Bulletin
British Museum |
(Natural History )
OLUME 60 NUMBER 7 25 APRIL 1991
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The Bulletin of the British Museum (Natural History ), instituted in 1949,
is issued in four scientific series, Botany, Entomology, Geology
(incorporating Mineralogy) and Zoology, and an Historical series.
The Entomology Series is produced under the editorship of the
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
© British Museum (Natural History), 1991
ISBN 0 565 06041 4 Entomology Series
ISSN 0524-6431 Vol 60, No. 1, pp. 1-204
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 April 1991
Typeset by Computer Typesetting Services, Redhill, Surrey
Printed in Great Britain by Henry Ling Ltd, Dorchester, Dorset
Bull. Br. Mus. nat. Hist (Ent.) 60 (1):1—204
Issued 25 April 1991
The bumble bees of the Kashmir Himalaya
(Hymenoptera: Apidae, Bombini)
PAUL H. WILLIAMS
clo Department of Entomology, British Museum (Natural History), Cromwell Road, South Kensington,
London SW7 5BD
CONTENTS
WERTROXG ECCI Et Ar Gin RET caer On ree Ee ICR BICEP ct Roca Stoner ace cee at a ten eee 2
Materialexamined|and/depositonies! 5.2.2. 000. satase snes sens sae sss dgers ne 8
Diasnosis of MimalayanBOmbiM. 2. aaa ence | See ee eee fey eee ee 10
G@haracters Studied). 7. cs sever cays agate) ne esc yaeya coepevebaseis pies ener ase ayerd wie le 2 11
Relationships among species by ancestry ............ 0. cc cece c eee ee 13
GeneraOMmBOml Dually arcccn! dese ce tsrte rae «alle Mae acdelakebegara pele ele ache a cet iais ae 14
Discrimination of species and intraspecific variation ................2...0.000005 16
The ‘biological’ species and the Recognition Concept ................0---0e eee 17
Mate-searching behaviour of male bumble beesin Kashmir..................... 18
Relationship between male mate-searching behaviour and habitat structure ....... 25
Inference of allopatric, conspecific taxa... 2.2... ccc eens Pa
Inference of allopatric, separate species ......... 0... cece cece eee eee eee ee 27
Characters of species not related to male mate-searching behaviour .............. 28
The description of intraspecific variation. ................. cee cece cece e eee eee 29
INomenclatuinalistimmmtiany) rac cits «is <ciers cian n, Soe Relea ame sells eee Deere ales ohare alahe 29
Taxonomy of the Kashmir Bombini..................... 000s cece eee ee eee eens 31
Key to species (females: queens and workers) ............-.--22 0000 eeeeeeeeee 31
ING Vi OSPEGIESi (MALES mye cas br cecco go sore x ap a DET ero ncn a AE RE eattte spa tela te 35
Genus/BomDusileatree: 5 vx, aprescteg:vsvaya:s. evn ve sacora gees cchesbel etn as Sarees aya Sind «Wat ees cle = 38
Distnibutionionthe fama rye chs te ack <2, decesnie sbeiae Ge oes asians eaux om e.s 107
Distributlomwithaltipudes «7. Gactaxs «yee. + «lcd mnehe. scavuce Aeiewmele Orie emesis hase as 107
Distnbutioniacrossthe Himalaya. (2... .254-005-q-0eeseeneesee esses ase anes 111
DistributionoutsideKashmin ). 2225.26. cece sees s ceca ees hone ene nase nna 112
Relationships between patterns at different spatialscales...................... 113
Colour patterns of the Kashmirfauna ................0.. cee cece eee e ee eee 115
WarlationonsyasiancusimKashmit a. 2... eet adsense nee tei e ss tees 2 2 115
Converpenticoloun patterns! sea. csc n serene eccrine acannon aie acre abies © 118
PNCKNOWIEU SCIIER Sine, Sette erty, Mess cies oeisnsiacse cote emote craeinen atte roe e nom anal: 120
IREFETLEN CES a yies eps es FM ach eh eieenvaape aks G/T icTs METAS Ae is ote detest 121
IURYSLOS2 EN ces, An? Rae create tet gk ae cee gene aR raaee i GAC OA ae 202
SYNOPSIS. Bumble bees are extraordinarily variable in the colour patterns of their
pubescence. This has contributed to the lack of agreement among authors concerning the
discrimination of species. Material from Kashmir has been listed previously under 80
names, but for the broadest interpretations of these species, more than 500 names have
been published for differing individuals from throughout their distributions. In this
review, all of the available voucher material from Kashmir (6312 museum specimens from
78 sites) is assigned to just 29 species. The reduction in the number of nominal taxa that are
inferred to represent separate species depends on two factors. First, variation is examined
among the larger samples of new material. Second, current species concepts require a re-
assessment of this evidence for the limits of reproductive cohesion in space and time. A
particular study is made of the variation in male mate-searching behaviour and of its
relationship to the different kinds of habitat. For Bombus asiaticus, patterns of colour
variation across the Great Himalaya are compared with those expected from simple
PAULH. WILLIAMS
genetic models. Type material is examined for 103 of the nominal specific and subspecific
taxa. Seventeen lectotypes are designated and 80 new synonyms and provisional new
synonyms are established. Keys to the species are accompanied by diagrams of the colour
patterns and by distribution maps.
INTRODUCTION
Bumble bees rank among the most abundant and
conspicuous of flower visitors in alpine, temperate
and arctic environments of the northern continents.
In the southern hemisphere they are native only in
the East Indies and South America (see Map 1),
where most species are associated with the high-
lands. Adults feed mainly on nectar, which
provides energy for flight. The larvae are fed on a
mixture of pollen and honey, which provides their
requirements for growth. Most species are social
(reviews of behaviour by e.g. Alford, 1975; Morse,
1982; Plowright & Laverty, 1984). The colonies
consist almost always of a single, mated queen,
which lays most of the eggs, and usually of no
more than a few hundred, unmated workers. The
establishment and development of colonies takes
place each summer and may take less than two
months (Richards, 1973), culminating with a
switch to the production of males and young
queens near the end of the season. After mating,
young queens normally diapause through the
winter, away from the nest, before they attempt to
found colonies on their own in the following year.
Only a few species from the tropics of South
America (Sakagami, 1976) and South East Asia
(Michener & Amir, 1977) may have colonies that
persist for more than one year. Bumble bees show
unusually well-developed endothermy, which can
be facultative when foraging (reviewed by Heinrich,
1979). Almost all species are generalists in their
choice of food-plants, visiting any remunerative
flowers. Their foraging follows a ‘scramble’ pattern,
without either recruitment of nest-mates to good
food sources or defence of food sources. These
characteristics of bumble bees may account for
their abundance in cool environments that have a
predictable season of adverse conditions, where
flowers are often fairly evenly dispersed. In
contrast, the other social, long-proboscis bees
(honey bees and stingless bees) have their centres
of diversity and abundance in the tropical lowlands.
The bumble bees of Kashmir are of particular
interest because this narrow corridor of mountains
is almost the only major, modern point of contact
between the large and divergent Oriental and
Palaearctic bumble bee faunas. These faunas are
otherwise separated by deserts in central Asia
and in China, except for another corridor of
contact near Beijing, which is occupied by relatively
few species (Panfilov, 1957). Greater Kashmir
encompasses almost the entire mountain system
that links the high Tibetan (Xizang-Qinghai)
plateau in the east with the Hindu Kush, Pamir
and Tien Shan mountain ranges to the west and
north. Hence Kashmir covers segments of the
Pir Panjal, Great Himalaya, Zanskar, Ladakh,
Karakoram and Hindu Raj ranges (Map 2), and
includes some of the highest peaks in the world
(altitudes range between about 400-8600 m; for
an account of the geological structure see Searle et
al., 1988). Kashmir is slightly larger in area than
the European Alps and lies at the same latitude
as Syria, Tunisia and Arizona. Since Indian
independence in 1947, the sovereignty of Kashmir
has been a matter of dispute and sections are now
administered by India (Jammu & Kashmir State),
Pakistan (Northern Areas) and China (as part of
the Xinjiang Autonomous Region).
The high relief of Kashmir provides a broad
range of habitats for bumble bees. Patterns in the
distributions of bumble bees in Europe have been
linked with climatic factors (e.g. Pekkarinen etal. ,
1981). The differing climates of Kashmir can be
summarised in three regions (e.g. Gurcharan
Singh & Kachroo, 1976). First is the subtropical
region of the Jammu foothills, which is subject to
the summer monsoon (Fig. 2). Second, beyond
the Pir Panjal range, is the more temperate Vale
of Kashmir, where most of the rain and snow fall
in the winter months (Fig. 1). Third, in the rain
shadow of the Great Himalaya, is the arid alpine
region of the Zanskar, Ladakh and Karakoram
ranges (Figs 3 & 4). But of equal importance to
climate, as an influence on whether or not a
species of bumble bee could persist at a particular
locality, may be the nature of the local vegetation
(e.g. Bowers, 1985; Williams, 1988, 1989), even
though bumble bees are seldom dependent on
particular species of food-plants. Map 3 shows a
summary of the vegetation in Kashmir, based on
the survey of the Himalaya by Schweinfurth
(1957). Not only do the three principal climatic
regions within Kashmir differ in their flora, but in
combination with the influences of local altitudinal
zonation and variation in local exposure (Troll,
1972), this contributes towards a particularly
broad range of habitats. In comparison with some
parts of the Himalaya, Kashmir retains relatively
THE BUMBLE BEES OFTHE KASHMIR HIMALAYA
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PAULH. WILLIAMS
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6 PAULH. WILLIAMS
Fig. 1 Subalpine zone of Mt Apharwat in the Pir Panjal range, viewed from 3000 m towards the peak at 4143 m (Map 4
locality 1: foreground pasture with Euphorbia wallichiana; lower slopes dominated by Betula utilis, Salix sp., Syringa
emodi, with a few Abies spectabilis; middle slopes dominated by B. utilis, Rhododendron campanulatum; higher slopes
dominated by R. anthopogon, Juniperus recurva). Of 17 species of bumble bees recorded in this immediate area, the
most restricted to this zone is B. biroi, and the most abundant is B. rufofasciatus.
Fig. 2 Lower montane coniferous forest zone on the Patnitop ridge in the Jammu foothills, viewed from 2000 m
towards the plains of India (Map 4 locality 11: Patnitop dominants, Cedrus deodara [foreground], Pinus wallichiana,
P. roxburghii). Of 3 species of bumble bees recorded in this area, the most restricted to this zone is B. haemorrhoidalis,
and the most abundant is B. trifasciatus.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 7
Fig. 3 High, arid alpine steppe zone at the terminal moraine of Nimaling plain in the Zanskar ranges, viewed from 4800
m across the southern lateral moraine towards a peak (Kang-y-sey) at 6400 m (Map 4 locality 51: foreground, shrubs of
Caragana versicolor). Of 9 species of bumble bees recorded in this immediate area, the most restricted to this zone are
B. personatus, B. oberti and B. ladakhensis, of which the most abundant is B. oberti.
Fig. 4 Subalpine semidesert zone at Lamayuru near the Indus valley, with a view of valley terraces at 3400 m, against a
background of mountain ridges at 4000-5000 m (Map 4 locality 42). Of 3 species of bumble bees recorded in this area,
the most restricted to this zone, and the most abundant, is Bombus semenovianus.
8
large forests and yet access to some of the varied
alpine areas is no longer difficult.
Previous studies of the bumble bees of Kashmir
have been based on a total sample of only a couple
of hundred specimens. The first important collec-
tion of bumble bees from Kashmir was made by
Lt.-Col. C. G. Nurse in 1901. A complete inven-
tory of this material was never published, although
it provided the specimens that have since been
described as the types of many nominal taxa
(Friese, 1909, 1918; Richards, 1928a, 19285, 1930;
Tkalct, 1974b). Other collections were mostly
small, but particularly important material was
obtained by A. Jacobson in Kashmir and Ladakh
during 1912 (Skorikov, 1914a) and by Col. R.
Meinertzhagen in Ladakh during 1925 (Richards,
19285). Skorikov (19335) compiled a preliminary
list of the fauna of Kashmir and discussed the
fauna of the entire Himalaya, though his work was
based on a different concept of the species from
that accepted at present. The only recent revision
of any part of the large Himalayan fauna is
Tkalct’s (19745) description of a collection of 73
bumble bees from Nepal. Although there are
many elements in common between the faunas of
Nepal and Kashmir, 16 species that are known
from Kashmir are not represented in this collection.
In the same paper Tkalci refers to his ‘Monographie
der Unterfamilie Bombinae des Himalaya’ as in
press. Unfortunately this has never been published,
although recently he has described several new
taxa from the region (Tkalcu, 1989). The only key
that has been intended to cover any part of the
Himalayan fauna is that published by Bingham
(1897). This key artificially subdivides four of the
species now recognised from Kashmir, whereas a
further 20 species are not included at all.
For this review, much of the older material is re-
examined and compared with the larger collections
of new material, in order to describe some of the
patterns of variation (and the discontinuities
in these patterns) among the bumble bees of
Kashmir. This evidence, together with informa-
tion from the habitat associations of the species in
Kashmir and information from their broader
distributions beyond Kashmir, is used to discuss
the likely relationships of ancestry and inter-
breeding among the nominal taxa.
MATERIAL EXAMINED AND
DEPOSITORIES
Large samples of undescribed material are avail-
able from western Kashmir in the collections
PAULH. WILLIAMS
Table 1 Localities that have been sampled for bumble
bees. Sites in close proximity are treated as combined
and represented by a single number on Map 4.
Localities from which samples of bumble bees have been
examined (numbers in circles on Map 4).
1 (Mt) Apharwat
2 Astor
3 Atro Sar
4 Babusar
5 Baltal
6 Baltit
7 Banidas
8 Banihal
9 Bara Deosai
10 Batakush
11 Batote
12 Bulandi
13 Burzil Chauki
14 Chakorkhand
15 Chhantir Gah
16 Chhota Deosai
17 Chittakatha Sah
18 Chogdo
19 Chushul
20 Dachhigam
21 Daksum
22 Dalti
23 Darkot
24 Dras
Gamelti = Darkot
25 Gilgit
26 Gulmarg
27 Gulmiti
28 Gumri
42 Lamayuru
43 Leh
44 Lilam
45 Luskum
46 Mahthantir Gah
Martselang = Chogdo
47 Maru
48 Matayan
49 Muzaffarabad
Nagar = Holshal
50 Naltar
Nigagar = Baltal
51 Nimaling
52 Nomal
Nowgam = Banihal
53 Pahalgam
54 Parkutta
Patnitop = Batote
55 Rawat
56 Rumbak
57 Saif-ul-Maluk Sar
58 Sangisfaid
59 Shamran
60 Shardi
61 Shigar
62 Shinghai Gah
63 Skardu
64 Sonamarg
65 Srinagar
Harwan = Dachhigam 66 Stakmo
29 Hemis 67 Sumbal
30 Holojut 68 Surgun
31 Holshal 69 Suru
Hunza = Baltit Tangmarg = Gulmarg
32 Imit 70 Taobat
33 Janwai 71 Teru
34 Kangan Thajiwas = Sonamarg
35 Kanur 72 Thonde
36 Karsa 73 Tiggur
37 Kel 74 Tso Morari
38 Khalsi Tungri = Karsa
39 Khardung La 75 Verinag
Khilanmarg = Apharwat 76 Wangat
40 Kishtwar 77 Yasin
41 Lal Pani 78 Yusmarg
Zoji La = Gumri
Localities at which no bumble bees could be found
during brief surveys in August-September 1980, 1985 or
1986 (letters in triangles on Map 4).
a Anantnag
b Baramula
c Chrar-i-Sharif
d Jammu
e Mulbekh
f Ramban
g Udhampur
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
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‘JOAR] BAS BAOgR WOOQOE ULY) B10W puke]
10
made during 1953 and 1954 by F. Schmid (BMNH
accession 1962-457; account of the expedition in
Schmid, 1958). In addition, I was able to make
large, selective collections from southern and
eastern Kashmir during 1980, 1985 and 1986.
Unfortunately, large areas near the borders,
especially to the north and east of the country, are
prohibited to foreign visitors, so only a few
specimens from older collections are available.
The sites in Kashmir from which material has
been examined are listed in Table 1 and shown on
Map 4. Geographical information is taken from
the United States Survey of India maps (scale 1 :
250,000) and from The Times atlas of the world
(1987). The world-wide distribution of species is
also shown by maps (see Map 1). These are
compiled with a preference for records from those
specimens that I have been able to examine,
supplemented by a list of selected references. The
source material is summarised briefly in the text
either by political region (for the Himalayan/
Tibetan mountain system in India and China) or
by major mountain system (for the more restricted
bumble bee faunas of central Asia), such as the
Elburz, Hindu Kush, Pamir, Alai— Tien Shan (in
its broadest sense) and Altai mountains. More
precise data can be found in the original references.
The specimens that have been examined are
deposited in collections as shown by the references
to the following abbreviations.
AB Dr A. A. Bhat, S.K. University of Agricultural
Sciences & Technology, Srinagar, India.
BMNH British Museum (Natural History), London,
U.K.
BPBM Bishop Museum, Honolulu, U.S.A.
BT Dr B. Tkalct, Prague, Czechoslovakia.
FA Faculté des Sciences Agronomiques de l’Etat,
Gembloux, Belgium.
INHS Illinois Natural History Survey, Champaign,
U.S.A.
ITZ Instituut voor Taxonomische Zodlogie,
Amsterdam, Netherlands.
IZ Institute of Zoology, Beijing, China.
LK Dr L. S. Kimsey, University of California,
Davis, U.S.A.
LSL Linnean Society, London, U.K.
MCSN Museo Civico di Storia Naturale, Genoa,
Italy.
MI Dr M. Ito, Hokkaido University, Sapporo,
Japan.
MNHN Muséum National d’Histoire Naturelle, Paris,
France.
MNHU Museum fiir Naturkunde an der Humbolt-
Universitat, Berlin, D.D.R.
NM Naturhistorisches Museum, Vienna, Austria.
NMP Narodni Muzeum, Prague, Czechoslovakia.
PAULH. WILLIAMS
NMS Natur-Museum Senckenberg, Frankfurt,
D.B.R.
NR Dr N. D. Rishi, University of Kashmir,
Srinagar, India.
PW Author’s collection, London, U.K.
RH Dr R. W. Husband, Adrian College, Adrian,
U.S.A.
SEMK Snow Entomological Museum, Lawrence,
Kansas, U.S.A.
TL Dr T. M. Laverty, University of Western
Ontario, London, Canada.
™ Természettudomanyi Muzeum, Budapest,
Hungary.
UM University Museum, Oxford, U.K.
USNM United States National Museum, Washington
ID(Cz, WES AN
ZI Zoological Institute, Leningrad, U.S.S.R.
ZM Zoologisk Museum, Copenhagen, Denmark.
ZMMU_ Zoological Museum of Moscow State Univer-
sity, Moscow, U.S.S.R.
ZS Zoologische Staatssammlung, Munich, D.B.R.
DIAGNOSIS OF HIMALAYAN BOMBINI
Among the Apidae, the Bombini have the labrum
at least twice as broad as long. The labrum lacks
a longitudinal median ridge, although for the
females it has a strong transverse basal depres-
sion. The clypeus has a transverse subapical
depression and the apico-lateral corners are curved
back towards the occiput. A malar area (= malar
space) separates the compound eye from the base
of the mandible, often by a distance greater than
the breadth of the mandible at its base. The hind
wings lack a jugal lobe (= anal lobe). The volsella
(= lacinia) of the male genitalia is greatly enlarged
and is produced apically beyond the gonostylus (=
squama).
Bumble bees are large (body length 7-27 mm)
robust insects. Their bodies have a dense covering
of variously-coloured long plumose hairs, although
these are few or absent on some parts of the ventral
surface of the gaster, on parts of the propodeum,
on parts of the anterior face of gastral tergum I,
and on parts of the head. The sclerites are usually
black, or lighter brown on the distal parts of the
limbs, but are never marked with bright yellow,
red, or metallic (= interference) colours. The
wings may be transparent (= hyaline) to strongly
darkened (= infuscated), but rarely show strongly
metallic reflections.
Female bumble bees have 12 antennal ‘segments’
(= scape, pedicel and 10 flagellomeres) and six
visible gastral terga and sterna (abbreviated to TI-
VI, SI-VI). Males have 13 antennal ‘segments’ (=
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
scape, pedicel and 11 flagellomeres) and seven
visible gastral terga and sterna (abbreviated to TI-
VII, SI-VI).
CHARACTERS STUDIED
Although this review aims to describe many
aspects of variation among the bumble bees of the
Kashmir Himalaya, inevitably those characters
that vary most among species and least within
species receive most attention. Many characters
of the females show more pronounced expression
and divergence among the queens (which are
usually the larger females) than among the workers
(smaller females).
The most variable characters of bumble bees
are the colours of their pubescence. These are so
variable that they can be very misleading if
used alone for the identification of species. For
instance, it is known that in some areas of Europe
and North America the local parts of populations
from even very divergent species-groups appear
to have converged closely in colour pattern (see
the comments on discrimination of species and
intraspecific variation, and the discussion of colour
patterns of the Kashmir fauna). Differences in the
shape and sculpturing of the sclerites have been
found to be more reliable indicators of relationship.
On the head, the shape and details of the
sculpturing of the LABRUM, CLYPEUS and
MALAR AREA (= oculo-malar space) are
frequently used as characters (Fig. 14). The super-
ficial structure and sculpturing of the labrum may
be particularly complex (Figs 23-30, 220-222).
The shape of the malar area can be measured as
the ratio of the oculo-malar distance to the
breadth of the mandible at its base (Fig. 15). This
shape is related to variation in proboscis length
(Medler, 1962), which is in turn related to varia-
tion in the behaviour governing flower choice,
both among and within species (reviewed by e.g.
Morse, 1982). However, within at least some of
the species with longer proboscides, the shape of
the malar area may be subject to allometric effects
among individuals of different sizes (e.g. Sakagami,
1972). Furthermore, because the boundaries of
the malar area require arbitrary definition,
measurements by different authors are not always
comparable. Therefore this character is used here
only when the differences in shape are relatively
large. The MANDIBLES provide many characters
including the number and shape of the ridges
known as KEELS, between which are grooves,
such as the SULCUS OBLIQUUS (Figs 31-38).
11
There is a BASAL TOOTH, variable numbers
of subsidiary or INTERCALARY TEETH and
often an APICAL TOOTH, which is defined at
its base by an anterior notch known as the
INCISURA. These characters vary within species
as well as among them and are subject to further
apparent variation caused by wear. Nonetheless,
they can still be useful in some species-groups
(e.g. the subgenera Psithyrus, Alpigenobombus).
The shape of ANTENNAL ‘SEGMENTS’ (=
scape, pedicel and flagellomeres), measured as
ratios of length to breadth, is also used here, but
only when the differences in shape are large (Figs
17-22, see the comments above on the shape of
the malar area). The OCELLI vary in size and in
position relative to the compound eyes. The
separation of the lateral ocellus from the neigh-
bouring margin of the compound eye can be
measured in relation to the diameter of the lateral
ocellus (Fig. 16). The strongly enlarged ocelli of
the females of some species are associated with
enlarged compound eyes among the conspecific
males, and hence with particular kinds of male
mate-searching behaviour (see the comments on
mate-searching behaviour of male bumble bees in
Kashmir). One of the characters most prone to
variation, both within and among species, is the
sculpturing of the OCELLO-OCULAR AREAS
of the VERTEX (= frontovertex), on the top of
the head (Figs 215-219). Just as for variation in
the sculpturing of the labrum and clypeus, some of
the elements in the patterns of punctures of the
ocello-ocular areas are constant among large
groups of species, whereas other details are
subject to variation among individual bees from
the same colony.
On the thorax, the majority of characters are
taken from the appendages (Fig. 43). Subtle
differences in the shape of the wing venation
have occasionally been used (e.g. Milliron, 1971;
Plowright & Stephen, 1973), although these are
not easy to measure (see the comments above on
the malar area). Above the wing bases are a pair
of small dome-shaped TEGULAE, which are
shown in the colour pattern diagrams. Between
the tegulae, on the thoracic dorsum, is the
SCUTUM (= mesoscutum), and immediately
posterior to it, the SCUTELLUM (= mesoscutel-
lum). These sclerites show some variation in
sculpturing (Figs 223 & 224), but are also useful
for relating to the distribution of pale or black
pubescence. The relative length of the pubescence
on various parts of the legs can be used to measure
the general length of the pubescence of the body
(e.g. by comparison with tibial breadth). The
outer surface of the hind TIBIA has large areas
without long pubescence for females of most
12
species, although to a lesser extent for females of
the subgenera Psithyrus and Mendacibombus and
for males of some other species (Sakagami & Ito,
1981). For these females the bare areas are
surrounded by dense fringes of long, stout hairs
that function as a CORBICULA for carrying
pollen. Variation in the length and extent of this
pubescence and in the sculpturing of the bare
areas have been used to distinguish taxa (Figs
225 & 226), although there can be considerable
differences between queens and workers within
the same species (see the comments on B. simil-
limus). Among some species-groups there is a
pronounced development of the disto-posterior
corners of the hind tibia and of the mid and hind
BASITARSI (= first tarsomeres) (Figs 39-42).
These can form spinose projections, but differences
between closely-related species are seldom discrete.
The shape, sculpturing and pubescence of the
hind basitarsus are also used (Figs 227-229).
On the abdomen, the most obvious characters
are found in the variation of the sculpturing of the
sclerites. Bumble bees, like other Apocrita, have
a waist between the first two abdominal segments,
so that the first abdominal segment (= propodeum)
is associated with the thorax, whereas the sub-
sequent segments form a separate unit, the
GASTER (= ‘metasoma’), which can be moved
independently. In this review all references to
segment numbers for the dorsal TERGA (=
tergites) and ventral STERNA (= sternites) refer
to gastral segments, so gastral tergum I is the
dorsal sclerite of the second abdominal segment.
For the females, variation in sculpturing is often
especially clear on gastral tergum VI (= epipygium
or pygidium) and sternum VI (= hypopygium),
which may also show single or double KEELS
(= ridges) (Figs 206-211), or even a rounded boss,
as well as variation in the shape of the apex (Figs
212-214, 230, 231). There is some variation in the
structures associated with the female STING (=
modified ovipositor), especially in the shape of
the ‘inner projections from the sting sheath’
(Richards, 1927b, 1968; see also Kopelke, 1982).
However, many of the ‘sting sheath’ characters
are parts of the folds that lie between the rami of
the left and right valvulae (see Williams, 1985:
fig. 5) and these folds are usually only weakly
sclerotised. Consequently they do not always
preserve well in dried specimens and may become
distorted, so these characters are not used here.
In contrast, the male GENITALIA (= genital
capsule) do preserve well because they are strongly
sclerotised and so require no treatment other than
a straightforward extraction from the gaster witha
pin. The male genitalia consist of a pair of
gonoforceps for clasping the female and an intro-
PAULH. WILLIAMS
mittant organ (Figs 44 & 45). During pairing, the
female is locked with her sting assembly pulled
posteriorly and dorsally away from the male
(Williams, 1985: fig. 5) by at least three points of
contact: (1) the VOLSELLAE (= laciniae, but
see the discussion below) press the sting base from
above and (2) the GONOSTYLI (= squamae)
hold the rami of the female valvulae with an
Opposing action from below, so that the sting base
is gripped; at the same time, (3) the apex of the
female’s sternum VI is gripped between the male’s
parapenial processes of the GONOCOXITES (=
stipes) and the base of the SPATHA, ensuring
that the sting assembly is clamped in a protruded
position, which gives access to the female gonopore
for the male intromittant organ. Kopelke (1982)
describes how the PENIS VALES (= sagittae)
anchor among the weakly sclerotised folds that lie
between the rami of the valvulae of the female
sting base. Many of these structures of the male
genitalia show complex elaborations that are
particularly valuable taxonomic characters (Figs
46-205, see the comments on relationships among
species by ancestry). For the males, gastral sternum
VIII and sternum IX (= subgenital plate) are found
below the genital capsule and are normally retracted
above sternum VII. Variation in their shape and
pubescence has been described by many authors.
This variation may also be great even within
species and it is often accompanied by a pronounced
overlap in variation among related species.
Consequently these characters are not used here.
There has been some disagreement as to the
homology of the ‘lacinia’ of male bumble bee
genitalia, concerning whether it is derived from
part of the volsella or from part of the gonostylus
(e.g. Snodgrass, 1941; Smith, 1970). As yet no
study of ontogeny has been sufficiently detailed to
establish its homology conclusively, so the inter-
pretation depends on comparative studies of the
morphology of adults. Zander (1900) did study
the ontogeny of male bumble bee genitalia and
concluded that a small scale (‘Schuppe’) on the
inner ventral surface of the gonocoxite is of
volsellar origin because it is in the expected
position and becomes strongly chitinised early in
development. However, he made no detailed
observations on the derivation of the larger ventral
structure known as the ‘lacinia’. Later Smith (1970)
concluded from their topological similarity that
the ventral scale is derived from the digitus of the
ancestral volsella, but that the larger ventral
structure known as the ‘lacinia’ is derived from the
cuspis of the ancestral volsella. Snodgrass (1941)
had argued that Zander’s scale is the only expres-
sion of the volsella and that the large ventral
‘lacinia’ is derived from the ventral lobe of the
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
gonostylus. He supported this argument with the
observation that the ‘lacinia’ is connected with the
dorsal gonostylus for part of its length by a
membrane. However, against this argument it
must be noted firstly that some bees, including
centrines such as Epicharis rustica (Olivier) and
euglossines such as Eufriesea pulchra (Smith),
clearly show both major lobes of the gonostylus as
well as a third large ventral sclerite in a similar
position to the base of the ‘lacinia’ of bumble bees,
below the inner ventral margin of the gonocoxite
(see Fig. 5). Therefore this ‘lacinia’ cannot be the
ventral lobe of the gonostylus, although it is in a
reasonable position to be part of the volsella
and appears to be distinct from Zander’s scale.
Secondly, among these bees with a bilobed gono-
stylus, the two lobes are clearly strongly fused
near their bases, which is not known between the
gonostylus and the ‘lacinia’ for any bumble bees,
which have only the weakly sclerotised membrane
noted by Snodgrass. Traces of the two lobes of the
gonostylus may be retained among bumble bees as
the distal lobe and the interio-ventral process,
although these are reduced among species of the
subgenus Mendacibombus and among many
euglossine bees. Thus in the absence of a more
detailed ontogenetic study, the weight of evidence
supports the argument that a volsellar derivation
for the ‘lacinia’ is most likely.
RELATIONSHIPS AMONG SPECIES BY
ANCESTRY
The bumble bees, including the ‘cuckoo’ bumble
bees, form an easily recognised and monophyletic
(sensu Hennig, 1981) group of about 240 species
(present estimate). Their closest relatives are
believed to be the South American orchid bees,
the Euglossini (Winston & Michener, 1977;
Kimsey, 1984), which have been used in the
cladistic analyses for out-group comparisons.
Opinions differ concerning the relationships
among the many described species-groups of
bumble bees, so that many specialists working
with bumble bees currently follow their own
individual supraspecific classifications.
The present ideas of relationship among species
of bumble bees are revised and developed from
those proposed in a preliminary cladistic analysis
of the functional characters of the male genitalia
(Williams, 1985). In practice, most authors since
Schmiedeknecht (1882) and Radoszkowski (1884)
have used differences in characters of the male
genitalia to recognise species-groups among
13
bumble bees. Richards (1968) reviewed the varia-
tion in morphology among these established groups
and provided diagnoses that also include other
characters from both sexes. More recently, Ito
(1983, 1985) compared similarities between the
male genitalia of species in these groups in a
detailed phenetic study.
Some criticism has been made of the classifica-
tion of bumble bees primarily by characters of the
male genitalia because it relies on such a restricted
character set. But the characters of the male
genitalia that function in clamping the male to the
female for copulation (see the comments on
characters studied) are especially suitable for use
in classification not only because they present
many different character states but also because
there appears to be a particularly high congruence
in the patterns of these character states among
taxa (Williams, 1981, 1985). The evolutionary
interpretation of the high congruence in the
patterns of character states of bumble bee genitalia
is that there is relatively little homoplasy among
these characters. The classification of flowering
plants has long been based on characters of their
genitalia for similar reasons.
Evolutionary explanations for why male genitalia
are often so strongly differentiated among species
have included those that depend on a principal
role for the genitalia as a species-isolating
mechanism (the lock-and-key hypothesis, reviewed
by Shapiro & Porter, 1989), or as a mechanism for
female arousal (the stimulation hypothesis, see
Eberhard, 1985), or as an impregnable holdfast
mechanism (see below). From a study of bumble
bee morphology, Richards (1927a, 1927b) con-
cluded that at least in their case mechanical
incompatibility is unlikely to have evolved as an
isolating mechanism (see also the comments
on the biological species and the Recognition
Concept). At that time Richards had apparently
not actually observed how the genitalia fit together
during pairing between bumble bees, because he
referred to Boulangé (1924) for a description of a
pinned pair of B. hortorum (L.): ‘[p. 290] ...
les branches du forceps sont vigoureusement
appliquées par leurs extré — [p. 291] mités sur les
parties latérales du VII° sternite de la femelle
(c’est-a-dire le dernier visible en surface).’. This
led Richards (1927b: 262) to conclude that ‘. . . the
stipes, volsella and squama, that is the most
complex part of the genitalia, do not enter the
abdomen of the female at all .. .’. In contrast to
this, present evidence confirms that for bumble
bees the apices of the volsellae and gonostyli
(= squamae) do grip tightly around the female
sting base in the manner shown for B. (Melano-
bombus) lapidarius (see Kopelke, 1982: figs 1-3;
14
Williams, 1985: fig. 5), even among apparently
rather divergent species (e.g. from direct examina-
tion of live paired B. (Bombus) lucorum, B.
(Sibiricobombus) asiaticus, B. (Melanobombus)
rufofasciatus and from a freshly-killed pair of B.
(Fervidobombus) niger Franklin [= B. atratus
Franklin, a name preoccupied by B. mucidus var.
atratus Friese]). In fact, whether these claspers
grip sternum VI or the sting base has little effect
on the substance of Richards’s arguments against
the lock-and-key hypothesis, namely that the
supposed ‘lock’ structures of the females are
relatively undifferentiated among species and that
inter-specific pairing is probably rarely attempted
in any case. Eberhard (1985) believed that similar
comments by Richards (1927a) showed that the
male genitalia of bumble bees could not function
as ‘holdfast devices’, although I have seen two
pairs in museum collections that were still locked
together firmly by their genitalia.
Eberhard (1985) suggested that an alternative
explanation for the elaborate structure of male
genitalia is that they function as female stimulaters,
which have diverged as a result of inter-sexual
selection by female choice. Of course this idea
does not depend on an absence of the hoildfast
function, but it does require that a female should
be able to discern differences in stimulation by
males with subtly different genital morphology.
However, at present there is little evidence from
bumble bees either for the existence of the sensilla
that would be required in those heavily sclerotised
areas of the sting base that are clasped by the
males, or for the definite rejection of some males
in preference to others by females.
Another likely evolutionary explanation of the
great range of morphology of the male genitalia of
bumble bees is that they function primarily
as impregnable holdfast devices, which have
diverged as a result of intra-sexual competitive
selection among males (Richards, 1927a; Williams,
1985). In Kashmir, male bumble bees were seen to
attempt to catch and mount the females in order
to clasp them with their genitalia, without any
obvious courtship (for further details see the
comments on mate-searching behaviour of male
bumble bees in Kashmir). Any radical changes in
the functional characters of the male clasping
mechanism might only be inherited in the unlikely
event that they appeared at the same time as the
necessary reciprocal changes in the anatomy of
the female. Consequently characters of the male
claspers might be more conservative than characters
that do not have to fit the female securely in order
to prevent any interruption of copulation. Yet the
competition from other males of the same species
to dislodge a paired male for access to the same
PAULH. WILLIAMS
female might still favour any slight elaborations of
structure if this would increase that male’s chances
of hanging on to reproduce. Competition among
males can be intense (e.g. Free, 1971; Lloyd,
1981; pers. obs.), probably because a male bumble
bee’s principal influence on its reproductive success,
other than through the quality of its ejaculate or
possibly through the removal of competitor’s
sperm, is likely to be by maximising the number of
matings that it can achieve. This may account in
part for why bumble bee mating systems are
usually variations of scramble competition polygyny
(see the comments on male mate-searching
behaviour).
Genera of Bombini
Many of the inferences of relationship among
bumble bees from the preliminary cladistic analysis
of characters of the male genitalia remain very
tentative (Williams, 1985), especially those
for some of the most morphologically divergent
species (e.g. the North American B. nevadensis
Cresson, see the comments on the Kashmir
species of Mendacibombus). Nevertheless, further
evidence from characters of the females (Ito &
Sakagami, 1985) and from enzyme mobilities
under electrophoresis (Pamilo et al. , 1987) supports
the conclusion that the socially parasitic or “cuckoo”
bumble bees (Psithyrus) constitute a monophyletic
group. It also appeared from the results of the
preliminary cladistic analysis that Mendacibombus
could be the monophyletic sister-group to all
other bumble bees. This group in turn appeared to
consist of two monophyletic sister-groups, the
social parasites (Psithyrus) and all remaining
bumble bees. Consequently the use of three
genera was recommended: Mendacibombus,
Psithyrus and Bombus.
Now that it has been possible to examine the
male genitalia of most of the described species,
the present interpretation is that Mendacibombus
is probably paraphyletic with respect to all other
bumble bees (Fig. 5, Psithyrus + Bombus
represented by B. exil, see the comments on the
Kashmir species of Mendacibombus and B.
(Psithyrus) bohemicus). So recognising Mendac-
ibombus as a genus for these peculiar, high
mountain bees is unsatisfactory because it is no
longer well supported by evidence of monophyly.
But then the traditional concept of the genus
Bombus that includes Mendacibombus as a sub-
genus, but not Psithyrus, is also apparently para-
phyletic and so recognisable only by phenetic
similarity.
A pragmatic solution is to return to an emphasis
of the more widely- shared characters and of the
va)
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THE BUMBLE BEES OF THE KASHMIR HIMALAYA
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INISSOTONA
16
more distant affinities for the generic concept, with
the result that one genus, Bombus, is recognised
for all bumble bees, including Psithyrus. Of
course the use of Bombus for all of these species
does bring certain names originally ascribed to the
genus Psithyrus into secondary homonymy with
names in the genus Bombus. It also requires that
qualifications concerning the socially parasitic
species will have to be made more often to any
generalisations about the species of Bombus. But
despite this break with the most widely-established
bi-generic tradition, the use of one genus for all
bumble bees has the advantage that it recognises a
group for which the evidence of monophyly is
particularly strong, so that the nomenclature is
most likely to remain stable. This single genus is
also very easy for non-specialists to recognise.
Within the genus Bombus, the established
subgeneric names (e.g. Richards, 1968) may be
found convenient by some specialists who wish to
label those assemblages of species that are more
closely similar to one another. Unfortunately,
Richards’s usage of subgeneric names does not
always communicate ideas of relationship well,
because some of these assemblages now appear to
be paraphyletic (e.g. Mendacibombus) or even
polyphyletic (e.g. Sibiricobombus, see the com-
ments on B. asiaticus and B. oberti). Of course
subgeneric names can simply be treated as
synonyms of Bombus Latreille and ignored by
those who prefer to do so. But when a comprehen-
sive cladistic revision is available, then perhaps
the nomenclature of the more reliably mono-
phyletic species-groups could be revised, if the
system is still found to be useful.
In this review, the bumble bee species of
Kashmir are listed in a sequence that is chosen to
reflect their relationships (after the sequencing
convention of Nelson, 1972) as these are currently
envisaged.
DISCRIMINATION OF SPECIES AND
INTRASPECIFIC VARIATION
Early work on the taxonomy of Asian bumble
bees showed little consistency in the delimitation
of the taxa named as ‘species’ in some species-
groups. In part this has been due to the changing
nature of the species category as applied to living
organisms. The species concept adopted for this
review is briefly summarised here so that it is
explicit.
