Bombus

B. impatiens queen resting.

Pyrobombus ecology and behaviour   HABITAT: Mountain-meadow, forest, grassland, semi-desert, and tropical montane forest. This subgenus includes species of both the arctic tundra and tropical hill forests.   FOOD-PLANTS: Short to medium tongue-length bumblebees visiting shallow to medium flowers. Workers often have particularly small body sizes and are more likely than many other bumblebee species to visit flowers where they have to hang upside down. They also provide 'buzz' pollination.   NESTING BEHAVIOUR: Nests underground or on the surface. Non-pocket makers. Colonies are often small with a short cycle. Some species have more than one colony cycle per year.   MATE-SEARCHING BEHAVIOUR: Males patrol circuits of scent marks.

Subgenus PYROBOMBUS Dalla Torre
Bombus (Pyrobombus) Dalla Torre, 1880:40, type-species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by monotypy
Bombus (Poecilobombus) Dalla Torre, 1882:23, type-species Bombus sitkensis Nylander by subsequent designation of Sandhouse, 1943:589
[Bombus (Pyrrhobombus) Dalla Torre, 1882:28, incorrect subsequent spelling]
Bombus (Pyrrhobombus) Dalla Torre, 1896:503, unjustified emendation
Bombus (Pratobombus) Vogt, 1911:49, type-species Apis pratorum Linnaeus (= Bombus pratorum (Linnaeus)) by subsequent designation of Frison, 1927:67
[Bombus (Pratibombus) Ball, 1914:78, incorrect subsequent spelling]
Bombus (Anodontobombus) Krüger, 1917:61,65 (proposed as a section name but stated by Milliron, 1961:53, to be synonymous with his concept of the subgenus Pyrobombus Dalla Torre), type-species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by subsequent designation of Williams, 1991:69
Bombus (Uncobombus) Vogt in Krüger, 1917:65 (proposed as a group name but stated by Milliron, 1961:53, to correspond to his concept of Pyrobombus Dalla Torre), type-species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by subsequent designation of Williams, 1991:69
Bombus (Lapponicobombus) Quilis-Pérez, 1927:19, type-species Apis lapponica Fabricius (= Bombus lapponicus (Fabricius)) by subsequent designation of Milliron, 1961:58
[Bombus (Hypnorubombus) Quilis-Pérez, 1927:19, incorrect original spelling]
[Bombus (Laponicobombus) Quilis-Pérez, 1927:63, incorrect original spelling]
Bombus (Hypnorobombus) Quilis-Pérez, 1927:97, type-species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by monotypy
Bremus (Pressibombus) Frison, 1935:342, type-species Bremus pressus Frison (= Bombus pressus (Frison)) by original designation
Pratibombus Skorikov, 1937:59, unjustified emendation
Bombus (Pressobombus) Kruseman, 1952:102, unjustified emendation

Part of the bumblebee phylogenetic tree including available Pyrobombus species from an analysis of DNA sequence data for five genes (Cameron et al. 2007 [pdf]). Values above branches are Bayesian posterior probabilities, values below branches are parsimony bootstrap values. Alternative resolution from parsimony analysis is shown with dotted lines.

vagans-group of species

Bombus (Pr.) vagans Smith
vagans Smith, 1854:399, examined
?cockerelli Franklin, 1913:356
5 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic, E Nearctic Regions.

Bombus (Pr.) centralis Cresson
centralis Cresson, 1864:41
5 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) flavifrons Cresson
pleuralis Nylander, 1848:231, examined
flavifrons Cresson, 1863:105
7 names

TAXONOMIC STATUS: B. pleuralis and B. flavifrons have usually been regarded as conspecific, but they were regarded as separate species by Franklin (1913), and more recently Poole (1996) lists them as separate species without explanation. In my opinion, the lectotype of B. pleuralis designated by Milliron (1960:95) is an individual of the dark form of B. flavifrons (see descriptions of variation by e.g. Stephen, 1957; Thorp et al., 1983). See also the comments on B. mixtus.

NOMENCLATURE: B. pleuralis is the oldest available name for this species.

Although B. pleuralis is the oldest available name for the present interpretation of this species, the name B. flavifrons has been in common use for the species since 1950 (e.g. Stephen, 1957; Thorp, 1969, 1970; Plowright & Stephen, 1973; Macior, 1975; Sakagami, 1976; Hurd, 1979; Plowright & Owen, 1980; Thorp et al., 1983; Laverty & Harder, 1988). I know of no publications using the name B. pleuralis since 1950, apart from the list by Poole (1996). It is suggested that, in the interests of stability (ICZN, 1999: Article 23), prevailing usage should be maintained (in prep.).

