHB). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	diagnosis	Morphological diagnosis Female Wings nearly clear with veins dark brown, hair medium to long, oculo-malar area shorter than broad, clypeus adjacent to the labrum slightly raised in the centre so that the transverse anterior groove is interrupted, this central area with few small punctures and more shining than the surrounding areas, oculo-ocellar area in almost its outer half with large and medium punctures (cf. B. rainai); hair of the side of the thorax in at least its upper half and the scutellum either white or yellow, hair of T 5 orange with white tips. Male Wings nearly clear with veins dark brown, hair medium to long, oculo-malar area shorter than broad; genitalia (Fig. 108) with the gonostylus long and distally near its midline axis convexly rounded, length on its outer side about a quarter as long as on its inner side with the two inner corners of the distal lobe rounded, the margin between them nearly straight, penis-valve head strongly recurved, the recurved hook much longer than broad, but broad and narrowing only just before the apex (cf. B. rainai); male eye distinctly enlarged relative to female eye; hair of the side of the thorax either yellow or white with the pale hair reaching to the midleg bases, hair of T 3 usually predominantly black, hair at the side of T 5 ‒ 6 orange at the base but white-tipped.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	materials_examined	Material sequenced in Fig. 12 CHINA • 1 ♀ (worker); Sichuan, Queer Mountain; 31.8969 ° N, 99.1485 ° E; 5 Aug. 2018; Z. Ren et al. leg.; KIB seq: QESM 302006; KIB: AG # 083 • 1 ♀ (worker); Xizang, Ailashan; 31.6142 ° N, 98.4803 ° E; 4 Aug. 2018; Z. Ren et al. leg.; KIB seq: ALSM 104001; KIB: AG # 085 • 1 ♀ (worker); Qinghai, Kunlun Pass road; 35.6917 ° N, 94.0533 ° E; 19 Aug. 2013; PW seq: PWK 1; IAR: AG # 084. PAKISTAN • 1 ♁; Baltistan [Olthingthang]; 34.7093 ° N, 76.1208 ° E; 16 Sep. 2016; S. Jaffar leg.; BOLD seq: 1555 G 02; PW: AG # 082. Additional sequences in Fig. 10 and haplotype duplicates CHINA • 1 ♀ (worker); Xizang, Yelanshan; 30.1489 ° N, 97.3091 ° E; 22 Jul. 2018; Z. Ren et al. leg.; KIB seq: YLSH 308001; KIB: AG # 130.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	distribution	Global distribution Himalaya and Hengduan: Pakistan, India (Kashmir, Himachal Pradesh, Uttarakhand, Sikkim), Nepal, China (Xizang, Qinghai, Sichuan, Gansu): IAR, IOZ, KIB, NHMUK, PW, RMNH, SC, ZMHB. This species is recorded at elevations of 2400 ‒ 4800 m in the Himalaya and trans-Himalaya, and of 2272 ‒ 4700 m in the Hengduan mountains (Williams 1991; Williams et al. 2009, 2010; An et al. 2014). In Kashmir, Sichuan, and Gansu, B. kashmirensis occurs primarily above the tree line and high into the alpine zone (queens have been seen to forage from flowers at 4000 m in Kashmir even when it was snowing heavily, PW pers. obs.) and even in the semi-arid trans-Himalaya (Williams 1991; Williams et al. 2009).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790F04179ECAFB43FDBFFD62.taxon	biology_ecology	Behaviour Male eye slightly enlarged relative to female eye: males show ‘ cruising’ behaviour when searching for mates in Kashmir (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	description	Figs 49 ‒ 53, 109	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	discussion	Species-taxon concept and variation The taxon concept of the species B. rainai here agrees with the recent interpretation (Williams 2022 a) that it is separate from the taxon concept of the species B. kashmirensis (see the comments above on B. kashmirensis), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). This taxon was discussed as divergent from B. kashmirensis by Williams (1991: 68 ‒ 69, figs 308 ‒ 310), although it was not then named as a separate species. Although the PTP and morphological results (Fig. 12, keys) support the interpretation that B. rainai and B. kashmirensis are separate species (Williams 2022 a), the available COI-barcode-like sequences may all be low-divergence neonumts (Fig. 11). Variation of B. rainai is shown in the colour-pattern diagrams in Figs 49 ‒ 53. Bombus rainai, with its white-banded red-tailed colour pattern in the Himalaya, appears to mimic the abundant B. (Sibiricobombus) longiceps Smith, 1878 (until recently regarded as part of B. asiaticus Morawitz, 1875), and similar species (Williams 2007: fig. 5 l).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	materials_examined	Type material Bombus (Alpigenobombus) rainai Williams 2022 a: 132. Holotype by original designation: ♀ (queen) Kashmir, India (ZSC). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	diagnosis	Morphological diagnosis Female Wings nearly clear with veins dark brown, hair medium length, oculo-malar area shorter than broad, clypeus adjacent to the labrum slightly raised in the centre so that the transverse anterior groove is interrupted, this central area with few small punctures and more shining than the surrounding areas, oculo-ocellar area in its outer half with only a few small shallow punctures, any larger punctures confined to the eye margin (cf. B. kashmirensis); hair of the side of the thorax in no more than its upper third white and the scutellum predominantly black, hair of T 5 entirely orange without obviously paler tips. Male Wings nearly clear with veins dark brown, hair medium length, oculo-malar area shorter than broad; genitalia (Fig. 109) with the gonostylus long and distally near its midline axis convexly rounded, length on its outer side about a quarter as long as on its inner side with the two inner corners of the distal lobe rounded, the margin between them convex, penis-valve head strongly recurved, the recurved hook much longer than broad and narrow, tapering gradually to the apex (cf. B. kashmirensis); male eye distinctly enlarged relative to female eye; hair of the side of the thorax with black and white intermixed, but often predominantly black in its lower half, hair of the sides of T 5 ‒ 6 orange, hair of T 3 often predominantly orange.