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B. pyrosoma worker visiting Lomatogonium. |
Melanobombus
ecology
and behaviour
HABITAT:
High
alpine grassland, mountain-meadow, open grassland,
semi-desert, together with a few very large species
(B. eximius, B. rufipes) of tropical hill
forests.
FOOD-PLANTS:
Short, medium, or occasionally long tongue-length
bumblebees, visiting correspondingly shallow to
deep flowers. Some species of this subgenus may
have a particular preference for compound inflorescences,
such as those of the Compositae. They also provide
'buzz' pollination.
NESTING
BEHAVIOUR:
Nests underground. Non-pocket makers. Melanobombus
includes a few species (e.g. B. rufipes)
of tropical hill forests that have colonies which
may persist for more than one year.
MATE-SEARCHING
BEHAVIOUR:
Males of many species patrol circuits of scent
marks. But males of some species have enlarged
compound eyes relative to the females and hover
or perch before racing after potential mates.
These species may be truly territorial (e.g. B.
rufofasciatus; Williams, 1991).
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Subgenus
MELANOBOMBUS Dalla Torre
Bombus (Melanobombus) Dalla Torre, 1880:40,
type-species Apis lapidaria Linnaeus (= Bombus
lapidarius (Linnaeus)) by subsequent designation
of Sandhouse, 1943:569
Bombus (Lapidariobombus) Vogt, 1911:58,
type-species Apis lapidaria Linnaeus (= Bombus
lapidarius (Linnaeus)) by subsequent designation
of Sandhouse, 1943:562
Kozlovibombus Skorikov, [1923]:152,
type-species Bombus kozlovi Skorikov, 1910b
(= Bombus keriensis Morawitz) in the sense of
Skorikov, [1923] (based on males = Bombus pyrosoma
Morawitz, a misidentification, see Reinig, 1934:169),
by subsequent fixation of Sandhouse, 1943:561
Rufipedibombus Skorikov, [1923]:156,
type-species Bombus rufipes Lepeletier by monotypy
[Atrocinctob.[ombus] Skorikov, 1933b:244,
published without description]
Bombus (Kozlowibombus) Bischoff, 1936:10,
unjustified emendation
Lapidariibombus Skorikov, 1938a:145,
unjustified emendation
?Bombus (Tanguticobombus) Pittioni, 1939d:201,
type-species Bombus tanguticus Morawitz by original
designation
Bombus (Rufipedobombus) Kruseman, 1952:102,
unjustified emendation
Pyrobombus (Festivobombus) Tkalcu, 1972:26,
27, type-species Bombus festivus Smith by original
designation
[Bombus (Lapedariobombus) Esmaili & Rastegar,
1974:52, incorrect subsequent
spelling]
TAXONOMIC
STATUS: For a discussion of why several former subgenera
have been synonymised within this subgenus see Williams
et al. (2008
[pdf]) .
Part
of the bumblebee phylogenetic tree including available
Melanobombus 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.
rufipes-group
of species
Bombus
(Ml.) rufipes Lepeletier
rufipes Lepeletier, [1835]:473
richardsi (Frison, 1930:6
[Bremus])
5 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Sumatran Region.
Bombus
(Ml.) eximius Smith
eximius Smith, 1852b:47,
examined
latissimus Friese, 1910:405
13 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region.
festivus-group
of species
Bombus
(Ml.) festivus Smith
festivus Smith, 1861:152,
examined
atrocinctus Smith in Horne, 1870:193,
examined
terminalis Smith in Horne, 1870:193,
examined
8 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region.
rufofasciatus-group
of species
Bombus
(Ml.) tanguticus Morawitz
tanguticus Morawitz, 1887:200
2 names
TAXONOMIC
STATUS: Queens of B. tanguticus are morphologicaly
very distinctive (discussed in Williams, 1991
[pdf]), so much so that Pittioni (1939d)
considered the species warranted a subgenus of its own.
The male remains apparently unknown (the species occurs
at high altitudes on the Tibetan Plateau and is very
rare in collections). Its precise relationships are
difficult to resolve at present and a separate subgenus
seems premature.
DISTRIBUTION:
Oriental Region.
Bombus
(Ml.) simillimus Smith
simillimis Smith, 1852b:48,
examined
[similis Smith, 1854:403,
incorrect subsequent spelling]
[simillimus Dalla Torre, 1896:548,
incorrect subsequent spelling]
grossiventris Friese, 1931:303,
examined
oculatus (Frison, 1933:335
[Bremus]) examined
tonsus (Skorikov, 1933b:248
[Sibiricobombus]) examined
simillimus Williams, 1991:99,
justified emendation
12 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region.