A particularly visible part of the problem of
discriminating species of bumble bees is that some
PAULH. WILLIAMS
of their populations are now known to include
individuals with very different colour patterns.
This was not always appreciated when only small
samples were available from widely-scattered
localities. Conspicuous colour patterns are likely
to be advantageous because they advertise a
warning to potential predators against the pain-
ful sting that female bumble bees can inflict.
Consequently, the theory of Miullerian mimicry
may explain not only the regional convergences in
colour pattern among unrelated species, but also
how parts of the same species may have diverged
in colour pattern in different regions of its
distribution (see the comments on characters not
related to male mate-searching and the discussion
of the colour patterns of the Kashmir fauna). It is
perhaps precisely because bumble bees are such
large and colourful insects that, like butterflies,
they have been widely collected and have received
so much attention in the literature. The consider-
able effort that has been invested in the descrip-
tion of subtleties of colour variation among some
bumble bees has left an unfortunate nomen-
clatural burden of over 3000 formal names (i.e.
about 12 names per species).
A large part of the more general problem in the
application of the category ‘species’ to living
organisms arises from confusion of the different
kinds and degrees of relationship among popula-
tions (reviewed by e.g. Queiroz & Donoghue,
1988). Sexually reproducing organisms have been
envisaged as living in populations that are united
by interbreeding, so that in the long term relation-
ships within the population tend to be reticulate.
This provides the possibility of some evolutionary
cohesion through gene flow, even though without
concerted contraction and expansion of popula-
tions, random ‘diffusion’ of genes may not be
sufficient to curb divergence (Barton, 1988). In
contrast, relationships among populations are
envisaged as tending to be more consistently
divergent. In the short term, parts of a population
may show restricted interbreeding with neighbour-
ing parts of the population and may even show
genetic and apomorphic divergence from them.
Only with the benefit of hindsight is it possible to
know whether divergence will be maintained or
whether it will subsequently be dissipated in the
broader population, should interbreeding become
reinstated when the parts of the population meet.
At one extreme, each of these parts of a popula-
tion that may be more or less reproductively
distinct in the short term could be regarded as a
terminal element and given a name. But it can be
seen that there is a broad range of possible
relationships among these parts of the popula-
tion, which may involve various degrees of
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
inter-breeding both past and present, in differing
combinations with various degrees of genetic
and apomorphic divergence. Consequently, no
absolute criterion can be devised that can univer-
sally define the species category.
Faced with the apparent impossibility of finding
a universal solution to the ‘species problem’, it
is still clearly desirable to try to minimise
the confusion in the application of the species
category. For the practice of discriminating species
in this review, I have attempted to explore the
application of species-defining criteria that place a
greater emphasis on the potential for interbreeding
(see the following section) than on general
apomorphic divergence. General apomorphic
divergence may have the appeal that it can be
considered to have resulted in an observable
pattern of monophyletic taxa, the discovery of
which may be independent of any ideas of the
process that caused it, which may not be discover-
able. However, this rejection of process robs
the ‘morpho-species’ category of any particular
theory-based significance and so allows it to be
defined only by a choice of some degree of
apomorphic divergence that, in consequence,
must be essentially arbitrary. Hence the cladistic
method is not appropriate for the discovery of
biological species. In contrast, the interbreeding
criterion should be most closely associated with
the process of gene flow within populations over
the long term, which is believed to be one of the
processes that can maintain some evolutionary
cohesion (see Templeton, 1989). Of course there
are also considerable problems with applying the
interbreeding criterion, which are discussed in the
next section. In practice, the application of the
interbreeding criterion to many cases requires a
similar degree of subjective judgement to that
required by the use of the general apomorphic
divergence criterion. Often the only available
evidence for interbreeding is very indirect and
must still be inferred from patterns of variation in
characters of morphology or of molecules.
The ‘biological’ species and the Recognition
Concept
Early authors may have regarded species as
convenient classes for the typological description
of a particular level of variation, but since Darwin
(1859), the evolutionary significance of biological
species has been stressed. This biological concept
defines species of sexually reproducing organisms
by a lack of interbreeding between populations,
where these overlap. One of the problems with
the biological species concept is that it does not
solve the problem of interpretation for populations
17
that do not overlap at present (reviewed by e.g.
Splitter, 1982), because in its simple form it does
not show how to obtain direct and representative
evidence of the ‘potential’ for interbreeding be-
tween individuals that do not meet under natural
conditions.
Dobzhansky (1937) suggested that isolating
mechanisms have been selected for because they
perform the function of preventing interbreeding
between species. Isolating mechanisms were said
to have the advantage of protecting their ‘more
harmonious’ gene assemblages from disruptive
recombination. But since isolating mechanisms
might be selected for only when populations with
such gene assemblages overlap, Dobzhansky’s
Isolation Concept provides little help in the
assessment of the potential for interbreeding
among non-overlapping (allopatric) populations.
Paterson (1980, 1985) criticised Dobzhansky’s
Isolation Concept of the species in response to
difficulties with the population genetics of the
selection of isolating mechanisms. Paterson’s
Recognition Concept defines a species by the
common ‘specific-mate recognition systems’ that
are shared by individuals in the population. From
this viewpoint, individuals of different species are
not expected to interbreed because they do not
recognise one another as potential mates (thus
‘postmating isolating mechanisms’ [Mayr, 1963]
are, strictly speaking, incidental to delimiting
species). Paterson stressed that the breakdown in
recognition at speciation might occur in allopatric
populations, but only indirectly, as a consequence
of other evolutionary divergences and not by
reinforcement of hybrid disadvantage (i.e. ‘pre-
mating isolating mechanisms’ [Mayr, 1963] evolved
as effects, and were not selected for their isolating
function). Evidence of this particular kind of
divergence in allopatry might help in the assess-
ment of the potential for interbreeding among
non-overlapping populations.
Paterson’s Recognition Concept is not entirely
satisfactory in that any allopatric populations with
effective postmating isolation but no premating
isolation would apparently be included in the
same species. If these populations subsequently
made contact, there would then be a selective
advantage for reinforcement by premating isola-
tion, as envisaged in the Isolation Concept.
Ultimately it is likely that both kinds of specia-
tion process have occurred, so that a major
contribution of the Recognition Concept may be
its emphasis on the role of premating barriers or
specific-mate recognition systems. Changes in
these specific-mate recognition systems could
evolve rapidly by sexual selection (see West-
Eberhard, 1983) in allopatry, despite stabilising
18
selection. Furthermore, neither stabilising selec-
tion nor sexual selection necessarily imply uniform
selection pressures across a geographically exten-
sive population and so they do not preclude the
possibility of clinal variation in the recognition
system (see Verrell, 1988). But if the characters
that are crucial to specific-mate recognition were
known to differ among individuals of at least some
of the problematic disjunct populations, then
the Recognition Concept might provide better
grounds for discounting a potential for inter-
breeding and gene flow in these cases (e.g. Vrba,
1985).
In principle, the most widely-applicable
morphological criterion for the discrimination of
species might therefore be found where there are
differences in the characters that function in
specific-mate recognition. However, the Recogni-
tion Concept of species does not require individuals
of separate species necessarily to differ at all in
morphology (or in ecology, see Hengeveld, 1988)
and the search for key characters for the discrim-
ination of species has indeed been unsuccessful
(reviewed by Templeton, 1981). This search can
be traced to the classical concept of species as
typological classes, whose members should share
some particular similarity or ‘essence’. More
recently species have been viewed as individuals
(Ghiselin, 1975), characterised instead by their
internal organisation, within the population (Hull,
1980). This organisation could be provided by their
shared specific-mate recognition systems and
potential for interbreeding. Yet from a cladistic
viewpoint, free interbreeding within a population
is a shared ancestral characteristic and so does not
define a taxon (Rosen, 1979). Cladists would
therefore be obliged to recognise some para-
phyletic terminal elements, because interbreeding
populations are not always distinguished by the
possession of convenient apomorphies by all
individuals (e.g. Ackery & Vane-Wright, 1984;
Queiroz & Donoghue, 1988). If apomorphic
characters were always to become fixed through-
out a population as a necessary part of the process
of speciation, then these character states might
fulfil the role of an essence. In contrast, character
states correlated with specific-mate recognition
systems are not essences because they may only
be expected to evolve as secondary consequences
of reproductive processes and they may occur in
just one of the sexes. So these characters can
provide no more than a guide for the inference
of the potential for interbreeding among in-
dividuals as parts of the same species in certain
cases.
PAULH. WILLIAMS
Mate-searching behaviour of male bumble
bees in Kashmir
The mate-searching behaviour of male bumble
bees represents some of the first stages in the
specific-mate recognition systems of these insects
(cf. Paterson, 1985: fig. 2). In Kashmir, differences
in male mate-searching behaviour among some
closely-related species are associated with dif-
ferences in some of their morphological character
states, such as eye size and shape, antennal length
and thoracic shape. Two contrasting examples are
summarised briefly here (see the comments on the
inference of allopatric, conspecific taxa & on
allopatric, separate species). By analogy, these
differences might provide a particularly strong
form of morphological evidence from which to
discriminate certain other likely species, even
among preserved specimens in collections.
The use of characters related to specific-mate
recognition systems for discriminating species still
requires support from the study of the subsequent
stages in the operation of these systems. Key
components in this may involve certain volatile or
contact pheromones. The head-gland secretions
of males of European ‘patrolling’ species are
believed to act as female arrestants (Svensson,
1979) and some of these secretions have been
characterised (e.g. Bergstrom et al., 1981). This
has not been undertaken for female head-gland
secretions, which may be just as important in the
subsequent stage of specific-mate recognition by
males (see Free, 1971; van Honk et al., 1978).
Some method to assay for behavioural discrimina-
tion between secretions of different composition
by the organisms themselves in the field is now
needed.
In 1985 I recorded the distribution of mate-
searching activity in time and space from transect
counts of numbers of male bumble bees around
Gulmarg. Distribution with respect to altitude is
described from altitudinal transects of the Pir
Panjal range (the mountains on which Gulmarg is
situated), between Tangmarg at 1800 m in the
Vale of Kashmir, and the peak of Mt Apharwat
above at 4143 m (13,592 ft) (total number of
bumble bees identified > 2000, see Fig. 6). The
habitat is described in more detail, together with
the altitudinal distribution of foraging females, in
the discussion of distribution with altitude. Spatial
and temporal pattern in the distributions of mate-
searching males can also be described at the
smaller scale of a 30 m-high hill at Gulmarg, from
vertical and horizontal transects (Figs 7 & 8).
I also recorded the activities of males that had
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Altitude
in feet (metres):
LES 13500 (4115)
Alpine 13000 (3962)
19
12500 (3810)
12000 (3658)
11500 (3505)
11000 (3353)
10500 (3200)
Subalpine
10000 (3048)
9500 (2896)
9000 (2743)
8500 (2591)
8000 (2438)
7500 (2286)
7000 (2134)
6500 (1981)
Upper forest
Gulmarg meadow
Lower forest
Terraces
trifasciatus
pyrosoma
hypnorum
tunicatus
FEMALES
13500 (4115)
Alpine 13000 (3962)
lucorum
rufofasciatus
himalayanus
kashmirensis
avinoviellus
lepidus
melanurus
astaticus
biroi
keriensis
12500 (3810)
12000 (3658)
11500 (3505)
11000 (3353)
10500 (3200)
10000 (3048)
9500 (2896)
9000 (2743)
8500 (2591)
8000 (2438)
7500 (2286)
7000 (2134)
6500 (1981)
Subalpine
Upper forest
Gulmarg meadow
Lower forest
Terraces
@ Patrolling males, same colour pattern as females
© Patrolling males, pubescence predominantly yellow
@ Foraging females
A Cruising males
@ Territorial males
* Racing males
Fig. 6 Distribution of mate-searching males (above) and foraging females (below) of the social bumble bees (i.e.
excluding species of the subgenus Psithyrus) with altitude on Mt Apharwat in the Pir Panjal Range around Gulmarg.
Altitudes are estimated from the 500 ft contours of the 1 : 63,360 British Survey of India Map (Oxford University
Library). Records are combined from transects made during July, August and September 1985. For a description of
the vegetation in each habitat zone, see the discussion of distribution of the fauna with altitude.
been individually labelled, using the plastic tags
that were developed for use with queen honey
bees (52 male bumble bees were labelled in the
vicinity of the 30 m hill in 1985; 108 males labelled
in the same area in 1986). Numbered tags are ideal
for the many perching males, which can be
approached for their tags to be read, but plain
discs of different colours are all that can be seen on
patrolling bees, which spend most of their time
in rapid flight. Labelling individuals did not
obviously affect them adversely. For instance, one
B. tunicatus male had resumed patrolling within
10 minutes of being released. Activities of males
were recorded in part to compare their time
20 PAULH. WILLIAMS
ee
oe Perches of racing B. asiaticus males (29.viii.1986).
@ _ _ Contested perch area of two territorial B. rufofasciatus males (20.viii.1985).
Cc] Circuit of places approached by a patrolling B. funicatus male (14.viii.1985).
Fig. 7 Sketch map of the 30 m hill on Gulmarg meadow, showing the distribution of mate-searching activity by
individually-labelled males of three species of bumble bees. Dotted lines show contours at 10 m intervals, black
rectangles show the positions of two huts.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
30°C Air temperature
20°C
sunrise
10°C
21
sunset
B. ferganicus
B. tunicatus
B. rufofasciatus
B. asiaticus
Time:
5.00 hrs 10.00
a Marking
Mate-searching
Feeding
10 individuals
15.00 20.00 hrs
Fig. 8 Distribution of activity by males of four species of bumble bees between dawn and dusk on the 30 m hill in
Gulmarg meadow. Counts were made every 30 mins during 19.viii.1985, along a transect of 450 paces around the 30 m
contour at the top of the hill (see Fig. 7).
budgets, but also to record the nature and out-
come of interactions between males. This included
mapping the circuits followed on the 30 m hill by
males of each of three species (B. ferganicus, B.
melanurus and B. tunicatus), and mapping the
perch positions of other species (B. avinoviellus,
B. rufofasciatus and B. asiaticus). The reactions of
males of all species to moving objects of different
sizes were observed and tested. Young queens of
B. asiaticus were tethered with lengths of thin
black cotton at different places on the 30 m hill
and the reactions of perching males were noted.
2D;
Number of records
oO
fo secs 10 secs
Number of records
PAULH. WILLIAMS
B. rufofasciatus
30 secs
LJ Flight durations
20 secs Perching durations
B Medians
30 secs ter
Fig. 9 Duration of consecutive flight and perching periods for mate-searching males of B. rufofasciatus (above) and B.
asiaticus (below). For each species, records of 20 consecutive flight and perching periods were made from each of five
males on the 30 m hill in Gulmarg meadow, during 11—12.viii.1985.
Four kinds of male mate-searching behaviour
were observed during 1985. These are referred to
here as ‘patrolling’, ‘racing’, true ‘territorial’
and ‘cruising’. The territorial behaviour can be
considered as an example of what has been termed
pure dominance or lek polygyny, whereas the
other three categories of behaviour represent
divergent forms of scramble competition polygyny
(see Thornhill & Alcock, 1983).
(A) PATROLLING behaviour has been described
previously from British bumble bees and from
most other European species (e.g. Darwin, see
Freeman, 1968; Awram, 1970; Schremmer, 1972;
Alford, 1975; Svensson, 1979). These males
appear to mark objects early each day with an
oral secretion, which presumably contains a
pheromone. The objects are numerous, spaced by
any distance from a few centimetres to some tens
of metres, and chosen so that they can be patrolled
as ‘approached places’ on a circuit flown regularly
by the male. It has been suggested that receptive
females are arrested at these pheromone ‘traps’
and that mating takes place when the male returns
on his next patrol of the circuit (see Svensson,
1979). Examples of species whose males perform
patrolling behaviour in Kashmir are B. pyrosoma
and B. tunicatus (e.g. records from the 30 m hill,
see Fig. 7). These males did not interact with one
another directly nor did they respond to moving
objects by pursuit, either near or between
approached places.
(B) RACING behaviour differs from patrolling in
that each male perches in one small area and will
then pursue potential mates from this look-out
position, but without competing with other males
for preferred perches.
Racing behaviour is described here from B.
asiaticus. These males perched on rocks, cowpats,
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
leaves or fragments of wood (a maximum of 107
males were recorded at their perches on the 30 m
hill on any one day, 29.viii.1986, Fig. 7). The
perches were not associated with flowers (i.e. food
resources) or nests (no nests of any species were
found on the 30 m hill in either 1985 or 1986).
Males would leave their perches in darting flights
in pursuit of rapidly moving objects, which included
conspecific queens and other males in flight, birds
flying overhead and even stones thrown past the
perch. Males rarely performed any slow inspec-
tion flights around perches (as shown by the short
durations of flights by B. asiaticus males in Fig. 9).
Successful pursuit flights with conspecific queens
ended with the male bringing the queen to the
ground, where he grappled with her, and attempted
to mount her and clasp her sting base with his
genitalia. There was no obvious courtship. When
a queen was tethered, males approached rapidly
and hovered for a few seconds, antennae almost in
contact, before attempting to clasp her. Once a
male had succeeded in clasping an untethered
queen with his genitalia, she ceased pushing at
him with her legs and the pair immediately flew
from the area in tandem.
When a pursuit flight with a conspecific male
resulted in an approach within a few centimetres,
they diverged without clashing and returned to
their perches without displacement of either male.
Males were rarely found to have damaged limbs
(2/108 labelled males, 1986, 30 m hill). Individual
males returned to the same perches on many days
(the maximum period spanned 27 days, 2-29. viii.
1986, male red-83, 30 m hill) and other males
rarely usurped these perches after they had
become vacant (occupancy is known to have
changed for 5/170 perches 2. viii-11.ix.1986, 30 m
hill, though only 108 males were individually
labelled). Marking of perches with oral secre-
tions was observed only occasionally, and only
immediately after males arrived at their perches in
the morning.
Some of these males were found to spend the
night torpid in tight clusters of up to 10 individuals,
deep among the short plant stems and roots on the
middle slopes of the 30 m hill. Some individuals
(e.g. male red-18, 1986) switched between clusters
on successive nights, although this could have
been the result of disturbance.
(C) TERRITORIAL behaviour differs from
patrolling in that each male perches in one small
area and will then pursue potential mates from
this look-out position, but unlike racing behaviour,
the males do compete for preferred perches.
Territorial behaviour is described here from B.
rufofasciatus (e.g. on the 30 m hill, see Fig. 7).
23
These males perched and pursued moving objects
much as racing males did, but they also responded
to conspecific males on their perches as well as in
the air, and clashed audibly with them during
extended chasing flights (Fig. 9). This may have
contributed to the higher incidence of damage
to appendages among these males (5/20 males
labelled on the 30 m hill, 1985, had lost at least half
of one antenna or leg) and did result in displace-
ment from perches. For instance, on 18.viii.1985
male green-7 was perched above the entrance to a
3 m-long, 1 m-deep gully at the edge of the flat-
topped 30 m hill. It left this perch to engage
yellow-7 as it entered the gully, but after the
ensuing chase, only yellow-7 returned to the gully,
where it then perched within 10 cm of where
green-7 had been. Later the same day green-7
returned, chased and replaced yellow-7 in this
gully (occupancy changed at least 14 times for 9
perches 9—20.viii.1985, with or without contest,
and 10 of these changes were for this gully).
Yellow-7 then moved to a less frequently occupied
perch area that was 10 m away on the side of the
hill. These perches were not associated with nests
or with concentrations of resources, but neverthe-
less continued to be highly preferred despite a
high turnover in individual males (only 9 perch
areas, each 2-4 m in diameter, could be found on
the entire 30 m hill, but 20 males were labelled in
these areas between 9-20. vili.1985). Males of B.
rufofasciatus marked their perches with oral
secretions much more frequently than did males
of B. asiaticus.
The frequent inspection tours of B. rufofasciatus
males around perch areas, such as the gully at the
top of the 30 m hill, have a characteristic
slow, stalling flight, with a low, interrupted buzz.
Pursuit flights, like those of B. asiaticus, are much
faster than the flight of females and have a higher-
pitched buzz.
(D) CRUISING behaviour differs from patrolling
in that each male uses one small area as a look-out
position from which to pursue potential mates.
Unlike racing behaviour, the males hover almost
stationary in the air rather than perch, and unlike
territorial behaviour the males do not compete for
preferred positions.
Cruising behaviour was not seen on the 30 m
hill and is described from observations of B.
kashmirensis from just a few days (most on
23.vili.1985) near the peak of Mt Apharwat.
These males maintained a very slow, almost
hovering flight about 1 m above the ground, but
each was confined within an area only 34 m
across. From these aerial positions they pursued
moving objects, much as racing and territorial
.ntennal Shape
24 PAULH. WILLIAMS
SEXUAL DIMORPHISM (males/queens)
asiaticus
melanurus
elongate
male antennae
kashmirensis
trifasciatus
avinoviellus
C) pyrosoma
himalayanus
75 |e
hypnorum rufofasciatus
birot @
keriensis
js
Fens ()
1.0
undifferentiated
male antennae
reduced
male eyes
narrowed
LS male thorax
enlarged
male eyes
Eye shape
Fig. 10 Sexual dimorphism in some characters that may be related to male mate-searching behaviour among the social
species of bumble bees (i.e. excluding species of the subgenus Psithyrus) of Mt Apharwat in the Pir Panjal Range.
Shape dimorphism is measured as the ratio of mean shape of males/mean shape of queens, so values of 1.0 show no
dimorphism in shape between males and queens. Eye shape is measured as the-ratio of the maximum right compound
eye length/minimum dorsal inter-compound eye distance; antennal shape is measured as right flagellum length/right
scape length; and thoracic shape is measured as distance between and including tegulae/right radial cell length. Sample
sizes (males/queens): B. avinoviellus (10/10), B. himalayanus (10/3), B. trifasciatus (8/5), B. melanurus (10/10), B.
kashmirensis (10/10), B. hypnorum (10/4), B. lepidus (2/6), B. biroi (6/5), B. tunicatus (10/10), B. lucorum (8/3), B.
asiaticus (10/10), B. keriensis (4/8), B. pyrosoma (10/10), B. rufofasciatus (10/10). Symbols show the different kinds of
male mate-searching behaviour and colour pattern dimorphism (see Fig. 6, strong divergence in male colour pattern
from worker colour pattern is only apparent among patrolling species): filled circles — patrolling males with similar
colour pattern to females; open circles — patrolling males with pubescence predominantly yellow; star — racing males;
squares — territorial males; triangle — cruising males.
America have also been seen to return to
perches between intermittent pursuit flights (e.g.
Schremmer, 1972; Haas, 1976; Alcock & Alcock,
males did on the 30 m hill. Cruising males also
interacted in flight, but no clashes with violent
physical contact were detected, no males were
Thoracic sha
found to have damaged limbs (0/12 of the males
collected) and no displacement of males was seen.
Marking of vegetation with oral secretions within
the cruised area was seen.
Patrolling behaviour is well known for bumble
bees, but the other three categories are distin-
guished here for the first time. Males of a few
species from central Europe and from North
1983). Pairs of males of some of these species were
seen to engage in protracted interactions, at least
occasionally, so their behaviour may correspond
to the territorial behaviour described here (e.g.
Schremmer, 1972; Alcock & Alcock, 1983).
Sexual dimorphism in relative eye size, antennal
length, wing shape and thorax shape is least strongly
developed in patrolling (‘marathon’) species and
more pronounced among the other ‘perch and
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
sprint’ categories (e.g. review by Schremmer,
1972), especially for the racing species (Fig. 10).
The species that patrol exclusively in the upper
forest show pronounced sexual dimorphism in
colour with similar, almost uniformly bright yellow
males (see the discussion of colour patterns of the
Kashmir fauna).
Relationship between male mate-searching
behaviour and habitat structure
It seemed likely from a comparison of the male
mate-searching behaviour among the bumble
bees observed during 1985 that each of the four
categories could have advantages for increasing a
male’s number of matings. But which of these
kinds of behaviour is most advantageous to an
individual male for maximising his mating success
is likely to depend on the particular conditions of
density and dispersion in the distributions of
receptive females. This scheme is summarised in
Table 2. Each species’ characteristic density and
dispersion of receptive females is expected to
reflect the particular responses by its individuals
to the different kinds of habitat in the mountains:
Table 2 Summary of the likely characteristics of the
distributions of receptive female bumble bees on Mt
Apharwat (altitudes 2700-4143 m), for the most
abundant species in the four principal kinds of
habitats (1985 survey). These different distributions
may each favour one of the four groups of male mate-
searching behaviour (in bold).
Dispersion
high low
(even) (clumped)
meadow high alpine
high RACING CRUISING
e.g. B. asiaticus e.g. B. kashmirensis
Density
forest subalpine
low PATROLLING TERRITORIAL
e.g. B. pyrosoma e.g. B. rufofasciatus
(A) PATROLLING. Males of all species found
around Gulmarg to be mate- searching only within
the forest (1900-3000 m) (e.g. B. pyrosoma), and
males of the species most widespread among the
different kinds of habitat (e.g. B. tunicatus),
were patrolling in search of mates (Fig. 6). The
densities of all bumble bees are expected to be
especially low in the coniferous forest, where
floral resources are scant (see the discussion of
distribution with altitude). Dense forest has a
complex physical structure, which reduces the
visibility of potential mates and obscures any long
25
range visual cues that might be used by individuals
to aggregate for pairing. A web of pheromone
traps, formed by the approached places that these
males patrol, should be especially well suited
for collecting mates at low, relatively uniform
densities. Patrolling behaviour is also likely to
work especially well in open habitats where young
queens are present at low density and may not be
predictably aggregated (e.g. arctic environments).
(B) RACING. Resources in the large meadow of
Gulmarg are apparently much richer, because it
appears to have the highest density of bumble
bees among all of the areas surveyed on Mt
Apharwat (see the discussion of distribution with
altitude). For species specialising in this open
habitat, patrolling behaviour might be expected to
break down at high density if some males were to
exploit the pheromone ‘traps’ at approached
places by perching there and waiting for the more
frequent females to arrive. At high density, males
that persisted in patrolling would be wasting
much of their time in flight between traps (when
patrolling males may not be responsive to recep-
tive females, see Free, 1971), without increasing
their chances of finding a receptive female.
Racing B. asiaticus were present at more
uniformly high densities than patrolling species, in
some open, grassy areas below the forest (1800—
1900 m), in parts of the meadow at Gulmarg
(2650-2700 m) and just above the upper edge of
the forest (3000-3100 m, Fig. 6). Males appeared
to be less aggregated than those of the cruising or
territorial species, which are associated with the
higher relief, alpine habitats. This presumably
reflects either a more even distribution of recep-
tive females, or the unpredictable spatial pattern
of this distribution in the meadow. Transect
counts show a tendency for males to concentrate
on steep ridges (vertical transects 16 & 23.viii.
1985: 53/86 males in steepest middle half of slope,
7.5—22.5 m up the 30 m hill; horizontal transect
18.vili.1985: 23/26 males on ridges rather than
gullies) of south and east-facing slopes (i.e. facing
the sun in the morning, see Fig. 7), but perch sites
were still relatively widely and evenly spread. The
mate-searching activities of B. asiaticus males
were also spread over a longer period of weeks
than those of the territorial or cruising species in
subalpine and alpine habitats. This may reflect the
longer season during which floral resources are
available to fuel colony development and the
rearing of young queens at the lower altitude of
Gulmarg.
(C) TERRITORIAL. Above the forest, in the
subalpine zone, the season of profitable foraging
26
is shorter, the levels of resources may be lower
than in the forest meadows, and so the bumble bee
density is expected to be lower (see the discussion
of distribution with altitude). The open subalpine
zone covers a large area on Mt Apharwat, although
it also provides abundant topological cues that could
be exploited to facilitate the meeting of the sexes
even if they are present at low density, for instance
by local ‘hilltopping’ (hilltopping could arise
through the evolution of female choice, see
Kirkpatrick, 1982).
The most abundant species above the forest is
B. rufofasciatus. Males were indeed seen to be
very closely aggregated in small groups on the tops
of shoulders protruding from the subalpine slopes
of Apharwat (3000-3800 m) and, more rarely, in
the alpine zone (c. 3900 m) and on top of the 30 m
hill in the meadow at Gulmarg (2700 m, two perch
areas shown in Fig. 7). At any one time these
males were few in number, but were all closely
associated with even fewer perch areas. So although
perch areas were widely spaced, they each had up
to seven males flying around them. If this does
reflect the pattern in the distribution of receptive
females, then territorial defence is likely to be
more advantageous for these males than for
species at higher densities, because it could win
exclusive access to females at an especially attrac-
tive site, while the probability of missing a female
during a male-male chase would be lower.
(D) CRUISING. Habitat conditions in the alpine
zone resemble those in the subalpine, but are
even more extreme in features such as the short
foraging season (see the discussion of distribution
with altitude). However, this open habitat has the
ultimate topological cues for potential mates at
low density to meet by hilltopping — the mountain
peaks.
The most abundant high alpine specialist is B.
kashmirensis. Workers forage throughout most of
the alpine and subalpine zones (3000-4000 m, Fig.
6), but cruising males were found to be aggregated
exclusively in the high alpine zone (4000-4100 m),
in the lee of the peak of Mt Apharwat (4143 m).
Moreover, males were seen on just a few of the
days during which this area was visited, although
in larger aggregations than the males of B.
rufofasciatus, so that mate-searching activity may
be highly concentrated in time as well as in space.
Both factors could contribute greatly to enhance
the encounter rate or effective local density of
mates. If the female arrival rate were sufficiently
high, then territorial chases between males might
result in more females being missed, or lost to
‘sneaky’ males, than would be lost to tolerated
competitors. The hovering flight of cruising males
PAULH. WILLIAMS
may be one way of gaining an advantage over
perching competitors, both by saving time on
take-off in pursuit of a potential mate and possibly
by increasing the field of view. Thus it may be
that, whereas racing males tolerate one another’s
proximity because no one perch site is predictably
much more attractive to females than any other, a
male cruising over a particularly attractive site
tolerates the proximity of another male because
displacing the competitor might cost more in
lost opportunities than could be gained through
attempts at sole possession.
The characteristics of density and dispersion in the
distributions of receptive female bumble bees
could not be measured directly. Young queens are
not often seen before winter. More particularly,
bumble bees have rarely been found in copula,
despite their abundance. Preliminary searches
had shown that male marking activity is greatest in
the early morning, so I had expected that searching
at this time might yield more observations of
pairing. In fact this was seen only four times, all in
the afternoon (e.g. Fig. 8). An explanation for the
rarity with which bumble bees are seen paired,
despite the length of time for which females
remain clasped by the males (usually in excess of
20 minutes, pers. obs.), may lie with the observa-
tion that paired females and males flew in tandem
from the areas where males were mate-searching
as soon as the males had clasped the females,
which took just a few seconds. This scattering to
inconspicuous places may help to avoid compet-
itive interference from other males of the same
species. Unfortunately it also precludes the use of
any observations of paired bumble bees as a
measure of the distribution of receptive females.
Less direct methods may nonetheless reveal the
distribution of receptive females. It can be assumed
that it would be to the advantage of each indivi-
dual male if the male were to invest its mate-
searching efforts in time and space in a pattern
that closely resembles the predictable component
in the distribution of receptive females. Therefore
the density and dispersion of mate-searching
males should reflect the pattern for receptive
females.
Estimates of density and dispersion of mate-
searching males were made from transect counts
in each of the principal habitats at Gulmarg during
the summer of 1986 (Tables 3 & 4). Only 54 mate-
searching males of B. pyrosoma, B. asiaticus, B.
rufofasciatus and B. kashmirensis were recorded
during this transect census. The spring thaw had
been unusually late and the summer was cold and
wet. The peak of mate-searching activity by males
of the alpine and subalpine bumble bee species
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Table 3 Estimates of mean/variance ratio in the
number of mate-searching males among 160 m
segments of transect on Mt Apharwat as a measure of
male dispersion (12 & 14.ix.1986). The figures were
calculated for each of the most abundant species only
within its associated habitat (see Table 2).
Dispersion
high low
(even) (clumped)
B. asiaticus 0.51|B. kashmirensis 0.21
B. pyrosoma 0.61 | B. rufofasciatus 0.14
Table 4 Estimates of mean number of mate-searching
males counted along each 1 km of transect on
Mt Apharwat as a measure of male density (12 &
14.ix.1986). The figures were calculated for each of
the most abundant species only within its associated
habitat (see Table 2).
high 8B. asiaticus 8.8 B.kashmirensis 4.4
Density ——
low B. pyrosoma 3.1 B. rufofasciatus 1.7
was about 20 days later in 1986 than it had been in
1985 and their density also appeared to be lower in
1986 (e.g. males of B. rufofasciatus were found on
the 30 m hill at Gulmarg during 1985 but not
during 1986). Further counts were prevented by
the first heavy fall of snow, which blanketed the
subalpine and alpine zones the morning after the
first complete transect census.
Despite the small size of the sample in 1986, the
figures in Tables 3 & 4 do appear to corroborate
the expected characteristics of the common bumble
bee distributions as postulated in Table 2, at least
insofar as the figures in adjacent cells of the tables
differ in the expected directions. Of course this
result is only an apparent correlation based on
observations of the bumble bees on one mountain
and does not constitute a test of the relationship.
The behaviour of one group of B. rufofasciatus
males on a high, steep-sided shoulder of Mt
Apharwat (c. 3700 m, 23.viii.85) provides stronger
support for the importance of distribution, although
it is still only circumstantial evidence. In this
situation their behaviour was apparently not
territorial but resembled the cruising of B.
kashmirensis. Conforming with the pattern in
Table 2, these B. rufofasciatus males had formed a
large group of unusually high density (estimated
to be 50-100 males). However, it must be noted
that individuals of most species are apparently
not this flexible in their behaviour. For instance,
B. asiaticus males were never found patrolling,
even where they were present at low density. No
males of patrolling species were found racing,
27
either at localities with unusually high density or
otherwise.
If differences in habitat structure between
separated (allopatric, peripatric or parapatric)
parts of an ancestral population may affect the
balance of advantage among the different kinds
of mate-searching behaviour, then the mate-
searching behaviour of the descendent populations
could diverge very rapidly by sexual selection.
This may be particularly likely to affect the
specific-mate recognition system and could result
in speciation (see Rice, 1987, for a similar model
for sympatric speciation).
Inference of allopatric, conspecific taxa
Several nominal taxa that are closely similar in
morphology show pronounced variation in the
colour pattern of the pubescence among adjacent
regions. One example is provided by individuals
with the two principal colour patterns that are
ascribed to B. asiaticus in Kashmir (see the
comments on B. asiaticus). Individuals with both
the unbanded yellow pattern (e.g. Figs 371-376)
and the banded grey-white pattern (e.g. Figs 386,
387, 389-391) of B. asiaticus occupy similar open
meadow or steppe habitats on either side of the
Great Himalaya range (Map 48). The mate-
searching behaviour of the males in the two
regions could not be distinguished and consists of
the racing behaviour that is unique to these
nominal taxa among the bumble bees of Kashmir.
This close similarity in mate-searching behaviour
is reflected in the similarity of the associated
morphological characters of the males, such as
their much larger eyes and longer antennae in
relation to those of the females (Fig. 10). Support-
ing evidence for interbreeding between individuals
with the different principal colour patterns is
provided by specimens with intermediate colour
patterns (Figs 377-382) from a few localities in
high valleys of the Great Himalaya. The frequencies
of the different states for two of the three variable
colour characters examined from these localities
is consistent with simple models of population
genetics (see the discussion of variation of B.
asiaticus in Kashmir). So as parts of what appears
to be a single, interbreeding population, these
nominal taxa are considered to be parts of a single
species.
Inference of allopatric, separate species
Discontinuous geographical variation has also
been found in the morphological characters that
are associated with the different kinds of male
mate-searching behaviour, between what are in
28
other respects very similar nominal taxa. B.
wurflenii is widespread in Europe among the
upper forests of mountains, where the males,
which patrol their scent-traps within the forests in
search of mates (pers. obs., Austrian Alps), have
eyes similar in size to those of the females. The
morphologically similar B. kashmirensis is wide-
spread in the Himalaya and eastern Tibet (Map
31) among the open alpine zones. The principal
difference from B. wurflenii is that the males
of B. kashmirensis possess slightly-enlarged eyes
relative to the females (Fig. 10). On Mt Apharwat
in Kashmir at least, these males apparently
aggregate in the neighbourhood of mountain
peaks, where they ‘cruise’ in search of females by
sight. If this behaviour is representative and
constant, then individuals of the two taxa would
be unlikely to interbreed even if they were to
occur on the same mountain, because they appear
to search for mates in different ways and in
different kinds of habitat. They may therefore
have diverged in at least the first stages of their
specific-mate recognition systems and so can
be considered to be separate species (see the
comments on B. kashmirensis, and on another
group of more or less parapatric sister-species, B.
simillimus, B. pyrosoma and B. rufofasciatus).
Characters of species not related to male
mate-searching behaviour
Unfortunately the mate-searching behaviour
component of the specific-mate recognition system
only differs sufficiently among species for diver-
gence in associated morphological characters to
be apparent in a few cases. So, of necessity,
discrimination of most species still depends on the
interpretation of even less direct evidence of
interbreeding. This is usually inferred from a
diagnosis by a combination of other morphological
characters. Specimens are now available from
many more localities in Kashmir so that a more
comprehensive examination of the variation in
these characters among individuals can be made.
At the beginning of this study, cladistic methods
were used to divide material in collections into
what are likely to be monophyletic taxa by
character state patterns of the male genitalia.
Material was further sorted by sample sites.
Patterns of variation in a broad range of characters
were then examined for description, both within
and among these groups of individuals.
For the discrimination of species, particular
attention was paid to any strictly coincident
discontinuities in the patterns of character variation
among. individuals, irrespective of whether they
occur between the supposedly monophyletic taxa
PAULH. WILLIAMS
or within them. These coincident discontinuities
may provide evidence for barriers to interbreed-
ing between populations and so may show the
presence of separate species, which need not
correspond to monophyletic taxa.
On the other hand, where individuals were
found that show intermediate or mixtures of
character states between the otherwise differen-
tiated taxa, this is interpreted as evidence in
favour of the occurrence of hybridisation between
them, as parts of a single population and a single
species. For instance, in Europe wide variation is
known to occur between parapatric parts of the
population of B. soroeensis (Fabricius). Individuals
of this species from much of Russia, Scandinavia,
Britain and Spain are banded yellow bumble bees
with white ‘tails’. Individuals from central and
south-eastern Europe are unbanded, red-‘tailed’
bees. But individuals from the intervening areas in
both France and eastern Europe show a continuous
range of variation in both characters between the
two, more widespread, colour patterns (Reinig,
1939: fig. 10). In some areas, individuals with both
principal colour patterns may even be reared from
the same nest, apparently as the offspring of the
same queen (Vogt, 1909). Therefore individuals
with either of these two colour patterns have been
considered to be interbreeding as parts of the
more or less continuous population of a single
species.
Zones of hybridisation between nominal taxa
can be very narrow, with steep clines in character
frequencies (see e.g. Mallet, 1986; Hewitt, 1988).
For instance, localised steep clines in hybrid zones
are well known for groups of colour pattern
characters within populations of certain European
bumble bees (e.g. Reinig, 1970). Yet as long as
some evolutionary cohesion could be maintained
by the passage of some alleles across a hybrid zone
through interbreeding, then the parts of the
population could be considered to be parts of a
single species. Among Sino-Himalayan bumble
bees, some of the most difficult decisions concerning
conspecific status involve bees from the montane
forests (e.g. see the comments on B. lepidus, B.
pyrosoma). These forests are confined to a long,
narrow ribbon around the southern and eastern
edges of the great Tibetan massif. In this situation,
any hybrid zones across this ribbon are likely to
be much shorter than those described for B.
soroeensis in Europe and so more difficult to
locate and study adequately.
