Mastrucatobombus Krueger, 1917
Nobilibombus Richards, 1968
Bumblebees with big teeth: revising the subgenus Alpigenobombus with the good, the bad and the ugly of numts (Hymenoptera: Apidae)
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.
European Journal of Taxonomy
2023
2023-09-27
892
1
1
65
Skorikov, 1914
Skorikov
1914
[659,1050,1746,1773]
Insecta
Apidae
Bombus
Animalia
Hymenoptera
19
20
Arthropoda
subGenus
Alpigenobombus
= Mastrucatobombus Krueger, 1917. = Nobilibombus Richards, 1968.
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. Fig. 18.Most likely ancestral ranges reconstructed for all extant currently recognised species from the dispersal model in Fig. 17, using the model DIVALIKE+J in S-BioGeoBEARS from a sample of 10000 trees from BEAST used to make the estimates of species phylogenies from the six genes in Fig. 16. Letters represent the area units in Fig. 17: letter combinations at terminals show species’ current distributions (key lower left show the colour codes used for the principal areas of endemism, other colours not in the key represent combinations of areas e.g., darker green for area combination CDE, with black for a mixture of other areas); letter combinations at nodes show the most likely reconstructions for ancestral distributions; pies at nodes indicate the percentage of solutions for that node in which solutions occur (area E does not occur alone in any ancestor distribution). Numbers on the x-axis are ages in Ma before present. Outgroup shown in grey. The subgenus MastrucatobombusKrueger, 1917is a synonym of Alpigenobombus. The subgenus Nobilibombus Richards, 1968has 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 Nobilibombuswas published initially without fixation of a typespecies ( Skorikov 1933b), or as a junior synonym ( Bischoff 1936; Milliron 1961), so the first valid publication is by Richards (1968). The species of Nobilibombusare recognised here as the nobilis-group of species.
wurflenii-group Bombus wurflenii Radoszkowski, 1860 s. str. = alpigenus Morawitz, 1874 = uralicus( Tkalců, 1969) Bombus mastrucatus Gerstaecker, 1869 stat. rev. = brevigena Thomson, 1870 = flavicantePérez, 1890 = pyrenaicus Vogt, 1909 = collaris Friese, 1909 = transitorius Friese, 1909 = lutescens Friese, 1909 = luteus Friese, 1909 = tirolensisFriese, 1911 = apfelbeckiReinig in Reinig & Rasmont, 1988 = knechteliReinig in Reinig & Rasmont, 1988 = balcaniensis Rasmont, Ghisbain & Terzo, 2021 kashmirensis-group Bombus kashmirensis Friese, 1909 = stramineus Friese, 1909 = tetrachromus Cockerell, 1909 = pulcherrimus( Skorikov, 1914) = meinertzhageni Richards, 1928 Bombus rainaiWilliams, 2022 nobilis-group Bombus sikkimi Friese, 1918 stat. rev. Bombus nobilis Friese, 1905 s. str. = xizangensis Wang, 1979 = chayaensis Wang, 1979 Bombus validus Friese, 1905 stat. rev. = morawitziides(Skorikov, 1933) breviceps-group Bombus genalis Friese, 1918 Bombus breviceps Smith, 1852 s. lat. = nasutus Smith, 1852 = dentatus Handlirsch, 1888 = simulus Gribodo, 1892 = channicus Gribodo, 1892 = laticeps Friese, 1905 = orichalceus Friese, 1916 = brevigenalis Friese, 1918 = rufocognitus Cockerell, 1922 = pretiosus Bischoff, 1936 = cantonensis Bischoff, 1936 = coloricontrarius Tkalců, 1968 = colorilaetus Tkalců, 1968 = vicinus Tkalců, 1968 = bischoffiellus Tkalců, 1977 Bombus grahami( Frison, 1933) s. lat. = beresovskii(Skorikov, 1933) syn. nov. = melani Wang & Yao, 1993syn. nov. Bombus angustus Chiu, 1948
Key to species for females of the subgenus Alpigenobombus Future identification of species of the subgenus Alpigenobombuswill 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.