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	materials_examined	Material sequenced in Fig. 12 INDIA • 1 ♀ (queen); Kashmir, Achoora; 34.636 ° N, 74.864 ° E; 26 Jun. 2009; R. Raina leg.; BOLD seq: 1552 E 01; ZSC: AG # 080 • 1 ♀ (queen); Kashmir, Apharwat; 34.055 ° N, 74.386 ° E; 21 Aug. 2009; R. Raina leg.; BOLD seq: 1552 D 12; ZSC: AG # 081.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	distribution	Global distribution Western Himalaya: Pakistan, India (Kashmir): NHMUK, PW, ZSC. This species is recorded at elevations of 2542 ‒ 4200 m, primarily above the tree line and high into the alpine zone (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790D04169EECFCC2FCA7FB24.taxon	biology_ecology	Behaviour Male eye distinctly enlarged relative to female eye: males are expected to show ‘ cruising’ behaviour when searching for mates in Kashmir (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	description	Figs 4, 54 ‒ 60, 110	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	discussion	Species-taxon concept and variation The taxon concept of the species B. sikkimi here agrees with the recent interpretation (Williams 2022 a) that it is separate from the taxon concept of the species B. nobilis (see the comments below on B. nobilis), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The morphological results (Fig. 12, keys) support the interpretation that the divergent colour patterns of the yellow-banded taxa sikkimi s. str. and individuals from the Himalaya with an unnamed white-banded colour pattern are conspecific. We have been able to obtain COI barcode sequences so far only from the white-banded individuals, not the yellow-banded individuals. The latter will be needed to confirm the conspecific relationship. Variation in the colour-pattern diagrams of B. sikkimi in Figs 54 ‒ 60 is arranged approximately from east to west within the Himalaya. Bombus sikkimi, with the yellow-banded colour pattern in the west (central Himalaya) of its range (Figs 56 – 57, 59 – 60), appears to mimic the common B. (Melanobombus) eurythorax Wang, 1982, and similar species (Williams 2007: fig. 5 j), whereas the white-and-yellow-banded colour pattern in the east (eastern Himalaya: Figs 54 – 55, 58) appears to mimic the abundant B. (Ml.) prshewalskyi (Williams 2007: fig. 5 p).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	materials_examined	Type material Bombus sikkimi Friese, 1918: 82. Syntype: ♀ (queen) Sikkim, India (ZMHB). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	diagnosis	Morphological diagnosis Female Wings nearly clear with veins dark brown, hair long, oculo-malar area longer than broad, oculo-ocellar areas densely and broadly punctured with scattered medium-sized punctures with many small punctures between them, anterio-laterally to the lateral ocellus with large, medium and many small punctures almost coalescing and without flat shining areas in between (cf. B. nobilis, B. validus); hair on the anterior dorsum of the thorax and of the side of the thorax and scutellum and T 1 either grey or olive-yellow, on the scutellum the pale hair anteriorly often deeply divided in the middle by a triangle of black, T 2 either predominantly yellow or with large anterior lateral patches yellow. Male Wings nearly clear with veins dark brown, hair long, oculo-malar area longer than broad; genitalia (Fig. 110) with the gonostylus short and distally near its midline axis nearly consistently weakly convexly rounded, its outer side about a half as long as its inner side with the two inner corners of the distal lobe rounded, the margin between them concave, penis-valve head weakly recurved, the recurved section only slightly longer than broad at the base (cf. B. nobilis, B. validus); hair on the anterior dorsum of the thorax and of the side of the thorax and scutellum and T 1 either grey-white or dull olive-yellow, T 2 anterio-laterally yellow, on the scutellum in the middle the grey or yellow nearly completely divided by black.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	materials_examined	Material sequenced in Fig. 12 INDIA • 1 ♁; Arunachal Pradesh, Donyi La; 28.9819 ° N, 95.2403 ° E; 4 Sep. 2017; NCBS seq: BE 961; NCBS: AG # 205 • 1 ♀ (worker); Arunachal Pradesh, Sango-Camp; 28.9814 ° N, 95.2482 ° E; 4 Sep. 2017; NCBS seq: BE 963; NCBS: AG # 206. Additional sequences in Fig. 10 and haplotype duplicates INDIA • 1 ♀ (worker); Arunachal Pradesh, Sela Lake; 27.5050 ° N, 92.1016 ° E; 24 Sep. 2015; J. Neumayer leg.; NCBS seq: BE 968; NCBS: AG # 005 • 1 ♀ (worker); Arunachal Pradesh, Sango-Camp; 28.9744 ° N, 95.2542 ° E; 2 Sep. 2017; NCBS seq: BE 942; NCBS: AG # 204 • 1 ♀ (queen); Arunachal Pradesh, Nagula; 27.6981 ° N, 91.8486 ° E; 30 May 2016; NCBS seq: BE 967; NCBS: AG # 207.