If some of the provisional decisions reached in
this review in cases where the available information
remains inconclusive seem unduly biassed in the
direction of ‘lumping’, it is because the burden of
proof of a divisive speciation event within an
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
ancestral population is considered to rest with the
‘splitter’. It has been argued that evidence from
subtle differences between two individuals is not
sufficient. For example, it can be seen that the
people from the different parts of Europe, of
Asia, or even from different areas within Kashmir,
also vary in shape and hair colour, and yet from
their breeding behaviour they would all be placed
in one biological species.
But irrespective of the decisions reached here,
as much of the available information as time has
allowed is presented in the comments on the
variation within Kashmir and in the figures and
maps, so that readers with a different concept of
the nature of species should be able to make their
own decisions regarding this fauna.
The description of intraspecific variation
Many examples of character variation among
samples of a single species can be related to where
the samples were collected within the species’
distribution. Often this geographic variation in
single characters can be regarded as belonging
to a continuum between extreme states, so that
defining two or more discrete states would require
threshold criteria that must be essentially arbitrary.
Furthermore, where comparisons are drawn
between two or more characters, these may show
geographically incongruent patterns of variation,
so that most combinations of states can be found
somewhere within the species’ distribution.
Therefore, because variation of characters among
individuals does not always delimit discrete sub-
units within populations (Wilson & Brown, 1953;
but see Mallet, 1986; Barton, 1988), previously
described nominal taxa of subspecific rank are not
attributed here with formal names. This removes
any requirement for a general subspecies concept
that would otherwise necessitate imposing more
assumptions about the genetic structure of
populations. The significance of intraspecific
variation is better judged separately for the different
cases that are presented by each species.
In the place of the earlier systems of subspecific
and infrasubspecific nomenclature, in this review
the most obvious character states are referred to
using shorthand descriptions. An example is an
‘unbanded (= no band of black pubescence
between the wing bases) yellow (= colour of the
pale pubescence, excluding any red hairs on the
apical terga) queen (= large female) of Bombus
asiaticus.. This terminology may appear less
elegant, but it has the advantage that it is easily
modified to give a more precise description of the
recognised attributes of a particular individual.
Intraspecific variation of selected characters is
29
described separately for each species as it is
represented in the samples available, together
with the frequencies of the various states in
these samples. These frequencies may have been
biassed at some sites by selective collecting, at
least for the later samples (1985, 1986). But for the
maps of the geographical distribution of variation
in the bees’ colour patterns, it is necessary
for clarity of presentation to resort to a more
typological approach, referring instead to the
predominant local phenotype among the workers
(see the figures accompanying the maps). This
emphasis is placed on the workers because they
are usually the most abundant sex and caste to be
seen by collectors. Species’ distributions within
Kashmir are shown by spot maps rather than by
grid maps, because sampling effort could not be
even nearly uniform across the country (Map 4;
see the comments on material examined and
depositories).
NOMENCLATURAL SUMMARY
Where possible type specimens have been
examined in order to assign specific nomenclature
more precisely. References in the text to ‘Art. n’
allude to the numbered articles of the /nternational
Code of Zoological Nomenclature (ICZN, 1985).
Synonyms are listed here only in cases where I
have examined the types or specifically discussed
their identity. Names of taxa whose status as part
of the same species is in particular doubt are cited
as provisional synonyms. Unavailable names are
cited in brackets. Nomenclatural details for taxa
in the genus group are not repeated in this section.
BOMBUS Latreille 1802a
avinoviellus (Skorikov, 1914a)
callophenax Cockerell, 1917 syn. n.
[subtunicatus Richards, 1930, infrasubspecific]
himalayanus (Skorikov, 1914a)
varius (Skorikov, 1914a) (provisional synonym)
marussinus Skorikov, 1910a
afghanus Reinig, 1940 syn. n.
bohemicus Seidl, 1837
nemorum (Fabricius, 1775)
novus (Frison, 1933) comb. n.
nepalensis (Tkalct, 1974b) (provisional synonym)
branickii (Radoszkowski, 1893) comb. n.
eriophoroides (Reinig, 1930)
elisabethae (Reinig, 1940)
ferganicus (Radoszkowski, 1893) comb. n.
indicus (Richards, 1929a)
morawitzianus (Popov, 1931) comb. n.
redikorzevi (Popov, 1931)
30
skorikovi (Popov, 1927b) comb. n.
[mesoxanthus (Richards, 1928c) infrasubspecific]
gansuensis (Popov, 1931)
trifasciatus Smith, 1852a
montivagus Smith, 18785 syn. n.
secundus Dalla Torre, 1890 syn. n.
ningpoensis Friese, 1909
wilemani Cockerell, 1911 (provisional synonym)
albopleuralis Friese, 1916 syn. n.
maxwelli Pendlebury, 1923 (provisional synonym)
mimeticus Richards, 1931 syn. n.
albolateralis Richards, 1931 syn. n.
gantokiensis Richards, 1931 syn. n.
turneri Richards, 1931 syn. n.
insidiosus Richards, 1931 syn. n.
geminatus Richards, 1931 syn. n.
magrettianus Richards, 1931 syn. n.
minshanicus Bischoff, 1936 syn. n.
quasibreviceps (Tkalct, 1968b) syn. n.
atropygus (Tkalci, 1989) syn. n.
haemorrhoidalis Smith, 1852a
orientalis Smith, 1854 syn. n.
buccinatoris Smith, 1879 syn. n.
assamensis Bingham, 1897 syn. n.
montivolans Richards, 19295 syn. n.
khasianus Richards, 1929b syn. n.
semibreviceps (Tkalcu, 19685) syn. n.
semicoloricontrarius (Tkalct, 1968b) syn. n.
cinnameus (Tkalct, 1989) syn. n.
melanurus Lepeletier, 1836
tschitscherini Radoszkowski, 1862 syn. n.
difficillimus Skorikov, 1912b (provisional synonym)
subdistinctus Richards, 1928b syn. n.
griseofasciatus Reinig, 1930 (provisional synonym)
maidli Pittioni, 19396 syn. n.
personatus Smith, 1879
roborowskyi Morawitz, 1886 syn. n.
kashmirensis Friese, 1909
stramineus Friese, 1909
tetrachromus Cockerell, 1909
pulcherrimus (Skorikov, 1914a)
meinertzhageni Richards, 1928b
[albohirtus Richards, 1930, infrasubspecific]
beresovskii (Skorikov, 19335) syn. n.
hypnorum (Linnaeus, 1758)
bryorum Richards, 1930 syn. n.
fletcheri Richards, 1934 syn. n.
subtypicus (Skorikov, 1914c) comb. n.
leucopygus Morawitz in Fedtschenko, 1875
leucurus Bischoff & Hedicke, 1931
kohistanensis (Tkalcu, 1989) syn. n.
Jemniscatus Skorikov, 1912b
flavopilosus Friese, 1918 syn. n.
peralpinus Richards, 1930
lepidus Skorikov, 1912b
genitalis Friese, 1913 syn. n.
tetrachromus Friese, 1918 syn. n.
yuennanicola Bischoff, 1936 (provisional synonym)
hilaris (Tkalct, 1989) syn. n.
biroi Vogt, 1911
PAULH. WILLIAMS
flavobistriatus Vogt, 1911 syn. n.
[flavostriatus Vogt, 1911, infrasubspecific]
[flavofasciatus Vogt, 1911, infrasubspecific]
nursei Friese, 1918 syn. n.
abbotti Cockerell, 1922
agnatus Skorikov, 1933b (provisional synonym)
kotzschi Reinig, 1940 (provisional synonym)
tunicatus Smith, 1852a
gilgitensis Cockerell, 1905 syn. n.
simlaensis Friese, 1909
fulvocinctus Friese, 1909
Jucorum (Linnaeus, 1761)
cryptarum (Fabricius, 1775) (provisional synonym)
modestus Cresson, 1863
moderatus Cresson, 1863
magnus Vogt, 1911 (provisional synonym)
jacobsoni Skorikov, 19126 syn. n.
lucocryptarum Ball, 1914 (provisional synonym)
longipennis Friese, 1918 syn. n.
alaiensis Reinig, 1930 syn. n.
mongolicus Kriiger, 1954 syn. n.
reinigi Tkalcu, 19746 syn. n.
asiaticus Morawitz in Fedtschenko, 1875
longiceps Smith, 1878a
regeli Morawitz, 1880
miniatocaudatus Vogt, 1911
[fuscocaudatus Vogt, 1911, infrasubspecific]
[albocaudatus Vogt, 1911, infrasubspecific]
[tenuifasciatus Vogt, 1911, infrasubspecific]
falsificus Richards, 1930 syn. n.
flavodorsalis (Skorikov, 1933b)
oshanini (Skorikov, 1933b)
oberti Morawitz, 1883
semenovi Morawitz, 1886 syn. n.
duanjiaoris Wang, 1982 syn. n.
ladakhensis Richards, 1928b
phariensis Richards, 1930 syn. n.
variopictus Skorikov, 19335 syn. n.
bianchii Skorikov, 1933b
reticulatus Bischoff, 1936 syn. n.
semenovianus (Skorikov, 1914a)
problematicus Bischoff, 1935
keriensis Morawitz, 1886
separandus Vogt, 1909 syn. n.
kohli Vogt, 1909 syn. n.
postzonatus Vogt, 1909 syn. n.
kozlovi Skorikov, 19105 syn. n.
incertoides Vogt, 1911 syn. n.
tenellus Friese, 1913 syn. n.
meridialis (Skorikov, 1914a) syn. n.
alpivagus Richards, 1930 syn. n.
karakorumensis (Tkalct, 1989) syn. n.
simillimus Smith, 18526
grossiventris Friese, 1931 syn. n.
oculatus (Frison, 1933) syn. n.
tonsus (Skorikov, 1933b) syn. n.
haemorrhous Richards, 1934 syn. n.
pyrosoma Morawitz, 1890
flavothoracicus Bingham, 1897 (provisional synonym)
miniatus Bingham, 1897 (provisional synonym)
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
[canosocollaris Skorikov, 1912b, infrasubspecific]
friseanus Skorikov, 1933a syn. n.
formosellus (Frison, 1934) (provisional synonym)
hoenei Bischoff, 1936 syn. n.
[flavocorbicularis Tkalct, 1961, infrasubspecific]
wutaishanensis (Tkalct, 1968a) syn. n.
rufofasciatus Smith, 1852b
prshewalskyi Morawitz, 1880
rufocinctus Morawitz, 1880
chinensis Dalla Torre, 1890
championi Richards, 1928a syn. n.
[rufior Richards, 1928b, infrasubspecific]
[intermedius Richards, 1930, infrasubspecific]
waterstoni Richards, 1934
TAXONOMY OF THE KASHMIR
BOMBINI
BOMBINI Michener
Bombini Michener, 1944: 290. Type genus:
Bombus Latreille, 1802a: 437.
In this review, formal descriptions of bumble bees
are not repeated because the literature already
contains many descriptions of particular individuals.
Instead I describe just a few characters that may
be useful to distinguish species. Tentative inter-
pretations of some of these character-state patterns
are included for the continuing discussion of
relationships among species.
In the keys, those characters that have been
found more reliably diagnostic have generally
been placed nearer the beginning of each couplet.
Characters of the genitalia are used frequently in
the key to males (see Figs 44 & 45), so male
specimens should be prepared for examination
by extraction of their genital capsules. Colour
patterns of the pubescence are used in the final
stages of the keys for the determination of species
within a species-group, where the range of varia-
tion does not overlap among these species within
Kashmir. But after a reference collection has been
prepared for a particular locality using the keys, it
may often be possible to identify the majority of
further specimens accurately by direct comparison,
using only selected details of their colour patterns.
Diagrams of the colour patterns are provided here
to illustrate some of the variation (Figs 232-439).
These diagrams should be coloured by hand to
render them more immediately intelligible (see
Fig. 258 for a colour key). Inevitably these
diagrams are a compromise between portraying
the general appearance of an insect and accurately
showing the position of every coloured hair. For
31
instance, the presence of a minority of black hairs
among pale pubescence could not be shown
without reducing the general resemblance of
the diagram to the insect. Consequently these
diagrams provide only a guide that should not be
used without the keys.
Records of food-plants are included from notes
made in the field during 1980, 1985 and 1986.
These plant species were identified by reference to
Polunin & Stainton (1984). The lists are by no
means exhaustive, but they do include at least
some of the preferred species.
Key to species (females: queens and workers)
1 Outer surface of hind tibia flat or partially concave,
without long hairs on posterior part of lower or distal
half of outer surface (corbicula), but with a comb of
stout spines (rastellum) along inner distal margin
(Figs 225 & 226); gastral sternum VI without ventro-
lateral keels; clypeus strongly protuberant, with
lateral margins curved back towards occiput (Fig.
ES) St. eee eres Mes Sone eet 2)
— Outer surface of hind tibia convex, with moderate to
long hairs throughout, but without a comb of stout
spines along inner distal margin; gastral sternum VI
with a pair of ventro-lateral keels (Figs 206-211);
clypeus nearly uniformly flat, only apico-lateral
corners curved back strongly towards occiput ..... 5
2 Outer (corbicular) surface of hind tibia usually
smooth and shining or only weakly sculptured (retic-
ulate coriaceous), without any long stout hairs aris-
ing from posterior part of outer surface below upper
or proximal quarter (Fig. 226); proximoposterior
corner of hind basitarsus strongly and acutely pro-
duced, projection longer than its own basal breadth
(Figs 41 & 42); labrum with basal transverse depres-
sion extending apically as a deep median furrow
between pronounced lateral tubercles, displacing
ridge between them to form a lamella that overhangs
apical margin (Figs 25 & 26) ................... 10
— Outer (corbicular) surface of hind tibia coarsely
sculptured (imbricate), appearing very rough, with
widely-spaced long stout hairs or bristles arising from
near middle of outer surface throughout upper or
proximal half (Fig. 225); proximo-posterior corner
of hind basitarsus rounded, projection no longer
than its own basal breadth (Fig. 43); labrum with a
narrow transverse basal depression, leaving a straight
transverse ridge joining weak lateral tubercles, so
that there is no median furrow and no apical lamella
(Rigs: 2392205222) er omen faethe See choc 3
3 (Subgenus Mendacibombus.) Ridge between labral
tubercles is, at its mid point, narrower than basal
depression (Figs 23 & 220), with few punctures and
shining; wings clouded with brown (moderately
imkuscated) Wan teaser eee eee cea avinoviellus
— Ridge between labral tubercles is, at its mid point,
broader than basal depression, with few or many
32
punctures (Figs 221 & 222); wings nearly clear
(subhyaline) 24555 Sjeeseee ee ere eee 4
Ridge between labral tubercles subsiding towards
middle, where there are often many punctures (Fig.
221); pubescence of thoracic dorsum grey-white
or lemon-yellow, but with a broad black band
between wings (Figs 242-245, 247, 248, 250, 251)
SE Re or ae Me et PR Ree) Sa EC, core himalayanus
Ridge between labral tubercles convex for its entire
length, with few punctures (Fig. 222); pubescence of
thoracic dorsum lemon-yellow, with only a very few
black hairs (Figs 254, 255, 257)......... marussinus
(Subgenus Psithyrus.) Lateral keels of sternum VI
projecting well beyond tergum VI from dorsal
aspect, crests of keels sharply acute and blade-like in
section, distinctly angled about midway along their
length (Figs 207 & 208); pubescence of tergum V
usually mostly orange-red, without yellow or white
hairs, only occasionally entirely black............ 6
Lateral keels of sternum VI not clearly projecting
beyond tergum VI from dorsal aspect, crests of keels
broadly rounded, both in section and along their
entire length (Figs 206, 209-211); pubescence of
tergum V black, yellow or white, but without orange-
red hairs
Crests of lateral keels of sternum VI, beyond project-
ing angle of mid-point, with a strongly concave margin
(Fig. 208); sternum II with transverse ridge sharply-
defined and nearly straight; labral furrow narrow,
about a fifth of total basal breadth of labrum; clypeus
with many large punctures spaced more closely than
their own widths, except in a well-defined narrowly-
unpunctured mid-apical area; pubescence of thoracic
dorsum without black hairs, except for a few hairs
adjacent to tegulae, tergum IT predominantly black
(lips:26326552660) = ease eee branickii
Crests of lateral keels of sternum VI, beyond project-
ing angle of mid-point, almost straight (Fig. 207);
sternum II with transverse ridge rounded and curved
unevenly towards anterior margin in middle; labral
furrow wide, about a third of total basal breadth of
labrum (Fig. 24); most of clypeus with scattered large
punctures spaced more widely than their own widths;
pubescence of thoracic dorsum usually with a poorly-
defined patch of black hairs between wings, tergum
II yellow (Figs 259 & 261)
Lateral keels of sternum VI strongly swollen almost
to their apices, where they are separated by a groove
that is narrower than their own breadths (Figs 209 &
210); basal depression of labrum deep and clearly
delimited by an absence of punctures, apex of
lamella pointed
Lateral keels of sternum VI declining from near
their mid-points so that strongly swollen parts are
separated by more than their own breadths (Figs 206
& 211); basal depression of labrum shallow and with
punctures, apex of lamella broadly rounded
Tergum VI with many large punctures, but shining;
10
1
—
PAULH. WILLIAMS
basal keel of mandible weak or absent; thoracic
dorsum without black pubescence (Fig. 267)
“oC ivArmir aes Vag Aa e Se oe ee cee ferganicus
Tergum VI with only fine punctures, strongly shining;
basal keel of mandible well developed but interrupted
before margin of mandible; pubescence of thoracic
dorsum black with an admixture of yellow hairs as an
anterior band (Fig. 269) ........... morawitzianus
Lateral keels of sternum VI small and converging to
a point well before apex, which is down-curved as a
narrow spinose hook that projects strongly beyond
apex of tergum VI (Fig. 211); basal keel of mandible
present but interrupted before reaching margin;
labral tubercles and lamella weakly developed and
rounded in section; oculo-malar distance nearly
equal to basal breadth of mandible; pubescence of
thoracic dorsum and tergum I predominantly yellow,
at most with a narrow band of black hairs between
wings (Biss 271(& 275) see skorikovi
[From material from Britain.] Lateral keels of sternum
VI strongly swollen and converging to a point only
just before apex of sternum, which is broadly
triangular and not down-curved or particularly
narrowed and does not project beyond apex of
tergum VI (Fig. 206); basal keel of mandible weak or
absent; labral tubercles pointed and lamella strongly
swollen; oculo-malar distance less than two-thirds of
basal breadth of mandible; pubescence of thoracic
dorsum and tergum I black, with only a broad yellow
band anteriorly onthorax............. bohemicus
Apex of mandible broadly rounded, with a basal
tooth, one pre-basal tooth and often an apical tooth
(e.g. Fig. 34); longest of erect hairs near anterior
margin of outer surface of hind basitarsus shorter
than greatest breadth of basitarsus ............. 11
Apex of mandible with six teeth, though these are
subject to wear (Fig. 33); longest of erect hairs near
anterior margin of outer surface of hind basitarsus
just longer than greatest breadth of basitarsus.
(Subgenus Alpigenobombus, oculo-malar distance
about half of basal breadth of mandible; pubescence
long, with a broad black band between wings, terga
III-V always with some reddish hair, remainder grey-
white, yellow or black, Figs 295, 296, 298, 299, 301-
30373052506) 3085309) eee ee eee kashmirensis
Hind basitarsus with dense pubescence of proximal
margin (auricle) continuing onto outer surface of
proximo-posterior projection as just a few sparse
hairs (Fig. 229); oculo-malar distance shorter or
longer than one and a fifth times basal breadth of
mandible, but anterior part of malar area smooth,
at most the larger individuals have only a narrow
diagonal band with a few very small punctures... . 12
Hind basitarsus with dense pubescence of proximal
margin (auricle) continuing onto outer surface of
proximo-posterior projection as a dense long brush
(Figs 227 & 228); oculo-malar distance more than
one and a fifth times longer than basal breadth of
mandible, and for larger individuals at least, anterior
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
12
13
14
15
part of malar area uneven with many small to
moderately-large but widely-spaced punctures almost
RERROOEL NOUR erates eee, Tee exc! aynins aie) ye, sate dan chewte' 3 23
Disto-posterior corner of mid basitarsus usually
broadly rounded, never produced as a spine (Fig.
40); oculo-malar distance shorter or longer than
one and a fifth times basal breadth of mandible . . 13
Disto-posterior corner of mid basitarsus spinose
(Fig. 39), or for small workers, at least acutely
produced; oculo-malar distance at least one and
a fifth times longer than basal breadth of
mandible
Ocello-ocular area with unpunctured and shining
areas large or small, but area anterior to ocelli
unpunctured for less than breadth of an ocellus;
labral lamella narrow, only about a third of basal
breadth of labrum; clypeus with many large punctures
scattered on flattened central area; pubescence of
thoracic dorsum with or without much pale
AME eft: Marts ctl css hacia pcte ete ayes wees wees 14
Ocello-ocular area with unpunctured and shining
areas very large and including most of area anterior
to ocelli for a distance of more than breadth of an
ocellus (Fig. 215); labral lamella broad, more than
half of basal breadth of labrum; clypeus with almost
no large punctures in flattened central area; pube-
scence of thoracic dorsum entirely black. (Subgenus
Orientalibombus, large individuals with very dark
wings; pubescence short and very even, terga I-II
bright yellow, III-V orange-red, Figs 279 &
DOU) Preeiictse cis crt seve dale Dees haemorrhoidalis
Band of punctures along eye margin in oculo-
ocellar area opposite lateral ocellus with few small
punctures, large punctures separated by more than
their own widths, whole band occupying about a
third of distance between lateral ocellus and eye
(Figs 216-219); tergum VI subapically nearly flat,
or at most with a poorly-defined apical swelling
or a narrow median ridge, but without a discrete
subapical rounded boss; wings clear (hyaline), or at
most only weakly clouded with brown (weakly
AUUSCAted) Pe. aah wits was Beak S Saya dea dene oo 18
Band of punctures along eye margin in oculo-
ocellar area opposite lateral ocellus occupying
nearly half of distance between lateral ocellus and
eye, with many small and large punctures all
separated by less than widths of large punctures, or
if punctures are mostly large and more widely
spaced, then tergum VI has a discrete subapical
rounded boss (e.g. Fig. 231, often weak or absent
for small workers); wings clear (hyaline) to nearly
black (strongly infuscated)................... 15
Oculo-malar distance about equal to or distinctly
longer than basal breadth of mandible; tergum VI
with a discrete subapical rounded boss (e.g. Fig.
231, often weak or absent for small workers);
mandible with only a shallow notch (incisura) and
tooth apically (Fig. 34) ...................4.. 24
Oculo-malar distance less than two-thirds of basal
16
17
18
19
20
33
breadth of mandible; tergum VI subapically nearly
flat, without a rounded boss; mandible with a deep
notch (incisura) and tooth apically
Clypeus without large punctures in central area;
antennal segment 4 longer than broad and more
than three-quarters of length of segment 5; pube-
scence of gaster white, cream, yellow, brown or
black, but without orange-red ................ 17
Clypeus with many large punctures throughout;
antennal segment 4 just shorter than broad and less
than three-quarters of length of segment 5 (Fig. 17);
pubescence of gaster with some orange-red. (Sub-
genus Diversobombus, pubescence of thoracic
dorsum black, terga I-II bright yellow, III-V
predominantly orange-red, Figs 275, 276, 278)
6b JW ie AOS OS SE HURCIC Bey ee ORAS trifasciatus
(Subgenus Subterraneobombus.) Central area of
clypeus almost without punctures, at most with only
a few very widely-spaced and fine punctures; pube-
scence of head, lower sides of thorax, legs, all sterna
and terga III-V black (Figs 282-284, 286-289)
ra Seas eas aR aR Rea s wlare at melanurus
Central area of clypeus with many fine punctures,
often separated by a distance of only about twice
their own diameter; pubescence of thorax almost
entirely cream-yellow, except for a black band
between wings, and with many pale hairs on head
around bases of antennae, on coxae and hind
femora, on gastral sterna and as posterior fringes of
terga III-V (Figs 291 & 292) .......... personatus
(Subgenus Pyrobombus.) Pubescence of thoracic
dorsum grey-white or yellow, but not brown, often
with many black hairs, terga [V-V orange-red . . .19
Pubescence of thoracic dorsum uniformly yellow-
brown, only occasionally faded to grey, terga IV-V
white (Figs 311, 312, 314,315) ......... hypnorum
Vertex, postero-laterally to ocello-ocular area,
shining, with few fine punctures between large
punctures (Figs 216 & 217), punctures becoming
more numerous and more closely spaced towards
posterior margin of vertex (occiput); apex of tergum
VI rounded, subacute (Fig. 214) .............. 20
Vertex, postero-laterally to ocello-ocular area, with
many close, fine punctures between large punctures
(Figs 218 & 219), punctures uniformly spaced
and numerous towards posterior margin of vertex
(occiput); apex of tergum VI broadly truncate (Figs
212 & 213)
Labral lamella narrow, less than one-third of breadth
of labrum; hind basitarsus with posterior margin
nearly straight in distal half (Fig. 42), stout bristles
on outer surface arising from sockets that are
scarcely raised from surface; pubescence long,
longest hairs of anterior margin of hind tibia much
longer than its distal breadth, longest hairs of face at
least three-quarters as long as segment 1 (scape) of
antenna; pale pubescence of thoracic dorsum either
yellow, grey-white or absent, but if it is yellow then
34
21
22
23
24
there are no black hairs intermixed (Figs 335, 336,
338, 339, 341, 342, 344-346) ............... biroi
Labral lamella broad, more than one-third of
breadth of labrum; hind basitarsus with posterior
margin usually distinctly concave just proximal to
disto-posterior corner (Fig. 41), stout bristles on
outer surface arising from sockets that are strongly
raised from surface on their proximal sides; pube-
scence short, longest hairs of anterior margin of
hind tibia only as long as its distal breadth, longest
hairs of face only two-thirds as long as segment 1
(scape) of antenna; pale pubescence of thoracic
dorsum yellow, with black hairs between wing bases
(Figs 317, 319, 320, 322,323) ......... subtypicus
Thoracic scutum mid-dorsally with an area as large
as tegula around posterior end of longitudinal
median groove smooth with few or no punctures
(Fig. 224); oculo-malar distance shorter than basal
breadth of mandible; hairs of ventral parts of thorax
and gaster predominantly grey-white, top and front
of head often with many pale hairs intermixed
Ppa Re Nan PEERS Cate eat AONE eve SHOES ee nN lepidus
Thoracic scutum mid-dorsally with punctures and
sculpturing almost throughout area around posterior
end of longitudinal median groove (Fig. 223); oculo-
malar distance equal to or just longer than basal
breadth of mandible (Fig. 15); hairs of ventral parts
of thorax and gaster predominantly black, top and
front of head without pale hairs ....... Jemniscatus
(Subgenus Bombus.) Pubescence of thoracic dorsum
grey-white, with a more or less well-developed black
band between wings, tergum V red (Figs 347, 348,
350 435143537354) s eect cheese ae tunicatus
Pubescence of thoracic dorsum black, with a broad,
lemon-yellow or cream anterior band, sometimes
with a narrow posterior band, and tergum V nearly
white (Figs 356, 357, 359-361, 363, 364, 366, 368,
BOO) oF tee cate ead swage cetera weet veneene ee Jucorum
(Subgenus Sibiricobombus.) Oculo-malar area with
many widely-spaced moderate to large punctures;
antennal segment 4 less than half of length of
segment 3 (Fig. 19); tergum VI coarsely sculptured
but without amedian groove ........... asiaticus
Oculo-malar area with many widely-spaced small
punctures almost throughout, surface uneven;
antennal segment 4 more than half of length of
segment 3 (Fig. 20); tergum VI coarsely sculptured,
with a well-defined long narrow median groove
(FIge230) isos eee a ee eee oberti
(Subgenus Melanobombus.) Ocello-ocular areas
laterally shining between few large punctures along
eye margin opposite ocelli, these punctures separated
by distances greater than their own widths; oculo-
malar distance longer than, or about same length
as, basal breadth of mandible; long hairs of head
interspersed with a dense cover of short branched
hairs, which are especially evident between ocelli
and occiput; pubescence of tergum V usually with
some white hairs, or if these are absent, then wings
are nearly black (strongly infuscated) .......... 25
25
26
27
28
PAULH. WILLIAMS
Ocello-ocular areas laterally matt, with a dense
band of large punctures along eye margin opposite
ocelli, many of these punctures separated by
distances less than their own widths; oculo-malar
distance just shorter than basal breadth of mandible;
only a few short branched hairs interspersed with long
hairs on head; wings nearly clear (subhyaline) and
pubescence of tergum V usually uniformly red, or if
white hairs are present then there are very few black
hairs intermixed with pale pubescence on anterior
Pant ofthoracte COTSunN yey sn tebe eee 27
(rufofasciatus-group.) Outer (corbicular) surface of
hind tibia with only a few short branched hairs,
which are confined to margins near disto-posterior
corner; pubescence of thoracic dorsum yellow or
grey-white, but with a black band between wings;
tergum V usually with some white hairs, at least
apically; wings nearly clear or lightly clouded with
brown (subhyaline to weakly infuscated) ....... 26
Outer (corbicular) surface of hind tibia with many
short to moderately long, branched hairs, at least in
proximal half (few or absent for small workers and
subject to loss for older females, Fig. 226); pube-
scence of thorax either entirely black or almost
entirely cream-white dorsally and without a black
band between wings, tergum V red without any
white hairs (Figs 417, 418, 420); wings nearly black
(strongly infuscated)................. simillimus
Boss on tergum VI nearly circular and evenly
convex, only narrowly pointed adjacent to apex of
tergum; pubescence of head entirely black, thoracic
dorsum and tergum I with white pubescence (Figs
431,432, 434, 435, 437, 438)........ rufofasciatus
Boss on tergum VI nearly triangular, for some
queens with a weak median groove, for workers it
may be weakly ridged; short hairs of head yellow,
thoracic dorsum and tergum I with yellow pube-
scence (Figs 422, 423, 425, 426, 428, 429)
wick Boatingnend. svahniisie, Geneon eee a eR ee pyrosoma
(lapidarius-group.) Ocello-ocular area with a
continuous broad band of fine punctures along eye
margin; pubescence of thoracic dorsum uniformly
lemon-yellow, without black hairs (Figs 397, 398,
AN. AQ) ie a2 cea ue Sine ans ange era semenovianus
Ocello-ocular area with band of fine punctures
along eye margin narrowly interrupted opposite
ocelli, so that there is a small shining area with only
large punctures; pubescence of thoracic dorsum
pale yellow, cream, or grey-white, but with a broad
black band between wings ..................- 28
Apex of tergum VI usually broadly, but shallowly,
notched; pubescence of tergum III predominantly
black, often with a posterior fringe of pale hairs,
and of tergum V predominantly red (Figs 403-409,
ALI—AN5)). casei hey eRe tak eee eee keriensis
Apex of tergum VI nearly straight; pubescence of
tergum III predominantly orange-red and of
tergum V white, although these hairs have very
dark bases (Figs 394 & 396) .......... Jadakhensis
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Key to species (males)
1
Volsella always strongly sclerotised, inner corner
usually much closer to apex than to base (to form an
interio-apical corner), often bearing two inwardly-
directed hooks or a single combined inwardly-
directed process (Figs 136-165); gonostylus usually
with a distinct interio-basal process or shelf, but not
associated with long hairs (Figs 176-205); head of
penis valve much modified, often curved strongly in
towards mid-line of body, as a sickle-shaped hook
(Figs 61-71, 76-85), otherwise curved outwards for
some individuals (Figs 72-75), but if it is nearly
straight from dorsal aspect (Figs 56-60), then volsella
has pronounced inner hooks (Figs 136-140)...... 10
Volsella weakly or strongly sclerotised, inner corner
located near mid-point of its length, without any
inwardly-directed hooks, so that volsella is usually
nearly triangular in the distal section (Figs 126-135);
gonostylus without an interio-basal process (Figs
166-169), or if present (Figs 170-175), then it is
associated with many long branched hairs; head of
penis valve nearly straight from dorsal aspect (Figs
AGS =SIN) Ge Soa OR ORC EC OE CREE Cee ee een 2
Volsella weakly sclerotised, yellowish in colour;
gonostylus with a pronounced interio-basal process
(Figs 170-175), associated with many long branched
hairs; head of penis valve, as defined by an outer
lateral ridge, less than a quarter of total length, not
strongly curved but shaped like a slender arrowhead
from lateral aspect (Figs 90-95) ................. 5
Volsella strongly sclerotised, dark brown in colour;
gonostylus without an interio-basal process or hairs
(Figs 166-169); head of penis valve, as defined by an
outer lateral toothed ridge, accounting for nearly
half of total length, ventrally curved and sabre-
shaped from lateral aspect (Figs 86-89) .......... 3
(Subgenus Mendacibombus.) Penis valve strongly
laterally flattened (Figs 46, 47, 86, 87); gonostylus
dorso-ventrally flattened (Figs 166 & 167); mandible
with basal tooth strongly developed (Fig. 35); wings
clouded with brown (moderately infuscated)
MRA Set st seicke Polar hive a) niet tN anwar avinoviellus
Penis valve nearly tubular (Figs 48, 49, 88, 89);
gonostylus stout and not strongly flattened (Figs 168
& 169); mandible without a distinct basal tooth;
wings nearly clear (weakly infuscated or subhyaline)
Volsella with interio-apical margin weakly concave
(Fig. 128); gonostylus from outer dorso-lateral
aspect with a distinct, though broadly-rounded,
exterio-apical corner (Fig. 168); pubescence of
thoracic dorsum anteriorly and of terga I-II grey-
white and/or lemon-yellow, with a broad black
band between wings (Figs 246, 249, 252, 253)
FON eT RTA COO 6 OIC ome oie himalayanus
Volsella with interio-apical margin strongly concave,
so that apex resembles a curved finger (Fig. 129);
gonostylus from outer dorso-lateral aspect with
exterio-apical margin evenly rounded, without any
35
trace of an exterio-apical corner (Fig. 169); pube-
scence of thoracic dorsum lemon-yellow, with few
blackihairs (Rig256) 79" saueteisees 20: marussinus
(Subgenus Psithyrus.) Volsella in its distal section,
beyond inner corner, greatly narrowed, almost
parallel-sided and finger-shaped (Fig. 135); gono-
stylus with only afew shorthairs ......... skorikovi
Volsella distally broader, in the form of a broad
nearly triangular plate (Figs 130, 131, 133, 134), or if
distal section is narrowed then inner corner is
strongly produced (Fig. 132); gonostylus usually with
many long hairs around interio-basal process. ..... 6
Ventro-basal angle of penis valve strongly and
broadly produced ventrally and outwardly, so as to
be clearly visible from dorsal aspect (Figs 51-54, 91-
94); inner corner of volsella well defined, strongly
produced for some individuals (Figs 131-134);
antennal segment 3 distinctly shorter than segment 5
(e.g. Fig. 22); pubescence of scutellum usually
almost entirely yellow or grey, terga [V-V with red or
yellow, or if terga IV-V are nearly white (Fig. 270)
then tergum II has a narrow apical band of pale
LA ETT RSE Tea Ro ae CR rar Ia ire an 7
Ventro-basal angle of penis valve much reduced and
not visible from dorsal aspect (Figs 50 & 90); inner
corner of volsella weak, almost unmarked for some
individuals (Fig. 130); antennal segment 3 just longer
than segment 5 (Fig. 21); pubescence of scutellum
black, with only a narrow band of yellow hairs, apical
margin of tergum II without a band of pale hair, terga
IV-V nearly entirely white (Fig. 258) ... bohemicus
Inner margin of distal section of volsella irregular
but not predominantly concave (Figs 133 & 134);
antennae very long, reaching back nearly to gaster;
pubescence of terga V-VII black, with lateral patches
OMGOMO WTO ECS sa a apace facaie ap cvaday ara alten cut secant 8
Inner margin of distal section of volsella predom-
inantly concave, with exception of a weak subapical
process (Figs 131 & 132); antennae short, reaching
back only to tegulae; pubescence of terga V-VII
OLAN PETER .Gracae eee a ee ae eR aie 9
Distal section of volsella as long as about one and a
half times its maximum breadth from ventral aspect
(Fig. 133); gonostylus broadly triangular (Fig. 173);
pubescence of thoracic dorsum predominantly
brownish-yellow, with only a few black hairs above
tepulae(bicn268) een ae cctcean esate oF ferganicus
Distal section of volsella only just longer than its
maximum breadth from ventral aspect (Fig. 134);
gonostylus reduced to a narrow transverse band at
apex of gonocoxite, but retaining a well-developed
interio-basal process (Fig. 174); pubescence of thoracic
dorsum light yellow with a broad black band between
Wines (HIG: 270). sects suri cee morawitzianus
Distal section of volsella almost twice as long as its
maximum breadth from ventral aspect (Fig. 132);
pubescence of thoracic dorsum yellow, with black hairs
intermixed between wings (Fig. 264)...... branickii
36
11
12
Distal section of volsella about as long as its maximum
breadth from ventral aspect (Fig. 131); pubescence
of thoracic dorsum grey-white, with a poorly-defined
black band between wings (Figs 260 & 262) . .novus
Penis valve from dorsal aspect turned inwards
before apex and dorso- ventrally flattened in the
form of a sickle (Figs 61-71, 76-85) ............ 11
Penis valve from dorsal aspect apically nearly
straight, turned outwards, or at least not strongly
inwardly directed and flattened as a sickle-shape
(Figs:$6=6057/2=75))... J aoatSs sonmavewe tings he 14
Ventro-basal angle of penis valve much reduced,
shown only as a weak curve, or absent (Figs 116-
125); spatha, beyond basal fusion with penis valves,
more than three times longer than its breadth
midway along this length, sides nearly parallel in
proximal al figs iced cece cyencveusts w ae cseuevssts ae aie 12
Ventro-basal angle of penis valve produced as
a basally-directed hook (Figs 101-111); spatha,
beyond basal fusion with penis valves, less than
three times longer than its breadth midway along
this length, sides weakly diverging in proximal half
(B. kashmirensis), or distinctly converging towards
apex throughout theirlength ................. 13
Gonostylus just longer than its greatest breadth,
excluding interio-basal process, which is pronounced
as a broad shelf, and broadening apically (Figs 196—
199); shaft of penis valve more than three-quarters
as broad from lateral aspect as greatest breadth of
gonostylus, excluding its interio-basal process (Figs
116-119); head of penis valve with a broad outer
ridge (Figs 76-79, 116-119) .................. 23
Gonostylus shorter than its greatest breadth,
excluding interio- basal process, which is reduced to
a small rounded projection or a right-angled corner,
and narrowing apically (Figs 200-205); shaft of
penis valve usually strongly narrowed from lateral
aspect, its breadth less than two-thirds as broad as
greatest breadth of gonostylus, excluding interio-
basal process (Figs 120-125); head of penis valve
with only a narrow outer ridge (Figs 80-85, 120—
125) Deze ony Serra eto evecesisl ane ota eae 24
Apex of mandible acutely pointed, with a basal
tooth and a pre-basal tooth (Fig. 37); gonostylus
with a pronounced interio-basal process (Figs 181—
183); volsella, from dorsal aspect, projecting beyond
gonostylus apically by about its own apical breadth.