1. Oculo-malar area (= malar area) shorter than broad (OM length measured as the minimum distance between the eye ventrally and concave margin between the mandibular condyles (hinges); OM breadth measured as the distance between and including the mandibular condyles) ....................... 2 – Oculo-malar area longer than broad ................................................................................................. 9 2. Clypeus adjacent to the labrum not raised in the centre but flat so that the transverse anterior groove is not interrupted, this central area uniformly with many small punctures and not shining (Europe, Turkey, Caucasus, Urals) .................................................................................................................. 3 – 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 (Asia east of Pakistan) ...................................................................................................................... 4 3. Ocello-ocular area (= ocellocular area) with the punctures anterior to the unpunctured area (adjacent to the lateral ocellus and between it and towards the inner ocular margin) less dense than posterior to it; hair of the thoracic dorsum eitherentirely black orsometimes with bands anteriorly and posteriorly yellow (Figs 26‒30) (Europe, in Scandinavia, Pyrenees, Alps, Carpathians, Greece) ..... ........................................................................................ B. mastrucatus Gerstaecker, 1869 stat. rev. – Ocello-ocular area with the punctures anterior to the unpunctured area at least as dense as posterior to it; hair of the thoracic dorsum black with bands anteriorly and usually posteriorly white (Figs 19‒22) ( Turkey, Caucasus, Urals) .................................................... B. wurflenii Radoszkowski, 1860 s. str. 4. Wings eitherclear orvery lightly clouded with brown, T1-2 with hair predominantly (nearly completely) eitherwhite oryellow (occurring at elevations above 3000 m) ................................... 5 – Wings eitherdarkly clouded with brown or ifpale yellow ornearly clear thenT1-2 with hair black (occurring at elevations below 3000 m) ........................................................................................... 6 5. Oculo-ocellar area in its outer half with large and medium punctures; hair of T5 orange with white tips, the side of the thorax in at least its upper half and the scutellum eitherwhite oryellow (Figs 35‒41) (Himalaya, Hengduan) ................................................... B. kashmirensis Friese, 1909 – Oculo-ocellar area in its outer half with only a few small shallow punctures, any larger punctures confined to the eye margin; hair of T5 entirely orange without paler tips, the side of the thorax in no more than its upper third white and the scutellum predominantly black (Figs 49‒51) (Kashmir) ..... ..................................................................................................................... B. rainaiWilliams, 2022 6. Wings very darkly clouded with brown, hair of T1 eitheryellow orgrey orwhite ......................... 7 – Wings eitherpale yellow ornearly clear, hair of T1 black .............................................................. 8 7. Clypeus in its central area with many small punctures, many spaced by only their own widths; hair of the thoracic dorsum black, often with orange bands anteriorly and posteriorly, T1 yellow (Figs 76‒84) ................................................................................................ B. breviceps Smith, 1852 – Clypeus in its central area with only a few large and small punctures, most spaced by more than their own widths; hair of the thoracic dorsum black, eitherusually with many grey hairs intermixed so as to appear silvery olive-grey, with the majority of the hair along the longitudinal midline white, or ifentirely black thenT1 predominantly grey-white (Figs 94‒100) ............. B. grahami( Frison, 1933) 8. Mid and hind tibiae with exoskeleton and hair bright orange; wings pale yellow with the veins orange; clypeus in its central area with widely-scattered large punctures and only a few micropunctures that are more widely-spaced than their own widths (Fig. 74) (Himalaya) ......................... ........................................................................................................................ B. genalis Friese, 1918 – Mid and hind tibiae with the exoskeleton and hair predominantly black; wings nearly clear with the veins black; clypeus in its central area with widely-scattered large punctures and many small punctures that are as widely spaced as their own widths (Fig. 104) ( Taiwan)................................... ....................................................................................................................... B. angustus Chiu, 1948 9. Oculo-ocellar area sparsely punctured with few scattered medium-sized punctures with few small punctures between them with broad smooth areas; hair of the side of the thorax and scutellum predominantly black, T2–3 with only a few black hairs along the midline (Figs 70‒71) ( Sichuan, Gansu) .............................................................................................. B. validus Friese, 1905 stat. rev. – Oculo-ocellar areas densely punctured with scattered medium-sized punctures with many small punctures between them and lacking smooth areas; hair of the side of the thorax and scutellum predominantly eithergrey oryellow, T2–3 usually with a few black hairs along the midline and a transverse band of black hairs (Himalaya, Hengduan) ................................................................... 10 10. Oculo-ocellar area anterio-lateral to the lateral ocellus with large and small punctures sufficiently separated to have flat shining areas in between (= interspaces) (at least for queens, differences weaker for workers); hair on the anterior dorsum of the thorax andof the side of the thorax andscutellum eithergolden yellow orgrey-white, on the scutellum the yellow hair anteriorly incompletely divided in the middle by black, T1 yellow, T2 eitherpredominantly black orwith anterior lateral patches yellow (Figs 61‒66) (Hengduan) .................................................................... B. nobilis Friese, 1905 – Oculo-ocellar area anterio-lateral to the lateral ocellus with large, medium and many small punctures almost coalescing and without flat shining areas in between (= interspaces) (at least for queens, differences weaker for workers); hair on the anterior dorsum of the thorax andof the side of the thorax andscutellum andT1 eithergrey orolive-yellow, on the scutellum the pale hair anteriorly often deeply divided in the middle by a triangle of black, T2 eitherpredominantly yellow orwith large anterior lateral patches yellow (Figs 54‒57) (Himalaya) ....... B. sikkimi Friese, 1918 stat. rev.