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	distribution	Global distribution Himalaya: India (Himachal Pradesh, Uttarakhand, Sikkim, Arunachal Pradesh), Nepal: NCBS, NHMUK, PW, ZMHB. This species is recorded (under the name B. nobilis) at elevations of 2700 ‒ 4300 m in the Himalaya, mostly above the tree line and into the subalpine zone (especially rich in Rhododendron L. species) (Williams et al. 2010; Streinzer et al. 2019). MS identified two nests, at 3860 m (near BE 963) and one near Sela Pass at 4230 m.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790C04109EB9FB05FDBAFEEF.taxon	biology_ecology	Behaviour Male eye not obviously enlarged relative to female eye: males are expected to show ‘ patrolling’ behaviour in search of mates (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	description	Figs 5, 61 ‒ 69, 111	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	discussion	Species-taxon concept and variation The taxon concept of the species B. nobilis here agrees with the recent interpretation (Williams 2022 a) that it is separate from the taxon concept of the species B. sikkimi (see the comments above on B. sikkimi), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that B. nobilis, B. sikkimi, and B. validus are separate species (Williams 2022 a). The available COI-barcode-like sequences may all be low-divergence neonumts (Fig. 11). From the morphology and colour pattern it appears likely that the yellow-banded holotype queen of the taxon chayaensis (Fig. 63, from Chagyab, north-eastern Xizang) and the white-banded holotype worker of the taxon xizangensis (Fig. 62, from Zayü, southern Xizang) are conspecific with B. nobilis. A small collection from northern Myanmar (Adung valley, 1931) of both yellow-banded (NHMUK: 3 ♀♀ (2 queens, 1 worker), Figs 65 – 66) and white-banded (NHMUK: 2 ♀♀ (workers), Fig. 61) individuals matches this species in morphology. Variation of B. nobilis is shown in the colour-pattern diagrams in Figs 61 ‒ 69. Bombus nobilis, with the yellow-banded colour pattern in the north (northern Hengduan) of its range (Figs 63 – 64, 69), appears to mimic the abundant B. (Melanobombus) friseanus Skorikov, 1933 (Williams 2007: fig. 5 j), whereas the white-and-yellow-banded colour pattern in the south (southern Hengduan: Figs 61 – 62, 67 – 68) appears to mimic the abundant B. (Ml.) prshewalskyi (Williams 2007: fig. 5 p).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	materials_examined	Type material Bombus nobilis Friese, 1905: 513. Type not found, but identity not in doubt (Williams et al. 2009).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	materials_examined	Holotype by original designation: ♀ (worker) Zay ue, Xizang, China (IOZ). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	materials_examined	Holotype by original designation: ♀ (queen) Chagyab, Xizang, China (IOZ). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	diagnosis	Morphological diagnosis Female Wings lightly clouded with brown with the veins dark brown, hair of medium length, oculo-malar area longer than broad, oculo-ocellar areas densely punctured with scattered medium-sized punctures with many small punctures between them, anterio-laterally to the lateral ocellus with large and small punctures sufficiently separated to have flat shining areas in between (cf. B. sikkimi, B. validus); hair on the anterior dorsum of the thorax and of the side of the thorax and scutellum either golden yellow or grey-white, on the scutellum the yellow hair anteriorly scarcely divided in the middle by a triangle of black, T 1 yellow, T 2 either predominantly black or with large anterior lateral patches yellow. Male Wings lightly clouded with brown with the veins dark brown, hair of medium length, oculo-malar area longer than broad; genitalia (Fig. 111) with the gonostylus short and distally near its midline axis deeply concavely rounded, its outer side about a half as long as its inner side with the two inner corners of the distal lobe rounded, the margin between them concave, penis-valve head weakly recurved, the recurved section about as long as broad at the base (cf. B. sikkimi, B. validus); hair on the anterior dorsum of the thorax and of the side of the thorax and scutellum either golden yellow or grey-white, T 1 either yellow or if nearly white then T 2 anterio-laterally also nearly white, on the scutellum in the middle the yellow only shallowly divided by black.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	materials_examined	Material sequenced in Fig. 12 CHINA • 1 ♀ (worker); Yunnan, Lijiang; 27.0156 ° N, 100.1714 ° E; 12 Aug. 2017; H. Liang leg.; KIB seq: # 2018064; KIB: AG # 089 • 1 ♁; Xizang, Zhowagoin; 28.3469 ° N, 98.0102 ° E; 3 Sep. 2014; Q. - T. Wu leg.; IOZ seq: FPW 30; IOZ: AG # 187 • 1 ♀ (worker); Sichuan, Queershan; 29.5718 ° N, 98.1847 ° E; 5 Aug. 2018; Z. Ren et al. leg.; KIB seq: QESH 201020; KIB: AG # 143 • 1 ♀ (worker); Xizang, Dongdashan; 29.5718 ° N, 98.1847 ° E; 21 Jul. 2018; Z. Ren et al. leg.; KIB seq: DDSM 202013; KIB: AG # 087 • 1 ♁; Xizang, Zhowagoin; 28.3469 ° N, 98.0102 ° E; 2 Sep. 2014; Q. - T. Wu leg.; IOZ seq: FPW 24; IOZ: AG # 183 • Yunnan; IAR seq: HJF 6; IAR: AG # 090. Additional sequences in Fig. 10 and haplotype duplicates CHINA • 1 ♀ (worker); Yunnan, Lijiang; 27.0156 ° N, 100.1714 ° E; 2 Sep. 2012; Y. Zhao leg.; KIB seq: # 218628241; KIB: AG # 108 • 1 ♀ (worker); Xizang, Zhowagoin; 28.3469 ° N, 98.0102 ° E; 3 Sep. 2014; Q. - T. Wu leg.; IOZ seq: FPW 31; IOZ: AG # 186 • 1 ♁; Xizang, Zhowagoin; 28.4853 ° N, 97.3634 ° E; 30 Aug. 2014; Q. - T. Wu leg.; IOZ seq: FPW 29; IOZ: AG # 188 • 1 ♀ (worker); same collection data as for preceding but 31 Aug. 2014; IOZ seq: FPW 28; IOZ: AG # 182 • 1 ♀ (worker); Xizang, Zhowagoin; 28.3469 ° N, 98.0102 ° E; 3 Sep. 2014; Q. - T. Wu leg.; IOZ seq: FPW 26; IOZ: AG # 189 • 1 ♀ (worker); same collection data as for preceding but 2 Sep. 2014; IOZ seq: FPW 25; IOZ: AG # 190 • 3 ♁♁; Xizang; 29.6114 ° N, 94.6441 ° E; 10 Aug. 2012; IAR seq: ZA 3 ZA 4 ZA 5; IAR: AG # 010 to AG # 012 • 3 ♀♀ (workers); Xizang, Lulang; 29.6414 ° N, 94.6982 ° E; 26 Jul. 2018; Z. Ren et al. leg.; KIB seq: SJLSM 108001, SJLSM 104001, SJLSM 104002; KIB: AG # 086, AG # 146, AG # 147 • 1 ♀ (worker); Xizang, Galonglashan; 29.8219 ° N, 95.7110 ° E; 24 Jul. 2018; Z. Ren et al. leg.; KIB seq: GLLSL 106001; KIB: AG # 148 • 1 ♀ (worker); Yunnan, Lijiang; 27.0156 ° N, 100.1714 ° E; 12 Aug. 2017; H. Liang leg.; KIB seq: # 2018063; KIB: AG # 103 • 1 ♀ (worker); Xizang, Markam; 29.7432 ° N, 95.6753 ° E; 2 Sep. 2014; Q. - T. Wu leg.; IOZ seq: FPW 32; IOZ: AG # 191.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	distribution	Global distribution Hengduan: China (Qinghai, Xizang, Yunnan, Sichuan, Gansu), Myanmar: IAR, IOZ, KIB, NHMUK, PW, SC, ZMHB. This species is recorded at elevations of 2600 ‒ 4500 m in Sichuan, mostly above the tree line and into the subalpine zone (especially rich in Rhododendron species) (Williams et al. 2009).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790A04139EC7FE7EFDBAF832.taxon	biology_ecology	Behaviour Male eye not obviously enlarged relative to female eye: males are expected to show ‘ patrolling’ behaviour in search of mates (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	description	Figs 70 ‒ 73, 112	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	discussion	Species-taxon concept and variation The taxon concept of the species B. validus here agrees with the recent interpretation (Williams 2022 a) that it is separate from the taxon concept of the species B. nobilis (see the comments above on B. nobilis), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that B. validus, B. sikkimi, and B. nobilis are separate species (Williams 2022 a). Morawitz’s taxon haemorrhoidalis, with the oculo-malar area longer than for B. mastrucatus, matches closely with B. validus. The collection locality for one worker was given by Morawitz as ‘ Gan-su’ (used in a much broader sense then than currently) and this individual was later referred to by Skorikov (1933 b) is consistent with B. validus. It is intriguing that Skorikov (1933 b) mentioned B. validus in his comments on morawitziides without giving any distinguishing characters. The other location for haemorrhoidalis given by Morawitz is ‘ Ordoss’ (= Mu Su Shamo, referring to the Ordos desert, in Shaanxi), which is unexpected because it is at a much lower elevation and mostly more arid. This raises the possibility that Morawitz also had some rare paler individuals of the taxon beresovskii (not seen; but see the comments on B. grahami) in front of him, although this species has a shorter oculo-malar area that does not match Morawitz’s description, which is more likely to refer to B. (Melanobombus) pyrosoma Morawitz, 1890. Variation of B. validus is shown in the colour-pattern diagrams in Figs 65 ‒ 68. Bombus validus, with its black-thorax and largely-orange-metasoma colour pattern in northern Hengduan, appears to mimic the abundant B. (Melanobombus) pyrosoma (Williams 2007: close to fig. 5 m).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	materials_examined	Type material Bombus validus Friese, 1905: 510. Syntype (Tkalců 1987: 60): ♀ (queen) ‘ Turkestan Kashgar’, but probably Sichuan (ZMHB). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	diagnosis	Morphological diagnosis Female Wings nearly clear with the veins dark brown, hair of medium length, oculo-malar area longer than broad, oculo-ocellar area sparsely punctured with few scattered medium-sized punctures with few small punctures between them with broad smooth areas (cf. B. sikkimi, B. nobilis); hair of the side of the thorax and scutellum predominantly black with only a few scattered grey hairs, T 2 ‒ 3 with only a few black hairs along the midline. Male Wings nearly clear with the veins dark brown, hair of medium length, oculo-malar area longer than broad; genitalia (Fig. 112) with the gonostylus short and distally near its midline axis concavely rounded, its outer side about a half as long as its inner side with the two inner corners of the distal lobe rounded, the margin between them nearly straight, penis-valve head weakly recurved, the recurved section about as long as broad at the base (cf. B. sikkimi, B. nobilis); hair of the side of the thorax and scutellum varying from predominantly black to predominantly grey-white, T 2 ‒ 3 with only a few black hairs along the midline.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	materials_examined	Material sequenced in Fig. 12 CHINA • 3 ♀♀ (workers); Gansu, Zhagana; 34.2477 ° N, 103.1329 ° E; 24 Jul. 2009; JH seq: PWB 11, PWB 14, PWB 16; IAR: AG # 039, 091, 092. Additional sequences in Fig. 10 and haplotype duplicates CHINA • 1 ♁; Gansu, Shangbaimu Xia; 36.9620 ° N, 102.8209 ° E; 14 Aug 2011; JH seq: PWZA 1; IAR: AG # 008.