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) caliginosus (Frison)
caliginosus (Frison, 1927:376 [Bremus]) examined
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) vandykei (Frison)
vandykei (Frison, 1927:375 [Bremus])
cascadensis (Milliron, 1970a:382 [Pyrobombus]) examined
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

lapponicus-group of species

Bombus (Pr.) melanopygus Nylander
melanopyge Nylander, 1848:236
Edwardsii Cresson, 1878:184
melampygus Handlirsch, 1888:231, unjustified emendation
[melanopygus Viereck, 1904:99, incorrect subsequent spelling]
melanopygus Franklin, 1913:334, justified emendation
10 names

TAXONOMIC STATUS: B. melanopygus and B. edwardsii were shown by Owen & Plowright (1980) to differ by a single pair of alleles at one locus controlling the colour of the pubescence on gastral terga II-III. There can be little doubt that they are conspecific (Owen et al., 2010).

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) sylvicola Kirby
sylvicola Kirby, 1837:272
5 names

TAXONOMIC STATUS: B. sylvicola is morphologically closely similar to B. lapponicus, and it has been suggested repeatedly that they are conspecific (e.g. Sladen, 1919; Skorikov, 1922a, 1937; Pittioni, 1942, 1943; Thorp, 1962; Thorp et al., 1983).

Evidence from comparisons of DNA sequences from the 16S gene is not strong but consistent with the two taxa being separate species (Cameron et al., 2007 [pdf]). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as two separate species. See also the comments on B. monticola.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Arctic, W Nearctic Regions.

Bombus (Pr.) lapponicus (Fabricius)
lapponica (Fabricius, 1793:318 [Apis])
zhaosu Wang, 1985:162, examined
44 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic, Arctic Regions.

Bombus (Pr.) monticola Smith
montanus Smith, 1844:549, not of Lepeletier, [1835]:463 (= B. ruderarius (Müller))
monticola Smith, 1849:lx, replacement name for montanus Smith, 1844:549
lugubris Sparre-Schneider, 1909:155, not of Kriechbaumer, 1870:159 (= B. maxillosus (Klug))
norvegicus Friese, 1911:571
scandinavicus Friese, 1912:684, replacement name for lugubris Sparre-Schneider, 1909:255
29 names

TAXONOMIC STATUS: B. scandinavicus (= B. monticola) and B. lapponicus are names that were applied initially to two colour forms in Scandinavia.

Løken (1973) reported that these two taxa overlap narrowly in distribution and intergrade. However, they have been found to differ consistently (for the samples analysed) in the composition of cephalic secretions (Bergström & Svensson, 1973; Svensson & Bergström, 1977). Svensson (1973, 1979) also described subtle differences in morphological characters, although other morphological studies by Løken (1973) and Pekkarinen (1979) found no distinct differences. Pekkarinen (1982, in litt.) now believes that they are separate species. However, from a study of sequence data from COI genes, Koulianos (1999) suggests that they are likely to be conspecific.

It remains possible that there is a hybrid zone in Scandinavia where the colour forms B. monticola and B. lapponicus intergrade, with some gene flow. In this case, depending on the species concept embraced, these taxa might be considered conspecific (see the comments on B. ruderatus). Until further evidence is available, I shall continue to treat them as separate species.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region.

INTRODUCTIONS: Until the twentieth century B. monticola was not known in Ireland, where it is now established (see references in Alford, 1975, 1980) (see comments on B. pratorum).

NOTES on this species in Britain.

Bombus (Pr.) bimaculatus Cresson
bimaculatus Cresson, 1863:92
4 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: E Nearctic Region.

Bombus (Pr.) bifarius Cresson
bifarius Cresson, 1878:185
andamanus Gribodo, 1882:268, examined
fernaldi Franklin, 1911:157, not a replacement name
9 names

TAXONOMIC STATUS: B. andamanus was described as originating from 'Andaman' (= Andaman Islands, Indian Ocean), but appears to be a mislabelled queen of B. bifarius (Tkalcu, 1966). I have examined this specimen and agree with this identification (i.e. contrary to Richards, 1929b, it is not a species of the subgenus Bombus s. str.).