Key to species for males of the subgenus Alpigenobombus
1. Oculo-malar area shorter than broad (OM length measured as the minimum distance between the eye ventrally and concave margin between the mandibular condyles (hinges); OM breadth measured as the distance between and including the mandibular condyles); genitalia with the gonocoxa with the distal (posterior) inner corner not projecting inwards ( Figs 106‒109, 113‒115) ....................... 2 – Oculo-malar area longer than broad; genitalia with the gonocoxa with the distal (posterior) inner corner projecting strongly inwards ( Figs 110‒112) .......................................................................... 9 2. From the region including European, Turkish, Caucasus and Ural mountains ................................ 3 – From the region including the mountains of Asia east of Pakistan.................................................. 4 3. Hair of the thoracic dorsum between the wing bases predominantly black, usually with few or no yellow hairs (Figs 31‒34) (Europe, especially in Scandinavia, Pyrenees, Alps, Carpathians, Greece) ........................................................................... B. mastrucatus Gerstaecker, 1869 stat. rev. – Hair of the thoracic dorsum between the wing bases eitherpredominantly yellow orwith many yellow hairs (Figs 23‒25) ( Turkey, Caucasus, Urals) .......... B. wurflenii Radoszkowski, 1860 s. str. 4. Genitalia with the penis valve distally recurved as a sickle by more than 90° with the hook much longer than broad ( Figs 108‒109) (occurring at elevations above 3000 m) ..................................... 5 – Genitalia with the penis valve distally recurved as a sickle by just 90° with the hook scarcely longer than broad ( Figs 113‒115) (occurring at elevations below 3000 m) ................................................ 6 5. Hair of the side of T5‒6 orange at the base but white-tipped, T3 predominantly black, the side of the thorax eitheryellow orwhite, the pale hair reaching to the midleg bases (Figs 42‒48); genitalia with the penis valve recurved head broad, the recurved section narrowing only just before the tip, gonostylus on its outer side less than half as long as on its inner side ( Fig. 108) (Himalaya including Kashmir, Hengduan) ............................................................................ B. kashmirensis Friese, 1909 – Hair of the side of T5‒6 orange, T3 predominantly orange, the side of the thorax with black and white intermixed, but often predominantly black in its lower half (Figs 52‒53); genitalia with the penis valve recurved head narrow, the recurved section tapering gradually to the tip, gonostylus on its outer side about half as long as on its inner side ( Fig. 109) (Kashmir) ......................................... ..................................................................................................................... B. rainaiWilliams, 2022 6. Wings very darkly clouded with brown (Figs 85‒93) ................................ B. breviceps Smith, 1852 – Wings eitherpale yellow ornearly clear .......................................................................................... 7 7. Hair of the thoracic dorsum predominantly black ............................................................................ 8 – Hair of the thoracic dorsum yellow, often between the wing bases with many black hairs intermixed (Figs 101‒103) .......................................................................................... B. grahami( Frison, 1933) 8. Wings pale yellow with the veins orange, the mid and hind tibiae with exoskeleton and hair bright orange (Fig. 75) (Himalaya) .......................................................................... B. genalis Friese, 1918 – Wings weakly clouded with brown with the veins brown, exoskeleton and hair of the mid and hind tibiae predominantly black (Fig. 104) ( Taiwan)........................................... B. angustus Chiu, 1948 9. Hair of the side of the thorax and scutellum predominantly black, T2‒3 with eithervery few orno black hairs along the midline and no transverse band of black hairs (Figs 72‒73); genitalia with the gonostylus on its outer side less than half its greatest length ( Fig. 112) ( Sichuan, Gansu) ................ .......................................................................................................... B. validus Friese, 1905 stat. rev. – Hair of the side of the thorax and scutellum predominantly eithergrey oryellow, T2‒3 usually with many black hairs along the midline and at least a narrow transverse band of black hairs eitherposteriorly on T2 and/oranteriorly on T3; genitalia with the gonostylus on its outer side more than half its greatest length ( Fig. 110‒111) (Himalaya, Hengduan) ....................................................... 10 10. Hair on the anterior dorsum of the thorax andof the side of the thorax andscutellum eithergolden yellow orgrey-white, T1 eitheryellow or ifnearly white thenT2 anterio-laterally also nearly white, on the scutellum in the middle the yellow only shallowly divided by black (Fig. 67‒69); genitalia with the gonostylus with the distal edge concave ( Fig. 111) (Hengduan) ..... B. nobilis Friese, 1905 s. str. – Hair on the anterior dorsum of the thorax andof the side of the thorax andscutellum andT1 eithergrey-white ordull olive-yellow, T2 anterio-laterally yellow, on the scutellum in the middle the grey or yellow nearly completely divided by black (Figs 58‒60); genitalia with the gonostylus with the distal edge weakly convex ( Fig. 110) (Himalaya) ........................... B. sikkimi Friese, 1918 stat. rev.