(Subgenus Alpigenobombus, pubescence with a
broad black band between wings, head predom-
inantly black, terga III-VI with some reddish
hair, remainder variously banded with grey-white,
yellow or black, Figs 297, 300, 304, 307, 310)
Weal phate GREET rad open b-t aoe nett a Bre kashmirensis
Apex of mandible more or less pointed, otherwise
with only a basal tooth; gonostylus without a
strongly projecting interio-basal process (Figs 184—
191); volsella, from dorsal aspect, projecting beyond
gonostylus apically by only about half of its own
apical breadth {i455 4c ei 18
14
15
17
PAULH. WILLIAMS
Penis valves strongly broadened in dorso-ventral
plane so as to form two halves of a tube, ends
splayed outwards as a broad funnel (Figs 72-75,
112-115); antennae short, reaching back only as far
aSitepulae ey nae elt ee ere oe oct eee eee 22
Penis valves dorso-ventrally narrow, at least in
apical third, which is further narrowed and more or
less ventrally-directed (Figs 56-60, 96-100); anten-
nae long, reaching back beyond tegulae ........ 15
Volsella apically not narrowed, with apical corner
forming a right angle, not produced, a projection
from inner corner terminating in a single small
serrated process, which is directed apically, without
any trace of a separate recurved basally-directed
hook (Figs 138-140); ventro-basal angle of penis-
valve strongly produced ventro-laterally, forming a
flattened paddle-shape, transverse to main axis of
penis valve (Figs 98-100); pubescence of thoracic
dorsum cream, yellow or brown, some individuals
with black bands between wings, terga III-VII
without oranpe-Ted =<). 6s ec. ity eee 17
Volsella apically much narrowed, or with apical
corner narrowly produced, inner corner terminating
in a pair of serrated hooks, one directed apically,
the other, which may be very much reduced for
some individuals, directed basally (Figs 136 & 137);
ventro-basal angle of penis valve produced ventrally,
but hardly laterally, as a single tooth (Figs 96 & 97);
pubescence of thoracic dorsum black, terga ITI-VII
predominantly orange-red ................... 16
Gonostylus with interio-basal process strongly
produced, with both an apically-directed hook and
a more ventrally-directed hook (Fig. 176); volsella
with inner hooks very close to apical corner, the
large apically-directed hook nearly straight and
spinose, the basally-directed hook strongly recurved
and broad with many teeth, both hooks projecting
beyond gonostylus from dorsal aspect, part of
volsella immediately proximal to inner hooks much
narrowed (Fig. 136); head of penis valve twice as
broad as shaft, from dorsal aspect, with pronounced
teeth along outer lateral ridge (Fig. 56) (Subgenus
DiversODOMmbDUS) ase eee eee trifasciatus
Gonostylus without an interio-basal process (Fig.
177); volsella with apical corner narrowly produced
distal to inner hooks and curled inwardly, inner
hooks close to inner corner and obscured from
dorsal aspect by gonostylus, part of volsella
immediately proximal to inner hooks broad (Fig.
137); head of penis valve scarcely broader than
shaft, from dorsal aspect, without distinct teeth along
lateral ridge (Fig. 57) (Subgenus Orientalibombus)
sue aes lay ale. Sa Ges et ae vee haemorrhoidalis
(Subgenus Subterraneobombus.) Curved head of
penis valve, from dorsal aspect, longer than broad
(Figs 58 & 59), paddle-like ventro-basal angle of
penis valve clearly tridentate, with a spinosely-
produced dorso-lateral tooth, which is longer than
tooth that some individuals have on exterio-basal
corner of penis valve head (Figs 98 & 99); tip of
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
19
20
aN
interio-basal process of gonostylus not projecting
apically as far as apex of gonostylus (Figs 178 &
179); pubescence of head around antennal bases,
of ventral parts of thorax and of terga III-VII,
predominantly black (Figs 285 & 290) .. melanurus
Curved head of penis valve, from dorsal aspect, as
broad as long (Fig. 60), paddle-like ventro-basal
angle of penis valve almost rounded, with only a
weak dorso-lateral tooth, which is shorter than
spinose tooth of exterio-basal corner of penis valve
head (Fig. 100); tip of interio-basal process of
gonostylus projecting apically as far as apex of
gonostylus (Fig. 180); pubescence of head around
antennal bases, of ventral parts of thorax and of
posterior fringes of terga III-VII predominantly
PASI SS 295 GC D94) rate w asasine eine aie personatus
(Subgenus Pyrobombus.) Inner margin of gono-
stylus weakly convex basally, longitudinal sub-
marginal groove arising close to apex of gonocoxite,
separated from it by a distance usually only about
equal to width of groove (Figs 185-191); pubescence
of terga V-VII orange-red ................... 19
Inner margin of gonostylus with a deep concavity or
notch basally, separating longitudinal submarginal
groove from apex of gonocoxite by a distance nearly
twice width of groove (Fig. 184); pubescence of
terga V-VII white (Figs 313 & 316) ..... hypnorum
Antennae short, reaching back only as far as
tegulae; head of penis valve with recurved hook not
distinctly narrower than adjacent penis valve shaft,
from dorsal aspect, and apically broadly rounded
(Figs 67-71); pubescence long, longest hairs arising
from outer surface of proximal part of mid basitarsus
nearly one and a half times breadth of mid basitarsus
Antennae moderately long, reaching back to scutel-
lum; head of penis valve with recurved hook
distinctly narrower than adjacent penis valve shaft,
from dorsal aspect, and apically acute (Figs 65 &
66); pubescence short, longest hairs arising from
outer surface of proximal part of mid basitarsus
shorter than breadth of mid basitarsus . subtypicus
Penis valve head with recurved hook no greater in
breadth than adjacent penis valve shaft, from dorsal
aspect (Figs 67-69), and ventro-basal angle of penis
valve close to mid-point between penis valve head
and its base (Figs 107-109); pale pubescence with
grey-white hair, or if this is lacking (i.e. pubescence
predominantly yellow), then tergum III is yellow or
orange-red with very few black hairs........... 21
Penis valve head with recurved hook more than one
and a half times breadth of adjacent penis valve
shaft, from dorsal aspect (Figs 70 & 71), and ventro-
basal angle of penis valve at less than a third of
length of penis valve shaft from its base to its head
(Figs 110 & 111); pale pubescence without grey-
white hair, tergum III predominantly black (Figs
331; SAN N343) ise Someries aes bale eae ea wee biroi
Pubescence of thorax and tergum I predominantly
22
23
24
25
37
yellow, without grey-white hairs (Figs 329 & 332);
volsella with interio-apical process narrow and
separated from apex by a distance nearly equal to its
own breadth (Figs 148 & 149)............ lepidus
[From material from Uttar Pradesh, India.]
Pubescence of thoracic dorsum and tergum I grey-
white, without yellow hairs, but with a broad black
band between wing bases; volsella with interio-
apical process arising directly from apex (Fig.
LATIONS. A Ss See tase oe adeno oon s lemniscatus
(Subgenus Bombus.) Pubescence of thoracic dorsum
grey-white, often with a more or less well-developed
black band between wings, terga V-VI red (Figs
FAQESS ZA SIME ae tee crite e aes see es ees tunicatus
Pubescence of thoracic dorsum with lemon-yellow
anteriorly, remainder with variable degrees of
replacement of black by yellow, terga V-VI nearly
white (Figs 358, 362, 365, 367,370) ...... Jucorum
(Subgenus Sibiricobombus.) Eyes greatly enlarged,
so that separation from lateral ocelli is by less than
an ocellar diameter; antennae very long, reaching
almost to gaster; volsella narrowed apically, interio-
apical process narrow and elongate but exterio-apical
corner scarcely acute (Figs 156-158); recurved
hook of penis valve head broad, tapering to tip (Figs
16-78); pubescence/short .............- asiaticus
[From material from the Pamir, U.S.S.R.] Eyes
similar in relative size to those of females, not
enlarged, and separated from lateral ocelli by more
than two ocellar diameters; antennae long, but
reaching only just past tegulae; volsella broadened
subapically, interio-apical process very short and
broad but exterio-apical corner strongly produced
as a broad tooth (Fig. 159); recurved hook of penis
valve head narrowly constricted (Fig. 79); pube-
SCENCEION De re tenets Fates akc aa tas oberti
(Subgenus Melanobombus.) Gonostylus with apical
margin concave, leaving an exterio-apical corner
and an interio-apical process, latter often with two
apical corners or bidentate (Figs 203-205); volsella
extending beyond gonostylus apically by at least
nearly twice its own breadth at the point where
it emerges from beneath gonostylus from dorsal
EY 0c C ROE MOLI Oe On oe CIE Ce oy ere Te 25
Gonostylus with apical margin broadly convex (Figs
200-202); volsella extending beyond gonostylus by
about its own breadth at the point where it emerges
from beneath gonostylus from dorsal aspect ....27
(rufofasciatus-group.) Gonostylus about a third as
long in middle as broad, excluding interio-basal
process, and reduced to a transverse band, with
interio-apical corner extended as a spinose or
bilobed process (Figs 204 & 205); pubescence of
tergum II extensively lemon-yellow ........... 26
Gonostylus more than half as long in middle
as broad, excluding exterio-basal process, nearly
rectangular, with interio-apical corner not strongly
produced (Fig. 203); pubescence of tergum II either
38
nearly white or chocolate brown, but not yellow
(Figsl9 G42) (se ees Apes ene ee simillimus
26 Eyes distinctly enlarged relative to those of females,
separated from lateral ocelli by one ocellar diameter;
interio-basal process of gonostylus with apex forming
nearly a right angle, interio-apical process with
sides diverging towards apex, which is broadly
bilobed (Fig. 205); pale pubescence of thorax
and tergum I grey-white (Figs 433, 436, 439)
PSOE: o Stic D Maas ema UE wate rufofasciatus
— Eyes not enlarged relative to those of females,
separated from lateral ocelli by two ocellar
diameters; interio-basal process of gonostylus with
apex distinctly acute, interio-apical process with
sides often converging towards apex (Fig. 204),
which may be spinose even from outer lateral
aspect; pale pubescence of thorax and tergum I
yellow (Figs 424, 427, 430) ............ pyrosoma
27 (lapidarius-group.) Eyes slightly enlarged relative
to those of females, separated from lateral ocelli by
less than two ocellar diameters; pubescence of
thoracic dorsum uniformly lemon-yellow, without
black hair (Figs 399 & 402) ......... semenovianus
— Eyes not enlarged relative to those of females,
separated from lateral ocelli by about two and a half
ocellar diameters; pubescence of thoracic dorsum
pale yellow with a black band between wings . . . .28
28 Free, recurved hook of penis valve head about two-
thirds as long as more apical part of penis valve head
(Fig. 82), penis valve shaft narrower at its mid-point
from lateral aspect (Fig. 122) than spatha is at
its mid-point from dorsal aspect; pubescence of
tergum III black, at most with only a narrow apical
and lateral fringe of pale hairs, terga ITV-VII orange-
red (Figs 410 & 416)................... keriensis
— Free, recurved hook of penis valve head about one-
third as long as more apical part of penis valve head
(Fig. 80), penis valve shaft broader at its mid-point
from lateral aspect (Fig. 120) than spatha is at its
mid-point from dorsal aspect; pubescence of terga
III-V predominantly orange-red, terga VI-VII
predominantly pale, hairs nearly white with orange
bases, black hairs mostly confined to basal margin of
tergum III, otherwise intermixed on lateral quarters
of remaining terga (Fig. 395) ......... ladakhensis
Genus BOMBUS Latreille
[Bremus [Jurine], 1801: 164. Type species: Apis
terrestris Linnaeus, 1758: 578 = Bombus terrestris
(Linnaeus), by subsequent designation of Morice
& Durrant (1915: 429). Suppressed by the
ICZN in Opinion 135 (ICZN, 1939).]
[Bremus Panzer, [1801]: pls. 19-21. Type species:
Apis agrorum Fabricius, 1787: 301 = Bombus
pascuorum (Scopoli), by subsequent designa-
tion of Sandhouse (1943: 532). Suppressed by
the ICZN in Opinion 220 (ICZN, 1954).]
PAULH. WILLIAMS
Bombus Latreille, 1802a: 437 [redescribed 1802b:
385]. Type species: Apis terrestris Linnaeus,
1758: 578 [cited as Apis terrestris F.| = Bombus
terrestris (Linnaeus), by monotypy.
Specialists often find it useful to be able to refer to
groups of closely related species by group-names.
The use of the established subgeneric names as
labels for assemblages of similar bumble bee
species is continued here solely for the sake of
nomenclatural continuity, because there is little
advantage at present in modifying the usage of
previous authors (see the introduction on genera
of Bombini). When it is possible to revise all of the
bumble bees thoroughly, this system could be
replaced or at least simplified. For this review,
only a few additions and revisions to a preliminary
cladistic study of relationships (Williams, 1985)
are included in the discussions of affinities for
each species. As little as possible is added to the
burden of subgeneric nomenclature, although the
application of certain names does require some
clarification.
Richards (1968) described the characteristics of
both sexes for species of the subgenera referred to
here, with the exception of Psithyrus, which is
described by Popov (1931) and by Lgken (1984).
Subgenus MENDACIBOMBUS Skorikov
Mendacibombus Skorikov, 1914a: 125. Type
species: Bombus mendax Gerstaecker, 1869:
323, by subsequent designation of Sandhouse
(1943: 572).
Mendacibombus Skorikov; Kriiger, 1917: 62 (asa
subgenus of Bombus Latrielle).
Species of the subgenus Mendacibombus have
been regarded as showing the most plesiomor-
phic forms of the male genitalia among bumble
bees (Williams, 1985). A brief review of most
species of this group is now possible thanks to
generous loans of material from other institutions
(ITZ, MNHU, ZI). These nominal taxa have
been described from female type specimens,
which are all closely similar in morphology (e.g.
mostly described as subspecies of B. mendax by
Skorikov, 1910a). The application of the names to
the more divergent males naturally depends on
correct association of the sexes. For these associa-
tions I have relied on reference material from A.
S. Skorikov’s collection (ZI, with a few specimens
in the BMNH), because he had access to relatively
large samples of material and had described many
of the taxa himself.
The results of a cladistic analysis show that
the subgenus Mendacibombus is likely to be
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
paraphyletic with respect to all other bumble bees
(Fig. 5, all other bumble bees represented by B.
exil, see the comments on B. avinoviellus, B.
himalayanus and B. bohemicus). This is a change
in the interpretation of relationships within this
species-group (see the introduction on generic
nomenclature) that results from a change in the
interpretation of the broadened state of the inner
lateral shelf of the penis valve. This character is at
best weakly developed in species of the out-group,
Euglossini, which have the shaft of the penis valve
much shorter. If the strong development of the
shelf is regarded as plesiomorphic rather than
apomorphic within the subgenus Mendacibombus,
then its compatibility with the additional character
state patterns in this analysis is maintained.
Holophyletic groups within the paraphyletic sub-
genus Mendacibombus are largely single species
(monotypic) or unresolved. For a general descrip-
tion of both sexes of the species of this subgenus
see Richards (1968) and the keys (couplets 1 & 2).
The North American B. (Bombias) nevadensis
Cresson is similar to species of the subgenus
Mendacibombus in the simple, straight form of
the penis valves (see Milliron, 1971: pl. 11) and in
the enlarged compound eyes of the male. How-
ever, at present the weight of evidence does
not support a particularly close relationship by
descent. On the one hand, the shared form of the
penis valves is interpreted as a symplesiomorphy
among the bumble bees and the enlarged eyes of
the males are likely to be a convergent character
state. Alcock & Alcock (1983) related the enlarged
eyes of male B. nevadensis to the particular kind of
mate-searching behaviour that they employ. Among
the Kashmir fauna, this behaviour is apparently
related in turn more to habitat structure than to
relationships by ancestry (see the introduction on
the relationship of male eye-size to mate-searching
behaviour and habitat structure, also see the com-
ments on the species of the rufofasciatus-group: B.
simillimus, B. pyrosoma and B. rufofasciatus). Thus
neither the shape of the penis valve nor the shape of
the eyes can be used as evidence of close relation-
ship. On the other hand, B. nevadensis has a
flattened form of the male gonostylus that has a
distinct interio-basal process. More particularly
this species has a strongly narrowed form of the
volsella that has a narrowly subapical and broad,
many-toothed, interio-apical process. B. nevaden-
sis appears to share these character states
(2synapomorphies) with species such as B.
(Alpinobombus) arcticus (Quenzel) (oldest avail-
able name for B. hyperboreus Schonherr), rather
than with any species of the subgenus Mendaci-
bombus. Consequently this species is not included
here in the subgenus Mendacibombus.
39
The males of the European and central Asian
B. (Confusibombus) confusus Schenck also have
relatively simple, straight heads of the penis
valves and enlarged compound eyes. Because of
this similarity, B. confusus, like B. nevadensis, has
sometimes been grouped with B. mendax (e.g.
Kruseman, 1952). However, both B. confusus
and B. nevadensis share the development of an
inwardly-directed process from the base of the
gonostylus with most bumble bees apart from
species of the subgenus Mendacibombus (synap-
omorphy of Psithyrus + all other subgenera of
Bombus excluding Mendacibombus). Ito (1985)
concluded from his detailed phenetic study that B.
nevadensis is most closely similar to B. confusus,
and then to B. (Kallobombus) soroeensis, followed
by species of the subgenera Rhodobombus or
Alpinobombus. Evidence from biochemical charac-
ters would be particularly useful in these cases.
Within the subgenus Mendacibombus, nests
have been described in detail for only the European
B. mendax (by e.g. Haas, 1976; see also a brief
footnote on a nest of B. makarjini by Reinig in
Bischoff, 1931). These nests appear to show a
greater resemblance in their structure to nests of
some stingless bees (cf. Wille & Michener, 1973)
than do those of other bumble bees. In particular,
the cocoons are almost completely torn down
soon after the emergence of the adults, so that
pollen and honey must be stored elsewhere. In
contrast to other bumble bees, these food reserves
are held exclusively outside the wax envelope of
the brood nest and the honey may be stored in
hexagonally arranged wax cells.
Bombus (Mendacibombus) avinoviellus
(Skorikov)
(Figs 14, 16, 23, 27, 31, 35, 43-47, 86, 87, 126, 127,
166, 167, 220, 225, 232-241, Maps 5 & 6)
Mendacibombus avinoviellus Skorikov, 1914a:
126. Lectotype by designation of Podbolotskaya
(in press) [not seen]. Paralectotype queen,
INDIA: Kashmir, Baltal, 11.vi.1912 (Jacobson)
(ZI) [examined].
Bombus niveatus subsp. callophenax Cockerell,
1917: 122. Holotype queen by monotypy (see
Note 2 below), INDIA: Kashmir, no further
data (Woglum) (USNM) [examined]. Syn. n.
Bombus (Mendacibombus) avinoviellus
(Skorikov); Richards, 1930: 635.
[Bombus (Mendacibombus) avinoviellus var.
subtunicatus Richards, 1930: 635 [examined].
Infrasubspecific (see Note 1 below) (Art. 45g
(ii)(1)), unavailable name (Art. 45e).]
NOMENCLATURE. Note 1. Richards’s use (1930:
40
634 etc.) of the term ‘Subsp.’ elsewhere in
his paper is taken to indicate that infrasub-
specific rank is meant for B. avinoviellus var.
subtunicatus.
TYPE MATERIAL. Note 2. Cockerell’s description of
B. niveatus callophenax gives no reason to believe
that he had based his description on more than
one specimen. Only a single queen in the USNM
collection agrees with the original description and
carries the data quoted (Dr B.B. Norden, in litt.)
and this specimen bears a red label ‘Type’. I
believe that this is the single specimen on which
the original description is based and regard it as
the holotype (Art. 73a(ii)).
AFFINITIES. B. avinoviellus is easily distinguished
from the other species of the subgenus Mendac-
ibombus (and from all other bumble bees) by the
laterally compressed and scythe-like heads of the
penis valves, which have the weakly-sclerotised
lateral channel relatively narrow (Figs 46, 47,
86, 87) (plesiomorphic within the bumble bees,
Bombini, and shared with the South American
orchid bees, Euglossini).
The closest relatives of B. avinoviellus are
probably B. mendax Gerstaecker and B. makarjini
Skorikov (Fig. 5). B. mendax has the head of the
penis valve not laterally compressed as for B.
avinoviellus, but rounded and finger-like, with a
narrowly subapical shoulder on the inner margin
from the dorsal aspect. The gonostylus is expanded
ventrally along its interio-apical margin, so that
it is particularly stout (synapomorphies of B.
mendax with all species of Mendacibombus
excluding B. avinoviellus; the gonostylus may be
secondarily slightly narrowed for B. makarjini and
the species of the defector-group, handlirschianus-
group and waltoni-group, see the comments on B.
himalayanus). B. mendax is known from the
European Alps and the Pyrenees (BMNH, PW).
B. makarjini has the head of the penis valve even
more strongly dorso- ventrally compressed than
for B. mendax and the penis valve head is relatively
shorter. The head of the penis valve is defined by
an outer toothed ridge (e.g. Fig. 86), which for B.
makarjini extends for only half of the distance
from the apex to the ventro-basal angle of the
penis valve. The apex of the volsella is strongly
concave on the inner margin so that the apex is
narrowed and inwardly curved (synapomorphies
of B. makarjini with all species of Mendacibombus
excluding B. avinoviellus and B. mendax). B.
makarjini is known from the region of the Pamir
and the Tien Shan ranges, where it is uncommon
(Reinig, 1930; Skorikov, 1931; Panfilov, 1957;
ITZ, ZI). All the other species of the subgenus
Mendacibombus have the penis valve narrowed
PAULH. WILLIAMS
basally, particularly by the reduction of the inner
lateral shelf (Figs 48 & 49) (synapomorphy; see
the comments on B. himalayanus).
B. niveatus subsp. callophenax was described by
Cockerell from a queen that is closely similar
in morphology and in its banded white colour
pattern to the paralectotype of B. avinoviellus.
There is no reason to believe that Cockerell was
aware of Skorikov’s description of B. avinoviellus.
These two nominal taxa have not previously been
recognised as conspecific because most authors
have followed the opinion of Richards (1930: 652)
that B. niveatus subsp. callophenax belonged to a
taxon of the subgenus Sibiricobombus that has a
similar, banded white colour pattern. This is
a misidentification (see the comments on B.
asiaticus).
A queen from the Karakoram range (Fig. 232),
kindly lent to me by B. Tkalci, resembles B.
marussinus in the unbanded yellow colour pattern
of the pubescence (Fig. 254). However, this queen
is otherwise closely similar in its morphological
characteristics (see the key), as well as in the
stronger clouding of the wings with brown, to the
paralectotype queen of B. avinoviellus (see the
comments on B. marussinus). It is therefore
considered likely to be part of the same species as
B. avinoviellus. More material, especially of the
males, is needed in order to establish its status
definitively.
DISTRIBUTION. B. avinoviellus is a west Himalayan
species (Map 5). It is known only from Uttar
Pradesh (BMNH), Himachal Pradesh (BMNH,
PW) and Kashmir.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. avinoviellus has been recorded from the
southern side of the Great Himalaya range, and
from the Pir Panjal and Karakoram ranges,
primarily in mesic subalpine scrub and in meadows
in the upper montane coniferous forest zone (Map
6, Fig. 6): 29 queens, 69 workers, 80 males, from
12 localities 1900-4600 m (BMNH, BT, IZ, LK,
MNHN, NR, PW, SEMK, USNM, ZI, ZM).
VARIATION WITHIN KASHMIR. The only specimen
of B. avinoviellus examined from the Karakoram
range, a queen from Banidas (Fig. 232), has the
thoracic dorsum, tergum I and the basal margin of
tergum II uniformly lemon-yellow, with only a
few black hairs above the wing bases.
Individuals from around the Kishanganga valley
have the thoracic dorsum dull yellow, with the
exception of a band of black hairs between the
wing bases (Figs 233-235). Individuals from around
the Vale of Kashmir have the pale pubescence of the
thoracic pubescence grey-white rather than yellow
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
(Map 6, Figs 236-241), although there is often
some yellow on terga I-II (Figs 236 & 237). Both
banded yellow individuals and banded white
individuals were taken together at Chittakatha
Sar (banded yellow: 2/4 males; banded white: 2/4
males) and at Lal Pani (banded yellow: 5/6
workers, 28/35 males; banded white: 1/6 workers,
7/35 males) in the upper Kishanganga valley, and
at Nigagar (1 banded yellow worker, 1 banded
white male, 2 banded cream males) in the upper
Sind valley.
The individuals with an unbanded yellow
thoracic dorsum (Fig. 232) are particularly closely
similar in colour pattern to those of B. marussinus
(Figs 254 & 255), which may occur with them.
They are also similar to some individuals of B.
branickii (Fig. 265), B. subtypicus (Fig. 319-324),
B. biroi (Figs 335-337), B. asiaticus (Figs 371-
376) and B. semenovianus (Figs 397-402). The
banded yellow individuals from the Kishanganga
valley (Figs 233-235) resemble the local, banded
yellow B. subtypicus (Figs 317 & 318), B. asiaticus
(Figs 377, 378, 382) and B. keriensis (Fig. 414—
416). The banded white individuals from around
the Vale of Kashmir (Figs 239-241) are closely
similar in colour pattern to the B. asiaticus that
occur with them (Figs 386, 387, 389-391), but also
show some similarity to some banded white
individuals of B. kashmirensis (Figs 306-310), B.
biroi (Fig. 342), B. tunicatus (Figs 350, 353, 355)
and B. keriensis (Figs 408 & 409, see also the
comments on B. subtypicus). B. avinoviellus can
be recognised by the presence of pale pubescence
on the face, by the clouded wings, by the sub-
generic and specific characters (see the key,
couplets 1—3) and by its small size.
FOOD PLANTS. (Ranunculaceae) Aconitum heter-
ophyllum Wallich ex Royle; (Balsaminaceae)
Impatiens glandulifera Royle; (Hippocastanaceae)
Aesculus indica (Colebr. ex Cambress.) Hook.;
(Leguminosae) Trifolium repens L.; (Compositae)
Cirsium falconeri (Hook.f.) Petrak, C. wallichii
DC., unidentified dandelion-like composites;
(Oleaceae) Syringa emodi Wallich ex Royle;
(Scropulariaceae) Scrophularia pauciflora Benth.,
Digitalis lanata Ehrh. [introduced]; (Labiateae)
Phlomis bracteosa Royle ex Benth., Stachys sericea
Wallich ex Benth., Prunella vulgaris L.
Bombus (Mendacibombus) himalayanus
(Skorikov)
(Figs 48, 88, 128, 168, 221, 242-253, Maps 7 & 8)
Mendacibombus varius Skorikov, 1914a: 125.
Lectotype by designation of Podbolotskaya (in
press) [not seen]. Paralectotype queen, INDIA:
41
Kashmir, Zoji La, above 3000 m, 12—15.vi.1912
(Jacobson) (Z1) [examined]. Junior secondary
homonym in Bombus of Psithyrus varius
Lepeletier, 1832 [= B. campestris (Panzer)].
Mendacibombus mendax subsp. himalayanus
Skorikov, 1914a: 127. Holotype queen by
monotypy (see Note below), INDIA: Kashmir,
Ladakh, Khardung La, 4500 m, 3.vii.1912
(Jacobson) (ZI) [examined]. Provisional
synonym.
Bombus_ (Mendacibombus) mendax subsp.
himalayanus (Skorikov); Richards, 1930: 635.
TYPE MATERIAL. Note. Skorikov specifies that he
made his original description of Mendacibombus
mendax subsp. himalayanus from a single female
specimen. Only a single queen in the ZI collection
agrees with the original description and carries the
data quoted. I believe that this is the single
specimen on which the original description is
based and regard it as the holotype (Art. 73a(ii)).
AFFINITIES. In addition to B. avinoviellus, B.
mendax and B. makarjini (see the comments on B.
avinoviellus), there are at least eight species of the
subgenus Mendacibombus that are known from
both sexes. These species share forms of the penis
valve that are narrowed at the base, particularly
by the reduction of the inner lateral shelf (Figs 48
& 49) (synapomorphy). This character state is also
shared with all the other bumble bees.
Three putative species, B. himalayanus, B.
marussinus and B. turkestanicus Skorikov, are
found in the mountain system of the western
Himalaya, the Hindu Kush, the Pamir and the
Tien Shan ranges (see below). Their relationships
are not fully resolved (Fig. 5). No synapomorphies
are known from which these bees can be recognised
as a true, monophyletic group, yet they are
morphologically closely similar and they lack the
synapomorphies of the remaining species of the
subgenus Mendacibombus. Among these other
species may be recognised the defector-group, a
possible handlirschianus-group and the waltoni-
group. The latter three groups share with all other
bumble bees (see the comments on B. bohemicus)
a further narrowing and straightening of the penis
valve base, with some development of a small
semi-circular projection from the inner basal part
of the penis valve by a more pronounced reduc-
tion of the inner lateral shelf, together with a
reduction of the outer lateral shelf (synapomor-
phies, least well developed for males of the
handlirschianus-group, also only weakly evident
for some species of other subgenera of Bombus).
The defector-group includes at least B. defector
Skorikov, for which the penis valve is elongated
beyond the volsella and the volsella is particularly
42
broad basally, so that its inner angle is only very
broadly rounded (?autapomorphies). B. defector
may be conspecific with similar females described
by Skorikov under the name B. mendax subsp.
margreiteri Vogt, which have the corbicular fringes
orange rather than black, and possibly also with
females described by Skorikov under the name B.
mendax subsp. altaicus, which have the pale
pubescence lighter in shade as well as the corbicular
fringes orange. I have not seen any males that can
definitely be associated with either of these two
nominal taxa. Bees of the defector-group are
known from the northern Pamir and Tien Shan
ranges (Skorikov, 1931; Panfilov, 1957; BMNH,
PW, ZI). They may also occur in some of the
mountains that extend north-eastwards as far as
the other localities known for these bees in
Kamchatka (Bischoff, 1930; Skorikov, 1931).
The handlirschianus-group may include two
species, for which the basal angle of the penis
valve is more pronounced and fully ventral rather
than lateral, and the gonostylus is narrowed
apically (synapomorphies of the handlirschianus-
group + waltoni-group + all other subgenera of
Bombus, for which they may be secondarily much
modified). This is not a true group in that no
exclusive synapomorphies are known for the
included species, yet they lack the apomorphic
character states of the waltoni-group and of the
other subgenera of Bombus. Females of B.
handlirschianus Vogt have the pale pubescence
grey-white. Other females, described by Skorikov
under the name B. mendax subsp. shaposhnikovi,
are known from the same area of distribution but
have the pale pubescence yellow. It is possible
that the white females are part of the same species
as those with yellow bands (cf. comments on
colour variation of B. keriensis). However, the
only male of this species- group that I have seen
with the pale pubescence white rather than yellow
(ITZ) also has the whole gonostylus and the apex
of the volsella distinctly broader than it is for the
yellow males (see also Skorikov, 1931: figs 14 &
15). Therefore the yellow and the white bees are
likely to belong to two separate species, B.
shaposhnikovi and B. handlirschianus respectively.
Both of these species of the handlirschianus-group
are known from Turkey, the Caucasus and the
Elburz mountains of Iran (e.g. Reinig, 1971;
BMNH, ITZ).
Species of the waltoni-group share with most
other groups of bumble bees the development of a
dorsal toothed ridge on the volsella (see Skorikov,
1931: fig. 20), which, from the ventral aspect,
projects beyond the inner margin (synapomorphy
of the waltoni-group + all other subgenera of
Bombus, possibly secondarily reduced for B.
PAULH. WILLIAMS
(Eversmannibombus) persicus Radoszkowski and
for species of Psithyrus). The species of the
waltoni-group share a very strongly curled apex
of the volsella (synapomorphy). The male gono-
stylus of B. waltoni Cockerell is uniquely much
broader at the apex than at the base and is dorso-
ventrally more flattened (autapomorphy). B.
waltoni is conspecific with B. mendax subsp.
chinensis Skorikov (junior secondary homonym in
Bombus of Apathus rupestris var. chinensis
Morawitz), and is probably also conspecific with
B. rufitarsus Friese (see Skorikov, 1914a@), and
with B. asellus Friese. B. waltoni is known from
Tibet (Cockerell, 1910; Friese, 1918; Richards,
1930; Wang, 1982; BMNH), Qinghai (Morawitz,
1886; Skorikov, 1910a, 19126; Bischoff, 1936;
Wang, 1982; ZI) and Sichuan (Wang, 1982). A
record for Mongolia (Friese, 1924) needs confirma-
tion. B. convexus Wang (oldest available name for
B. lugubris Morawitz, 1880, syn. n., which
is a junior secondary homonym in Bombus of
Psithyrus lugubris Kriechbaumer, 1870 [= B.
maxillosus Klug]) has the gonostylus narrow and
finger-like (?plesiomorphic within this group), but
the apex of the volsella is strongly elongated
(autapomorphy). B. convexus is known from Tibet
(Wang, 1979, 1982; PW), Qinghai (Wang, 1982),
Sichuan (Morawitz, 1890), Gansu (Morawitz,
1880, 1890; Bischoff, 1936; MNHU, ZI) and
Beijing (Panfilov, 1957). Another species that
may belong to this group was described from a
single female from the mountains of northern
Mongolia (Mendacibombus superbus Tkalci,
1968a). This individual is unusually large for a
species of the subgenus Mendacibombus.
B. himalayanus, B. marussinus and B. turk-
estanicus appear to differ from one another only in
colour pattern (Skorikov, 1910a, 1914a), in subtle
differences in the male genitalia (Figs 48, 49, 168,
169, 128, 129) and in details of female sculpturing
(Figs 220-222). Nonetheless these differences are
apparently maintained in combination at sites
where they are recorded together. For instance,
two queens and three workers from Chhantir Gah
in the Hindu Raj range are assigned in this review
to B. marussinus. If these workers were considered
together with the material from the same locality
that is ascribed to B. himalayanus, all of the
workers could be interpreted as representing
almost a continuum in colour variation between
the extreme individuals with an unbanded yellow
thorax (originally described under the name B.
mendax subsp. marussinus) and those with a
banded white thorax (described under the name
Mendacibombus varius). However, the workers
with a black band on the thoracic dorsum (both
Mendacibombus_ varius and Mendacibombus
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
mendax subsp. himalayanus) do have more
punctures on the labral ridge, which, although
broad, is also particularly weakly raised near its
mid-point (Figs 220-222). This combination of
character states is strictly maintained where the two
taxa occur together at Holojut as well as at Chhantir
Gah. Therefore there is no evidence of hybrids from
interbreeding between B. himalayanus and B.
marussinus, so they are considered to be separate
species (see the comments on B. marussinus).
There is a similar lack of obvious hybrids as
evidence for interbreeding between B. himalaya-
nus and B. turkestanicus, although I have not seen
any samples from single localities that include
both taxa. From the material available, B. himal-
ayanus always retains the particularly weak labral
ridge with many punctures in combination with a
tendency towards black pubescence on the lower
sides of the thorax and on the legs, both of which
distinguish it from B. turkestanicus.
A single male with a yellow thorax with black
hairs between the wing bases from Kilian
(Raskam range, Xinjiang, China) was ascribed by
Skorikov (1914a :127) to Mendacibombus mendax
subsp. himalayanus s.str. Unfortunately this
specimen could not be found in the ZI collection
(M. Podbolotskaya, in litt.). This could be
the same individual from the ‘Raskemkette,
Nordhang des Kilieng’ that Skorikov later listed
under the name Mendacibombus makarjini (see
Skorikov, 1931: 215). I have seen only two queens
of B. himalayanus s.str. from Ladakh. These do
differ from queens of B. makarjini, but resemble
queens of Mendacibombus varius (Fig. 222), in
the more coarsely punctured middle section of the
labral ridge.
A comparison of much more material of B.
himalayanus s.str. with Mendacibombus varius,
B. turkestanicus and B. makarjini, especially for
the males, is still needed. From the meagre
evidence available at present, I consider that B.
himalayanus s.str. is most likely to be part of the
same interbreeding population and species as
Mendacibombus varius, which is likely to be
separate from B. turkestanicus and B. makarjini.
DISTRIBUTION. B. himalayanus is a west Himalayan
species (Map 7). It is known from Himachal
Pradesh (BMNH, PW) and Kashmir.
MATERIAL EXAMINED FROM KASHMIR. B. himalay-
anus is widespread in Kashmir, where it is recorded
from the Hindu Raj, Ladakh, Zanskar, Great
Himalaya, and Pir Panjal ranges, in alpine scrub
and steppe (Map 8, Fig. 6): 15 queens, 70 workers,
12 males, from 13 localities 2700-4800 m (BMNH,
NR, PW, SEMK, ZI, ZM; 1 male photographed
in the Sangisfaid valley, C. A. Chadwell).
43
VARIATION WITHIN KASHMIR. Two queens from the
Ladakh and Zanskar ranges (Map 8) have the pale
pubescence yellow, with a band of black hairs
between the wing bases and black basally on
tergum III. The queen from Nimaling has a
pronounced black band between the wing bases
(Fig. 242), but Skorikov’s queen from the Khardung
La has only a poorly-defined black band on the
thorax (cf. B. marussinus). Five workers from
Chhantir Gah in the Hindu Raj range have the
thoracic band well defined, but the black band on
the gaster is much reduced (Fig. 243). In common
with many workers in this species-group (but not
among queens, except for B. turkestanicus), these
workers have many pale hairs ventrally.
Also from Chhantir Gah, another queen and
eight workers have a similar pattern, although the
yellow of the thorax, and to a variable extent
on tergum II, is replaced by grey-white. These
specimens, and other similar individuals from
elsewhere in the Hindu Raj range (Figs 244, 245,
247, 248), have tergum III almost entirely orange-
red. This banded white and yellow pattern is
predominant among individuals in the samples
from the mountains around the Kishanganga
valley and the Vale of Kashmir (Figs 246 & 249).
However, many of these individuals show some
replacement of the pale band at the rear of the
thorax by black hairs and black is also more
extensive on tergum III, so that it may form a
distinct black band on the gaster (Figs 250-252).
The single male from Sangisfaid valley is even
more extreme, in that all of the orange-red
pubescence of the gaster appears to have been
replaced by black (Fig. 253).
The yellow individuals of this species (Figs 242
& 243) are similar in colour pattern to the B.
kashmirensis (Figs 295-300), B. lepidus (Figs 327
& 328), B. oberti (Figs 392 & 393), B. ladakhensis
(Figs 394 & 396) and B. keriensis (Figs 412 & 413)
that occur with them. The white and yellow
individuals (Figs 244-252) are similar to the B.
novus males (Figs 260 & 262), B. kashmirensis
(Figs 301-304), B. lemniscatus (Figs 325 & 326),
B. lepidus (Figs 330, 331, 333, 334), B. biroi (Figs
338 & 339) and B. rufofasciatus (Figs 431-436,
438, 439) that occur with them, although the yellow
pubescence is on tergum I for B. himalayanus and
on tergum II for the other species. B. himalayanus
is easily distinguished from all other species of the
genus Bombus with similar colour patterns by
using the subgeneric characteristics (see the key,
couplets 1 & 2).
FOOD PLANTS. Kashmir: unidentified pink clover-
like legume; (Compositae) Cirsium falconeri
(Hook.f.) Petrak, unidentified dandelion-like
44
composites; (Gentianaceae) Swertia petiolata D.
Don; (Scrophulariaceae) Picrorhiza kurrooa
Royle ex Benth.; (Labiateae) Thymus linearis
Benth. ex Benth.
Ladakh: (Leguminosae) Caragana versicolor
(Wallich) Benth.
Bombus (Mendacibombus) marussinus
Skorikov
(Figs 49, 89, 129, 169, 222, 254-257, Maps 9 & 10)
Bombus mendax subsp. marussinus Skorikov,
1910a: 330. Lectotype queen by designation
of Podbolotskaya (in press), U.S.S.R.:
Tadzhikistan S.S.R., Shugnan, 3000 m, 24—
27.vi.1909 (Makarjin) (Z1) [examined].
Bombus (Mendacibombus) marussinus subsp.
afghanus Reinig, 1940: 230. Syntype queen and
male, presumed lost (Tkalct, 1969a, see also
Note 4 on B. biroi), AFGHANISTAN:
Badakhshan, Khwaja Muhammed, 3800-4000
m, 25.vii-10.viii.1936 (Kotzsch) [not seen].
Syn. n.
AFFINITIES. B. marussinus is closely similar to B.
himalayanus and B. turkestanicus in morphology
(Fig. 5, see the comments on B. himalayanus).
Female B. marussinus have fewer fine punc-
tures on the labral ridge than do those of B.
himalayanus (Figs 221 & 222) or B. turkestanicus.