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) ternarius Say
ternàrius [ternarius] Say, 1837:414
ornatus Smith, 1854:398, examined
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic, E Nearctic Regions.

Bombus (Pr.) huntii Greene
Huntii Greene, 1860:172
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic, S Nearctic Regions.

Bombus (Pr.) vosnesenskii Radoszkowski
Vosnesenskii Radoszkowski, 1862:589
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

INTRODUCTIONS: In 1999 I was shown a photo of a specimen of this species that had been collected in Queensland, Australia, where it must have been introduced. See Thorp (2003).

Bombus (Pr.) impatiens Cresson
impatiens Cresson, 1863:90
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: E Nearctic Region, W Nearctic border.

INTRODUCTIONS: This species has been introduced into Mexico and California, which are outside its native range (Thorp, 2003).

Bombus (Pr.) ephippiatus Say
ephippiàtus [ephippiatus] Say, 1837:414
formosus Smith, 1854:403, examined
lateralis Smith, 1879:134, examined
wilmattæ [wilmattae] Cockerell, 1912:21, examined
alboniger Franklin, 1915:409, examined
folsomi (Frison, 1923:322 [Bremus]) examined
13 names

TAXONOMIC STATUS: B. folsomi was described as originating from 'Kina Bala / N. Borneo' (= Gunung Kinabalu, Sabah), but appears to be a mislabelled queen of B. ephippiatus, probably from Costa Rica or Panama (Starr, 1989). I have examined this specimen and agree with this identification.

The taxa wilmattae, alboniger and B. ephippiatus have been regarded both as conspecific and as separate species. B. ephippiatus and wilmattae were regarded as separate species by Labougle et al. (1985) and Labougle (1990), who described diagnostic characters of colour pattern and morphology. However, D. Yanega (in litt.) and G. Chavarría (pers. com.) believe that all of these nominal taxa are part of the widespread and variable B. ephippiatus.

Very extensive studies of DNA variation by Duennes et al. (2016) show these taxa (1) separately not all to be monophyletic and (2) bGMYC analysis of the COI gene shows the entire Mesoamerican complex to be unambiguously a single species. I shall treat them as parts of one species.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: S Nearctic, N Neotropical Regions, W Neotropical border.

hypnorum-group of species

Bombus (Pr.) haematurus Kriechbaumer
haematurus Kriechbaumer, 1870:157
7 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region.

Bombus (Pr.) subtypicus (Skorikov)
leucopygus Morawitz in Fedtschenko, 1875:3, not of Illiger, 1806:172 (= B. hypnorum (Linnaeus))
[leucopygos (Skorikov, 1914b:294 [Pratobombus]) incorrect subsequent spelling]
subtypicus (Skorikov, 1914b:294 [Pratobombus]) examined
leucurus Bischoff & Hedicke, 1931:391, replacement name for leucopygus Morawitz in Fedtschenko, 1875:3
kohistanensis (Tkalcu, 1989:49 [Pyrobombus]) examined
11 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region, Oriental border.

Bombus (Pr.) hypnorum (Linnaeus)
Hypnorum (Linnaeus, 1758:579 [Apis]) examined
leucopygus Illiger, 1806:172
calidus Erichson in Middendorff, 1851:65
fletcheri Richards, 1934:90, examined
insularis Sakagami & Ishikawa, 1969:180, not of Smith, 1861:155 (= B. insularis (Smith))
koropokkrus Sakagami & Ishikawa, 1972:610, replacement name for insularis Sakagami & Ishikawa, 1969:180
29 names

TAXONOMIC STATUS: B. hypnorum is a broadly distributed species with a fairly easily recognised brown-black-white colour pattern (e.g. Reinig, 1939; Williams, 1991 [pdf]). It is possibly a complex of similar cryptic species.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic, Japanese, Oriental Regions, Arctic border. The first definite record of this species from Britain was in 2001 (Goulson & Williams, 2001 [pdf]). It has since been recorded from several sites in southern England.

PHOTOGRAPH: The first definite British specimen, in the collection of the Natural History Museum, London.

NOTES on this species in Britain.