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	distribution	Global distribution Northern Hengduan: China (Sichuan, Gansu): IAR, IOZ, PW, ZIN, ZMHB. This species has an unusually restricted distribution, recorded at elevations of 2196 ‒ 3524 m in Sichuan and Gansu, mostly above the tree line and into the subalpine zone (Williams et al. 2009; An et al. 2014).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867908041D9EBAFEEBFDBAFB3C.taxon	biology_ecology	Behaviour Male eye not obviously enlarged relative to female eye: males are expected to show ‘ patrolling’ behaviour in search of mates (Williams 1991).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	description	Figs 6, 74 ‒ 75, 113	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	discussion	Species-taxon concept and variation The taxon concept of the species B. genalis here agrees with the long-standing interpretation (Williams 2022 a) that it is separate from the taxon concepts of other species in the subgenus Alpigenobombus, based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that B. genalis and B. breviceps are separate species. The available COI-barcode-like sequences may all be low-divergence neonumts (Fig. 11). No substantial colour-pattern variation is known for B. genalis (Figs 74 ‒ 75). Bombus genalis, with its black with orange-legged colour pattern, appears to mimic the common B. (Pyrobombus) flavescens Smith, 1852, and B. (Melanobombus) eximius Smith, 1852 (Williams 2007: fig. 5 a).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	materials_examined	Type material Bombus genalis Friese, 1918: 84. Syntypes: ♀♀ (queen and worker) Sikkim, India (ZMHB). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	diagnosis	Morphological diagnosis Female Wings pale yellow with the veins light orange (unique within the subgenus Alpigenobombus), hair short, oculo-malar area shorter than broad, clypeus in its central area shining with only a few small and large widely-spaced punctures (cf. B. breviceps); mid and hind tibiae exoskeleton and hair bright orange, hair of the thoracic dorsum and T 1 black. Male Wings pale yellow with the veins light orange (unique within the subgenus Alpigenobombus), hair short, oculo-malar area shorter than broad; genitalia (Fig. 113) with the gonostylus nearly equally short on both its outer side and its inner side but with the distal lobe projecting inwards as a long broadly triangular acutely pointed process, triangular in section (cf. B. breviceps, B. grahami); mid and hind tibiae exoskeleton and hair bright orange, the hair of the scutellum and T 1 predominantly black.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	materials_examined	Material sequenced in Fig. 12 CHINA • 1 ♀ (worker); Xizang, Linzhi; 29.5710 ° N, 95.4633 ° E; 29 Aug. 2015; Q. - T. Wu leg.; IOZ seq: FPW 12; IOZ: AG # 184. Additional sequences in Fig. 10 and haplotype duplicates CHINA • 1 ♀ (worker); Yunnan, Guanshizhai; 25.9334 ° N, 98.8008 ° E; 18 Aug. 2011; PW seq: PWB 9; IAR: AG # 040 • 1 ♀ (worker); Xizang, Miyuecun; 29.4065 ° N, 95.3721 ° E; 18 Aug. 2015; Q. - T. Wu leg.; IOZ seq: FPW 13; IOZ: AG # 185. INDIA • 2 ♀♀ (workers); Arunachal Pradesh, Salaya; 27.5969 ° N, 93.851 ° E; 22 Jun. 2018; J. Narah leg.; NCBS seq: BG 463 BG 471; NCBS: AG # 200, AG # 201 • 1 ♀ (worker); Arunachal Pradesh, Hari; 27.5838 ° N, 93.8446 ° E; 27 Sep. 2015; NCBS seq: BE 637; NCBS: AG # 202.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	distribution	Global distribution Himalaya and borders of southern Hengduan: India (Sikkim, Arunachal Pradesh), China (Xizang, Yunnan), Myanmar: IAR, IOZ, NCBS, NHMUK, PW, ZMHB. This is an uncommon species and few individuals have been recorded. This species is recorded at elevations of 1561 ‒ 1852 m in the Himalaya in the middle forest zone (Streinzer et al. 2019).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867907041C9EE7FB2DFAD4F993.taxon	biology_ecology	Behaviour Male eye very slightly enlarged relative to female eye, male mate-searching behaviour unknown.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	description	Figs 7 – 8, 76 ‒ 93, 114	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	discussion	NOTE 1. Acceptance of B. breviceps as the valid name in preference to B. nasutus follows Tkalců (1968) as the First Reviser (ICZN 1999: Article 24). Species-taxon concept and variation The taxon concept of the species B. breviceps here agrees with the long-standing interpretation (Williams 2022 a) that it is separate from the taxon concepts of other species in the subgenus Alpigenobombus, based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that B. breviceps and B. genalis are separate species. The available COI-barcode-like sequences may all be low-divergence neonumts (Fig. 11). The PTP and morphological results (Fig. 12, keys) support and confirm the interpretation that the divergent colour patterns of the taxa breviceps s. str., channicus, and dentatus are conspecific as parts of the species B. breviceps (Tkalců 1968; Williams et al. 2009; Hines & Williams 2012). Variation in the colour-pattern diagrams of B. breviceps in Figs 76 ‒ 93 is arranged for the black-thorax Himalayan individuals approximately from west (Kashmir) to east (Myanmar) and for the orange-banded Chinese and Southeast Asian individuals from north (Gansu) to south (Thailand). Bombus breviceps, with the black-thorax colour pattern in the west (Himalaya) and centre (Hengduan) of its range (Figs 76 ‒ 80, 85 ‒ 89) and the orange-banded colour pattern in the south-east (Southeast Asia: Figs 81 ‒ 84, 90 ‒ 93), appears to mimic the abundant B. (Orientalibombus) haemorrhoidalis Smith, 1852, and B. (Megabombus) montivagus Smith, 1878 (Williams 2007: fig. 5 h – i). The black-tailed colour patterns occur on the India / Myanmar border (Figs 80, 89) and in southern China (Figs 82 ‒ 83, 91 ‒ 92).