This difference appears to be retained in combina-
tion with the unbanded yellow thoracic dorsum
where B. marussinus and B. turkestanicus occur
together in the Tarest mountains of the Hindu
Kush (BMNH) and where B. marussinus and B.
himalayanus occur together in the Hindu Raj
range (see the comments on B. himalayanus). I
infer from this precise coincidence between the
discontinuities in the patterns of variation, for
both colour and sculpturing, that individuals of
these three taxa are unlikely to be interbreeding.
However, it is possible that there are narrow
hybrid zones, so their specific status is still not
certain. More information would be desirable
concerning their mate-searching behaviour,
where they occur together, in order to ascertain
more directly whether or not they interbreed. The
distribution of B. turkestanicus includes the Tien
Shan (Skorikov, 1910a, 1931; Panfilov, 1957; ZI),
the Pamir and the Hindu Kush ranges (BMNH).
B. turkestanicus may occur in Kashmir, but I have
yet to see specimens from this area that can be
attributed definitely to this species (but cf. yellow
workers assigned to B. himalayanus).
According to Tkalcu (1969a), the syntypes of B.
marussinus subsp. afghanus appear to have been
lost. The original description contains nothing to
PAULH. WILLIAMS
distinguish this material from ‘Chodja-Mahomed’
[= Khwaja Muhammed] in the Hindu Kush from
a slight variation of the colour pattern of the
lectotype of B. marussinus. Tkalci ascribed
material with a similar colour pattern from the
‘Nachbargebiet Afghanistans (Karakorum)’ to B.
afghanus Reinig. From his detailed description
and from an examination of a queen from Banidas
kindly lent to me by Dr Tkalci, it is likely that
these specimens from the Karakoram are part of
the same species as the banded white B. avino-
viellus, despite their uniformly yellow thoracic
dorsa (see the comments on B. avinoviellus).
DISTRIBUTION. B. marussinus is a central Asian
species (Map 9). It is known from the Hindu Kush
(Reinig, 1940; Tkalct, 1969a; BMNH), the Pamir
(Skorikov, 1910a; Reinig, 1930; Skorikov, 1931;
ZI), Pakistan (PW) and Kashmir.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. marussinus has been recorded only from the
Hindu Raj and western Karakoram ranges, in the
drier subalpine scrub (Map 10): 5 queens, 7
workers, 2 males, from 5 localities 2700-3200 m
(BMNH).
VARIATION WITHIN KASHMIR. All of the individuals
of B. marussinus have some yellow hairs in the
pubescence of terga I-II, although there is varia-
tion in the extent of this pale pubescence on
tergum II (Figs 254-257). Workers have more
pale pubescence ventrally, as for workers and
queens of B. turkestanicus.
This species is closely similar in colour pattern
to some of the B. avinoviellus (Fig. 232), B.
branickii (Fig. 265), B. subtypicus (Figs 319-324),
B. biroi (Fig. 335-337), B. asiaticus (Figs 371-376)
and B. semenovianus (Figs 397-402) that may
occur with it. It is, however, easily recognised
using the subgeneric characteristics (see the key,
couplets 1 & 2). B. marussinus can be separated
from any B. avinoviellus with a similar colour
pattern (Fig. 232) of the pubescence by its clear
(hyaline) wings and by the specific characters (see
the key, couplet 3).
FOOD PLANTS. No records.
Subgenus PSITHYRUS Lepeletier, stat. n.
Psithyrus Lepeletier, 1832: 373. Type species:
Apis rupestris Fabricius, 1793: 320 = Bombus
rupestris (Fabricius), by subsequent designa-
tion of Sandhouse (1943: 572).
Apathus Newman, 1835: 404 [footnote k]. Replace-
ment name for Psithyrus Lepeletier, incorrectly
stated to be a junior homonym of Psithyros
Hubner (Lepidoptera).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Laboriopsithyrus Frison, 1927: 69 (as a subgenus
of Psithyrus Lepeletier). Type species: Bombus
laboriosus Fabricius, 1804: 352 [?= Emphoropsis
laboriosa (Fabricius)] in the sense of Frison [=
Bombus citrinus (Smith), a misidentification,
see Milliron (1960: 99), requiring designation
by the ICZN (see Note below)] by original
fixation. Provisional synonym.
Ashtonipsithyrus Frison, 1927: 69 (as a subgenus
of Psithyrus Lepeletier). Type species: Apathus
ashtoni Cresson, 1864: 42 = Bombus ashtoni
(Cresson), by original designation [p. 70].
Synonymised with Psithyrus Lepeletier by
Milliron (1961).
Fernaldaepsithyrus Frison, 1927: 70 (as a sub-
genus of Psithyrus Lepeletier). Type species:
Psithyrus fernaldae Franklin, 1911: 164 =
Bombus fernaldae (Franklin), by original
designation. Synonymised with Psithyrus
Lepeletier by Milliron (1961).
Eopsithyrus Popov, 1931: 134 (as a subgenus of
Psithyrus Lepeletier). Type species: Apathus
tibetanus Morawitz, 1886: 202 = Bombus
tibetanus (Morawitz), by original designation
[p. 135]. Synonymised with Psithyrus Lepeletier
by Milliron (1961).
Metapsithyrus Popov, 1931: 135 (as a subgenus of
Psithyrus Lepeletier). Type species: Apis
campestris Panzer, 1801: 74 = Bombus camp-
estris (Panzer), by original designation. Synon-
ymised with Psithyrus Lepeletier by Milliron
(1961).
Allopsithyrus Popov, 1931: 136 (as a subgenus of
Psithyrus Lepeletier). Type species: Apis bar-
butella Kirby, 1802: 343 = Bombus barbutellus
(Kirby), by original designation. Synonymised
with Psithyrus Lepeletier by Milliron (1961).
Ceratopsithyrus Pittioni, 1949: 271 (as a subgenus
of Psithyrus Lepeletier). Type species: Psithyrus
klapperichi Pittioni, 1949: 273 = Bombus
cornutus (Frison) (see Tkalct, 1989), by original
designation. Synonymised with Psithyrus
Lepeletier by Milliron (1961).
Citrinopsithyrus Thorp [in Thorp et al.], 1983: 50
(as a subgenus of Psithyrus Lepeletier). Type
species: Apathus citrinus Smith, 1854: 385 =
Bombus citrinus (Smith), by original designa-
tion. Syn. n.
Note. Application of Laboriopsithyrus is a matter
that should mandatorially be referred to the
ICZN (Art. 70b). Since this name is treated here
as a synonym of Psithyrus, I intend to take no
action.
Descriptions of the diagnostic characters of both
sexes for species of this subgenus are provided in
the keys (couplet 1 for the females and couplet 2
45
for the males; for further details see Popov, 1931;
Loken, 1984).
The many separate subgeneric names for
species-groups of parasitic bumble bees are not
used here, because the male genitalia of their type
species are more similar to one another than is
general among the subgenera of the genus
Bombus (e.g. Pittioni, 1939a; Ito, 1985; Williams,
1985).
Knowledge of the systematics of species of the
subgenus Psithyrus from Asia has been limited by
the scarcity of material in collections. These bees
are permanent social parasites in colonies of other
species of the genus Bombus (e.g. reviews by
Alford, 1975; Fisher, 1987; no records from
colonies of species of the subgenus Mendaci-
bombus). Consequently they are usually much
less abundant than individuals of the host species.
Many of the nominal taxa have been described
from very small samples, collected at widely
separated localities. There is therefore relatively
little information available to describe the varia-
tion of many of the nominal taxa from Asia.
Bombus (Psithyrus) bohemicus Seidl
(Figs 21, 50, 90, 130, 170, 206, 258, Maps 11 & 12)
Apis nemorum Fabricius, 1775: 380. Lectotype
female by designation of Lgken (1984: 8) (see
Note 1 below), DENMARK: Copenhagen, no
further data (BMNH) [examined]. Junior
primary homonym of Apis nemorum Scopoli,
1763.
Bombus bohemicus Seidl, 1837: 73. Type not
found (Tkalct, 1969b) (see Note 2 below),
CZECHOSLOVAKIA: Prague, no further
data [not seen]. Synonymy with Apis nemorum
Fabricius suggested by Blithgen (1918), con-
firmed here.
TYPE MATERIAL. Note 1. In her revision of the
Scandinavian Psithyrus fauna, Lgken (1984) states
that the ‘type’ of Apis nemorum F. is a female in
the Banks collection (BMNH). This had been
examined for her by I. H. H. Yarrow, who had
ascribed it to the subgenus Ashtonipsithyrus. In
this collection, above an original, handwritten
drawer label that begins ‘Apis nemorum ?’, stand
two female Psithyrus that carry Yarrow’s deter-
mination labels. One is a female labelled Psithyrus
sylvestris Lepeletier and the other a female labelled
Psithyrus bohemicus by Yarrow, identifications
that I confirm. Lg@ken’s use of the word ‘type’ in
reference to the second specimen, which has had
terga and sterna V-VI mounted on card on a
separate pin, is taken to be a valid lectotype
designation (Art. 74b).
46
Note 2. There is no reason to doubt the usual
interpretation of B. bohemicus, so it is neither
necessary nor valid to designate a neotype (Art.
75b).
AFFINITIES. The rare Mongolian (NMP) and
Siberian (BMNH) 8B. (Exilobombus) exil
(Skorikov) (justified emendation of ‘Mucidobombus
exiln. nov.’ Skorikov by Milliron, 1961: 56) shares
a pronounced interio- basal process of the male
gonostylus (Fig. 5) with all groups of bumble
bees (synapomorphy), apart from the species
of the subgenus Mendacibombus (secondarily
partly reduced for species of the subgenera
Orientalibombus, Kallobombus Dalla Torre,
Alpinobombus Skorikov, Pyrobombus and Cul-
lumanobombus Vogt). A general dorso-ventral
flattening of the male gonostylus against the
dorsal surface of the volsella is shared by most
groups of bumble bees (?synapomorphy), apart
from species of the subgenera Mendacibombus,
Exilobombus_ Skorikov, Confusibombus and
possibly Bombias (but see the comments on the
subgenus Mendacibombus). The absence of an
interio-apical process on the volsella is also shared
by species of the subgenera Eversmannibombus
and Psithyrus (possibly not a symplesiomorphy
with Mendacibombus, see Williams 1985, but
a secondary reduction and synapomorphy of
Eversmannibombus + Psithyrus, Figs 130-132).
The species of the subgenus Psithyrus share the
development of a particularly dense brush of long,
branched hairs associated with the interio-basal
process of the male gonostylus (synapomorphy).
Within the subgenus Psithyrus, B. bohemicus
belongs to a group of species (the ashtoni-group,
formerly subgenus Ashtonipsithyrus) that share a
reduction of the ventro-basal angle of the penis
valve (Fig. 90) (synapomorphy). Most females of
this group have the keels of sternum VI broad,
narrowing apically where they converge before
the apex of the sternum (Fig. 206) (the keels are
more pronounced and blade-like in the North
American B. suckleyi Greene).
Within the ashtoni-group, B. bohemicus shares
with the North American (BMNH, PW) B. ashtoni
(Cresson) and the Korean (Kim & Ito, 1987;
BMNH) B. coreanus (Yasumatsu) the develop-
ment of a pronounced shoulder on the outer side
of the penis valve shaft, immediately behind the
penis valve head (Fig. 50) (?synapomorphy). This
shoulder is weakly developed in the European and
North African (Popov, 1927a, 1931; Loken, 1984;
BMNH, PW) B. vestalis (Geoffroy in Fourcroy),
which can also be distinguished by its broader
male gonostylus. The male of B. coreanus has the
head of the penis valve shorter so that it is almost
PAULH. WILLIAMS
semi-circular in dorsal aspect, with the apex
inwardly directed.
B. bohemicus is closely similar to the North
American B. ashtoni and it is possible that they
are parts of the same species. However, subtle
differences in shape and sculpturing could be
taken to reflect a lack of interbreeding as separate
species. These differences include the broader
male gonostylus, relative to the apex of the
gonocoxite, and a slightly smaller ventro-basal
angle of the penis valve for B. ashtoni.
DISTRIBUTION. B. bohemicus is one of the most
widespread of all socially parasitic bumble bee
species (Map 11). It is known from Europe (e.g.
Popov, 1927a, 1931; Alford, 1975; Lg@ken, 1984;
BMNH, PW), south-eastwards to Turkey and the
Caucasus (BMNH), and north-eastwards across
the U.S.S.R. (Popov, 1927a, 1931; BMNH) to
Kamchatka (Popov, 1927a; Bischoff, 1930). In
central Asia it reaches southwards to the Tien
Shan (Popov, 1927a, 1931), the Pamir (Popov,
1931) and Kashmir; and in the far east it reaches
southwards to Mongolia (Popov, 1927a, 1931;
Tkalci, 1974a; BMNH, PW), Inner Mongolia
(Reinig, 1936), Heilongjiang (Popov, 1931),
North Korea (Kim & Ito, 1987), Shanxi (Maa,
1948), Gansu (Bischoff, 1936) and Sichuan
(Popov, 1927a).
MATERIAL EXAMINED FROM KASHMIR. B. bohemicus
has been collected in Kashmir only at Lal Pani (Map
12): 10 males, 3000-3700 m, 2-3.ix.1953 (Schmid)
(BMNH).
In Britain this species is known to breed in the
nests of B. lucorum (Alford, 1975; see the
comments on B. lucorum).
VARIATION WITHIN KASHMIR. The males from
Kashmir are closely similar in colour pattern (Fig.
258) to individuals from Britain. Among the fauna
of Kashmir, they are most similar to some males of
B. morawitzianus (Fig. 270), but are easily distin-
guished by the narrow volsella. The females are
likely to have a similar colour pattern to the males,
but with very much less of the yellow pubescence
at the rear of the thorax and on tergum I. This
pattern is quite distinctive among the Kashmir
fauna.
FOOD PLANTS. No records.
Bombus (Psithyrus) novus (Frison), comb. n.
(Figs 24, 28, 32, 36, 51, 91, 131, 171, 207, 259-262,
Maps 13 & 14)
Psithyrus (Psithyrus) novus Frison, 1933: 340.
Holotype female by original designation [p.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
341], INDIA: Kashmir, Nagaberan, 10000-
10500 ft [3000-3200 m] (Bion) (Calcutta) [not
seen].
Psithyrus (Psithyrus) novus subsp. nepalensis
Tkalct, 1974b: 318. Holotype female by original
designation, NEPAL: Khumbu, Pheriche,
4300 m, 5.vii.1964 (Dierl) (ZS) [examined].
Provisional synonym.
AFFINITIES. B. novus belongs to a group of species
(the rupestris-group, formerly subgenus Psithyrus
s.str.) for which the volsella has an acutely
produced inner corner (Figs 131 & 132) (synapo-
morphy). Females of this group have the keels of
sternum VI blade-like and converging close to the
apex of the sternum (Figs 207 & 208). The females
have the mandibles conspicuously narrowed in the
distal half (Fig. 32).
Within the rupestris-group, males of B. rupestris
and B. branickii (see the comments on B. branickii)
are distinguished by a slight elongation of the
penis valve shaft between the penis valve head and
the ventro-basal angle (Figs 51 & 52) and by the
narrower distal part of the volsella (Figs 131 &
132) (?synapomorphies). The females have the
clypeus closely and coarsely punctured almost
throughout.
In contrast, other material from Kashmir runs
to the name Psithyrus morawitzi Friese in Popov’s
(1931) key, although it agrees closely with the
more recent description of B. novus. Tkalct
(1974b) considered B. novus to belong to the
‘chinensis-Gruppe von Psithyrus s. str.’. From
Popov’s key this includes B. chinensis (Morawitz)
and B. morawitzi, which were both described
from Chinese material (from Gansu & Sichuan
respectively). I have seen no specimens of either
of these nominal taxa for comparison with B.
novus. Bischoff (1936) also described material
belonging to this group from Sichuan and Yunnan.
His material showed the colour patterns both with
and without yellow bands, as described for B.
chinensis and B. morawitzi respectively, although
he found that these individuals scarcely differed
in morphology (‘Morphologisch ist diese Form
[Psithyrus chinensis subsp. hoenei Bischoff, with
yellow pubescence] kaum von dem typischen
chinensis zu trennen, weicht aber in der Farbung
sehr stark ab.’) and Bischoff identified them all as
Psithyrus chinensis. More recently, Tkalct (1987)
has regarded Psithyrus morawitzi as a subspecies
of Psithyrus chinensis.
The holotype of Psithyrus novus subsp.
nepalensis differs from B. novus s.str. not only in
colour (which could be due in part to the fading of
any yellow or orange pubescence to whitish for
this abraded, old specimen), but also in several
47
morphological details. In particular, none of the
specimens from Kashmir has the basal keel of the
mandible reaching the margin (Fig. 32, including
the holotype, see Frison, 1933: fig. Sb). The weak
incision in this part of the margin is present for the
specimen from Nepal, as noted by Tkalci, but is
not at all distinct for the specimens from Kashmir.
The dorsal surface of the labral tubercles of the
Nepalese specimen has many large punctures,
which are few or lacking for the specimens from
Kashmir. The unpunctured part of the ocello-
ocular area extends nearly three-quarters of
the breadth of the ocello-ocular distance for the
specimen from Nepal but no more than half of
the distance for the specimens from Kashmir. The
weakly-arched transverse ridge of sternum
II shows a median anterior deviation for the
specimens from Kashmir that is not marked
for the specimen from Nepal. However, both
nominal taxa share a reduction in the density of
the punctures on the clypeus and on antennal
segment 1 (scape). With so little material avail-
able from the eastern Himalaya and from the
intervening areas, especially of the males, I
consider it premature to conclude that these
nominal taxa represent separate species. More
material is needed to establish the relationships of
all of the nominal taxa in this group.
DISTRIBUTION. The present, provisional concept
of B. novus is of a Himalayan species (Map 13).
It is known from Nepal (Tkalct, 1974b; ZS),
Himachal Pradesh (BMNH, PW) and Kashmir.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. novus is recorded from the southern side of the
Great Himalaya range and from the Pir Panjal
range, in mesic subalpine scrub (Map 14): 13
females, 31 males, from 2 localities 3000-3700 m
(BMNH, BT, NR, PW).
During July and August 1986, I saw many
females investigate gaps among large boulders
that were lying in gullies in the subalpine zone
of Mt Apharwat. Queens of several species of
bumble bees had been seen at this site, many
apparently searching for nest sites. Most abundant
among these bees were queens of B. rufofasciatus.
VARIATION WITHIN KASHMIR. The colour pattern
of only one of the females agrees closely with the
original description, in that it has a broad black
band between the wing bases (Fig. 261). Other
females have only a few black hairs along the
posterior margin of the scutum (Fig. 259). The
putative males are associated with these females
because, among material from Kashmir, they also
belong to the rupestris-group but, unlike B.
branickii, resemble female B. novus in having
48
some form of black band between the wing bases
(Figs 260 & 262). The males differ from the
females in that the pale pubescence of the thorax
and tergum I is grey-white, as for the female from
Nepal (Tkalci, 19745).
The colour pattern of the pubescence of the
males is black, with the sides of the thorax
as far as the leg bases, the rather poorly-
defined anterior and posterior bands on the
thoracic dorsum and tergum I grey-white;
terga II-III pale yellow; terga IV-VII orange-
red, with variable degrees of replacement by
black basally on tergum IV (Fig. 260), which
may extend over the entire tergum (Fig. 262);
sterna and legs predominantly black. The
wings are weakly to moderately strongly
clouded with brown (infuscated).
The colour pattern of the males (Figs 260 &
262) is similar to that of some males of B.
himalayanus (Figs 246, 249, 252), B. kashmirensis
(Fig. 304), B. rufofasciatus (Figs 433, 436, 439),
and probably B. lemniscatus (cf. Figs 325 & 326),
that occur in the same localities.
FOOD PLANTS. (Ranunculaceae) Aconitum laeve
Royle; (Gentianaceae) Swertia petiolata D. Don;
(Scrophulariaceae) Scrophularia paucifiora Benth.
Bombus (Psithyrus) branickii
(Radoszkowski), comb. n.
(Figs 52, 92, 132, 172, 208, 263-266, Maps 15 &
16)
Psithyrus Branickii Radoszkowski, 1893: 241.
Lectotype female by designation of Tkalct
(1969a: 204), U.S.S.R.: ?Kirgiziya S.S.R.,
‘Kara-Karyk’ mountains, 10000 ft [3000 m]
(Barei) (MNHU) [examined].
Psithyrus rupestris var. eriophoroides Reinig,
1930: 110 (see Note 1 below). 4 syntype males,
U.S.S.R.: Tadzhikistan S.S.R.; KirgiziyaS.S.R.;
CHINA: Xinjiang (ITZ) [not seen]. Synon-
ymised with Psithyrus branickii Radoszkowski
by Tkalct (1969a).
Psithyrus (Psithyrus) rupestris subsp. elisabethae
Reinig, 1940: 231. Holotype female by monotypy
(see Note 2 below), AFGHANISTAN: Hindu
Kush, north side of ‘Nuksan’ pass, 3500-4000
m, mid vii.1936 (Kotzsch) (ZS) [examined].
Synonymised with Psithyrus _ branickii
Radoszkowski by Tkalct (19692).
NOMENCLATURE. Note 1. Reinig’s use (1930: 83
etc.) of the term ‘ssp.’ elsewhere in his paper is
taken to indicate that infrasubspecific rank is
meant for Psithyrus rupestris var. eriophoroides.
However, this name has subsequently been
PAULH. WILLIAMS
treated as a name in the species group by Reinig
(1935), so it is deemed to be of subspecific rank
(Art. 45g(11)(1)).
TYPE MATERIAL. Note 2. Reinig’s description of
Psithyrus rupestris subsp. elisabethae specifies that
only a single female was examined. A single
female in the ZS collection agrees with the
original description and bears labels with the data
quoted. It also bears a red typed label “Holotypus’
and a handwritten label ‘Psithyrus (Ps.) / rupestris
Fabr. / ssp. elisabethae / [female] n. Reinig /
det.W.F.Reinig 1939’. I believe that this is the
single specimen on which the original description
is based and regard it as the holotype (Art.
73a(ii)).
AFFINITIES. B. branickii belongs to the rupestris-
group and is closely similar to B. rupestris (see the
comments on B. novus), a Eurosiberian species
(e.g. Popov, 1931; Reinig, 1935; Lgken, 1984). B.
rupestris is known from an area that reaches south-
eastwards from Europe to Turkey and the Caucasus
(BMNH); in central Asia it reaches south to the
Altai (Popov, 1931); and in the far east it reaches
southwards to Mongolia (Popov, 1931; Tkalcu,
1974a; BMNH), Inner Mongolia (Reinig, 1936),
and Gansu (Popov, 1931; Bischoff, 1936). B.
branickii has shallower and less well-defined post-
ocular punctures, has a weaker post-ocular groove
on the vertex (the ‘dorsal furrow of the gena’ in
Richards, 1968) and has longer pubescence. This
is especially evident in the posterior fringe of the
hind basitarsus, where the length of the hairs
exceeds the breadth of the basitarsus for B.
branickii, but not for B. rupestris. The wings of B.
branickii are also only moderately, rather than
very strongly, darkened (infuscated).
DISTRIBUTION. B. branickii is a central Asian and
Tibetan species from the mountains around the
deserts of China (Map 15). It is known from
Transbaikal, Mongolia and the Altai (Popov,
1931), the Tien Shan (Radoszkowski, 1893;
Morawitz, 1894; Popov, 1931; Maa, 1948; MNHU),
the Pamir (Reinig, 1930), the Hindu Kush (Reinig,
1940; Tkalci, 19692; BMNH, ZS), Kashmir,
Himachal Pradesh (BMNH, PW), Tibet (BMNH),
Sichuan (Reinig, 1935), Shanxi (Yasumatsu,
1951) and North Korea (Kim & Ito, 1987). Reinig
(1935: chart 1) mapped part of this distribution
under the name Psithyrus rupestris subsp.
eriophoroides.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. branickii is recorded from the Hindu Raj,
Great Himalaya and Zanskar ranges, in the drier
alpine scrub and steppe (Map 16): 21 females, 71
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
males, from 9 localities 1500-4800 m (BMNH, IZ,
MI, NM, PW).
During July 1980, I saw females of this species
investigate gaps under boulders and holes in dry-
stone walls on Nimaling plain. Although queens
of 8 species of the genus Bombus were flying in the
area, only B. asiaticus, B. kashmirensis and B.
keriensis had workers flying at this time.
VARIATION WITHIN KASHMIR. One of the females
from Atro Sar (Fig. 266) and another from
Sonamarg agree closely in colour pattern with the
original description, in that the pubescence of
terga V-VI is entirely black. The remaining females
have red pubescence on these terga (Figs 263,
265). All females have tergum I yellow and often
have a few pale hairs apico-laterally on tergum II.
Two females from Nimaling have more of these
pale hairs on tergum II, so as to form an apical
band that is only interrupted in the middle by
black. Another two females from the same locality
also have a pale, apical band on tergum III (Fig.
263). The males associated with these females
belong to the rupestris-group, but unlike B.
novus, they lack a black band in the pubescence
between the wing bases, although they do have a
few black hairs intermixed on the thoracic dorsum
(Fig. 264).
Some individuals (Fig. 265) are closely similar
in colour pattern to some B. avinoviellus (Fig.
232), B. marussinus (Figs 254-257), B. subtypicus
(Figs 319-324), B. biroi (Figs 335-337), B. asiaticus
(Figs 371-376) and B. semenovianus (Figs. 397-402)
that may occur with them. B. branickii is easily
distinguished by the subgeneric characters (see
the key, couplets 1 & 2).
FOOD PLANTS. (Leguminosae) Caragana versicolor
(Wallich) ex Benth.; unidentified yellow thistle-like
composites.
Bombus (Psithyrus) ferganicus
(Radoszkowski), comb. n.
(Figs 22, 53, 93, 133, 173, 209, 267, 268, Maps 17
& 18)
Psithyrus ferganicus Radoszkowski, 1893: 241.
Lectotype female by designation of Tkalct
(1969a: 206), U.S.S.R.: ?Kirgiziya S.S.R.,
“Kara-Karyk’ mountains (MNHU) [examined].
Psithyrus indicus Richards, 1929a: 139. Holotype
male by original designation [p. 141], INDIA:
Kashmir, Gulmarg, 1913 (Thomson) (BMNH)
[examined]. Synonymised with Psithyrus fergan-
icus Radoszkowski by Griitte (1937).
AFFINITIES. B. ferganicus belongs to a group of
species (the campestris-group, formerly subgenus
49
Metapsithyrus) for which the apical part of the
volsella is greatly broadened (Figs 133 & 134) and
the ventro-basal angle of the penis valve is apically
rounded, although this angle retains its curved,
hook-like form (Figs 53 & 54) (synapomorphies).
Females of this group have the keels of sternum
VI broad throughout their length, although they
converge distinctly before and below the apex of
the sternum (Figs 209 & 210).
Within the campestris-group, B. ferganicus
shows the least broadening of the volsella (Figs 133
& 134) (see the comments on B. morawitzianus).
The most similar European species is the Euro-
siberian (e.g. Popov, 1931; Alford, 1975; Lgken,
1984) B. campestris (Panzer). From Europe B.
campestris reaches south-eastwards to Turkey
(BMNH), the Caucasus (Popov, 1931; BMNH)
and the Elburz (BMNH); from central Asia it
reaches southwards to the Tien Shan (Popov,
1931); and in the far east it reaches southwards to
Mongolia (Popov, 1931; Tkalct, 1974a; BMNH),
Inner Mongolia (Reinig, 1936), Ningxia (Popov,
1931), and Gansu (Popov, 1931; Bischoff, 1936).
The unique female of B. susteraianus (Tkalct)
from North Korea is closely similar to B. campestris,
but has slightly less pronounced keels of sternum
VI (Tkalci, 1959) and slightly more pronounced
labral tubercles. The male of this nominal taxon
remains unknown. The Chinese B. pieli (Maa)
(synonymised with Psithyrus tajushanensis Pittioni
by Tkalci, 1987) is known from Sichuan (Tkalcu,
1987; BMNH), Hubei (Sakagami, 1972), Anhui
and Zheijiang (Maa, 1948) and Fujian (Pittioni,
1949; Tkalct, 1987; BMNH). The male genitalia
of B. ferganicus, in ventral aspect, have the distal
half of the volsella narrower than for B. campestris
or B. morawitzianus. I have not seen males of B.
pieli, but Maa (1948: fig. 16) described its volsella
as slightly shorter than for B. campestris. Females
of B. ferganicus can be distinguished from those of
the other three species by the shallower punctures
of the post-ocular area. These punctures are
widely spaced for B. ferganicus, as for B.
morawitzianus, but in contrast they are very
shallow and poorly-defined, so that the area is
nearly smooth and strongly shining. The females
of B. campestris and B. pieli have smaller punctures
in the post-ocular area, separated by less than their
own widths, though these punctures are slightly
closer and better defined for B. pieli. B. pieli can
also be distinguished from B. campestris by their
labral tubercles, which are more strongly angled.
DISTRIBUTION. B. ferganicus is a central Asian
species (Map 17). It is known from the Tien Shan
(Radoszkowski, 1893; Morawitz, 1894; Griitte,
1937; MNHU), the Hindu Kush (Reinig, 1937;
50
Tkalci, 19692; BMNH, PW), Pakistan (PW),
Kashmir and Himachal Pradesh (BMNH, PW).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. ferganicus is recorded from the western
Karakoram, Great Himalaya and Pir Panjal ranges,
in both montane coniferous forest and subalpine
scrub (Map 18): 14 females, 115 males, from 5
localities 2400-3700 m (BMNH, BT, NR, PW,
ZM, ZS).
VARIATION WITHIN KASHMIR. Only minor variation
is evident in the replacement of pale hair on the
gaster by black, especially on terga I-II, and in the
colour of the pale hair, which varies from lemon-
yellow to brown (Figs 267 & 268). This species is
quite distinctive in colour pattern among the
fauna of Kashmir.
FOOD PLANTS. (Leguminosae) Trifolium repens
L.; (Compositae) Cirsium falconeri (Hook.f.)
Petrak, C. wallichii DC.
Bombus (Psithyrus) morawitzianus (Popov),
comb. n.
(Figs 54, 94, 134, 174, 210, 269, 270, Maps 19 &
20)
Psithyrus (Metapsithyrus) morawitzianus Popov,
1931: 148, 183 (see Note 1 below). Lectotype
female by designation of Podbolotskaya (in
press), U.S.S.R.: Kirgiziya S.S.R., Naryn, 19.
vii.1904[?] (ZI) [examined].
Psithyrus (Metapsithyrus) redikorzevi Popov,
1931: 160, 181. 5 syntype males, U.S.S.R.:
Tadzhikistan S.S.R.; Kirgiziya S.S.R. (ZI)
[not seen]. Synonymised with Psithyrus mora-
witzianus Popov by Griitte (1937) (see Note 2
below).
NOMENCLATURE. Note 1. Popov described Psithyrus
morawitzianus with particular reference to a female
that bore Vogt’s manuscript label “Ps. campestris
var. morawitzianus typicus O.V. Cotype’ (Popov,
1931: 183, although the label actually reads ‘Ps.
campestris / var. morawitzianus O.V./typicus O.V.
/ Cotype’). Therefore Popov is deemed to be the
author (Art. 50a).
Note 2. Following the Principle of the First
Reviser (Art. 24), B. morawitzianus is accepted in
precedence to B. redikorzevi, by the action of
Griitte (1937).
AFFINITIES. B. morawitzianus, like B. ferganicus,
belongs to the campestris-group (see the comments
on B. ferganicus). The male genitalia of B.
morawitzianus clearly differ from those of B.
ferganicus, B. campestris and B. pieli by the great
breadth of the distal half of the volsella, in ventral
PAULH. WILLIAMS
aspect (Figs 133 & 134), and by the extremely
short, or transverse, gonostylus (Figs 173 & 174).
The punctures of the post-ocular area of the
female B. morawitzianus are larger, more distinct
and more widely spaced than for individuals of B.
ferganicus, B. campestris or B. pieli. The punctures
of tergum VI are finer for B. morawitzianus than
for individuals of B. ferganicus, B. campestris or
B. pieli.
DISTRIBUTION. B. morawitzianus is acentral Asian
species (Map 19). It is known from the Tien Shan
(Morawitz, 1875; Popov, 1931; Griitte, 1937; ZI),
the Pamir (Popov, 1931; Griitte, 1937), the Hindu
Kush (Reinig, 1940; Tkalci, 1969a) and Kashmir.
A record from the Kopet Dag (Rasmont, 1983)
needs to be confirmed.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. morawitzianus is recorded from the Hindu Raj
and Great Himalaya ranges (Map 20): 1 female, 7
males, from 4 localities 2700-4000 m (BMNH).
VARIATION WITHIN KASHMIR. The colour patterns
of both sexes are similar to those described by
Popov (1931) (Figs 269 & 270). There is some
variation in the colour of the pale pubescence of
the male gaster between white and dull yellow.
This species is quite distinctive in colour pattern
among the fauna of Kashmir.
FOOD PLANTS. No records.
Bombus (Psithyrus) skorikovi (Popov),
comb. n.
(Figs 55, 95, 135, 175, 211, 271-274, Maps 21 &
22)
Psithyrus skorikovi Popov, 1927b: 267. Holotype
female by original designation [p. 268], CHINA:
Gansu [?or Qinghai], northern slopes of the
Xining mountains, before 1.vii.1910 (Grum-
Grzhimailo) (Z1) [examined].
[Psithyrus skorikovi var. mesoxanthus Richards,
1928c: 360 [examined]. Infrasubspecific (see
Note 1 below) (Art 45g(ii)(1)), unavailable
name (Art. 45e).]
Psithyrus (Fernaldaepsithyrus) gansuensis Popov,
1931: 168, 202. Holotype male by monotypy
(see Note 2 below), CHINA: Qinghai, ‘Ui-ju’
to ‘Choto’, 15-16.viii.1908 (Kozlov) (ZI) [not
seen]. Synonymy with Psithyrus skorikovi
suggested by Popov (1931), confirmed here.
NOMENCLATURE. Note 1. In the introduction to his
paper, Richards (1928c: 345) contrasts his use of
the term ‘varieties’, for ‘sporadically’ occurring
colour variations, with the term ‘races’, which he
used for ‘. . . populations inhabiting definite, usually
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
continuous, areas and characterised by peculiarities
of sculpture, hair-length or average colour-
pattern.’. This is taken to show that infrasub-
specific rank is meant for Psithyrus skorikovi var.
mesoxanthus.
TYPE MATERIAL. Note 2. Popov’s description of
Psithyrus gansuensis specifies that only a single
male was examined. If only a single male with the
data quoted can be found, then this specimen
should be regarded as the holotype (Art. 73a(ii)).
AFFINITIES. B. skorikovi belongs to a group of
morphologically similar species (the fernaldae-
group, formerly subgenus Fernaldaepsithyrus) of
mainly Eurosiberian distribution (e.g. Popov,
1931; Lg@ken, 1984), with one North American
species (B. fernaldae (Franklin)). These species
share a great reduction in the breadth of the male
volsella (Fig. 135) and a narrowing of the apex of
the gonocoxite (synapomorphies). Females of this
group have the keels of sternum VI narrowed
apically and converging a long way before the
apex of the sternum (Fig. 211). This apex is itself
narrowed into a ventrally-curved spine, which
projects distinctly beyond the apex of tergum VI.
Within the fernaldae-group, B. skorikovi is
most similar to the Eurosiberian (e.g. Popov,
1931; Loken, 1984) B. flavidus Eversmann in that
the ventro-basal angle of the penis valve remains
acute and strongly hook-shaped (Figs 55 & 95). B.
flavidus is otherwise unique within the fernaldae-
group for its restricted distribution in the sub-
alpine and subarctic zones (see Pittioni, 1942: map
4; Loken, 1984). In southern Europe, B. flavidus
is known from the Pyrenees, the Alps and western
Turkey (Pittioni, 1942; BMNH). In a separate
northern area of distribution it reaches eastwards
from Scandinavia to Kamchatka (Popov, 1931;
Pittioni, 1942), without records from further
south than Lake Baikal. The females of B. skorikovi
differ from those of B. flavidus in the weaker basal
keel of the mandible, in the less extensively and
less densely punctured ocello-ocular area and in
the more strongly marked labral tubercles and
lamella. The male of B. skorikovi has the ventro-
basal angle of the penis valve much narrower. The
single male from Qinghai described by Popov
under the name Psithyrus gansuensis appears
from the description of the penis valve (see
Popov, 1931: fig. 26D) to be a male of B.
skorikovi. A single female from Qinghai described
by Tkalci (1961) under the name Psithyrus kuani
also appears to be closely similar to B. skorikovi. I
have not seen either of these specimens.
The females from Kashmir that are ascribed
here to B. skorikovi differ from specimens from
Qinghai and Gansu in that tergum VI has, at most,
51
only a very weak median ridge and there are more
large punctures with fewer small punctures. For
some of the females from Kashmir the sculpturing
of the surface of tergum VI is weaker so that it
appears more strongly shining. However, some of
the females from southern Tibet appear to be
intermediate, at least in the reduction of the
median ridge (Richards, 1928c; BMNH). It is
considered premature to regard the Kashmir
material as part of a separate species.
DISTRIBUTION. B. skorikovi is a peri-Tibetan
species (Map 21). It is known from Kashmir,
southern Tibet (Richards, 1928c; BMNH), Qinghai
(Popov, 1931; Tkalci, 1961; ZS) and Gansu
(Popov, 1927b, 1931; Bischoff, 1936; PW, ZI).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. skorikovi is recorded from the southern side of
the Great Himalaya range and from the Pir Panjal
range (Map 22): 15 females, 301 males, from 2
localities 2700-3700 m (BMNH, NR, PW).
VARIATION WITHIN KASHMIR. Females from Mt
Apharwat are similar in colour pattern to the holo-
type, although the apical margin of tergum II is
yellow, whereas the median part of tergum IV and
the entirety of tergum V are black-haired (Fig. 273).
The majority of the females from Lal Pani have
the black pubescence of the thoracic dorsum
reduced to a small, indistinct patch at the rear of
the scutum and have more yellow hairs on the
lateral parts of terga II and IV (Fig. 271). The
putative males are associated with these females
because they are the only males of the fernaldae-
group of species among the material from Kashmir
and occur together with the known females at Lal
Pani, in the Kishanganga valley.
The colour pattern of the pubescence of the
male (Figs 272 & 274) is dull yellow, except for
black on the front and sides of the head, in a
poorly-defined band between the wing bases, on
terga V-VII and as a very few hairs in the middle
of the posterior margin of tergum IV; the dark
hairs of terga VI-VII are pale-tipped and often
distinctly orange; the hairs of the legs are
predominantly black, with some pale hairs inter-
mixed. The wings are weakly clouded with brown
(infuscated). Some males have the black band
between the wing bases much reduced (120/301
males) (Fig. 272), as described for the female. A
few males have a strong admixture of black hairs
basally on tergum II (25/301 males, Fig. 274).
This species is quite distinctive in colour pattern
among the fauna of Kashmir.
FOOD PLANTS. (Compositae) Cirsium falconeri
(Hook.f.) Petrak; (Scrophulariaceae) Scrophularia
pauciflora Benth.
52
Subgenus DIVERSOBOMBUS Skorikov
Diversobombus Skorikov, 1914d: 406 (as a sub-
genus of Bombus Latreille). Type species:
Bombus diversus Smith, 1869: 207, by sub-
sequent designation of Sandhouse (1943: 546).
Diversibombus Skorikov; Skorikov, 19385: 1.
Unjustified emendation (Art. 33b(i, iii).
For a general description of both sexes see
Richards (1968).
Bombus (Diversobombus) trifasciatus Smith
(Figs 11, 17, 39, 56, 96, 136, 176, 275-278, Maps
23 & 24)
Bombus trifasciatus Smith, 1852a: 43. LECT-
OTYPE queen by present designation (see
Note 1 below), CHINA: Zhejiang, Zhoushan
(Fortune) (BMNH) [examined].