Bombus (Pr.) perplexus Cresson
perplexus Cresson, 1863:91
3 names

TAXONOMIC STATUS: Although long regarded as a separate species (but see Williams, 1991 [pdf]:71), on the basis of DNA-sequence data Hines et al. (2006 [pdf]) and Cameron et al. (2007 [pdf]) have questioned whether B. perplexus might be conspecific with B. hypnorum. However, there are consistent morphological differences in the available samples.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic, E Nearctic Regions.

lepidus-group of species

Bombus (Pr.) abnormis (Tkalcu)
abnormis (Tkalcu, 1968a:33 [Pyrobombus]) examined
1 name

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) mirus (Tkalcu)
mirus (Tkalcu, 1968a:37 [Pyrobombus]) examined
?tibetanus Friese, 1913:86, examined, not of Morawitz, 1887:202 (= B. tibetanus (Morawitz))
2 names

DISTRIBUTION: Oriental Region.

Bombus (Pr.) lemniscatus Skorikov
lemniscatus Skorikov, 1912:607, examined
flavopilosus Friese, 1918:84, examined
peralpinus Richards, 1930:646, examined
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) lepidus Skorikov
lepidus Skorikov, 1912:606, examined
genitalis Friese, 1913:85, examined
tetrachromus Friese, 1918:85, examined, not of Cockerell, 1909:397 (= B. kashmirensis Friese)
yuennanicola Bischoff, 1936:7, examined
8 names

TAXONOMIC STATUS: B. lepidus and B. yuennanicola have been considered both as separate species (Bischoff, 1936) and as conspecific (Williams, 1991 [pdf]). Evidence from comparisons of DNA sequences from five genes is consistent with the two taxa being conspecific (Cameron et al., 2007 [pdf]).

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) infirmus (Tkalcu)
leucurus Bischoff, 1936:8, examined, not of Bischoff & Hedicke, 1931:391 (= B. subtypicus (Skorikov))
infirmus (Tkalcu, 1968a:24 [Pyrobombus]) replacement name for leucurus Bischoff, 1936:8
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) pressus (Frison)
pressus (Frison, 1935:342 [Bremus])
1 name

TAXONOMIC STATUS: The peculiar B. pressus was first recognised to be a species of the subgenus Pyrobombus by Cameron et al. (2007 [pdf]).

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) picipes Richards
flavus Friese, 1905:517, examined, not of Pérez, 1884:265 (= B. campestris (Panzer))
picipes Richards, 1934:90, examined
klapperichi Pittioni, 1949:266, examined
6 names

NOMENCLATURE: With Psithyrus regarded as being a subgenus of the genus Bombus (Williams, 1991 [pdf], 1995 [pdf]), B. pratorum subsp. flavus Friese (1905) becomes a junior secondary homonym in Bombus of Psithyrus campestris var. flavus Pérez (1884) (deemed to be subspecific, see ICZN, 1999: Article 45.6), and therefore B. flavus Friese is invalid (ICZN, 1999: Article 57). For this species, the oldest available name of which I am aware is B. parthenius var. picipes Richards, 1934 (deemed to be subspecific, see ICZN, 1999: Article 45.6), which becomes the valid name, B. picipes. The only publications using the name B. flavus Friese since 1950 of which I am aware are by Sakagami (1972), Ito (1993) and Yao & Luo (1997), so this change of valid name is not a serious disruption of common usage.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental, Palaearctic Regions.

Bombus (Pr.) parthenius Richards
parthenius Richards, 1934[14 April, Williams & Cameron, 1993]:89, examined
1 name

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) luteipes Richards
luteipes Richards, 1934:89, examined
signifer (Tkalcu, 1989:52 [Pyrobombus]), examined
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) infrequens (Tkalcu)
infrequens (Tkalcu, 1989:56 [Pyrobombus]) examined
1 name

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) avanus (Skorikov)
avanus (Skorikov, 1938b:2 [Pratibombus])
1 name

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) sonani (Frison)
sonani (Frison, 1934[30 April, Williams & Cameron, 1993]:175 [Bremus]) examined
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

pratorum-group of species

Bombus (Pr.) biroi Vogt
biroï [biroi] Vogt, 1911:51, examined
nursei Friese, 1918:84, examined
14 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic, Oriental Regions.

Bombus (Pr.) kotzschi Reinig
agnatus Skorikov, 1933b:248, examined, not of Skorikov, 1912:97 (= B. monticola Smith)
kotzschi Reinig, 1940:227, examined
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic, Oriental Regions.