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	materials_examined	Type material Bombus breviceps Smith, 1852: 44. Holotype by monotypy: ♀ (worker) Zhejiang, China (NHMUK). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	diagnosis	Morphological diagnosis Female Wings very darkly clouded with brown with the veins dark brown, hair short, oculo-malar area shorter than broad, clypeus in its central area with many small punctures (cf. B. genalis, B. grahami); hair of the thoracic dorsum black, often with orange bands anteriorly and posteriorly, hair of T 1 yellow. Male Wings very darkly clouded with brown with the veins dark brown, hair short; genitalia (Fig. 114) with the gonostylus nearly equally short on both its outer side and its inner side but with the distal lobe projecting inwards as a long narrowly pointed almost spine-like process, rounded in section (cf. B. genalis, B. grahami).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	materials_examined	Material sequenced in Fig. 12 BHUTAN • 1 ♀ (worker); Trashigang, Doksum; 27.4377 ° N, 91.5833 ° E; 23 May 2017; W. Klein leg.; PW seq: NBC 10098; RMNH: AG # 149. CHINA • 1 ♁; Zhejiang, [Huangjiaotang]; 28.805 ° N, 120.961 ° E; GenBank seq: FJ 175356; SC: AG # 069. THAILAND • 1 ♀ (worker); Chiang Mai, Doi Inthanon; 18.545 ° N, 98.516 ° E; 10 Aug. 2016; C. Sinpoo leg.; GenBank seq: MF 582612; BEEP: AG # 155 • 1 ♀ (worker); Chiang Mai, Doi Inthanon; 18.554 ° N, 98.582 ° E; 19 Jul. 2016; C. Sinpoo leg.; GenBank seq: MF 582613; BEEP: AG # 156. Additional sequences in Fig. 10 and haplotype duplicates CHINA • 1 ♀ (worker); Zhejiang, Gaolaoshan; 28.9043 ° N, 120.5496 ° E; 4 Apr. 2021; Y. Chen leg.; CAU seq: TL 01; CAU: AG # 192 • 1 ♀ (worker); Gansu, Qingyugou; 32.7734 ° N, 105.4137 ° E; 23 Jun. 2021; Y. Chen leg.; CAU seq: TL 02; CAU: AG # 193 • 1 ♀ (worker); Guangdong, Nanxiang; 24.7677 ° N, 113.5932 ° E; 18 Jul. 2021; Y. Chen leg.; CAU seq: TL 03; CAU: AG # 194 • 1 ♀ (worker); Zhejiang, Pingding cun; 29.6615 ° N, 120.0088 ° E; 1 Oct. 2020; L. Tian leg.; CAU seq: TL 04; CAU: AG # 195 • 1 ♀ (worker); Hunan, Dazhushan; 26.1662 ° N, 110.1746 ° E; 23 Jul. 2021; L. Tian leg.; CAU seq: TL 05; CAU: AG # 196 • 1 ♀ (worker); Hunan, Nanshan; 26.1513 ° N, 110.1644 ° E; 22 Jul. 2021; L. Tian leg.; CAU seq: TL 06; CAU: AG # 197 • 1 ♀ (worker); Gansu, Liushuwan; 33.0474 ° N, 105.2562 ° E; 21 Jun. 2021; Y. Chen leg.; CAU seq: TL 07; CAU: AG # 198 • 1 ♀ (worker); Guangdong, Nanxiang; 24.7677 ° N, 113.5932 ° E; 18 Jul. 2021; Y. Chen leg.; CAU seq: TL 08; CAU: AG # 199 • same collection data as for preceding; GenBank seq: KP 259094; IOZ: AG # 152 • same collection data as for preceding; GenBank seq: MF 478986; IAR: AG # 153 • Ningxia; IAR seq: NX 2; IAR: AG # 093. INDIA • 1 ♀ (worker); Arunachal Pradesh, Jang; 27.5821 ° N, 91.9704 ° E; 24 Sep. 2015; J. Neumayer leg.; NCBS BE 653 seq: PWB 6; NCBS: AG # 041. THAILAND • 1 ♀; Chiang Mai, Doi Inthanon; 18.5893 ° N, 98.4859 ° E; 21 Sep. 2006; Y. Areeluck leg.; BOLD seq: 3261 H 02; PCYU: AG # 102 • 1 ♀ (worker); Chiang Mai, Doi Suthep; 18.815 ° N, 98.4920 ° E; 26 Jun. 2016; C. Sinpoo leg.; GenBank seq: MF 582617; BEEP: AG # 157 • 2 ♀♀ (workers); Chiang Mai, Doi Inthanon; 18.543 ° N, 98.548 ° E; 19 Jul. 2016; C. Sinpoo leg.; GenBank seq: MF 582618, MF 582619; BEEP: AG # 158, AG # 159 • 1 ♀ (worker); Chiang Mai, Doi Inthanon; 18.545 ° N, 98.516 ° E; 10 Aug. 2016; C. Sinpoo leg.; GenBank seq: MF 582622; BEEP: AG # 160 • 1 ♀ (worker); Chiang Mai, Doi Inthanon; 18.5892 ° N, 98.48872 ° E; 21 Jul. 2015; N. Warrit et al. leg.; CT seq: CT 540; CUNHM: AG # 177 • 1 ♀ (worker); Chiang Mai, Doi Inthanon; 18.5447 ° N, 98.5179 ° E; 21 Jul. 2015; N. Warrit et al. leg.; CT seq: CT 552; CUNHM: AG # 178 • 1 ♀ (worker); Chiang Mai, Khun Chang Khian; 18.8387 ° N, 98.8965 ° E; 18 Jul. 2015; N. Warrit et al. leg.; CT seq: CT 607; CUNHM: AG # 179 • 1 ♀ (worker); Chiang Mai, Doi Kiew Lom; 19.3175 ° N, 98.6006 ° E; 20 Nov. 2017; N. Warrit et al. leg.; CT seq: CT 623; CUNHM: AG # 180 • 1 ♀ (worker); Chiang Mai, Doi Ang Khang; 19.8823 ° N, 99.0437 ° E; 21 Nov. 2017; N. Warrit et al. leg.; CT seq: CT 625; CUNHM: AG # 181.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	distribution	Global distribution One of the most widespread Oriental species, in the Himalaya, Hengduan, Central and southern China, and Southeast Asia: India (Kashmir, Himachal Pradesh, Uttarakhand, Sikkim, Arunachal Pradesh), Nepal, Bhutan, China (Xizang, Yunnan, Sichuan, Gansu, Shaanxi, Chongqing, Hubei, Jiangxi, Hunan, Guizhou, Zhejiang, Fujian, Guangdong, Guangxi), Myanmar, Vietnam, Laos, Thailand: BEEP, CAU, CUNHM, IAR, IOZ, MCSN, NCBS, NHMUK, PCYU, PW, RMNH, SC, USNM, ZMHB. This species is recorded at elevations of 484 ‒ 3000 m in the Himalaya (Williams et al. 2010; Streinzer et al. 2019), and at 270 ‒ 3350 m in the Hengduan and in Shaanxi (Williams et al. 2009; An et al. 2014). It is one of the few bumblebee species that occurs down to lower elevations in subtropical habitats, where it may be relatively common, even inside cities. However, in the warmer habitats it may be active only in the early morning and evening (Yunnan and Guangdong, PW pers. obs.; Thailand, C. Thanoosing pers. obs.).