Bombus montivagus Smith, 1878b: 168 [pos-
thumous re-proposal in error, 1879: 131].
Lectotype queen by designation of Tkalct
(1968b: 23), BURMA: Karen, Moolaiyet,
3000-6000 ft [900-1800 m] (BMNH) [examined].
Syn. n.
Bombus_ secundus Dalla Torre, 1890: 139.
Replacement name for Bombus montivagus
Smith, 1879, junior primary homonym of
Bombus montivagus Smith, 1878b. Synonymised
with Megabombus montivagus montivagus
(Smith, 1878b) by Tkalct (19685). Syn. n.
Bombus ningpoénsis Friese, 1909: 676. 2 syntype
workers, CHINA: Zhejiang, Ningbo (MNHU)
[not seen]. Synonymised with Bombus trifasci-
atus Smith by Tkalct (1961).
Bombus wilemani Cockerell, 1911: 100. Holotype
worker by original designation [p. 101],
TAIWAN: Arizan, 7500 ft [2300 m], 11.ix.1906
(Wileman) (BMNH) [examined]. Provisional
synonym.
Bombus haemorrhoidalis var. albopleuralis Friese,
1916: 108. Lectotype queen by designation of
Tkalct’ (1974b: 344), INDIA: Uttar Pradesh,
Kumaun, 21.vi.1910 (MNHU) [examined].
Syn. n.
Bombus maxwelli Pendlebury, 1923: 67. Holotype
queen by monotypy (see Note 2 below), WEST
MALAYSIA: Pahang, Gunong Berumbun,
4800 ft [1500 m], 4.ix.1922 (BMNH) [examined].
Provisional synonym.
Bombus (Hortobombus) mimeticus Richards, 1931:
529. Holotype queen by original designation
[p. 530], INDIA: West Bengal, Darjiling,
Tukvar, 4000 ft [1200 m], iv.1894 (Bingham)
(BMNH) [examined]. Syn. n.
Bombus (Hortobombus) mimeticus var. albol-
PAULH. WILLIAMS
ateralis Richards, 1931: 530. Holotype worker
by monotypy (see Note 3 below), NEPAL: no
further data (BMNH) [examined]. Syn. n.
Bombus (Hortobombus) mimeticus var. ganto-
kiensis Richards, 1931: 530. Holotype queen by
monotypy (see Note 4 below), INDIA: Sikkim,
Gantok, 24.vi.1903 (BMNH) [examined]. Syn.
n.
Bombus (Hortobombus) mimeticus var. turneri
Richards, 1931: 530. LECTOTYPE queen by
present designation (see Note 5 below), INDIA:
Meghalaya, Shillong, v.1903 (Turner) (BMNH)
[examined]. Junior secondary homonym in
Bombus of Psithyrus turneri Richards, 1929a [=
B. turneri (Richards)]. Syn. n.
Bombus (Hortobombus) mimeticus var. insidiosus
Richards, 1931: 531. Holotype queen by original
designation, PAKISTAN: Baltistan, no further
data (BMNH) [examined]. Syn. n.
Bombus (Hortobombus) mimeticus var. geminatus
Richards, 1931: 531. Holotype worker by
monotypy (see Note 6 below), INDIA: Kashmir,
Gulmarg, 1913 (Thomson) (BMNH) [exam-
ined]. Synonymised with Megabombus albo-
pleuralis albopleuralis (Friese) by Tkalct, 1974b.
Syn. n.
Bombus (Hortobombus) mimeticus var. magret-
tianus Richards, 1931: 531. Holotype male by
original designation [p. 533], 7BURMA [not
Sikkim (see Note 7 below)]: no further data
(MNHU) [examined]. Syn. n.
Bombus (Diversobombus) ningpoensis subsp.
minshanicus Bischoff, 1936: 19. LECTOTYPE
worker by present designation (see Note 8
below), CHINA: Gansu/Sichuan border, ‘Ma-
tou-shan’, 600-1000 m, 2.vi.1930 (Hummel)
(MNHU) [examined]. Syn. n.
Megabombus (Diversobombus) montivagus subsp.
quasibreviceps Tkalci, 1968b: 27. Holotype
queen by original designation, CHINA: Yunnan,
no further data (Genf) [not seen]. Syn. n.
Megabombus_ (Diversobombus) albopleuralis
subsp. atropygus Tkalct, 1989: 58. Holotype
worker by original designation, BURMA: Nam
Tamai valley, 27°48’N 97°48’E, 3500 ft [1067
m], 12.ix.1938 (Kaulback) (BMNH) [examined].
Syn. n.
TYPE MATERIAL. Note 1. There are two queens of
B. trifasciatus in the BMNH collection labelled
‘Shang / hai’ that may have been among Smith’s
syntypes in 1852. The original description quotes
neighbouring ‘Chusan’ [= Zhoushan] as the type-
locality. Much of Fortune’s material is labelled
‘Shang / hai’, although Smith described it as from
‘Chusan’ on Fortune’s advice (Smith, 1852a: 33,
43), even when some of it must have originated
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
from India (see the comments on B. haemor-
rhoidalis). In this case there is no reason to doubt
Zhoushan as the area of origin of the material.
The queen that agrees most closely with the
description, by the absence of a distinct black
band between the wing bases, carries a purple-
edged label (1) ‘Lecto- / type’; (2) a blue label
‘Shang / hai’, reverse side ‘52 / 28’; (3) ‘trifasciatus
/ Type Sm.’, in handwriting identical to that of
Smith; (4) ‘Bombus / trifasciatus / TYPE. Smith’;
(5) a red-edged, printed label ‘Type’; (6) ‘B.M.
TYPE / HYM. / 17B.1028.’. According to the
accessions catalogue, the number 52/28 refers
to 17 Hymenoptera collected at ‘Shanghai’ by
Fortune and purchased from him between February
28th and March 6th, 1852, which predates the
reading of Smith’s paper at the meeting (3.v. 1852)
of the Entomological Society of London by two
months. This specimen, which lacks both hind
legs, the distal joints of all the remaining tarsi and
part of the right antennal flagellum, is designated
as lectotype (Art. 74a).
Note 2. Pendlebury stated that he had seen two
specimens of B. maxwelli: a queen, which he
described, and a ‘similar’ worker. The “Type and
paratype’ were then said to have been deposited in
the Federated Malay States Museum, Kuala
Lumpur. A single pre-1923 queen in the BMNH
agrees with the original description and bears
labels with the data quoted by Pendlebury, together
with a label ‘Bombus / maxwelli Pendl. / Type.
Pahang.’ and an accession number 1926-167. This
number refers to a collection that is specified to
have included eight Hymenopteran types, which
was presented to the BMNH from the museum at
Kuala Lumpur in 1926. I believe that this is the
single specimen on which the original description
is based and regard it as the holotype (Art.
73a(ii)).
Note 3. Richards’s description of B. mimeticus
var. albolateralis specifies that only one worker
was examined. A single pre-1931 worker in the
BMNH collection agrees with the original descrip-
tion and bears labels with the data quoted. It also
carries a label with the name B. mimeticus var.
albolateralis and ‘TYPE’, in handwriting identical
to that of Richards. I believe that this is the single
specimen on which the original description is
based and regard it as the holotype (Art. 73a(ii)).
Note 4. Richards’s description of B. mimeticus
var. gantokiensis specifies that only a single queen
was examined. A single pre-1931 queen in the
BMNH collection agrees with the original descrip-
tion and bears labels with the data quoted. It also
carries a label with the name B. mimeticus var.
gantokiensis and ‘TYPE’, in handwriting identical
to that of Richards. I believe that this is the single
53
specimen on which the original description is
based and regard it as the holotype (Art. 73a(ii)).
Note 5. The original description of B. mimeticus
var. turneri lists one queen and one worker as
typical. A single pre-1931 queen in the BMNH
collection bears a purple-edged label (1) ‘Lecto- /
type’; (2) ‘Assam / R. Turner. / 1905-175’; (3)
‘Shillong / 5.03’; (4) ‘B. orientalis var.’; (5) ‘B.
MIMETICUS / RICHARDS / VAR. TURNERI
/ RICHARDS. / Type [female]’ in handwriting
identical to that of Richards; (6) a red-edged label
‘Type’; (7) “B.M: TYPE / HYM. / 17B.1027.’.
This specimen, which lacks the distal tarsal joints
of all legs, the tibia of the right mid leg, all of the
right front leg and most of both antennal flagella,
is designated as lectotype (Art. 74a).
Note 6. Richards’s description of B. mimeticus
var. geminatus lists two workers, the second of
which is specified to be the paratype. A single pre-
1931 worker in the BMNH collection agrees with
the original description and bears labels with the
data quoted for the first worker. It also carries a
label with the name B. mimeticus var. geminatus
and ‘TYPE’, in handwriting identical to that of
Richards. I believe that this is the single specimen
on which the original description is based and
regard it as the holotype (Art. 73a(ii)).
Note 7. Tkalci (1968b: 27) questioned the
validity of the locality data ‘Sikkim’ associated
with the holotype of B. mimeticus var. magret-
tianus Richards. The other material with this
colour pattern that Tkalci had examined bears
labels for “Tenasserim’ in Burma.
Note 8. Bischoff’s description of B. ningpoensis
subsp. minshanicus lists two workers from Gansu,
dated ‘14.5’ and ‘2.6’. A worker in the MNHU
collection bears Bischoff’s labels with this name,
the date 14.5.1930 and a red label ‘Para-/Typus’.
A second worker in the MNHU collection agrees
with the description and bears a printed label (1)
“Sven Hedins / Exp. Ctr. Asien / Dr Hummel’; (2)
‘Kina/S. Kansu’; (3) ‘ningpoensis / minshanicus /
n. subsp. [worker] / det. Bischoff’; (4) ‘(Bombus)
loubouen- / sis Friese m. minshani- / cus (Bisch.) /
Skorikov det.’; (5) a red label ‘68 / 61’; (6) a
pencilled label ‘loubouensis Friese / var. nov.’; (7)
a pencilled label ‘2/6’; (8) ‘M. (D.) tri- / fasciatus
(Sm. / Tkalci det.’; (9) ‘Zool. Mus. / Berlin’. This
second specimen, which lacks only the tarsi of the
left mid leg, is designated as lectotype (Art. 74a).
AFFINITIES. The species of the subgenera Mega-
bombus Dalla Torre, Senexibombus Frison and
Diversobombus share a displacement of the two
hooks on the inner margin of the volsella to a
narrowly subapical position (Fig. 136), and the
interio-basal process of the male gonostylus is
54
extended as a strongly recurved cusp (Fig. 176)
(synapomorphies). The species of the subgenus
Diversobombus share an enlargement of the inner
hooks of the volsella, of which the inner basal
hook has become broadened and carries many
small teeth (?synapomorphy).
Within the subgenus Diversobombus, two
species-groups have been recognised, the trifasci-
atus-group and the diversus-group (Sakagami,
1972). Species of the diversus-group have the
recurved part of the cuspate interio-basal process
of the gonostylus reduced to a single, long, slender
spine (?synapomorphy) (see Tkalct, 1965: figs 46
& 47). These species include the Japanese B.
diversus Smith (distribution mapped by Sakagami,
1975: fig. 14; see also Ito, 1987), the Chinese B.
longipes Friese (synonymised with B. hummeli
Bischoff by Tkalci, 1987, and possibly also con-
specific with Diversobombus malaisei Skorikov
[not seen]) and the more widespread east Asian B.
ussurensis Radoszkowski (distribution mapped by
Sakagami, 1975: fig. 14).
For males of the trifasciatus-group, the recurved
part of the cuspate interio-basal process of the
gonostylus bifurcates near its base, so that
it forms two shorter, strongly curved spines
(Fig. 176) (?plesiomorphy within the subgenus
Diversobombus). Females can be distinguished
from those of the diversus-group by their well-
defined punctures of the post ocello-ocular area,
whereas these punctures are embedded in coarse
sculpturing for the species of the diversus-group.
Three widespread nominal taxa have been
recognised in the trifasciatus-group, which have
been described under the names B. trifasciatus
s.str., B. montivagus and B. haemorrhoidalis var.
albopleuralis. They differ principally in colour
pattern, possibly as members of regional groups of
Miillerian mimics (Fig. 11, see the discussion of
the colour patterns of the Kashmir fauna). These
nominal taxa of the subgenus Diversobombus are
very similar in colour pattern to the individuals
of B. (Orientalibombus) haemorrhoidalis, B.
(Alpigenobombus) breviceps Smith and B. (Pyro-
bombus) rotundiceps Friese that occur with them
(see the comments on B. haemorrhoidalis, B.
kashmirensis and B. lemniscatus respectively;
Sakagami & Yoshikawa, 1961; Tkalci, 1968b,
1989).
B. haemorrhoidalis var. albopleuralis is the
oldest available name for the Himalayan nominal
taxa of the trifasciatus-group ( the ‘montivagus-
Gruppe’ of Tkalci, 1974b), which were first
described as a species separate from B. haemor-
rhoidalis by Richards (1931), under the name B.
mimeticus. The colour pattern of the pubescence
on the thoracic dorsum is usually entirely black.
PAULH. WILLIAMS
There is variation in the amount of grey-white
pubescence on the sides of the thorax (pleura).
This pale pubescence is most often extensive in
males. Among the material examined in the
BMNH, the highest proportions of individuals
with extensively pale-haired pleura appear to be
from localities at lower altitudes in the eastern
Himalaya. The pubescence of gastral terga I-II
is usually bright lemon-yellow. The remaining
gastral terga III-VI are predominantly red for
individuals from the western Himalaya to as far
east as Nepal, although the same extensively red
pattern is also shown by two workers labelled
‘Manipur’ (females with terga III-VI red and
without the white pubescence on the thorax were
described under the name B. mimeticus var.
insidiosus by Richards, whereas he described
similar workers with the white pubescence under
the name B. mimeticus var. geminatus). The red
pubescence is replaced by black on tergum III or
even on tergum IV for individuals from eastern
Nepal eastwards to Arunachal Pradesh (these
females with more extensive black on terga III-VI
and without the white pubescence on the thorax
were described under the name B. mimeticus var.
gantokiensis by Richards for those with terga I-II
pale yellow, or B. mimeticus s.str. for those
with terga I-II creamy-white; Richards described
similar females with the white pubescence under
the name B. mimeticus var. turneri for those with
terga I-II pale yellow, or B. mimeticus var.
albolateralis for those with terga I-II creamy-
white). Further east, the pubescence of terga III-
VI is entirely black for individuals from northern
Burma (described under the name Megabombus
albopleuralis subsp. atropygus by Tkalcu, 1989).
Material from further south in the highlands of
South East Asia was described under the name B.
montivagus by Smith (1878b). These individuals
usually have the pubescence of the thorax bright
orange, often paler yellow or grey-white at the
sides, with a black band between the wing bases.
The pubescence of tergum I and of the base of
tergum II is bright lemon-yellow. Some specimens
from above Hanoi have gastral terga III-VI black
(described under the name Megabombus mont-
ivagus subsp. quasibreviceps by Tkalci, 1968b), as
for the north Burmese Megabombus albopleuralis
subsp. atropygus, but individuals from most of
South East Asia have red pubescence on the
apical terga, as for the east Himalayan B. haemor-
rhoidalis var. albopleuralis.
Material from further to the north and east, in
China, was described by Smith (1852a) under the
name B. trifasciatus s.str. These individuals also
have pale pubescence on the thorax and on terga
I-II, although for this nominal taxon it is uniformly
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
light yellow. There is a general trend for the black
band between the wings to cover most of the
thoracic dorsum for individuals from western
China and to be reduced or absent for individuals
from near the east coast. The apical terga of all of
these individuals have at least some red pubescence.
B. albopleuralis, B. montivagus and B. trifasci-
atus s.str. have been regarded as three separate
species (e.g. Tkalct, 1968b, 1974b, 1989). No two
of these three principal nominal taxa of the
trifasciatus-group are known to occur together at
any locality without intermediates, so that direct
evidence for barriers to interbreeding is not avail-
able. Tkalct (19685) stated that in comparison
with the most similar taxon, B. albopleuralis, B.
montivagus differs slightly by its longer oculo-
malar distance and by the stronger protrusion of
its clypeus. However, individual differences
between selected specimens may not be easy to
interpret. For instance, the shape of the malar
area at least is subject to large allometric variation
with body size (Sakagami, 1972). Therefore large
random samples of material from a range of
localities across Asia are needed in order to
examine trends in these differences in detail.
Other characters of the morphology (especially of
the sculpturing of the ocello-ocular areas of the
females and of the sizes of the terminal process
and inner hooks of the male volsellae) also vary
considerably within each of these nominal taxa as
well as among them, so I have been unable to
find reliable morphological criteria to distinguish
them. Thus in practice all three nominal taxa are
still recognised by differences in the colour patterns
of their pubescence.
Some individuals from intermediate areas,
particularly from western China and eastern
Burma, appear to show intermediate colour
patterns of the pubescence between those of B.
albopleuralis, B. montivagus and B. trifasciatus
s.str. (Fig. 11). Sakagami (1972) described varia-
tion in the colour pattern of B. trifasciatus s.str.
from western China towards that of B. montivagus,
by replacement of the yellow hairs by black hairs
at the apex of tergum II (Sakagami, 1972: fig.
6bD), and towards that of B. albopleuralis by
replacement of the pale hairs by black hairs on the
thoracic dorsum (Sakagami, 1972: fig. 6aA; a
similar colour pattern was also described under
the name B. ningpoensis subsp. minshanicus by
Bischoff, 1936). Furthermore Richards (1931)
described individuals with a colour pattern (B.
mimeticus var. magrettianus, probably from central
Burma, see Note 7 on the type material) that is
intermediate between B. montivagus and B.
albopleuralis, in that both the thoracic dorsum
and most of tergum II are black-haired. I consider
55
it likely that at least some of these intermediate
colour patterns are evidence of clines between the
more widespread regional colour patterns (Fig.
11). The general pattern is for individuals from the
west of the range in the Himalaya to have most
black hair on the thorax, to have least black hair
on the gaster and to have the darkest wings,
whereas individuals from the east of the range
generally have the least black hair on the thorax,
more on the gaster and have lighter wings. This
interpretation would place all the individuals
within a single interbreeding population as parts
of a single species.
The names B. wilemani and B. maxwelli have
been applied to populations that are isolated from
other B. trifasciatus s.1. by sea and by tropical
lowland forest respectively (Fig. 11). Both
nominal taxa were originally described primarily
using colour pattern characters and without
comparison to any species of the subgenus
Diversobombus. B. wilemani is characterised by
an absence of yellow or orange-brown pubescence
and by its orange-brown to nearly clear (subhyaline)
wings. The tendency of individuals of B. maxwelli
towards nearly uniformly orange pubescence
converges closely with another very distantly-
related species that also occurs in the Cameron
Highlands of Malaysia, B. (Pyrobombus) rufoflavus
Pendlebury (possibly conspecific with the east
Himalayan B. flavescens Smith). The pubescence
of the three workers of B. maxwelli in the BMNH
varies from orange, with dark brown hairs inter-
mixed on tergum II, to orange with most of terga
II-III black. This dark specimen is therefore more
similar in appearance to the lectotype of B.
montivagus. Otherwise it differs by the lighter
brown sclerites and slightly shallower punctures of
the sculpturing of the head. I know of no evidence
from which to suggest that individuals from these
disjunct populations on either the island of Taiwan
or in the Cameron highlands would not interbreed
with those from the more widespread, northern
population if they were to occur together. So B.
maxwelli and even B. wilemani could be parts of a
single species, B. trifasciatus s.1. However, I have
not yet seen any males of B. wilemani from which
to examine their characters.
DISTRIBUTION. B. trifasciatus is a widespread
Oriental species (Map 23, Fig. 11). It is known
from Taiwan (Cockerell, 1911; Skorikov, 1933a;
Frison, 1934; Chiu, 1948; BMNH, PW, SEMK),
Jiangsu (BMNH), Zhejiang (Smith, 1852a; Friese,
1909; Tkalci, 1960; BMNH, SEMK), Fujian
(Pittioni, 1949; BMNH), Guangdong (Sakagami,
1972), Jiangxi (MNHN, PW), Hubei (Tkalci,
1960; Sakagami, 1972; PW), Gansu (Bischoff,
56 PAULH. WILLIAMS
Fig. 11 Distribution of the colour patterns of the ¢rifasciatus-group (for a colour key see Fig. 258). These individuals may all be
considered to be parts of a single species, B. trifasciatus, depending on which species-defining criterion is accepted. The dashed line
shows the 1000m contour above sea level and the solid line shows the 4000m contour. Most records are from material examined,
supplemented from descriptions by Sakagami (1972: Leizhou) and by Tkalcii(1968b: 5 localities in Southeast Asia).
Si
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
LP
58
1936; MNHU), Sichuan (Panfilov, 1957; Tkalci,
1960; Sakagami, 1972; BMNH), Yunnan (Panfilov,
1957; Tkalci, 1968b), Vietnam (Tkalct, 1968),
Laos (Tkalci, 1968b; BMNH), Thailand (Tkalct,
1968b; BMNH, PW, SEMK, ZM), Peninsular
Malaysia (Pendlebury, 1923; BMNH, PW), Burma
(Smith, 18785; Friese, 1918; Richards, 1931;
Frison, 1935; Tkalci, 1968b, 1989; BMNH),
Manipur (BMNH), Meghalaya (Richards, 1931;
Frison, 1933, 1935; BMNH), Arunachal Pradesh
(BMNH), south-eastern Tibet (Wang, 1982),
Sikkim (Friese, 1918; Richards, 1931; Frison,
1935; BMNH, MCSN, MNHU, UM), Darjiling
Bengal (Richards, 1931; BMNH), Nepal (Richards,
1931; Tkalci, 1974b; BMNH, NMS, PW), Uttar
Pradesh (Friese, 1916; Frison, 1935; BMNH,
MNHU, PW, SEMK), Himachal Pradesh (Richards,
1931; Frison, 1933, 1935; BMNH, SEMK), Kash-
mir and Pakistan (Richards, 1931; Frison, 1933).
A disjunct distribution between Burma and the
mountains of Peninsular Malaysia is also known
for B. (Pyrobombus) flavescens (Tkalct, 1974b;
see the comments on B. lemniscatus).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. trifasciatus is recorded from the lower slopes in
the Vale of Kashmir and from the foothills of the
Pir Panjal range, in the more mesic habitats in and
around the lower montane coniferous forests
(Map 24, Fig. 6): 6 queens, 72 workers, 9 males,
from 6 localities 1000-2700 m (AB, BMNH, IZ,
NR, PW, ZM, ZS).
VARIATION WITHIN KASHMIR. Almost all of the
specimens from Kashmir that have been examined
have the sides of the thorax black-haired and terga
III-VI red. There is a small patch of yellow hairs in
the middle of the basal margin of tergum III, often
with a few black hairs intermixed (Figs 275-277).
The only exception to this colour pattern is the
holotype worker of B. mimeticus var. geminatus,
labelled ‘Gulmarg’, which has grey-white hairs on
the sides of the thorax (Fig. 278). Among the
sample I collected at and around this locality
during 1985 and 1986 (5 queens, 29 workers, 5
males), all specimens have the pleura entirely
black-haired.
This species is closely similar in colour pattern
to the local B. haemorrhoidalis, which occur with
it at some localities outside the Vale of Kashmir
(Figs 279-281, see the comments on B. haemor-
rhoidalis). It can usually be recognised by the
presence of a few yellow hairs basally on tergum
III, and for the females in particular, by the
spinose mid basitarsus (Fig. 39) and by the broad
band of punctures in the ocello-ocular area of the
head. There may also be some similarity in colour
pattern to some individuals of B. biroi (Figs 344 &
PAULH. WILLIAMS
345), although the two species have not been
found together. B. biroi has the oculo-malar
distance much shorter and lacks a disto-posterior
spine on the mid basitarsus.
FOOD PLANTS. (Balsaminaceae) Impatiens glandu-
lifera Royle; (Leguminosae) Lupinus sp. [intro-
duced], unidentified pink clover-like legume;
(Compositae) Cirsium wallichii DC.; (Acanth-
aceae) Pteracanthus urticifolius (Kuntze) Bremek.;
(Labiateae) Stachys sericea Wallich ex Benth.
Subgenus ORIENTALIBOMBUS Richards
Orientalibombus Richards, 1929b: 378 (as a sub-
genus of Bombus Latreille). Type species:
Bombus orientalis Smith, 1854: 402 = Bombus
haemorrhoidalis Smith, by original designation.
Orientalobombus Richards; Kruseman, 1952: 102
(as a subgenus of Bombus Latreille). Unjustified
emendation (Art. 33b(i, iii)).
For a general description of both sexes see
Richards (1968).
Bombus (Orientalibombus) haemorrhoidalis
Smith
(Figs 18, 40, 57, 97, 137, 177, 215, 279-281, Maps
25 & 26)
Bombus haemorrhoidalis Smith, 1852a: 43. Types
presumed lost (see Note 1 below) [not seen].
Bombus orientalis Smith, 1854: 402. Lectotype
queen by designation of Richards (1929b:
383) (see Note 2 below), INDIA: West Bengal,
Darjiling (Pearson) (BMNH) [examined].
Change of status to Orientalibombus haemor-
rhoidalis orientalis (Smith) by Tkalcé (1974b).
Syn. n.
Bombus buccinatoris Smith, 1879: 132. Lectotype
worker by designation of Richards (1929b: 384),
INDIA: Himachal Pradesh, Kinnaur (BMNH)
[examined]. Synonymised with Bombus orient-
alis Smith by Bingham (1897). Syn. n.
Bombus assamensis Bingham, 1897: 550. Lectotype
male by designation of Richards (1929b:
384), INDIA: Assam, Margherita (Bingham)
(BMNH) [examined]. Change of status to
Orientalibombus haemorrhoidalis _ subsp.
assamensis (Bingham) by Tkalct (1989). Syn.
n.
Bombus (Orientalibombus) montivolans Richards,
1929b: 382. Holotype male by original designa-
tion [p. 383], LAOS: ‘Tintoe’, 1.xii.1918
(Salvaza) (BMNH) [examined]. Syn. n.
Bombus (Orientalibombus) orientalis var. khasi-
anus Richards, 1929b: 384. Holotype worker
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
by original designation, INDIA: Meghalaya,
Khasi Hills (BMNH) [examined]. Synonymised
with Orientalibombus haemorrhoidalis pectoralis
(Friese) by Tkalct (19745). Syn. n.
Orientalibombus montivolans subsp. semibreviceps
Tkalci, 19686: 10. Holotype worker by original
designation, VIETNAM: Tonkin, Chiem Hoa,
Viii-ix (Fruhstorfer) (Ziirich) [not seen]. Syn. n.
Orientalibombus montivolans subsp. semicolor-
icontrarius Tkalci, 1968b: 10. Holotype worker
by original designation [p. 11], BURMA:
?Karen, ‘Cheba’, 900-1100 m, vi.1888 (Fea)
(Zurich) [not seen]. Syn. n.
Orientalibombus haemorrhoidalis subsp. cinnameus
Tkalct, 1989: 47. Holotype queen by original
designation, BURMA: Arakan Yoma, Mount
Victoria, 1400 m, iv.1938 (Heinrich) (MNHU)
[examined]. Syn. n.
TYPE MATERIAL. Note 1. The abundant material
from the western Himalaya agrees with the original
description of B. haemorrhoidalis, although the
type-locality is stated to be ‘Chusan’ [= Zhoushan,
Zhejiang, eastern China]. I have not seen any
specimens of the subgenus Orientalibombus from
the region of Zhoushan. However, by analogy
with the pattern of geographical variation described
here for B. trifasciatus between Kashmir and
eastern China (which closely resembles the
Kashmir species of Orientalibombus throughout
its known distribution between Kashmir and
Vietnam, see the comments on B. trifasciatus), |
would not expect any species from Zhoushan to
agree in colour pattern with the original descrip-
tion of B. haemorrhoidalis. Therefore the type-
locality cited is considered to be inconsistent with
the description given by Smith.
Smith described B. haemorrhoidalis from syn-
types in the Fortune collection, which was sold in
1872 without any material going to the BMNH or
UM, and all syntypes are believed to be lost (D. B.
Baker, pers. comm.). The Indian specimens,
referred to separately by Smith in the original
publication as being ‘In the British Museum’,
cannot be considered as syntypes (Art. 72b(vi)).
There is a single queen in the BMNH collection
that had been acquired early enough to be one of
these specimens. It bears a label ‘Shang / hai’,
reverse side ‘51 / 26’, and a pale blue (drawer)
label ‘haemorrhoidalis Smith / MSS’, in hand-
writing identical to that of Smith. This accession
number refers to 19 Hymenoptera that had been
obtained in exchange with Baly for duplicates in
March 1851. The area of origin is given in the
accession catalogue as ‘China (Shanghai)’ (close
to Zhoushan), but this has been crossed out and
‘Northern Ind.’ added in identical handwriting.
59
Therefore this specimen is almost certainly of
Indian origin and agrees with the original descrip-
tion and the previous interpretation of the taxon.
It lacks the distal joints of all tarsi except that of
the left mid leg, the apical gastral segments are
curled ventrally and the pubescence has been
abraded from the central area of tergum III. At
some earlier stage, the gaster has also been glued
back into place.
Despite the inconsistency within the original
description of B. haemorrhoidalis, the more recent
authors such as Richards (19296, 1931), Frison
(1933, 1935) and Tkalci (1968b, 1974b, 1989)
have agreed on the identity of Smith’s B. haemor-
rhoidalis. Richards (1929b) and Frison (1933,
1935) continued to apply B. haemorrhoidalis var.
albopleuralis Friese to certain individuals of B.
(Orientalibombus) haemorrhoidalis, although
Tkalct’s (1974b) lectotype of B. haemorrhoidalis
var. albopleuralis belongs to a species of the
subgenus Diversobombus (conspecific with B.
trifasciatus). Nonetheless, I regard the identity of
B. haemorrhoidalis s.str. as not in doubt, so that it
is neither necessary nor valid to designate a
neotype (Art. 75b).
Note 2. Two queens in the BMNH collection
are candidates for having been among Smith’s
syntypes of B. orientalis in 1854. One is simply
labelled ‘India’. The other is labelled ‘Darjeeling’,
reverse side ‘Dr. / Pearson’, with a label ‘orientalis
/ Type Sm.’, in handwriting identical to that of
Smith, and with two identical accession labels ‘60—
15 / E.I.C.’. The accessions catalogue lists this
material as presented in 1860 by the Secretary
of the India Board and as having included
94 Hymenoptera from Java. This might have
explained the type-locality statement ‘East Indies’,
although it has a colour pattern that is characteristic
of the eastern Himalaya and the species is not
known from the East Indies. However, this dona-
tion also contained 83 Hymenoptera from ‘India
+ asiatic Islands’ that is specified to have included
‘Several of the Hymenoptera type specimens
described by Mr Smith’. According to D. B.
Baker (pers. comm.), not all of Smith’s ‘B.M.’
syntypes were deposited in the BMNH at the time
of publication. I agree with Richards (1929b: 383
footnote) that it is reasonable to regard the
Darjiling queen as one of Smith’s syntypes.
Richards’s use of the word ‘type’ in reference to
this specimen is then taken to be a valid lectotype
designation (Art. 74b).
AFFINITIES. The species of the subgenus Oriental-
ibombus share a reduction of the interio-basal
process of the male gonostylus (Fig. 177) and an
extended and twisted apex of the volsella (Fig.
60
137) (synapomorphies). This is a morphologically
divergent group with no obviously close relatives.
The present interpretation is that its affinities lie
with the more northern species of the subgenus
Thoracobombus Dalla Torre rather than with
those of the subgenus Diversobombus (in contrast
to the interpretation in Williams, 1985). Species of
the subgenus Orientalibombus share with the
species of the subgenus Thoracobombus an apical
extension of the part of the volsella beyond the
pair of hooks on its inner margin (?synapomorphy).
The volsella of both is also relatively broad and
short, with the inner hooks placed basally on the
inner apical margin, close to the inner corner.
Apart from B. haemorrhoidalis, the only other
species of the subgenus Orientalibombus from the
Himalaya is B. funerarius Smith, which is known
from Sichuan (Frison, 1935; Panfilov, 1957),
Yunnan (Panfilov, 1957), Burma (Skorikov, 19385;
Tkalct, 1989; BMNH), Darjiling Bengal and
Sikkim (Friese, 1918; Richards, 1929b; Frison,
1935; BMNH, UM), Nepal (PW) and Uttar
Pradesh (BMNH). The males of B. funerarius can
be distinguished by their gonostyli, which are
much shorter than broad in dorsal aspect, whereas
those of B. haemorrhoidalis are nearly square,
with a very much reduced interio-basal process
(Fig. 177) (autapomorphy). The females of B.
funerarius can be distinguished by the tubercles of
the labrum (Richards, 19296), which are unusually
pronounced so that they form a transverse ridge
that is only very narrowly interrupted by the
median furrow. The apical impression of the
clypeus is also deeper and less strongly punctured
than for B. haemorrhoidalis.
There may be just one other species in the
subgenus Orientalibombus, B. braccatus Friese
(synonymised with Bremus metcalfi Frison by
Tkalct, 1987). This is known only from Sichuan
and is similar in colour pattern to the B. trifasciatus
that also occur in Sichuan, in that the thorax and
terga I-II are extensively yellow-haired (Friese,
1905). The gonostylus of the male genitalia (holo-
type male of Bremus metcalfi examined) is inter-
mediate in outline between that of B. funerarius
and that of B. haemorrhoidalis s.1. (see Frison,
1935: fig. 7c). The volsella is unique for species of
this subgenus because the interio-basal hook is
longer than broad and broadens slightly towards
its serrated apex. I have not examined the females
of B. braccatus.
Three principal nominal taxa have been recog-
nised among the remaining haemorrhoidalis-
group of the subgenus Orientalibombus and these
were described under the names B. montivolans,
B. assamensis and B. haemorrhoidalis. They
differ from each other in colour pattern, possibly
PAULH. WILLIAMS
as members of regional groups of Millerian
mimics (see the discussion of the colour patterns
of the Kashmir fauna). These nominal taxa are
very similar in colour pattern to individuals of B.
(Diversobombus) trifasciatus, B. (Alpigeno-
bombus) breviceps Smith and B. (Pyrobombus)
rotundiceps Friese that occur with them (see the
comments on B. trifasciatus, B. kashmirensis
and B. lemniscatus respectively; Sakagami and
Yoshikawa, 1961; Tkalct, 1968b, 1989).
B. haemorrhoidalis is the name that has been
used most recently for all of the Himalayan
nominal taxa in the haemorrhoidalis-group (the
‘haemorrhoidalis-Gruppe’ of Tkalct, 1974b, 1989).
The pubescence of the thoracic dorsum is usually
entirely black. As in the case of B. trifasciatus, the
replacement of black by grey-white pubescence
on the sides of the thorax appears to be most
common among males from lower altitudes in
the eastern Himalaya, at least among BMNH
material. The pubescence of terga I-II is usually
pale yellow. Individuals from the western Himalaya
to as far east as Nepal have terga III-VI pre-
dominantly red. Individuals from eastern Nepal to
as far east as Arunachal Pradesh have black
pubescence on tergum III and even on tergum IV
(females with more extensive black on terga III-
IV and without the white pubescence on the
thorax were described under the name B. orientalis
by Smith for those with terga I-II cream-yellow, or
B. buccinatoris for those with terga I-II more
distinctly lemon-yellow; whereas similar females
with yellow on terga I-II and white pubescence on
the thorax were described under the name B.
orientalis var. khasianus by Richards). Tkalci
(1989) described a queen from Burma under the
name Orientalibombus haemorrhoidalis subsp.
cinnameus that has the pubescence of the thoracic
dorsum black, of the sides of the thorax white, of
terga I-II nearly white, of tergum III predominantly
black, and of the legs orange.
The name B. assamensis has been applied to
some specimens from Assam, which like those
from northern Burma have very little or no red
pubescence on the gaster. These individuals have
the thoracic dorsum black, the sides of the thorax
grey-white and terga I-II pale yellow.
Material from further south in the highlands of
South East Asia was described under the name B.
montivolans by Richards (1929b). These individuals
usually have the pubescence of the thorax bright
orange, though often paler at the sides, with a
black band between the wing bases. The pube-
scence of tergum I and of the base of tergum II is
pale yellow. Some specimens from above Hanoi
have gastral terga III-VI black (described under
the name Orientalibombus montivolans subsp.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
semibreviceps by Tkalci, 1968b) as for B. assam-
ensis, but individuals from most of South East
Asia have red pubescence on the apical terga, as
for the east Himalayan B. orientalis.
B. haemorrhoidalis and B. montivolans have
been regarded as separate species (e.g. Tkalct,
19686, 1974b, 1989). However, males of B.
haemorrhoidalis s.str. differ only slightly from
those of B. assamensis and B. montivolans, for
instance in the shape of the inner hooks of the
volsella (Richards, 1929b: figs 4-7, although
another B. assamensis male (BMNH) has the
volsella as shown in Richards’s fig. 5) and of the
distal margin of the gonostylus. These characters
also vary within each nominal taxon. The three
principal nominal taxa are not known to occur
together at any locality, so that direct evidence for
interbreeding is not available. Yet Tkalct (19685)
described an individual from central Burma under
the name Orientalibombus montivolans subsp.
semicoloricontrarius that could represent an inter-
mediate between B. orientalis var. khasianus of
India and B. montivolans of Laos. This specimen
has the pubescence of both the thoracic dorsum
and of tergum II almost entirely black. Further-
more, some individuals of B. orientalis var.
khasianus from Assam have the red hair of terga
III-VI largely replaced by black, so that they
appear similar to B. assamensis. Therefore I
consider it likely that these intermediate colour
patterns are evidence of clines between the more
widespread regional colour patterns. This inter-
pretation places all of the individuals within a
single interbreeding population as parts of a single
species.
DISTRIBUTION. B. haemorrhoidalis is a Himalayan
and South East Asian species (Map 25). It is
known from Vietnam (Tkalci, 19685), Laos
(Richards, 1929b; Tkalci, 19685; BMNH),
Thailand (Sakagami & Yoshikawa, 1961; Tkalct,
1968b; BMNH, PW, SEMK, ZM), Burma (Frison,
1933, 1935; Skorikov, 1938; Tkalci, 1968),
1989; BMNH, MNHU), Yunnan (Wang, 1987),
Meghalaya (Richards, 1929b; Frison, 1935;
BMNH), Assam (Bingham, 1897; Richards, 1929b;
BMNH), south-eastern Tibet (Wang, 1988),
Arunachal Pradesh and Bhutan (BMNH), Sikkim
(Friese, 1918; Richards, 1929b, 1930; Frison,
1935; BMNH, MCSN, PW, UM), Darjiling Bengal
(Friese, 1918; Richards, 1929b; Frison, 1933,
1935; BMNH), Nepal (Richards, 1929b; Frison,
1935; Tkalct, 1974b; BMNH, NMS, PW), Uttar
Pradesh (Richards, 19295; Frison, 1935; BMNH;
PW, UM, ZM), Himachal Pradesh (Smith, 1879;
Richards, 1929b; Frison, 1933, 1935; BMNH),
Kashmir and Pakistan (Richards, 1929b; Frison,
61
1933). There is also a dubious record from the
Indian plains at Calcutta (Dover, 1922).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. haemorrhoidalis is recorded from the foothills
of the Pir Panjal range (at Patnitop it was found in
humid gullies in the coniferous forest) (Map 26): 8
queens, 102 workers, 1 male, from 3 localities
1000-2000 m (BMNH, NR, PW).
VARIATION WITHIN KASHMIR. All of the females
from Kashmir have the sides of the thorax black-
haired and terga III-VI red, with only a few black
hairs intermixed in the middle of tergum III, near
the base (Figs 279-281). Only the male has grey-
white hairs intermixed on the lower sides of the
thorax (Fig. 281).