Bombus
(Ml.) rufofasciatus Smith
rufo-fasciatus [rufofasciatus] Smith,
1852b:48, examined
Prshewalskyi Morawitz, 1880:342
rufocinctus Morawitz, 1880:343,
examined, not of Cresson, 1863:106
(= B. rufocinctus
Cresson)
chinensis Dalla Torre, 1890[June
25]:139, replacement name for rufocinctus Morawitz,
1880:343; not of Morawitz,
1890[April 30]:352 (=
B. chinensis
(Morawitz))
waterstoni Richards, 1934:88,
examined
11 names
MORPHOLOGY:
photos of male
genitalia.
DISTRIBUTION:
Oriental Region, Palaearctic border.
Bombus
(Ml.) richardsiellus (Tkalcu)
richardsiellus (Tkalcu, 1968a:42
[Pyrobombus]) examined
2 names
MORPHOLOGY:
photos of male
genitalia.
DISTRIBUTION:
Oriental Region.
Bombus
(Ml.) miniatus Bingham
flavothoracicus Bingham, 1897:552,
examined, not of Hoffer, 1889:49
(= B. campestris
(Panzer))
miniatus Bingham, 1897:553,
examined
eurythorax Wang, 1982:435,
examined
stenothorax Wang, 1982:439,
examined
4 names
TAXONOMIC
STATUS: B. miniatus has been considered conspecific
with B. pyrosoma, B. formosellus and B. friseanus
(Williams, 1991
[pdf]).
Evidence of intermediates between B. miniatus
and B. friseanus is not strong, but perhaps not
least because so little material is available from where
these taxa occur in close proximity in the eastern Himalaya.
The few workers and males from this area that I have
seen are difficult to assign to either taxon with any
confidence, although the queens are closer to the colour
pattern of B. miniatus (Williams, 1991
[pdf]). More evidence is awaited.
B.
eurythorax and B. stenothorax are closely
similar in morphology and colour pattern to B. miniatus.
I know of no reason why these nominal taxa should not
be considered conspecific.
NOMENCLATURE:
With Psithyrus regarded as being a subgenus of
the genus Bombus, B. flavothoracicus Bingham
(1897) becomes a junior
secondary homonym in Bombus of Psithyrus campestris
var. flavothoracicus Hoffer (1889)
(deemed to be subspecific, see ICZN, 1999:
Article 45.6), and therefore B. flavothoracicus
Bingham is invalid (ICZN, 1999:
Article 57).
For
this species, the oldest available name is B. miniatus,
which becomes the valid name. The only subsequent publications
of which I am aware that use the name B. flavothoracicus
for this taxon as a species are by Tkalcu (1974b),
Wang (1982) and Macior
(1990), so this change
of valid name is not a serious disruption of common
usage.
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region.
Bombus
(Ml.) friseanus Skorikov
friseanus Skorikov, 1933a:62,
examined
hönei [hoenei] Bischoff, 1936:10,
examined
2 names
TAXONOMIC
STATUS: B. friseanus has been considered
conspecific with B. pyrosoma (Bischoff, 1936;
Tkalcu, 1961b;
Sakagami, 1972) and
has been considered conspecific with B. pyrosoma,
B. formosellus, and B. flavothoracicus (=
B. miniatus) (Williams, 1991
[pdf]). From a preliminary survey of colour variation,
these taxa appear to remain discrete and are likely
to be separate species. S.-f. Wang and J. Yao (in
litt.) also believe that the taxa may be separate.
More evidence is awaited.
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region.
Bombus
(Ml.) pyrosoma Morawitz
pyrosoma Morawitz, 1890:349,
examined
pyrrhosoma Dalla Torre, 1896:544,
unjustified emendation
wutaishanensis (Tkalcu, 1968a:39
[Pyrobombus]) examined
7 names
TAXONOMIC
STATUS: B. pyrosoma has been considered conspecific
with B. friseanus (Bischoff, 1936)
and has been considered conspecific with B. formosellus,
B. friseanus, and B. flavothoracicus (=
B. miniatus) (Williams, 1991
[pdf]). From a preliminary survey of colour variation,
these taxa appear to remain discrete and are likely
to be separate species. S.-f. Wang and J. Yao (in
litt.) also believe that the taxa may be separate.
More evidence is awaited.
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region, Palaearctic border.
Bombus
(Ml.) formosellus (Frison)
formosellus (Frison, 1934:163
[Bremus]) examined
3 names
TAXONOMIC
STATUS: B. formosellus has been considered
conspecific with B. pyrosoma, B. friseanus,
and B. flavothoracicus (= B. miniatus)
(Williams, 1991
[pdf]), as a disjunct peripheral population on Taiwan.