Bombus (Pr.) flavescens Smith
flavescens Smith, 1852a:45, examined
mearnsi Ashmead, 1905:959, examined
?baguionensis Cockerell, 1920:631, examined
tahanensis Pendlebury, 1923:65, examined
?rufoflavus Pendlebury, 1923:66, examined
34 names

TAXONOMIC STATUS: Several of these nominal taxa have been treated as separate species. B. rufoflavus [Peninsular Malaysia] and B. baguionensis [Philippines] are particularly distinct in colour pattern. They may prove to be separate species, but from the material available from a few sites, they appear to be closely similar in morphology to B. flavescens (Williams, 1991 [pdf]). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region, Sumatran border.

Bombus (Pr.) rotundiceps Friese
rotundiceps Friese, 1916:108, examined
montivolanoides Sakagami & Yoshikawa, 1961:431
shillongensis (Tkalcu, 1974b:334 [Pyrobombus]) examined
7 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental Region.

Bombus (Pr.) ardens Smith
ardens Smith, 1879:133, examined
andreae Friese, 1910:405, examined
8 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental, Japanese, Palaearctic Regions.

Bombus (Pr.) pyrenaeus Pérez
pyrenaeus Pérez, [1880, see Baker, 1996d:300]:127, not of Lepeletier, 1832:375 (= B. rupestris (Fabricius))
tenuifasciatus Vogt, 1909:49
[pyreneus Pagliano, 1995:23, incorrect subsequent spelling]
16 names

NOMENCLATURE: With Psithyrus regarded as being a subgenus of the genus Bombus (Williams, 1991, 1995), B. pyrenaeus Pérez (1880) becomes a junior secondary homonym in Bombus of Psithyrus pyrenaeus Lepeletier (1832), and therefore B. pyrenaeus Pérez is invalid (ICZN, 1999: Article 57). The next available name, tenuifasciatus, was used by Vogt (1909) for individuals with particular colour patterns from both B. pyrenaeus Pérez and B. sichelii. The choice of which of these two homonyms should have precedence depends on the Principle of the First Reviser (ICZN, 1999: Article 24). As far as I have been able to discover, Tkalcu (1973:266) is the first author to have recognised this problem. He recognised precedence for B. pyrenaeus ssp. tenuifasciatus Vogt. Consequently, the oldest available name for this species, and therefore the valid name, is B. tenuifasciatus.

Although B. tenuifasciatus is the oldest available name for this species, the name B. pyrenaeus has been in common use for the species since 1950 (e.g. Krusemen, 1958; Tkalcu, 1969, 1973, 1975; Reinig, 1972, 1981; Delmas, 1976; Rasmont, 1983; Ornosa, 1986; Williams, 1991; Rasmont et al., 1995). It is suggested that, in the interests of stability, an application be made to ICZN to use its Plenary Power to suppress the senior homonym (ICZN, 1999: Article 78) (see the comments on B. muscorum) (in prep.). However, the consequence of this action would be that pyrenaeus (Lepeletier) would no longer be available for a subspecies of B. rupestris.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region.

Bombus (Pr.) modestus Eversmann
modestus Eversmann, 1852:134, examined
Baïkalensis [baikalensis] Radoszkowski, 1877b:203
nymphae Skorikov, 1910b:409
eversmanni Skorikov, 1910c:581, not infrasubspecific after Skorikov, 1922a:149
13 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Oriental, Palaearctic Regions.

Bombus (Pr.) wangae Williams et al.
wangae Williams et al., 2009:159, examined
1 name

DISTRIBUTION: Oriental Region.

Bombus (Pr.) pratorum (Linnaeus)
pratorum (Linnaeus, 1761:424 [Apis]) examined
55 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region, Arctic border.

INTRODUCTIONS: This species was deliberately introduced into Sydney, Australia, but is not known to have persisted (Oliff, 1895). Until the twentieth century, B. pratorum was not known in Ireland, where it is now well established (see references in Alford, 1975, 1980) (see comments on B. monticola).

NOTES on this species in Britain.

Bombus (Pr.) brodmannicus Vogt
Brodmannicus Vogt, 1909:49, examined
2 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region.

Bombus (Pr.) sitkensis Nylander
Sitkensis Nylander, 1848:235
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) mixtus Cresson
Praticola Kirby, 1837:274
mixtus Cresson, 1878:186, not of Kriechbaumer, 1870:160 (= B. maxillosus Klug)
3 names

TAXONOMIC STATUS: The identity of B. praticola has remained uncertain (e.g. Cresson, 1863; Franklin, 1913). Recently, Poole (1996) has listed B. praticola, B. mixtus and B. flavifrons as separate species without explanation.