This species is closely similar in colour pattern
to the B. trifasciatus that occur with it (Figs 275—
277). It can usually be recognised by the absence
of any yellow hairs at the base of tergum III, and
for the females in particular, by the absence of the
spine on the mid basitarsus (Fig. 40), and by the
broad area without punctures on the ocello-ocular
area of the head (Fig. 215). At the lower altitude
of Muzaffarabad (1000 m, 15.x.1953), B. haemor-
rhoidalis was apparently far more abundant (93/94
workers) than B. trifasciatus (1/94 workers).
Higher up, at Patnitop (2000 m, 6-8.ix.1986), B.
haemorrhoidalis was less abundant (8/31 workers)
than B. trifasciatus (23/31 workers).
FOOD PLANTS. (Balsaminaceae) /mpatiens gland-
ulifera Royle; (Acanthaceae) Pteracanthus urtici-
folius (Kuntze) Bremek.
Subgenus SUBTERRANEOBOMBUS Vogt
Subterraneobombus Vogt, 1911: 62 (as asubgenus
of Bombus Latreille). Type species: Apis
subterranea Linnaeus, 1758: 579 = Bombus
subterraneus (Linnaeus), by subsequent desig-
nation of Frison (1927: 68).
Subterraneibombus Vogt; Skorikov, 1938a: 145.
Unjustified emendation (Art. 33b(i, iii)).
For a general description of both sexes see
Richards (1968).
Bombus (Subterraneobombus) melanurus
Lepeletier
(Figs 58, 59, 98, 99, 138, 139, 178, 179, 282-290,
Maps 27 & 28)
Bombus melanurus Lepeletier, 1836: 469. Lecto-
type queen by designation of Tkalci (1969a:
202), SYRIA: no further data (UM) [examined].
62
Bombus Tschitscherini Radoszkowski, 1862: 591
[by indication of Radoszkowski, 1859: 485].
Holotype queen by monotypy (see Note 1
below), U.S.S.R.: ‘Transoural’, no further data
(ZI) [not seen]. Recombined as Bombus
melanurus tschitscherini Radoszkowski by Vogt
(1909). Syn. n.
Bombus difficillimus Skorikov, 1912b: 609. Lecto-
type queen by designation of Podbolotskaya (in
press), U.S.S.R.: Tadzhikistan S.S.R., Pamir,
3700 m, 15.vi.1909 (Makarjin) (Z1) [examined].
Change of status to Bombus melanurus difficil-
limus Skorikov by Reinig (1934). Provisional
synonym.
Bombus (Subterraneobombus) melanurus subsp.
subdistinctus Richards, 1928b: 333. Holotype
queen by original designation, INDIA: Kashmir,
8000-9000 ft [2400-2700 m], vi.1901 (Nurse)
(BMNH) [examined]. Syn. n.
Bombus (Subterraneobombus) melanurus subsp.
griseofasciatus Reinig, 1930: 83. 36 syntype
queens, 38 syntype workers, 16 syntype males,
U.S.S.R.: Tadzhikistan S.S.R., Pamir, 3700—
4500 m, 5.vii- 29.viii.1928 (ITZ) [not seen].
Synonymised with Bombus melanurus difficil-
limus Skorikov by Reinig (1934). Provisional
synonym.
Bombus (Subterraneobombus) maidli Pittioni,
1939b: 246. LECTOTYPE worker by present
designation (see Note 2 below), MONGOLIA:
north, no further data, 1892 (Leder) (NM)
[examined]. Synonymised with Megabombus
subdistinctus (Richards) by Tkalct (1969a).
Syn. n.
TYPE MATERIAL. Note 1. Radoszkowski described
B. tschitscherini from a single queen sent to him by
E. Eversmann that had a band of black pube-
scence between the wings. If just a single specimen
can be found in the ZI collection that bears labels
with the appropriate data, this would be regarded
as the holotype (Art. 73a(ii)).
Note 2. Pittioni described B. maidli from one
queen, one male and three workers. There is a
single worker in the NM collection with (1) a red
printed label ‘Type’; (2) a printed label ‘N.
Mongolei/ Leder 92’; (3) ‘melanurus / det. Kohl.’;
(4) ‘Subterraneob. / maidli Pitt. [worker] / det.
Pittion’’ in handwniting identical to that of Pittion1;
(5) ‘B. MAIDLI / MAIDLI PITT. / DET.
PITTIONI, 1936’; (6) ‘LECTOTYPE / B. (S.) /
maidli Pitt. / Tkalci det.’ [designation not
published]; (7) a red printed label ‘Lecto- / typus’.
This specimen, which lacks the right mid basitarsus,
is designated as lectotype (Art. 74a).
AFFINITIES. A strongly inwardly-curved apex of
the penis valve head is shared by a large group of
PAULH. WILLIAMS
bumble bees, which in Kashmir includes the
species of the subgenera Subterraneobombus,
Alpigenobombus, Pyrobombus, Sibiricobombus
and Melanobombus (Figs 58-71, 76-85) (synap-
omorphy, secondarily much modified for Bombus
s.str., Figs 72-75). The species of the subgenus Sub-
terraneobombus share a particularly pronounced
broadening of the penis valves in lateral aspect, in
which the ventro-basal angle is produced and
broadened across the penis valve axis, into
a toothed paddle-like form (Figs 98-100) (synap-
omorphy). The male gonostylus has the interio-
basal process broadened antero-posteriorly
(synapomorphy) and, for most species (?second-
arily reduced for B. fedtschenkoi Morawitz),
curved ventrally (Figs 178-180).
The male genitalia of B. melanurus may
be unique for males of the subgenus Subter-
raneobombus, in that the head of the penis valve
lacks a strong posteriorly-directed process from
the dorsal exterior part of its base (Figs 58 & 59)
and the interio-basal process of the gonostylus
does not become broader distally (Figs 178 & 179)
(plesiomorphies within the subgenus Subterrane-
obombus) (see Reinig, 1930: fig. 5). This distin-
guishes B. melanurus from B. fragrans Pallas,
which also occurs in the central Asian steppes, but
with a more western and northern distribution
(distribution maps for both are provided by
Skorikov, 1931: figs 5, 9; see also Panfilov, 1984:
map 192). The head of the penis valve is not
shortened and broadened for either of these
species as it is for B. fedtschenkoi Morawitz,
B. personatus (Fig. 60) and the species of
the subterraneus-group (synapomorphy of B.
fedtschenkoi + B. personatus + subterraneus-
group, see the comments on B. personatus).
Neither is the head of the penis valve strongly
directed ventrally as it is for species of the
subterraneus-group alone (synapomorphy of the
subterraneus-group).
The females of B. melanurus can be distinguished
from B. fragrans, B. amurensis Radoszkowski
from Siberia and from B. fedtschenkoi (which is
closely similar in colour pattern) from the Tien
Shan ranges, by its virtually unpunctured central
area of the clypeus (Skorikov, 1914b). The oculo-
malar distance is much longer than the breadth of
the mandible at its base for B. melanurus, but
scarcely longer than this breadth for B. fragrans or
B. amurensis. I have seen no males of B. amurensis.
B. flaviventris is another species that has
been placed in the subgenus Subterraneobombus
together with these species with a relatively short
oculo-malar distance (Richards, 1930), although
the material in the BMNH collection shows many
character states in common with species of the
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
subgenus Sibiricobombus (see the comments on
B. oberti).
B. melanurus s.str. was originally described
from a queen with a yellow thoracic dorsum that
lacks a band of black hair between the wing bases.
Individuals with this colour pattern, which also
have the wings clouded with brown (infuscated),
occur in Turkey, the Tien Shan ranges and Kashmir.
The pale pubescence varies from brown to lemon-
yellow in colour (e.g. Figs 283 & 288). Richards
(1928b) described the individuals from Kashmir
with this unbanded colour pattern, but with wings
that ‘are, on the average, much less dark than
usual’ (see Tkalci, 1969a: plate facing page 200),
under the name B. melanurus var. subdistinctus.
Tkalct (1969a) concluded that this is a separate
species, noting the presence of a small posteriorly-
directed tooth on the posterior part of the penis
valve head as another distinguishing character.
However this tooth is not present for all individuals
(Figs 58 & 59), or even for both penis valves of
some individuals. Pittioni (1939b) also described
some unbanded individuals from Mongolia that
have the yellow pubescence particularly extensive
on the sides of the thorax under the name B.
maidli. All of these individuals from Kashmir and
Mongolia are otherwise closely similar to other B.
melanurus, so there is no reason to believe that
they are not all parts of the same species.
Individuals of the ‘melanurus-Gruppe’ (Tkalcu,
1974a, 1974b) from further east in Mongolia and
Gansu (e.g. Skorikov, 1931: fig. 9) usually have a
band of black hair between the wing bases. The
name B. tschitscherini was applied to these banded
individuals by Radoszkowski (1859, 1862). The
pale pubescence and the wings are similar in shade
to the darker individuals of B. melanurus s.str.
The male genitalia (Reinig, 1930: fig. 5; Tkalci,
1974a: figs 28-32) are very similar to those of B.
melanurus s.str., but the females have a broader
band of punctures in the oculo-ocellar area, there
are more micropunctures in the middle of the
clypeus, and the pubescence is particularly short
and even.
Some of the individuals from Mongolia and
from further south in the Tibetan massif and the
higher Pamir ranges resemble B. tschitscherini in
that they have a black band between the wings,
although the pale pubescence is lemon-yellow or
cream and the wings are nearly clear (subhyaline).
These individuals were first described under the
name B. difficillimus by Skorikov (resembling
Fig. 282). Reinig apparently redescribed this
taxon under the name B. melanurus subsp.
griseofasciatus, because he subsequently synon-
ymised the two (Reinig, 1934). I have not seen any
males, but the male genitalia (Reinig, 1930: fig. 5,
63
under the name ‘B. mel. griseofasciatus’) appear
to be very similar to those of B. melanurus s.str.
The females have the band of punctures in the
ocello-ocular area intermediate in breadth between
the other two nominal taxa. However, the inner
corners of the labral tubercles are usually more
narrowly acute so that the sides of the longitudinal
median furrow converge throughout its length,
the micropunctures are completely absent from
the middle of the clypeus, the disto-posterior
angle of the mid basitarsus is less strongly produced,
and the pubescence is particularly long and uneven.
There is considerable and possibly continuous
variation among individuals within each of these
nominal taxa in the colour pattern, in the shade of
the pale pubescence and of the wings, and in the
morphological characters. Therefore there is no
good evidence as yet from which to suggest that
these nominal taxa are not just divergent parts of a
single interbreeding population and so parts of a
single species. B. melanurus and B. difficillimus
have not been found together in the Pamir ranges
(Reinig, 1930: fig. 2) or in Kashmir. However,
they do occur at low density on opposite sides of at
least one of the ridges of the Zanskar mountains
(at Chogdo & Nimaling), where B. difficillimus
occupies the higher valley. Of course more
information is needed concerning interbreeding
between them in order to establish their status
more definitely.
Nests from the Pamir were described by Reinig
(1930) and by Bischoff (1931).
DISTRIBUTION. B. melanurus is a widespread, but
primarily central Asian species (Map 27). It is
known from Mongolia (Pittioni, 19396; Skorikov,
1933a; Bischoff, 1936; Tkalct, 1974a; BMNH,
NM, PW), the Altai (BMNH), the Tien Shan
(Skorikov, 1931; Bischoff, 1936; Panfilov, 1957;
BMNH), the Pamir (Skorikov, 1912b, 1931;
Reinig, 1930, 1934; Bischoff, 1931; ZI), the Hindu
Kush (Reinig, 1940; Richards, 1951; Tkalci,
1969a; BMNH), Pakistan (Frison, 1935; BMNH,
PW), Kashmir, Xinjiang [Kunlun Shan] (Morawitz,
1886), Himachal Pradesh (PW), Uttar Pradesh
(BMNH), Nepal (Tkalci, 1974b; BMNH, NMS),
Sikkim (Richards, 19286, 1930; BMNH), Tibet
(Richards, 1928b, 1930; Wang, 1982; BMNH),
Qinghai (Morawitz, 1886; Skorikov, 19125;
Panfilov, 1957; Tkalct, 1961; Wang, 1982; ZS),
Gansu (Bischoff, 1936; MNHU), Inner Mongolia
(Skorikov, 1933a) and Shanxi (Yasumatsu, 1951).
In the west it is also known from western Iran
(BMNH), the Caucasus (Skorikov, 1931), Turkey
(Skorikov, 1931; Reinig, 1971; BMNH), Syria
(UM) and Lebanon (BMNH). Part of this distribu-
tion is mapped by Skorikov (1931: fig. 9). A
64
similar disjunct distribution between the Hindu
Kush ranges on the one hand and the Elburz and
Armenian highlands on the other is shown by B.
keriensis.
MATERIAL EXAMINED FROM KASHMIR. B. melanurus
is widespread in Kashmir, recorded from the Hindu
Raj, Karakoram, Ladakh, Zanskar, Great
Himalaya and Pir Panjal ranges, in montane
coniferous forest, alpine scrub and steppe and in
high, subtropical semi-desert (Map 28, Fig. 6): 85
queens, 186 workers, 135 males, from 29 localities
1800-4800 m (AB, BMNH, FA, MI, NR, PW,
RH, ZM).
VARIATION WITHIN KASHMIR. Queens and workers
are often poorly differentiated in size. Although
the colour pattern of this species is apparently
stable over large areas of its distribution, there is
considerable variation within Kashmir. Queens
from the upper forest on Mt Apharwat (2700—
3400 m, 9/9 queens) have the pale pubescence
brown rather than yellow (Fig. 288). These queens
are all likely to be old, over-wintered individuals
because they have very torn wing margins. Young
queens seen at Gulmarg, all of the queens from
Leh (3500 m, 5/5 queens, Fig. 283), and workers
generally (Figs 284, 287, 289), have the pale
pubescence slightly paler and more sand-yellow to
lemon-yellow.
Queens from Nimaling (4800 m, 3/3 queens)
have the pale pubescence lemon-yellow to almost
cream. These queens, and another from Burzil
Chauki, also have well-defined and broad bands
of black hairs between the wing bases (Fig. 282,
the pattern described under the name B. difficil-
limus, Map 28). The presence of a few black hairs
on the thoracic dorsum, especially just above the
wing bases, is common among workers from
Gulmarg. The black hairs sometimes form
a diffuse band between the wings, which is
particularly obvious for a few, fresh workers from
Gulmarg (3/26 workers, Fig. 287), and for a queen
from Chittakatha Sar (4600 m, Fig. 286), above
the Kishanganga valley. There is also some varia-
tion in the extent of the yellow pubescence on the
sides of the thorax. This yellow pubescence usually
occupies the dorsal half of the area between the
wing bases and the leg bases, but may occupy
nearly all of this area for some workers and even
extends onto the legs in males.
This species is distinctive in its appearance at
most localities. At high altitudes, workers with a
black band between the wing bases (cf. Fig. 282)
may resemble B. personatus (Fig. 292, see the
comments on B. personatus). At a few localities in
the Great Himalaya range, it may be closely
similar in colour pattern to some individuals of B.
PAULH. WILLIAMS
asiaticus that lack red pubescence on the apical
terga (e.g. Fig. 278), but it is recognisable by the
shiny outer surface of the hind tibia and by the
lack of dense, short hair at the base of the hind
basitarsus (see the key, couplet 11).
FOOD PLANTS. Kashmir: (Ranunculaceae) Aconi-
tum hookeri Stapf; (Balsaminaceae) Impatiens
glandulifera Royle; (Leguminosae) Trifolium
repens L.; (Compositae) Cirsium falconeri
(Hook.f.) Petrak, C. wallichii DC.; (Gentiana-
ceae) Swertia petiolata D. Don; (Scrophulariaceae)
Verbascum thapsus L., Digitalis lanata Ehrh.
[introduced], D. purpurea L. [introduced];
(Labiateae) Stachys sericea Wallich ex Benth.,
Thymus linearis Benth. ex Benth.; (Polygonaceae)
Bistorta viviparia (L.) Gray.
Ladakh: (Leguminosae) Medicago falcata L.,
Caragana versicolor (Wallich) Benth.; (Composi-
tae) Echinops cornigerus DC.; (Scrophulariaceae)
Verbascum thapsus L.; (Labiateae) Stachys tibetica
Vatke, Nepeta podostachys Benth.
Bombus (Subterraneobombus) personatus
Smith
(Figs 60, 100, 140, 180, 291-294, Maps 29 & 30)
Bombus personatus Smith, 1879: 132. Lectotype
queen by designation [Art. 74b] of Richards
(1930: 656), INDIA: Himachal Pradesh, Kinnaur
(BMNH) [examined].
Bombus Roborowskyi Morawitz, 1886: 197. Lecto-
type queen by designation of Podbolotskaya (in
press), CHINA: Qinghai, Burhan Budai Shan,
14000 ft [4300 m] (Prshewalski) (ZI) [exam-
ined]. Syn. n.
AFFINITIES. Within the subgenus Subterraneo-
bombus (see the comments on B. melanurus), B.
personatus belongs to a group of species for which
the penis valve head is shortened and broadened
(Fig. 60) and the interio-basal process of the
gonostylus becomes broader distally (Fig. 180)
(synapomorphies of B. fedtschenkoi + B. per-
sonatus + subterraneus-group; this entire group
is the broader, more inclusive ‘subterraneus-
Gruppe’ described from other characters by
Tkalci, 1974a). However, for B. personatus and
B. fedtschenkoi the penis valve head is not as
strongly directed ventrally as for members of the
subterraneus-group (Fig. 100) (synapomorphy of
the subterraneus-group).
B. fedtschenkoi is known from the Tien Shan
region (Skorikov, 19146; Skorikov, 1931; BMNH).
B. fedtschenkoi lacks an apically directed interio-
basal process of the male gonostylus (autapo-
morphy; see Reinig, 1930: fig. 5; Skorikov, 1931:
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
fig. 35). B. personatus, like B. melanurus, has no
large punctures in the middle of the clypeus, but
there are more micropunctures. These punctures
are not nearly as large as those of B. fragrans, B.
amurensis, or B. fedtschenkoi. The oculo-malar
distance of females of B. personatus is relatively
longer than that of even the larger B. melanurus
(see the comments on B. melanurus) and has
fewer punctures.
Morawitz described queens from Qinghai
under the name B. roborowskyi. The lectotype is
closely similar to the lectotype of B. personatus
and is believed to be part of the same species. It is
possible that B. personatus may even be conspeci-
fic with B. difficillimus var. pamirus Skorikov
from the Pamir ranges (Skorikov, 1912b, 1931),
although the oculo-malar distance is apparently
shorter for this nominal taxon (for a discussion of
the identity of this nominal taxon, see also Reinig,
1930, 1934; and the comments on B. oberti). I
have seen no material of B. difficillimus var.
pamirus.
DISTRIBUTION. B. personatus is a Tibetan species
(Map 29). It is known from Kashmir, Himachal
Pradesh (Smith, 1879; BMNH), Tibet (Richards,
1930; Wang, 1982; BMNH), Qinghai (Morawitz,
1886; Panfilov, 1957; Tkalci, 1961; Wang, 1982;
ZI, ZS) and Gansu (Morawitz, 1890).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. personatus is recorded from the Zanskar
ranges, in the high, dry alpine steppe (Map 30):
12 queens, 2 workers, 2 males, from 2 localities
3500-4800 m (BMNH, FA, PW).
I found what could have been the site of a nest
on Nimaling plain (4800 m, 23.ix.1986). A queen
was seen to fly from, and later to return to, a hole
at the base of a dry-stone wall in a shepherd’s
summer hut.
VARIATION WITHIN KASHMIR. The pale pubescence
of the thorax and of terga I-II varies from lemon-
yellow (only on terga I-II) to pale sand-yellow to
almost cream. For queens and males, the pubes-
cence of terga ITI-V has many cream-white hairs
along the posterior margin (Figs 291, 293, 294),
but pale hairs may be very few in this position in
workers (Fig. 292).
This species is usually distinctive in its appear-
ance, although workers (Fig. 292) may resemble
the individuals of B. melanurus with a black band
between the wing bases, which are known to occur
with them (cf. Fig. 282). These dark workers of B.
personatus can be recognised by their pale hairs on
the leg bases and on the sterna.
FOOD PLANTS. (Leguminosae) Caragana versicolor
(Wallich) Benth.; (Labiateae) Stachys tibetica Vatke.
65
Subgenus ALPIGENOBOMBUS Skorikov
Alpigenobombus Skorikov, 1914a: 128. Type
species: Alpigenobombus __ pulcherrimus
Skorikov, 1914a: 128 = Bombus kashmirensis
Friese, by present designation.
Mastrucatobombus Kriiger, 1917: 66 (as a sub-
genus of Bombus Latreille). Type species:
Bombus mastrucatus Gerstaecker, 1869: 326 =
Bombus wurflenii Radoszkowski, by monotypy.
Synonymised with Alpigenobombus Skorikov
by Richards (19285).
Alpigenobombus Skorikov; Frison, 1927: 64 (as a
subgenus of Bombus Latreille).
[Nobilibombus Skorikov, 1933a: 62. Published
without fixation of type species, unavailable
name (Art. 13b).]
[Nobilibombus Bischoff, 1936: 12 (as a sub-
genus of Bombus Latreille). Type species:
Nobilibombus morawitziides Skorikov, 1933a:
62 = Bombus nobilis Friese, by monotypy.
Published as a junior synonym of Alpigenobom-
bus Skorikov, unavailable name (Art. 11e).]
Alpigenibombus Skorikov; Skorikov, 1938b: 1.
Unjustified emendation (Art. 33b(i, iii)).
[Nobilibombus Milliron, 1961: 54 (as a subgenus
of Pyrobombus Dalla Torre). Type species:
Bombus nobilis Friese, 1905: 513 [cited as
Bombus nobilis Skorikov], by original designa-
tion. Published as a junior synonym of Pyro-
bombus Dalla Torre, unavailable name (Art.
lle).]
Nobilibombus Richards, 1968: 222 (as a subgenus
of Bombus Latreille) (see Note 2 below). Type
species: Bombus nobilis Friese, 1905: 513, by
original designation. Syn. n.
Note 1. Skorikov (1914a) originally included ’B.
lefebvrei Lep. (= B. mastrucatus auct.)’ and A.
pulcherrimus Skorikov in his genus Alpigeno-
bombus. Unfortunately he did not clearly and
unambiguously designate a type species (Art. 67c)
and B. alpigenus cannot be the type species of
Alpigenobombus by absolute tautonomy. Frison’s
(1927: 64) subsequent designation of Bombus
alpigenus Morawitz as type species is invalid
because this species was not specifically included
by Skorikov in the original publication (Art.
67g). Skorikov’s original reference to B. lefeb-
vrei Lepeletier is a misidentification, because
Lepeletier’s holotype belongs to a species (oldest
available name B. pomorum (Panzer)) of the
distantly related subgenus Rhodobombus Dalla
Torre (Lgken, 1973), whereas Skorikov cites B.
mastrucatus auct. (conspecific with B. mastrucatus
Gerstaecker, for which the oldest available name
is B. wurflenii Radoszkowski) as a synonym.
66
Designation of Skorikov’s Alpigenobombus lefeb-
vrei as the type species would require the ICZN to
decide the identity of this nominal taxon (Art.
70b). Therefore I designate Alpigenobombus
pulcherrimus Skorikov as the type species of
Alpigenobombus Skorikov because its identity is
clear and this fixation maintains the stability of
usage of Alpigenobombus Skorikov.
Note 2. Skorikov (1933a) originally included
Nobilibombus nobilis (Friese), Nobilibombus
validus (Friese) and Nobilibombus morawitziides
Skorikov in his genus Nobilibombus, but did not
designate a type species. Richards (1968) consi-
dered B. nobilis Friese to be the type species of
Nobilibombus Skorikov by virtual tautonymy,
although this is not sufficient under the present
code, which requires absolute tautonymy (Art.
68e). However, Richards’s reference to B. nobilis
Friese as the type species is sufficient as an original
designation for the subgenus Nobilibombus
Richards (Art. 68b). Unfortunately no putative
syntype of B. nobilis has yet been found that
agrees with the original description in having four
or five teeth on the mandibles (see the comments
on B. pyrosoma; Richards, 1968: 222). Nonethe-
less the identity of B. nobilis is not in doubt, so the
designation of a neotype is neither necessary nor
valid (Art. 75b).
For a general description of both sexes see the
descriptions of the subgenera Alpigenobombus
and Nobilibombus by Richards (1968). These two
groups were originally distinguished primarily on
the basis of the greater oculo-malar distance
for the individuals ascribed to Nobilibombus. I
agree with Bischoff (1936) that placing all the
species together in a single group, by considering
Nobilibombus Richards as a junior subjective
synonym of Alpigenobombus Skorikov, empha-
sises their many similarities (e.g. Ito, 1985;
Williams, 1985).
Bombus (Alpigenobombus) kashmirensis
Friese
(Figs 25, 29, 33, 37, 61-63, 101-103, 141-143,
181-183, 295-310, Maps 31 & 32)
Bombus mastrucatus var. kashmirensis Friese,
1909 [September, see Tkalct, 1974b]: 673
[redescribed by Friese & Wagner, 1910: 47].
Lectotype queen by designation of Tkalcu
(1974b: 327), INDIA: Kashmir, 8000-9000 ft
[2400-2700 m], vi.1901 (Nurse) (MNHU)
[examined].
Bombus mastrucatus var. stramineus Friese, 1909:
673 [redescribed by Friese & Wagner, 1910:
47]. Type worker presumed lost (Tkalcu,
1974b: 327), INDIA: Kashmir, no further data
PAULH. WILLIAMS
[not seen]. Synonymised with Alpigenobombus
kashmirensis (Friese) by Tkalct (1974b) (see
Note 1 below).
Bombus tetrachromus Cockerell, 1909 [November,
see Tkalci, 19746]: 397. Holotype queen by
original designation, PAKISTAN: Baltistan
(BMNH) [examined]. Synonymised with AI-
pigenobombus kashmirensis (Friese) by Tkalca
(1974b).
Alpigenobombus pulcherrimus Skorikov, 1914a:
128. Holotype queen by monotypy (see Note 3
below), INDIA: Kashmir, Zoji La, above 3000
m, 12-15.vi.1912 (Jacobson) (ZI) [not seen].
Synonymised with Bombus tetrachromus
Cockerell by Richards (1930).
[Alpigenobombus (Alpigenobombus) beresovskii
Skorikov, 1922: 156. Published without descrip-
tion or indication, unavailable name (Art.
12a).]
Alpigenobombus (Alpigenobombus) kashmirensis
(Friese); Skorikov, 1922: 156
Bombus (Mastrucatobombus) mastrucatus subsp.
meinertzhageni Richards, 1928b: 335. Holotype
queen by monotypy (see Note 4 below), INDIA:
Kashmir, Ladakh, Chushul, 1925 (Meinertz-
hagen) (BMNH) [examined]. Synonymised
with Alpigenobombus kashmirensis (Friese) by
Tkalci (19745).
[Bombus (Alpigenobombus) tetrachromus vat.
albohirtus Richards, 1930: 637 [examined].
Infrasubspecific (see Note 2 below) (Art. 45g(i1)
(1)), unavailable name (Art. 45e).]
[[Alpigenobombus] beresovskii Skorikov, 1931:
204. Published without description, unavailable
name (Art. 13a).]
Alpigenobombus_ beresovskii Skorikov, 1933b:
248. Lectotype by designation of Podbolotskaya
(in press) [not seen]. Paralectotype queen,
CHINA: Sichuan, ‘Yach Zhou’, 28.i1i—S.iv.
1893 (Potanin) (ZI) [examined]. Syn. n.
NOMENCLATURE. Note 1. Following the Principle
of the First Reviser (Art. 24), B. kashmirensis is
accepted in precedence to B. stramineus by the
action of Skorikov (1922).
Note 2. Richards’s use (1930: 634 etc.) of the
term ‘Subsp.’ elsewhere in his paper is taken to
indicate that infrasubspecific rank is meant for B.
tetrachromus var. albohirtus. No subsequent
author has either adopted the name albohirtus for
a taxon in the species group or regarded it as a
senior homonym, so Richards’s taxon is deemed
to be of infrasubspecific status (Art. 45g(ii)(1)).
TYPE MATERIAL. Note 3. Skorikov’s description of
Alpigenobombus pulcherrimus s.str. specifies
that only a single queen was available. If a single
queen with the appropriate data can be found then
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
this should be regarded as the holotype (Art.
73a(il)).
Note 4. Richards described B. mastrucatus
meinertzhageni from ‘One female from Shusal’ [=
Chushul]. A single queen in the BMNH collection
agrees with the original description and carries the
data quoted. It also bears a label ‘B. tetrachromus
/ var. meinertz- / -hageni, / Type. Richards’, in
handwriting identical to that of Richards. I believe
that this is the single specimen on which the
original description is based and regard it as the
holotype (Art. 73a(ii)).
AFFINITIES. A dorso-ventrally flattened and sickle-
shaped head of the penis valve is shared by a large
group of bumble bees, which in Kashmir includes
the species of the subgenera Alpigenobombus,
Pyrobombus, Sibiricobombus, and Melanobom-
bus (Figs 61—71, 76-85) (synapomorphy, second-
arily much modified for Bombus s.str., Figs
72-75). Females of species of the subgenus Al-
pigenobombus share the development of six teeth
on the distal margin of the mandible (Fig. 33,
?synapomorphy).
Within the subgenus Alpigenobombus, females
of the breviceps-group share a particularly pro-
nounced development of the two pre-apical teeth
of the mandible, which are acute, and the adjacent
incisura is deeper than its own width (?synapo-
morphy of the breviceps-group). This group
includes B. grahami (Frison), B. genalis Friese
and B. breviceps. The males of the Chinese and
east Himalayan (Richards, 1930; Frison, 1935;
Skorikov, 1938b; Wang, 1987, 1988; BMNH, IZ,
PW) B. genalis are not known. The males of B.
breviceps have the gonostylus reduced apically to
a transverse band, although this retains a strongly
produced, interio-basal process and a strongly
produced interio-apical corner (see Frison, 1935:
fig. 5). The recurved hook of the penis valve head
is short, as for B. nobilis, but more broadened.
The males of the Chinese and east Himalayan
(Frison, 1933, 1935; Wang, 1982; BMNH) B.
grahami, which I have not seen, are apparently
intermediate in these character states between B.
nobilis and B. breviceps (Frison, 1935: fig. 6).
The Chinese B. breviceps Smith is very similar
in morphology to the west Chinese and Himalayan
B. dentatus Handlirsch, which has the orange
pubescence of the thoracic dorsum replaced by
black. Individuals with intermediate colour patterns
have been described (see Tkalct, 1968b). There-
fore these nominal taxa are likely to be parts of a
single species that is widespread from China
throughout much of South East Asia and the
Himalaya (records in e.g. Smith, 1852a; Frison,
1935; Bischoff, 1936; Skorikov, 19385; Tkalct,
67
1960, 1968b; Sakagami, 1972; Wang, 1987;
BMNH, MCSN, MNHN, PW, SEMK, UM). This
species, which resembles B. trifasciatus and B.
haemorrhoidalis in colour pattern (see the com-
ments on these species and Fig. 11), may also be
present in the foothills of south-eastern Kashmir
(oldest available name for B. orichalceus Friese,
unconfirmed record from Kashmir by Skorikov,
1933b; there is also a queen in the BMNH from
Dalhousie, just across the border from Kashmir in
Himachal Pradesh).
The females of B. wurflenii, B. kashmirensis
and B. nobilis have the incisura wider than deep
and the two pre-apical teeth of the mandible are
weak and obtuse (Fig. 33, ?plesiomorphic among
species of the subgenus Alpigenobombus). Males of
these species have the interio-apical process of the
volsella reduced (Figs 141-143) (?plesiomorphic
among species of the subgenus Alpigenobombus).
The males of B. nobilis can be distinguished from
those of B. kashmirensis by their shorter gono-
stylus, which has the apical margin concave rather
than straight or convex (Figs 181-183), and by the
shorter recurved hook of the penis valve head
(these character states appear to be intermediate
between B. kashmirensis and B. breviceps). The
females of B. nobilis have the oculo-malar dis-
tance just greater than the breadth of the man-
dible at its base, rather than distinctly shorter
than this breadth as for B. kashmirensis and B.
wurflenii. I have seen material of the banded yellow
B. nobilis Friese from Sichuan (BMNH, PW),
Yunnan, northern Burma, south-eastern Tibet
and Sikkim (BMNH, UM), Nepal (PW) and Uttar
Pradesh (BMNH). B. validus Friese (synony-
mised with Nobilibombus morawitziides Skorikov
by Tkalci, 1987), which has the pale pubescence
of the thorax grey-white rather than yellow, is
otherwise closely similar to B. nobilis and may be
part of the same species. I have seen material of
this nominal taxon from Gansu (ZI), northern
Burma, south-eastern Tibet and Sikkim (BMNH)
and Nepal (PW). Females of two further nominal
taxa from Qinghai, Tibet and Yunnan (B.
xizangensis and B. chayaensis), both very similar
to B. nobilis, have been described by Wang (1979,
1982, 1987, 1988) from slight differences in the
colour pattern (not seen).
Apart from species of the breviceps-group and
B. nobilis, the subgenus Alpigenobombus may
otherwise include just two species that show great
variation in the colour patterns of the pubescence.
B. wurflenii Radoszkowski [the correct original
spelling of B. wurfleini of authors] is a European
species that reaches eastwards as far as the Caucasus
and the Urals (e.g. Reinig & Rasmont, 1988: fig.
1; BMNH). The females of B. wurflenii are closely
68
similar to those of B. kashmirensis in morphology.
The clypeus of B. wurflenii has widely-spaced,
moderate or small punctures throughout, whereas
for B. kashmirensis the punctures are slightly
more widely-spaced in the middle of the clypeus,
so that there is an almost unpunctured, shining
area. However, the males can be separated more
easily because, from the anterior aspect, the eyes
of B. wurflenii (not noticeably enlarged relative to
the female) are at their broadest in their ventral
half, whereas the (enlarged) eyes of B. kashmiren-
sis are broadest in their dorsal half. The genitalia
of males of B. kashmirensis and of B. wurflenii
are closely similar, but the recurved hook of the
penis valve head is more narrowed to a point for
B. wurflenii and the apex of the gonocoxite is
narrower.
Differences in eye size between males of B.
kashmirensis and B. wurflenii are associated with
differences in mate-searching behaviour (see the
introduction on male mate-searching behaviour).
B. wurflenii is widespread in Europe among the
upper montane forests, where males patrol cir-
cuits of scent-marked sites within the forests to
find mates (pers. obs.). B. kashmirensis is wide-
spread in the Himalaya and Tibet among the high
alpine zones. On Mt Apharwat in Kashmir, the
males hover near the peak and watch for potential
mates. Individuals from these populations of B.
wurflenii and B. kashmirensis would be unlikely to
interbreed even if they were to occur on the same
mountain, because they appear to search for
mates in different ways and in different kinds of
habitat (cf. comments on B. rufofasciatus, B.
asiaticus and B. hypnorum).
The queen from Ladakh described by Richards
(19285) under the name B. mastrucatus subsp.
meinertzhageni differs from B. kashmirensis s.str.
principally in that the pale pubescence is entirely
yellow (resembling Fig. 295), rather than both
white and yellow (Fig. 302). The male genitalia of
the bees of this group from Kashmir show limited
variation, although this is more pronounced in the
shape of the gonostylus (Figs 181-183). The apex
of the gonostylus is formed from a thin plate,
which has an irregular margin. The variation in
the extent of this plate is not associated with the
variation in colour pattern and appears to be of
only individual significance. The yellow males are
otherwise closely similar to those of B. kashmiren-
sis S.str., even in the size and shape of their eyes.
All of the workers collected at Gumri near the
Zoji La in the Great Himalaya range are inter-
mediate between the B. mastrucatus subsp.
meinertzhageni of Ladakh and the B. kashmiren-
sis s.str. from the mountains around the Vale of
Kashmir in that they have the pale pubescence of
PAULH. WILLIAMS
the thorax grey-white and that of both abdominal
terga I-II bright yellow (Fig. 301).
Skorikov described females from Sichuan that
have the pale pubescence reduced to a few grey-
cream hairs mixed with the black pubescence on
the anterior thoracic dorsum under the name
Alpigenobombus_ beresovskii. The pattern of
punctures on the female clypeus does not associ-
ate them definitely with either B. wurflenii or B.
kashmirensis, but Skorikov’s description of the
male genitalia (Skorikov, 1933b: 248) agrees more
closely with B. kashmirensis. From the meagre
evidence available at present, I consider that
Alpigenobombus beresovskii is most likely to be
part of the same species as B. kashmirensis.
DISTRIBUTION. B. kashmirensis is primarily a
Tibetan species (Map 31). It is known from Gansu
(Morawitz, 1880; Bischoff, 1936; PW, ZS), Qinghai
(Morawitz, 1886; Bischoff, 1936; Wang, 1982;
PW, ZS), Sichuan (Morawitz, 1890; Skorikov,
1933b; Wang, 1982; ZI), Guangxi [no precise
locality cited] (Wang, 1982), Tibet (Richards,
1930; Wang, 1982; BMNH, PW), Nepal (Tkalci,
1974b; NMS), Uttar Pradesh (BMNH), Himachal
Pradesh (BMNH, PW) and Kashmir. A record
from Mongolia (MNHU) needs to be confirmed.
MATERIAL EXAMINED FROM KASHMIR. B. kash-
mirensis is widespread in Kashmir, recorded from
the Hindu Raj, Ladakh, Zanskar, Great Himalaya
and Pir Panjal ranges, in alpine scrub and steppe
(Map 32, Fig. 6): 34 queens, 172 workers, 67
males, from 15 localities 2400-4800 m (BMNH,
FA, ITZ, MI, MNHN, MNHU, NR, PW, TL; 1
worker photographed in the Sangisfaid valley, C.
A. Chadwell).
A colony was found at the western end of
Nimaling plain (4800 m, 15.vii.1980). The entrance
to the nest was under a 2 m boulder in the derelict
terminal moraine.
VARIATION WITHIN KASHMIR. Individuals from the
Ladakh and Zanskar ranges have the pale pubes-
cence yellow, with only a narrow band of black
hairs basally on tergum III (Figs 295-300). The
pubescence of terga III-VI is orange-red, with at
most only a few hairs of terga V-VI pale-tipped.
Workers from Gumri in the Great Himalaya
range are similar, but the pale hairs of the thorax
are grey-white rather than yellow (terga I-II
yellow, or 1/4 workers with some grey-white hair
antero-laterally on tergum I) and the orange-red
hairs of terga IV-VI are white-tipped (Fig. 301).
Males and females from the Hindu Raj range are
similar, but the pale hairs of tergum I are also
grey-white (tergum II yellow). Material from around
the Kishanganga valley includes individuals with
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
both yellow and white pubescence (4/10 queens,
16/123 workers, 10/43 males), and some individuals
for which the yellow is completely replaced by
white, even on tergum II (6/10 queens, 107/123
workers, 33/43 males). On Mt Apharwat in the Pir
Panjal range, the yellow and white individuals
predominate (9/10 queens, 24/27 workers, 16/17
males). Another worker from Apharwat has very
few pale hairs on the thorax and many black hairs
intermixed on terga I-III (cf. B. beresovskii). The
yellow and white individuals usually have a dis-
tinct black band at the base of tergum III that may
extend over most of tergum III for queens and
males (Figs 302-304). These queens have the
yellow very pale and the white-tipped hairs of
terga III-VI are only orange-red at their bases. In
contrast, the white individuals often have few
black hairs on tergum III, the hairs of terga III-VI
are often completely orange-red and the white
pubescence at the rear of the thorax may be
almost completely replaced by black hairs (Figs
305-310).
The yellow individuals of this species (Figs
295-300) from Ladakh are closely similar in
colour pattern to B. oberti (Figs 392 & 393), which
is known to occur with them. These individuals
may also resemble some of the B. himalayanus
(Figs 242 & 243) and B. ladakhensis (Figs 394 &
396) that occur with them. B. kashmirensis can be
recognised by its 6-toothed mandibles (Fig. 32).