From a preliminary survey of colour variation, these
taxa appear to remain discrete and are likely to be
separate species. More evidence is awaited.
DISTRIBUTION:
Oriental Region.
lapidarius-group
of species
Bombus
(Ml.) ladakhensis Richards
ladakhensis Richards, 1928a:336,
examined, not infrasubspecific after Tkalcu, 1974b:335
phariensis Richards, 1930:642,
examined, not infrasubspecific after Tkalcu, 1974b:336
variopictus Skorikov, 1933b:248,
examined
reticulatus Bischoff, 1936:7,
examined
6 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental Region, Palaearctic border.
Bombus
(Ml.) semenovianus (Skorikov)
semenovianus (Skorikov, 1914a:127
[Lapidariobombus]) examined
3 names
MORPHOLOGY:
photos of male
genitalia.
DISTRIBUTION:
Oriental, Palaearctic Regions.
Bombus
(Ml.) keriensis Morawitz
keriensis Morawitz, 1887:199,
examined
separandus Vogt, 1909:61,
examined
kohli Vogt, 1909:61,
examined, not of Cockerell, 1906:75
(= B. morio (Swederus))
kozlovi Skorikov, 1910b:413,
replacement name for kohli Vogt, 1909:61
tenellus Friese, 1913:86
[alagesianus (Skorikov, [1923]:152
[Lapidariobombus]) published without description]
?alagesianus Reinig, 1930:89
pamirus (Skorikov, 1931:226
[Lapidariobombus]) not of Skorikov, 1931:232
[= B. oberti Morawitz]
richardsi Reinig, 1935:341,
not of Frison, 1930:6
(= B. rufipes Lepeletier)
tibetensis Wang, 1982:439,
replacement name for richardsi Reinig, 1935:341
trilineatus Wang, 1982:441,
examined
42 names
TAXONOMIC
STATUS: Several of these nominal taxa have been
treated as separate species (e.g. Skorikov, 1931),
although B. keriensis has also long been considered
to be a broadly-distributed and variable species, including
both yellow-banded and white-banded individuals throughout
much of its range (Reinig, 1935,
1939; Williams, 1991
[pdf]).
B.
alagesianus is morphologically closely similar to
B. keriensis. Evidence from comparisons of DNA
sequences from the 16S gene is not strong but could
be consistent with the two taxa being separate species
(Cameron et al., 2007
[pdf]). Until more evidence to the contrary is available
from more detailed studies of patterns of variation,
I shall continue to treat them as parts of a single
variable species.
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental, Palaearctic Regions.
Bombus
(Ml.) sichelii Radoszkowski
Sichelii Radoszkowski, 1860:481,
examined
[Sicheli Radoszkowski, 1877b:213,
incorrect subsequent spelling]
tenuifasciatus Vogt, 1909:49,
not of Vogt, 1909:49
(= B. pyrenaeus
Pérez) after Tkalcu, 1973:266
chinganicus Reinig, 1936:6,
not of Reinig, 1936:8
(?= B. bohemicus
Seidl)
erzurumensis (Özbek, 1990:209
[Pyrobombus]) examined
27 names
TAXONOMIC
STATUS: Until recently, the white-banded form of
B. sichelii has been known from west of the Caspian
Sea only from the Caucasus (Reinig, 1935).
Now that B. erzurumensis (morphologically closely
similar to B. sichelii and with white bands)
has been described from Turkey, it could be interpreted
as another white-banded western colour form. By analogy
(see the comments on B. melanopygus)
the difference in colour could be the effect of a just
two alleles at a single locus for pigment. Evidence
from comparisons of DNA sequences from five genes is
consistent with the two taxa being conspecific (Cameron
et al., 2007
[pdf]).
Until more evidence to the contrary is available from
critical studies of patterns of variation, I shall continue
to treat them as parts of a single variable species.
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Oriental, Palaearctic Regions.
Bombus
(Ml.) incertus Morawitz
incertus Morawitz, 1881:229
2 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Palaearctic Region.
Bombus
(Ml.) lapidarius (Linnaeus)
Lapidaria (Linnaeus, 1758:579
[Apis]) examined
Strenuus (Harris, 1776:131
[Apis])
eriophorus Klug, 1807:265,
examined
caucasicus Radoszkowski, 1860:482,
examined
37 names
MORPHOLOGY:
photos of male genitalia.
DISTRIBUTION:
Palaearctic Region.
INTRODUCTIONS:
This species was deliberately introduced into Christchurch,
New Zealand, but is not known to have persisted (Sladen,
1912).
NOTES
on this species in Britain.
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