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A438786790604199ED3F9D2FAD4FCA4.taxon	biology_ecology	Behaviour Male eye very slightly enlarged relative to female eye, male mate-searching behaviour unknown.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	description	Figs 9, 94 ‒ 103, 115	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	discussion	NOTE 1. Acceptance of B. grahami as the valid name in preference to B. beresovskii follows Williams (2022 a) as the First Reviser (ICZN 1999: Article 24). Bombus grahami is the name in most widespread use for this species in the literature. Species-taxon concept and variation The taxon concept of the species B. grahami here is modified from the long-standing interpretation (Williams 2022 a), in that it includes the taxon beresovskii, based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that the divergent colour patterns of the taxa grahami s. str. and beresovskii are conspecific (Williams 2022 a). The available COIbarcode-like sequences may all be low-divergence neonumts (Fig. 11). From morphology, the taxon melani (holotype queen Fig. 99) is closely similar to the taxon beresovskii (syntype queen Fig. 100). For the typically coloured individuals of B. grahami, the wings of the males are (unusually among bumblebees) much paler than the wings of the females. The wings are also slightly paler for females of the taxon beresovskii (males not seen). Some females with the beresovskii colour pattern (from northern Sichuan: Figs 99 – 100) have a dark colour pattern that resembles B. angustus, but others (from Shaanxi: Fig. 98) have a light colour pattern that resembles B. validus. One worker (Fig. 98) has a distinct band of grey hairs fringing the thoracic dorsum anteriorly and posteriorly with some grey hairs along the midline. Males of the taxa beresovskii and melani are unknown to us. Bombus grahami females, with the grey-thorax colour pattern in the west (Himalaya) and centre (Hengduan) of its range (Figs 94 ‒ 97), appear to mimic the commoner B. (Orientalibombus) funerarius Smith, 1852 (Williams 2007: fig. 5 g). In contrast, the more extensively black-thorax colour pattern in the north-east (in the hills to the north of the Sichuan Basin: Figs 98 ‒ 100) appears to mimic the abundant B. (Melanobombus) pyrosoma (Williams 2007: close to fig. 5 m). Some of these females (Fig. 98) resemble closely B. validus (Figs 70 ‒ 71) in colour pattern, but can be distinguished easily by the shorter oculo-malar distance of B. grahami.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	materials_examined	Type material Bremus (Alpigenobombus) grahami Frison, 1933 [[ 30] September]: 334. Holotype by original designation: ♀ (worker) Sichuan, China (USNM). Examined. Alpigenobombus beresovskii Skorikov, 1933 a [30 September: Pesenko & Astafurova 2003]: 248. Syntype (unpublished Podbolotskaya lectotype): ♀ (queen) Sichuan, China (ZIN). Examined. Bombus (Alpigenobombus) grahami [subsp.] melani Wang & Yao, 1993: 422. Holotype by original designation: ♀ (queen) Henan, China (IOZ). Examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	diagnosis	Morphological diagnosis Female Wings very darkly clouded with brown with the veins dark brown, hair short, oculo-malar area shorter than broad, clypeus in its central area with only a few large and small widely-spaced punctures (cf. B. breviceps); hair of the thoracic dorsum black, either usually with many grey hairs intermixed so as to appear silvery olive-grey, with the majority of the hair along the longitudinal midline white, or if entirely black, then T 3 predominantly orange-red. Male Wings nearly clear with the veins dark brown, hair short; genitalia (Fig. 115) with the gonostylus half as long on its outer side than on its inner side with the distal lobe slightly reduced and broadly flattened, appearing as a triangular pointed inwardly-projecting process only from the inner aspect (cf. B. genalis, B. breviceps); hair of the thoracic dorsum yellow, often between the wing bases with many black hairs intermixed.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	materials_examined	Material sequenced in Fig. 12 CHINA • 1 ♀ (queen); Sichuan, Taoyuan; 32.6924 ° N, 106.8336 ° E; 21 Sep. 2011; PW seq: PWB 01; IAR: AG # 001 • 1 ♀ (worker); Yunnan, Lijiang; 27.0156 ° N, 100.1714 ° E; 12 Jul. 2017; H. Liang leg.; KIB seq: KIB 030; KIB: AG # 076. Additional sequences in Fig. 10 and haplotype duplicates BHUTAN • 1 ♀ (worker); Thimpu, Dochu-La; [27.4898 ° N, 89.7504 ° E]; 7 Jul. 1996; L. Blommers leg.; BOLD seq: 6880 H 10; RMNH: AG # 094. CHINA • 2 ♀♀ (workers); Yunnan, Lijiang; 27.0156 ° N, 100.1714 ° E; 2 Aug. 2017; H. Liang leg.; KIB seq: KIB 092, KIB 097; KIB: AG # 075, AG # 077 • 2 ♀♀ (workers); same collection data as for preceding but 12 Aug. 2017; KIB seq: KIB 116, KIB 032; KIB: AG # 078, AG # 079 • 1 ♀ (worker); Sichuan, Meigu; 28.329 ° N, 103.129 ° E; GenBank seq: FJ 175354; SC: AG # 096 • Yunnan, Yulong Snow Mountain; [27.047 ° N, 100.261 ° E]; H. Liang leg.; GenBank seq: MT 906009; KIB: AG # 150. INDIA • 1 ♀ (worker); Arunachal Pradesh, 1300 m E of Lama Tukkut; 28.9949 ° N, 95.2926 ° E; 30 Aug. 2017; NCBS seq: BE 782; NCBS: AG # 203. NEPAL • 1 ♁; Janakpur, Dongo Kharka; 27.9022 ° N, 86.2865 ° E; 23 Aug. 1983; I. Kudo leg.; BOLD seq: 1551 H 05; SEHU: AG # 119.