Although I know of no type material, Kirby provided a description of B. praticola from northern Canada (latitude 65° North) with a colour pattern (including anterior half of abdomen yellow, posterior ferruginous) that for individuals from this area is most likely to be conspecific with either B. mixtus (some individuals have few black hairs on gastral terga II-III), or conspecific with B. flavifrons (which has terga V-VI black, although this is not always apparent from the dorsal view).

In his original description of B. flavifrons, Cresson (1863) conceded that this might be the same species as Kirby's B. praticola, and he went on to write (p. 106) that he had not yet identified B. praticola. Franklin (1913:371) wrote that he had 'been unable to decide whether the original description of B. praticolus [sic] referred to this species [B. flavifrons] or to the colour variant of pleuralis [intermediate colour patterns between B. flavifrons and B. pleuralis].' Milliron (1971:42) subsequently listed Pyrobombus praticola flavifrons (Cr.) as a member of his 'Praticola Group'.

However, from the original description I believe that the original material was more likely to have been of the species that has come to be known as B. mixtus. See the comments on B. flavifrons.

NOMENCLATURE: B. praticola is probably the oldest available name for this species. Any remaining confusion could be resolved by the designation of an appropriate neotype (e.g. see the comments on B. subterraneus).

Although B. praticola is probably the oldest available name for this species, the name B. mixtus has been in common use for the species since 1950 (e.g. Stephen, 1957; Thorp, 1970; Plowright & Stephen, 1973; K. W. Richards, 1973; Macior, 1975; Sakagami, 1976; Hurd, 1979; Plowright & Owen, 1980; Thorp et al., 1983; Laverty & Harder, 1988; Macfarlane et al., 1994). It is suggested that, in the interests of stability, an application be made to ICZN to use its Plenary Power to suppress the senior homonym (ICZN, 1999: Article 78) (see the comments on B. muscorum) (in prep.). However, the consequence of this action would be that mixtus (Kriechbaumer) would no longer be available for a subspecies of B. maxillosus.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic Region.

Bombus (Pr.) sandersoni Franklin
sandersoni Franklin, 1913:353
1 name

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic, E Nearctic Regions.

Bombus (Pr.) beaticola (Tkalcu)
beaticola (Tkalcu, 1968a:28 [Pyrobombus]) examined
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Japanese Region.

Bombus (Pr.) frigidus Smith
frigidus Smith, 1854:399, examined
3 names

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: W Nearctic, E Nearctic Regions, Arctic border.

Bombus (Pr.) jonellus (Kirby)
Jonella (Kirby, 1802:338 [Apis]) examined
alboanalis Franklin, 1913:385, examined
28 names

TAXONOMIC STATUS: B. alboanalis has been regarded both as a separate species (Franklin, 1913; Frison, 1927) and as conspecific with either B. frigidus (Burks, 1951; Hurd, 1979; Poole, 1996) or B. jonellus (Williams, 1991 [pdf]:78 [as B. jonellus from western Canada]; Scholl et al., 1995).

Recently, Scholl et al. (1995) concluded from studies of enzyme mobility morphs that B. alboanalis and B. frigidus have separate gene pools, but that B. alboanalis and B. jonellus show a low level of genetic differentiation. They also noted the lack of colour gradation between sympatric B. alboanalis and B. frigidus.

From the limited amount of material I have examined, I believe that B. alboanalis and B. jonellus are morphologically closely similar. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic, Arctic, W Nearctic Regions.

NOTES on this species in Britain.

Bombus (Pr.) cingulatus Wahlberg
cingulatus Wahlberg, 1854:208
11 names

TAXONOMIC STATUS: See comments on B. oceanicus.

MORPHOLOGY: photos of male genitalia.

DISTRIBUTION: Palaearctic Region, Arctic border.

?Bombus (Pr.) oceanicus Friese
oceanicus Friese, 1909:675, examined
oceanicus Friese & Wagner, 1910:52, redescribed
2 names

TAXONOMIC STATUS: B. oceanicus is known only from the Kurile Islands. A particularly close relationship with the otherwise broadly distributed B. cingulatus (absent from the Kuriles, but present in Kamchatka, Reinig, 1939; Ito & Sakagami, 1980) has been suggested by Ito & Sakagami (1980) and it is possible that they are conspecific. More evidence is awaited.

DISTRIBUTION: Japanese, Palaearctic Regions.