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	distribution	Global distribution Himalaya, Hengduan, and into Central China: Nepal, India (Sikkim, Arunachal Pradesh), China (Xizang, Yunnan, Sichuan, Gansu, Shaanxi, Chongqing, Hubei, Henan, Guizhou): IAR, IOZ, KIB, NCBS, NHMUK, PW, RMNH, SEHU, USNM, ZIN. This species is recorded at elevations of 2650 ‒ 2800 m in the Himalaya (Williams et al. 2010; Streinzer et al. 2019), and 858 ‒ 3040 m in the Hengduan and in Shaanxi (Williams et al. 2009; An et al. 2014). Bombus grahami is active in open areas in the middle and upper forest zones, but also inside the forest. Individuals with a black-thorax colour pattern (taxon beresovskii) occur in the hills to the north of the Sichuan basin, where they appear to be extremely rare (only three queens and three workers have been examined: IAR, IOZ, PW) and are not known to co-occur with individuals with the olive-thorax colour pattern (although many more data are needed to clarify the situation).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867903041B9EDCFC85FB10FBF5.taxon	biology_ecology	Behaviour Male eye slightly enlarged relative to female eye, male mate-searching behaviour unknown.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	description	Figs 104 ‒ 105	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	discussion	Species-taxon concept and variation Evidence for the taxon concept of the species B. angustus here provides further support for the interpretation (Williams et al. 2022 b) that this is separate from the taxon concept of the species B. breviceps (Williams 1998), based on: (1) our PTP analysis supports independent species-level coalescents in the COI gene (Fig. 12); corroborated by (2) diagnostic morphological character states (see the keys). The PTP and morphological results (Fig. 12, keys) support the interpretation that B. angustus and B. breviceps are separate species (Williams et al. 2022 b). The available COI-barcode-like sequences may all be low-divergence neonumts (Fig. 11). The female mandible of B. angustus has been imaged by Starr (1992: fig. 6 a). No substantial colour-pattern variation of B. angustus is known (Figs 104 ‒ 105). Bombus angustus, with its black with red-tailed colour pattern from Taiwan, appears to mimic the commoner local B. (Megabombus) trifasciatus Smith, 1852 (Williams 2007: fig. 5 e).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	materials_examined	Type material Bombus (Alpigenobombus) angustus Chiu, 1948: 59. Holotype by original designation: ♁ Taiwan (TARI). Photographic images examined.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	diagnosis	Morphological diagnosis Female Wings very weakly clouded with brown, nearly clear, with the veins dark brown (cf. B. genalis, B. breviceps, B. grahami), hair of medium length, oculo-malar area shorter than broad, clypeus in its central area with many small punctures (cf. B. genalis, B. grahami); mid and hind tibiae with the exoskeleton and hair predominantly black, hair of the thoracic dorsum and T 1 ‒ 3 black, T 4 ‒ 6 orange-red. Male Examined from photographic images, also described and genitalia illustrated by Chiu (1948: fig. 1) (genitalia of the type not found by C. - F. Lee, pers. com.): wings very weakly clouded with brown, nearly clear, with the veins light brown; genitalia with the gonostylus nearly equally short on both its outer side and its inner side but with the distal lobe projecting inwards as a long broadly triangular pointed process; hair of the thoracic dorsum and T 1 ‒ 3 ‘ black’ (on the type specimen, which is in poor condition, hair of the thorax in part? faded to brown), T 4 ‒ 7 orange-red.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	materials_examined	Material sequenced in Fig. 12 TAIWAN • 1 ♀ (worker); Chiayi, Alishan; [23.446 ° N, 120.794 ° E]; 14 Jul. 2020; Y. Lin leg.; GenBank seq: MZ 831894; TFRI: AG # 074.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	distribution	Global distribution Taiwan endemic (Williams et al. 2022 b): PW, TARI, TFRI. This is a rare species with few individuals recorded: Chiu (1948) lists three individuals from three localities; Starr (1992) lists 21 individuals from eight localities (mapped in Starr: fig. 7) among a total sample of 4555 Taiwanese bumblebees (<0.5 %; from his map fig. 7, most B. angustus appear to be from elevations around 1000 ‒ 2000 m, although one individual in the north may be from ca 100 m).	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
1A4387867901041A9EECFB77FE02FA8A.taxon	biology_ecology	Behaviour Male behaviour not seen.	en	Williams, Paul H., An, Jiandong, Dorji, Phurpa, Huang, Jiaxing, Jaffar, Saleem, Japoshvili, George, Narah, Jaya, Ren, Zongxin, Streinzer, Martin, Thanoosing, Chawatat, Tian, Li, Orr, Michael C. (2023): Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae). European Journal of Taxonomy 892 (1): 1-65, DOI: https://doi.org/10.5852/ejt.2023.892.2283, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2283/9851