The white and yellow individuals (Figs 302-304)
from the mountains around the Vale of Kashmir
are particularly closely similar in colour pattern to
the B. rufofasciatus (Figs 434436, 438, 439) that
occur with them. B. kashmirensis can be recog-
nised by the longer and white-tipped, red hairs on
terga III-VI, where the hairs of B. rufofasciatus
are short and either red or white. These indivi-
duals may also resemble some B. himalayanus
(Figs 244-252), B. novus males (Figs 260 & 262),
B. lemniscatus (Figs 325 & 326), B. lepidus (Figs
330, 331, 333, 334) and B. biroi (Figs 338 & 339).
The white individuals of this species (Figs 305—
310) from the mountains around the Kishanganga
valley and the Vale of Kashmir are particularly
closely similar in colour pattern to some of the B.
biroi (Figs 341 & 342) that occur with them
at some localities. These individuals may also
resemble some B. avinoviellus (Figs 238-241), B.
himalayanus (Figs 250-252), B. tunicatus (Figs
350, 353, 355), B. asiaticus (Figs 386 & 387) and B.
keriensis (Figs 408 & 409). B. kashmirensis is
easily recognised by its 6-toothed mandibles (Fig.
32):
FOOD PLANTS. Kashmir: (Ranunculaceae) Aconi-
tum laeve Royle, A. heterophyllum Wallich ex
69
Royle, A. hookeri Stapf, A. violaceum Jacquem.
ex Stapf; (Compositae) Lactuca lessertiana (DC.)
C. B. Clarke, unidentified dandelion-like compo-
sites; (Gentianaceae) Swertia petiolata D. Don;
(Scropulariaceae) Scrophularia pauciflora Benth..,
Digitalis lanata Ehrh. [introduced], Pedicularis
oederi Vahl, P. pectinata Wallich ex Benth., P.
punctata Decne.
Ladakh: (Leguminosae) Caragana versicolor
(Wallich) ex Benth.; (Labiateae) Stachys tibetica
Vatke.
Subgenus PYROBOMBUS Dalla Torre
Pyrobombus Dalla Torre, 1880: 40 (as a subgenus
of Bombus Latreille). Type species: Apis
hypnorum Linnaeus, 1758: 579 = Bombus
hypnorum (Linnaeus), by monotypy.
Poecilobombus Dalla Torre, 1882: 23 (as a sub-
genus of Bombus Latreille). Type species:
Bombus sitkensis Nylander, 1848: 235, by sub-
sequent designation of Sandhouse (1943: 589).
Synonymised with Pyrobombus Dalla Torre by
Milliron (1961).
[Pyrrhobombus Dalla Torre; Dalla Torre, 1882:
28 (as a subgenus of Bombus Latreille). In-
correct subsequent spelling, unavailable name
(Art. 33c).]
Pyrrhobombus Dalla Torre; Dalla Torre, 1896:
503 (as a subgenus of Bombus Latreille). Un-
justified emendation (Art. 33b(i, iii)).
Pratobombus Vogt, 1911: 49 (as a subgenus of
Bombus Latreille). Type species: Apis pra-
torum Linnaeus, 1761: 424 = Bombus pratorum
(Linnaeus), by subsequent designation of Frison
(1927: 67). Synonymised with Pyrobombus
Dalla Torre by Milliron (1961).
Anodontobombus Kriiger, 1917: 61, 65 (as a
subgenus of Bombus Latreille). Type species:
Apis hypnorum Linnaeus, 1758: 579 = Bombus
hypnorum (Linnaeus), by present designation
(see Note below). Synonymised with Pyro-
bombus Dalla Torre by Milliron (1961).
Uncobombus Vogt in Kriger, 1917: 65 (as a
subgenus of Bombus Latreille). Type species:
Apis hypnorum Linnaeus, 1758: 579 = Bombus
hypnorum (Linnaeus), by present designation
(see Note below). Synonymised with Pyro-
bombus Dalla Torre by Milliron (1961).
Lapponicobombus Quilis-Pérez, 1927: 19 (as a
subgenus of Bombus Latreille). Type species:
Apis lapponica Fabricius, 1793: 318 = Bombus
lapponicus (Fabricius), by subsequent desig-
nation of Milliron (1961). Synonymised with
Pyrobombus Dalla Torre by Milliron (1961).
[Hypnorubombus Quilis-Pérez, 1927: 19 (as a
70
subgenus of Bombus Latreille). Incorrect original
spelling (Art. 32c(iv)), unavailable name (Art.
19a).]
[Laponicobombus Quilis-Pérez, 1927: 63 (as a
subgenus of Bombus Latreille). Incorrect original
spelling (Art. 32c(iv)), unavailable name (Art.
19a).]
Hypnorobombus Quilis-Pérez, 1927: 97 (as a
subgenus of Bombus Latreille). Type species:
Apis hypnorum Linnaeus, 1758: 579 = Bombus
hypnorum (Linnaeus), by monotypy. Synony-
mised with Pyrobombus Dalla Torre by Milliron
(1961).
Pratibombus Vogt; Skorikov, 1938a: 146. Unjus-
tified emendation (Art. 33b(i, iii)).
Note. The names Anodontobombus Kriger
and Uncobombus Vogt were originally pro-
posed for a section of the genus Bombus and a
group of subgenera respectively, but they must
both be treated as subgeneric names (Art. 10e).
Milliron (1961: 53) considered Anodontobom-
bus Kriiger to be synonymous with his concept
of the subgenus Pyrobombus Dalla Torre (with-
in his genus Pyrobombus Dalla Torre). Milliron
also considered Uncobombus Vogt to ‘corres-
pond’ to Pyrobombus Dalla Torre. The type
species of both subgenera are designated as
Apis hypnorum Linnaeus, in common with
Pyrobombus Dalla Torre, in order to stabilise
an unnecessarily complex nomenclature.
For a general description of both sexes see
Richards (1968). His descriptions include many
exceptions for the characters of B. festivus Smith,
which was subsequently placed in a separate
subgenus Festivobombus by Tkalci (1972).
Bombus (Pyrobombus) hypnorum
(Linnaeus)
(Figs 64, 104, 144, 184, 311-316, Maps 33 & 34)
Apis hypnorum Linnaeus, 1758: 579. Lectotype
queen by designation of Day (1979: 64),
SWEDEN: no further data (LSL) [examined].
[Bombus hypcorum (Linnaeus); Walckenaer,
1802: 148. Incorrect subsequent spelling, un-
available name (Art. 33c).]
Bombus (Pratobombus) hypnorum var. bryorum
Richards, 1930: 650 (see Note below). Holo-
type queen [not the male] by original designa-
tion [p. 651], INDIA: Kashmir, 8000-9000 ft
[2400-2700 mJ], vi.1901 (Nurse) (BMNH)
[examined]. Syn. n.
Bombus fletcheri Richards, 1934: 90. Holotype
queen by original designation, INDIA: Kashmir,
Gulmarg, 7.vi.1931 (Fletcher) (BMNH) [exam-
ined]. Syn. n.
PAULH. WILLIAMS
Pyrobombus (Pyrobombus) bryorum (Richards);
Tkalct, 1974b: 328.
NOMENCLATURE. Note. Richards’s use (1930: 634
etc.) of the term ‘Subsp.’ elsewhere in his paper
could be taken to indicate that infrasubspecific
rank is meant for B. hypnorum var. bryorum,
although his comment ‘this Indian form’ does
associate the nominal taxon with a particular
geographical area (Art. 45f(ii)). However, this
has subsequently been treated as a name in the
species group by Tkalci (1974b), so Richards’s
taxon is therefore deemed to be of subspecific
status (Art. 45g(ii)(1)).
AFFINITIES. Species of the subgenus Pyrobombus
share a reduction and ventral displacement of the
interio-basal process of the male gonostylus, with
an apically-directed extension that forms a longi-
tudinal ridge, just ventral and parallel to the inner
margin of the gonostylus (Figs 184-191) (synapo-
morphy). An intermediate form of this structure
may be seen in the gonostylus of B. (Festivobom-
bus) festivus Smith, which has the interio-basal
process only partially reduced and rotated ven-
trally, but with a weakly sclerotised fold running
apically, below the inner margin of the gonosty-
lus. The subgenus Pyrobombus has more species
(at least 42) than any other subgenus of bumble
bees and many of these species are morphologi-
cally closely similar. Consequently ideas of the
relationships among the species are at present
particularly tentative.
Himalayan B. hypnorum have often been con-
fused with the workers of B. (Festivobombus)
festivus, which have a similar colour pattern. B.
festivus is common in southern China and the
eastern Himalaya to as far west as Himachal
Pradesh (BMNH, PW, UM). It may also be
present in the foothills of south-eastern Kashmir
(unconfirmed record by Skorikov, 1933b). The
workers of B. festivus have the disto-posterior
angle of the hind tibia spinosely produced, where-
as for B. hypnorum this process is no longer than
broad. The pubescence of B. festivus is generally
shorter and more even, with a particularly dense
covering of short, branched hairs on the head,
posterior to the eyes. The male genitalia of B.
festivus show similarities to those of B. hypnorum,
but have a distinct interio-basal process of the
gonostylus (see Tkalci, 1974b: fig. 114), lack a
well-defined ventro-basal angle of the penis valve
(see the comments on B. asiaticus), and have the
volsella strongly constricted just before its apex.
Within the subgenus Pyrobombus, the rare B.
abnormis (Tkalct) from Sikkim (Tkalcu, 19684;
BMNH) also resembles B. hypnorum in colour
pattern, but with fewer pale hairs on the apical
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
terga. The ocello-ocular area is, however, almost
completely and densely covered with fine punc-
tures for both sexes of B. abnormis. The male
genitalia differ from those of B. hypnorum in that
the volsella is weakly, but distinctly, constricted
before its apex (more like B. festivus), of which
the outer corner is acute, and the recurved hook of
the penis valve head is shorter than for B.
hypnorum (more like B. festivus). The interio-
apical angle of the gonocoxite of B. abnormis is
broadly rounded, rather than acutely pointed, and
the shape of the gonostylus differs (see Tkalci,
1968a: fig. 52-56).
B. hypnorum may be one of the most plesio-
morphic species of the subgenus Pyrobombus, at
least insofar as the the gonostylus retains a well-
defined inner basal notch (Fig. 184) and the
recurved hook of the penis valve head remains
broad but narrowed to a rounded point (Fig. 64).
At present I am not aware of any synapomorphic
characters from which to recognise a mono-
phyletic ‘hypnorum-group’ (cf. the ‘hypnorum-
Gruppe’ of Tkalci, 19682, 1989). Many other Old
World species of the subgenus Pyrobombus have
the recurved hook broadened apically and more
spatula-shaped (e.g. the pratorum-group, see the
comments on B. biroi, Figs 70 & 71), or more or
less shortened (e.g. the parthenius-group, see the
comments on B. lepidus, Figs 67 & 68).
B. hypnorum is closely similar to the North
American B. perplexus Cresson. In the Old
World, its closest relative may be B. haematurus
Kriechbaumer, which is known from south-
eastern Europe, Turkey and the Caspian coast of
Iran (D. B. Baker, pers. comm.; BMNH, PW).
Females of B. haematurus have the posterior half
of the thoracic dorsum and tergum I black-haired
and the post ocello-ocular area is more smooth
and more finely punctured than for B. hypnorum.
The distal margin of the male gonocoxite is more
nearly at a right angle to its longitudinal axis for B.
haematurus (se Kriiger, 1943: p. 540 figs 1-12)
and the gonostylus is shorter than for B. hyp-
norum (see Krier, 1943: p. 538 figs 1-7).
B. hypnorum appears to be widespread between
northern Europe and Kashmir. Both females and
males differ slightly in colour pattern between the
two regions (by extension of the area of pale
pubescence onto at least tergum II among indivi-
duals from Asia; see Reinig, 1939: fig. 8). Indivi-
duals from the Himalaya have been regarded as a
separate species, B. bryorum, by Tkalct (1974b),
although I can find no reliable morphological
differences. The queen described under the name
B. fletcheri by Richards is closely similar to other
faded queens of B. hypnorum from Gulmarg.
Individuals of all of these nominal taxa occupy
71
similar habitats in the upper montane coniferous
forest. The males search for mates by patrolling
the forest in a similar way and I know of no
evidence to suggest that individuals from Kashmir
would not interbreed with those from Europe if
they were to occur on the same mountain (cf.
comments on B. asiaticus, B. rufofasciatus and B.
kashmirensis). Therefore they are all considered
to be parts of the same species.
DISTRIBUTION. B. hypnorum is widespread in the
Old World (Map 33). It is known from Europe
(but not Britain e.g. Reinig, 1939; Lgken, 1973)
and eastwards across the U.S.S.R. (Reinig, 1939;
Panfilov, 1984; BMNH) to as far as Kamchatka
(Bischoff, 1930; Reinig, 1939; BMNH) and
Hokkaido in Japan (Tkalct, 1965; Sakagami &
Ishikawa, 1969; BMNH, PW). In central Asia it
reaches southwards to the Caucasus and the Altai
(Panfilov, 1984) and Mongolia (Skorikov, 1931,
1933a; Reinig, 1939; PW); and in the south-east it
is recorded from North Korea (Kim & Ito, 1987),
Gansu (Bischoff, 1936; Reinig, 1939), Sichuan
(Morawitz, 1890), Yunnan (Wang, 1987), Burma
and Arunachal Pradesh (BMNH), Tibet (Richards,
1930; Wang, 1987), Sikkim (Friese, 1918), Nepal
(Tkalct, 19746; BMNH, NMS) and Kashmir.
Part of this distribution is mapped by Reinig
(1939: fig. 8) and also by Panfilov (1984: map 186).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. hypnorum is recorded from the Hindu Raj
range, from the southern side of the Great
Himalaya range and from the Pir Panjal range, in
the upper montane coniferous forest (Map 34,
Fig. 6): 6 queens, 43 workers, 22 males, from 5
localities 2400-3700 m (BMNH, NR, PW).
VARIATION WITHIN KASHMIR. The colour pattern
of the females resembles that of individuals from
northern Asia, in that pale brown hairs often
cover most of tergum II (Figs 311, 314, 315). Most
workers have some pale hairs intermixed on
tergum III as well, and extreme workers from Mt
Apharwat have just a few black hairs intermixed
on terga III-IV (Fig. 312, 5/26 workers). On the
thorax the pale brown pubescence extends down
to the leg bases and femora in most individuals.
The males are much lighter in colour pattern
than those from Europe (but see the discussion of
the colour patterns of the Kashmir fauna). Their
pubescence is almost uniformly lemon yellow,
with the exception of a few black hairs intermixed
on the head and on terga IV-V, and of white hairs
on terga V-VII (Figs 313 & 316).
Females of this species are distinctive in their
appearance. The yellow males with white hairs on
the apical terga (Figs 313 & 316) may be similar in
2
colour pattern to the males of B. lucorum (Figs
365 & 370), which occur with them at some
localities. B. hypnorum males can be recognised
by their smaller size and by the inwardly hooked
apices of the penis valves (Fig. 64).
Host PLANTS. (Ranunculaceae) Cimicifuga foetida
L., Aconitum laeve Royle; (Balsaminaceae)
Impatiens glandulifera Royle, unidentified small
yellow balsam; (Sambucaceae) Lonicera quinque-
locularis Hardw.; (Compositae) Cirsium falconeri
(Hook.f.) Petrak, C. wallichii DC.; (Scrophulari-
aceae) Scrophularia pauciflora Benth., Digitalis
lanata Ehrh. [introduced]; (Labiateae) Prunella
vulgaris L., Salvia hians Royle ex Benth.
Bombus (Pyrobombus) subtypicus
(Skorikov), comb. n.
(Figs 41, 65, 66, 105, 106, 145, 146, 185, 186, 216,
317-324, Maps 35 & 36)
Bombus leucopygus Morawitz in Fedtschenko,
1875: 3. Lectotype female by designation of
Skorikov (1914c: 293) (see Note 2 below),
U.S.S.R.: Tadzhikistan $.S.R. or Uzbekistan
S.S.R. (ZI) [not seen]. Junior primary homo-
nym of Bombus leucopygus Illiger, 1806 [= B.
hypnorum (Linnaeus)].
[Pratobombus leucopygos Skorikov, 1914c: 293.
Incorrect subsequent spelling, unavailable name
(Art. 33c).]
Pratobombus leucopygos var. subtypicus Skorikov,
1914c: 294. Lectotype by designation of Pod-
bolotskaya (in press) [not seen]. Paralectotype
queen, U.S.S.R.: ?Uzbekistan S.S.R., ‘Bukan-
Khapa’, 27.v.1911 (Topbek) (ZI) [examined].
Bombus leucurus Bischoff & Hedicke, 1931: 391.
Unnecessary replacement name for B. leucopy-
gus Morawitz. Synonymised with Pyrobombus
subtypicus (Skorikov) by Tkalcti (1968a).
Pyrobombus (Pyrobombus) subtypicus (Skorikov);
Tkalct,, 1968a: 27 (see Note 1 below).
Pyrobombus (Pyrobombus) kohistanensis Tkalci,
1989: 49. Holotype queen by original designa-
tion, PAKISTAN: Indus Kohistan, Kagan
valley, Naran, 2400-3000 m, 3—13.vi.1977 (de
Freina) (ZS) [examined]. Syn. n.
NOMENCLATURE. Note 1. Bombus subtypicus
(Skorikov) replaces Bombus leucopygus Morawitz
because it is the oldest available name (Art. 60b)
(Tkalcti, 1968a, 1969a).
TYPE MATERIAL. Note 2. Skorikov (1914c) des-
cribed the variation among Morawitz’s syntypes
of B. leucopygus in the ZMMU. He stated that he
considered that the first specimen bearing a large
label with the name of the species was the type.
PAULH. WILLIAMS
This use of the word type in reference to a single
specimen is taken to be a valid lectotype designa-
tion (Art. 74b).
AFFINITIES. Within the subgenus Pyrobombus
(see the comments on B. hypnorum), close rela-
tives of B. subtypicus, as judged from the similar-
ity in the form of the male genitalia, may include
both B. haematurus and B. lemniscatus (see the
comments on B. hypnorum and B. lemniscatus).
However, for B. subtypicus the recurved hook of
the penis valve head is more acutely pointed (Figs
65 & 66) than for either of the other two species
(e.g. Fig. 67) (see Kriiger, 1943: p. 541 figs 1-11).
Unlike B. haematurus, B. subtypicus has virtually
no interio-basal notch on the gonostylus above the
interio-apical corner of the gonocoxite near their
point of contact (Figs 185 & 186). The females of
B. haematurus can be distinguished by their long
pubescence and by their many close and regular
fine punctures in the post ocello-ocular area of the
head.
Skorikov (1914c) described the local variation
in the colour pattern of the pubescence among
Morawitz’s series of B. leucopygus Morawitz from
‘Turkestan’. Some of these individuals had the
black band between the wing bases partially or
completely replaced by yellow pubescence.
Recently Tkalci (1989) described a single
queen under the name Pyrobombus kohistanensis
from across the Pakistan border in the Indus-
Kohistan range that has a colour pattern similar to
the banded grey- white individuals of B. avinoviel-
lus. This specimen (Tkalcu, 1989: figs 14, 18, 22) is
closely similar in morphology to B. subtypicus and
is likely to be part of the same species. It differs
from individuals of B. subtypicus from Kashmir
that have a black band between the wings prin-
cipally in that the pale pubescence is nearly white,
rather than yellow. However, a single queen from
nearby on the Kashmir side of the border at Astor
is intermediate in these characters insofar as it has
both black hairs intermixed between the wing
bases and the pale pubescence is a particularly
pale yellow. More specimens, especially of the
males, are needed in order to establish definitively
the relationships of this nominal taxon.
DISTRIBUTION. B. subtypicus is a central Asian
species (Map 35). It is known from the Tien Shan
(Morawitz, 1880; Skorikov, 1914c, 1931; Kriiger,
1943; BMNH, Z1), the Pamir (Reinig 1930, 1934;
Skorikov, 1931; BMNH), the Hindu Kush (Reinig,
1934, 1940; Richards, 1951; Tkalct, 1969a;
BMNH, PW), Pakistan (Tkalci, 1989; BMNH,
PW, ZS) and Kashmir. The record from Gansu
(Bischoff, 1936) is based on a misidentification
(Tkalci, 19682).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. subtypicus is recorded from the Hindu Raj
range and from the Great Himalaya range, in dry
subalpine scrub (Map 36): 2 queens, 36 workers,
108 males, from 9 localities 2700-3700 m (BMNH,
NR, PW).
A colony was found near Dras (3200 m, 10. viii.
1986). The nest entrance was at the base of a dry-
stone wall.
VARIATION WITHIN KASHMIR. Material from
Kashmir is relatively homogeneous in colour
pattern, at least insofar as all specimens have the
pubescence of the apical terga orange-red rather
than white (Figs 317-324). Females from the
Hindu Raj range and from Matayan and Dras in
the Great Himalaya range have yellow hairs
on the head and to a more variable extent on
terga I-II (Figs 319, 320, 323). These individuals
have only a few black hairs on the thoracic
dorsum.
The single queen from Astor has the head, the
ventral parts of the thorax and terga I-II black
(Fig. 322). Black hairs are also more numerous
between the wing bases, where they form an
indistinct band. The yellow pubescence of this
individual is particularly pale.
The single worker from Lal Pani has a broad,
black band between the wing bases (Fig. 317). The
hairs of its head are black but those of terga I-II
are yellow. Some of the males from the same
locality also have a black band between the wing
bases (Fig. 318, 10/25 males).
The individuals of this species that lack a black
band between the wing bases (Figs 319-324) are
closely similar to some B. biroi (Figs 335-337),
although they are not known to occur together.
They are also similar to some of the B. avinoviel-
lus (Fig. 232), B. marussinus (Figs 254-257), B.
branickii (Fig. 265), B. asiaticus (Figs 371-376)
and B. semenovianus (Figs 397-402) that may
occur with them at some localities. Unbanded B.
subtypicus can be recognised by the characters
given in the key (couplets 19 & 20). The indivi-
duals with the black thoracic band (Figs 317 &
318) resemble some of the B. avinoviellus (Figs
233-235), B. asiaticus (Figs 377 & 378) and B.
keriensis (Fig. 414416) that occur with them. B.
subtypicus can be recognised by its small size, by
its shiny outer surface to the hind tibia, and by the
small number of fine punctures in the ocello-
ocular area of the head (Fig. 216).
FOOD PLANTS. (Leguminosae) Melilotus officinalis
(L.) Pallas, Trifolium repens L.; (Scrophulariaceae)
Verbascum thapsus L.; (Labiateae) Nepeta podo-
stachys Benth., Prunella vulgaris L.
73
Bombus (Pyrobombus) lemniscatus Skorikov
(Figs 15, 67, 107, 147, 187, 212, 218, 223, 325, 326,
Maps 37 & 38)
Bombus lemniscatus Skorikov, 19126: 606. Holo-
type queen by monotypy, CHINA: Qinghai,
northern slopes of the Xining mountains, 30.v.
1890 (Grum-Grzhimailo) (ZI) [examined].
Bombus nursei var. flavopilosus Friese, 1918: 84.
LECTOTYPE queen by present designation
(see the Note below), no data (Bingham)
(MNHU) [examined]. Syn. n.
Bombus (Lapidariobombus) peralpinus Richards,
1930: 646. Holotype queen [not a worker] by
original designation [p. 647], CHINA: Xizang
[= Tibet], Rongshar valley, Tasam, 12000 ft
[3700 ml], 20.vi.1924 (Hingston) (BMNH)
[examined]. Synonymised with Pyrobombus
lemniscatus (Skorikov) by Tkalct (19746).
TYPE MATERIAL. Note. Tkalct' (1968) described
Pyrobombus mirus from a queen that is the
only one of Friese’s three syntype queens of B.
nursei var. flavopilosus to bear one of the red
‘Type’ labels. Tkalct (1968a: 39) had noted that
there are some discrepancies between this indivi-
dual and Friese’s original description, but con-
trary to Tkalct’s conclusion (‘kann infolge der
Nachtiibereinstimmung mit der Urbeschreibung
nicht die Lectotype dieses Taxons [B. nursei var.
flavopilosus| darstellen’), this does not prevent
this specimen from being a syntype (Art. 73b), so
it remains a candidate for designation as the
lectotype of B. nursei var. flavopilosus. However,
another one of Friese’s syntypes is designated
here as lectotype of B. nursei var. flavopilosus in
order to maintain the current usage of the name B.
mirus (Tkalca). A queen in the MNHU collection
agrees with the original description of B. nursei
var. flavopilosus and carries a label (1) ‘Coll.
Bingham’; (2) ‘B. nursei / flavopilosus / [female]
1914 Friese det. / Fr.’; (3) a red printed label
‘Paratypus’; (4) ‘LECTOTYPUS/ B. nursei var. /
flavopilosus / Friese Tkalct det [female]’ [designa-
tion not published]. This specimen, which is badly
damaged with Anthrenus holes and lacks the left
front tarsi and the right mid and hind tarsi, is
designated as lectotype (Art. 74a).
AFFINITIES. Within the subgenus Pyrobombus
(see the comments on B. hypnorum), B. lemnis-
catus is likely to be related to the parthenius-group
of species. All other species of this group, with the
possible exception of B. mirus (Tkalct), have the
ventro-basal angle of the penis valve displaced
closer to the penis valve apex than to its base (Figs
107-109), with the shoulder between this angle
and the base of the penis valve head strongly
74
curved, and the head of the penis valve broadened
basally (Figs 67-69, 107-109) (synapomorphies).
This group includes Tkalcu’s (1974b, 1989)
‘parthenius-Gruppe’ together with the additional
species B. lepidus, B. infirmus (Tkalca) and
probably B. lemniscatus and B. mirus (see the
comments on B. lepidus).
There are two more-distantly related species of
the subgenus Pyrobombus that occur at lower
altitudes in the eastern Himalaya, to at least as far
west as Uttar Pradesh (BMNH): B. rotundiceps
Friese and B. flavescens. It is possible that B.
rotundiceps may also be present in Kashmir
(unconfirmed record by Skorikov, 1933b). This
species closely resembles the local B. trifasciatus
and B. haemorrhoidalis in the colour pattern of
the pubescence (see the comments on those
species). Females of both B. rotundiceps and B.
flavescens from Uttar Pradesh have the pubes-
cence of the thoracic dorsum entirely black. The
legs are extensively bright orange-brown for B.
flavescens but black for B. rotundiceps.
Another Himalayan species, which resembles
B. lemniscatus more closely and which may belong
to the parthenius-group in its broadest sense, is B.
mirus. Females of B. mirus have a colour pattern
similar to some of the darker, white B. biroi from
Kashmir (Fig. 341), but with the pubescence of
tergum III predominantly black. They can be
distinguished from most species by the profuse
punctures that cover more than half of the ocello-
ocular area of the head (though not as extensive
as for B. abnormis, see the comments on B.
hypnorum). Females of B. mirus are known from
the alpine zone of Nepal (BMNH, PW), Sikkim
(Tkalci, 19684; BMNH, MNHU) and Tibet
(Wang, 1982). Wang also described a male from
Zayii in Tibet (not seen). This male is apparently
similar in its genitalia (Wang, 1982: fig. 32) and in
its white and yellow colour pattern to another
male from Tibet described by Friese (1913) under
the name B. pratorum var. tibetanus (junior
secondary homonym in Bombus of Apathus
tibetanus Morawitz). The genitalia of both appear
to differ from those of B. lemniscatus primarily in
their strongly concave interior margin of the
gonostylus.
B. lemniscatus (redescribed independently
under the name B. nursei var. flavopilosus by
Friese, see the Note above, and under the name
B. peralpinus by Richards) is closely similar to B.
lepidus not only in morphology, but also in the
colour pattern of the pubescence and in its dis-
tribution. Skorikov (19125) distinguished females
of B. lemniscatus by their longer pubescence,
slightly longer oculo-malar distance, more sharply
defined band of black hairs between the wing
PAULH. WILLIAMS
bases and by their generally more extensive black
pubescence. Tkalct has listed several other subtle
differences between the females (Tkalci, 1989:
table 3) and noted that the scutum, on the thoracic
dorsum, is punctured almost throughout for B.
lemniscatus (Fig. 223) but broadly unpunctured in
the middle for B. lepidus (Fig. 224). A single male
in the BMNH collection from ‘Rata, 11,000 ft.’ [=
3350 m], Uttar Pradesh, shows similar hair length,
colour pattern and sculpture of the scutum to
females of B. lemniscatus. Although the genitalia
of this male are closely similar to those of B.
lepidus, they do differ in that, unlike B. lepidus
from Kashmir, the recurved head of the penis
valve is slightly narrower (Fig. 67), the ventro-
basal angle of the penis valve is just closer to the
base of the penis valve shaft than to the penis valve
head (Fig. 107), and the interio-apical process of
the volsella is broader and extends without a
concave angle to the exterio-apical corner of the
volsella (Fig. 147). All of the available males of B.
lepidus can be distinguished by their extensively
yellow colour pattern of the pubescence, which
has few white or black hairs (see the discussion
of convergent colour patterns). More material,
especially of the males, of B. mirus, B. lemnis-
catus and B. lepidus is needed from Nepal, where
all three species may occur in proximity, in order
to clarify their relationships.
DISTRIBUTION. B. lemniscatus is primarily a
Tibetan species (Map 37). It is known from Gansu
(Bischoff, 1936), Qinghai (Skorikov, 19126; PW,
ZI, ZS), Sichuan (BMNH, PW), Tibet (Richards,
1930; Wang, 1982, 1988; BMNH), Sikkim (BMNH,
MNHU), Nepal and Uttar Pradesh (BMNH) and
Kashmir.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. lemniscatus is recorded only from Mt Apharwat
in the Pir Panjal range, from above the montane
coniferous forest (Map 38): 2 workers, Apharwat,
above 3000 m, viii-ix.1986 (Williams) (PW).
VARIATION WITHIN KASHMIR. The workers from
Mt Apharwat differ from one another in the
extent of the pale pubescence on the scutellum, at
the rear of the thoracic dorsum, and on tergum III
(Figs 325 & 326).
This species is particularly closely similar in
colour pattern to some of the white and yellow B.
lepidus (Figs 330 & 334) and B. biroi (Figs 338 &
339) that occur with it. They also resemble some
of the B. himalayanus (Figs 247-252), B. novus
males (Figs 260 & 262), B. kashmirensis (Figs
302-304) and B. rufofasciatus (Figs 434-436, 438,
439) from the same area. B. lemniscatus can be
recognised by the combination of the dense patch
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
of small punctures in the ocello-ocular area of the
head (Fig. 218) together with the very narrowly
unpunctured median area of the thoracic scutum
(Fig. 223; see the key, couplets 19 & 21).
FOOD PLANTS. No records.
Bombus (Pyrobombus) lepidus Skorikov
(Figs 68, 69, 108, 109, 148, 149, 188, 189, 213, 219,
224, 327-334, Maps 39 & 40)
Bombus lepidus Skorikov, 1912b: 606. Lectotype
by designation of Podbolotskaya (in press) [not
seen]. Paralectotype queen, CHINA: Qinghai,
northern slopes of the Xining mountains, 30.v.
1890 (Grum-Grzhimailo) (ZI) [examined].
Bombus genitalis Friese, 1913: 85. Holotype male
by monotypy (see Note 1 below), CHINA:
Tibet, no further data (Staudinger) (MNHU)
[examined]. Syn. n.
Bombus nursei var. tetrachromus Friese, 1918: 85.
LECTOTYPE queen by present designation
(see Note 2 below), INDIA: Kashmir, 8000—
9000 ft [2400-2700 m], vi.1901 (Nurse) (MNHU)
[examined]. Junior primary homonym of
Bombus tetrachromus Cockerell, 1909 [= B.
kashmirensis Friese]. Syn. n.
Bombus (Pratobombus) yuennanicola Bischoff,
1936: 7. Holotype queen by orginal designation,
CHINA: Yunnan, Lijiang, 15.v.1934 (Hone)
(MNHU) [examined]. Change of status to Pyro-
bombus lepidus subsp. yuennanicola (Bischoff)
by Tkalct (1989). Provisional synonym.
Bombus (Pyrobombus) yunnanicola Bischoff;
Wang, 1982: 435. Unjustified emendation (Art.
33b(i, iii)).
Pyrobombus (Pyrobombus) lepidus subsp. hilaris
Tkalci, 1989: 48. Holotype queen by original
designation, PAKISTAN: Gilgit, Banidas,
36°11'N 74°33’E, 2600 m, 30.vi-2.vii.1959
(Lobbichler) (ZS) [examined]. Syn. n.
TYPE MATERIAL. Note 1. Friese’s description of B.
genitalis specifies that only a single male was
examined. A single male in the MNHU collection
agrees with the original description and carries the
data quoted, together with a red printed label
‘Type’, and one of Friese’s determination labels
‘Bombus / genitalis / [male] 1912 Friese det /
n. Fr.’. This is believed to be the single individual
on which the original description is based and is
regarded as the holotype.
Note 2. Friese’s description of B. nursei var.
tetrachromus lists two queens. A single queen in
the MNHU collection agrees with the original
description and carries a label (1) ‘Kashmir /
8-9000ft / 6.01’; (2) ‘Bombus / tetrachromus /
75
[female] 1915 Friese det. / Fr.’; (3) a red printed
label ‘Type’. This specimen, which lacks both
front tarsi and the mid left tarsus, is designated as
lectotype (Art. 74a).
AFFINITIES. Within the subgenus Pyrobombus
(see the comments on B. hypnorum), B. lepidus
belongs to the parthenius-group of species (see the
comments on B. lemniscatus).
Within the parthenius-group, the recurved
hook of the penis valve head remains broad and
scarcely shortened for males of B. lepidus (Figs
68 & 69) (plesiomorphy within the parthenius-
group), although males of most of the other
species have the recurved hook much shortened
(synapomorphy of B. infirmus + B. parthenius +
B. luteipes). Individuals of B. infirmus (Tkalct)
are banded with yellow pubescence like indivi-
duals of B. lucorum, but have fewer white hairs on
the apical terga. B. infirmus is known from
northern Burma (BMNH) and Sichuan (BMNH,
PW). Females of the two Himalayan species,
B. parthenius Richards (14.iv.1934) (which is
probably conspecific with Bremus sonani Frison,
30.iv.1934) and B. luteipes Richards (conspecific
with Pyrobombus signifer Tkalci, 1989, syn. n.),
can be recognised by their generally dull grey-
yellow pubescence (lighter for the males), with no
distinct black band between the wing bases. The
apical parts of the sterna and the distal sclerites of
the legs are usually a lighter orange-brown colour
for females of B. luteipes (see Tkalci, 1989: fig.
26) than for females of B. parthenius. Males of B.
luteipes have very distinctive large genitalia,
which have the interio-apical corner of the gono-
coxite spinosely produced rather than forming a
right angle as for B. parthenius. B. luteipes and B.
parthenius are known from China and the eastern
Himalaya to as far west as Nepal (BMNH, NMS,
PW) and Uttar Pradesh (BMNH). Individuals of
B. parthenius tend to occur at higher altitudes
in the montane forests than do those of B.
luteipes.
Another nominal taxon, Pyrobombus infre-
quens, has been described recently (Tkalcu, 1989)
from four workers from Burma and China. These
are similar to workers of B. parthenius, but
the oculo-malar distance is described as slightly
longer. Other workers from near Gongga Shan in
Sichuan (PW) are similar to the holotype of
Pyrobombus infrequens in that they have few
large punctures near the middle of the clypeus,
although these workers have most of the orange-
red pubescence of terga IV-V replaced by black.
A male from the same locality also differs from
Nepalese B. parthenius in that the orange-red
pubescence of terga IV-V is replaced by black.
76
The genitalia of this male are closely similar to
those of males from Nepal, although the gonosty-
lus projects inwards distinctly beyond the interio-
apical corner of the gonocoxite, whereas this
corner of the gonocoxite is longer for both B.
parthenius and B. luteipes, and the volsella is
slightly broader from the ventral aspect. Much
more material of the nominal taxa of the parthenius-
group is required to establish their relationships
definitively.
Most females of B. lepidus from Kashmir have
the pale bands of the thoracic pubescence grey-
white and have relatively few black hairs on the
gastral terga (e.g. Fig. 333; redescribed indepen-
dently under the name B. nursei var. tetrachromus
by Friese). A queen from north-western Kashmir
with the pale pubescence of the thorax and tergum
I yellow as well as on tergum II (Fig. 327) was
described by Tkalci (1989) under the name
Pyrobombus lepidus subsp. hilaris. Some indivi-
duals of both sexes from further to the east in
Tibet (Wang, 1982, 1987; BMNH, PW), Yunnan
(Bischoff, 1936; Wang, 1987; BMNH), Sichuan
(BMNH) and Shanxi (Yasumatsu, 1951) also have
all of the pale pubescence yellow. These indivi-
duals were described by Bischoff (1936) under
the name B. yuennanicola, which Tkalcu (1989)
has regarded as a subspecies of B. lepidus. For
females of B. yuennanicola, the many close, fine
punctures of the post-ocular area of the head do
not extend as far anteriorly among the large
punctures of the lateral part of the ocello-ocular
area, although the difference is subtle. The male
genitalia have the shoulder of the penis valve
particularly convex and the inner margin of the
gonostylus is more deeply and broadly concave
(Wang, 1982: fig. 39; PW). Both the yellow
individuals and the white individuals apparently
occur together at the same localities in the far east
of Tibet (Wang, 1982: Jomda & Chagyab) and in
Yunnan (Wang, 1987: Dongchuan). However, a
male from Tibet described by Friese (1913) under
the name B. genitalis is more extensively yellow
like the males of B. lepidus, although the genitalia
are more similar to those of B. yuennanicola. It is
possible that these bees are all part of the same
interbreeding population, in which the shape of
the gonostylus and the colour of the pale pubes-
cence show more or less correlated patterns of
variation. Much more information on interbreed-
ing and variation is needed to establish the status
of the nominal taxa in this group.
DISTRIBUTION. B. lepidus is a peri-Tibetan species
(Map 39). It is known from Shanxi (Yasumatsu,
1951), Gansu (Skorikov, 19126; Bischoff, 1936;
Wang, 1982; ZS), Qinghai (Skorikov, 19125;
PAULH. WILLIAMS
Wang, 1982; BMNH, ZI), Yunnan (Bischoff,
1936; Wang, 1987; BMNH, MNHU), Tibet
(Friese, 1913; Wang, 1982; BMNH, MNHU,
PW), Nepal (Tkalct, 19745; BMNH), Himachal
Pradesh (BMNH, PW) and Kashmir.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. lepidus is recorded from the Hindu Raj range,
from the southern side of the Great Himalaya
range and from the Pir Panjal range, primarily
from around the upper limit of the montane
coniferous forest (Map 40, Fig. 6): 23 queens, 79
workers, 22 males, from 11 localities 2400-4600 m
(BMNH, MNHU, NR, PW, ZS).
VARIATION WITHIN KASHMIR. Females from the
Hindu Raj and Karakoram ranges have the pale
pubescence of the thorax and terga I-II yellow and
have very few black hairs on tergum III (Figs 327
& 328).
Queens and workers from the Kishanganga
valley have a variable amount of grey-white hair
intermixed on the head. The pale pubescence of
the thorax and of tergum I is grey-white (tergum IT
remains yellow), and there are very few or no
black hairs on tergum III (Figs 330 & 331).
Females from Mt Apharwat are similar, although
the lateral margins of tergum II are fringed with
more of the white hairs (Figs 333 & 334), and one
queen has all of the pubescence of tergum II
cream-white.
Yellow females (Figs 327 & 328) are similar to
some of the B. himalayanus (Figs 242 & 243) and
B. keriensis (Fig. 404 & 407) that may occur with
them. White and yellow workers of this species
are particularly closely similar in colour pattern
(Figs 331 & 334) to the B. lemniscatus (Figs 325 &
326) and B. biroi (Fig. 339) workers that occur
with them at some localities in the Kishanganga
valley and around the Vale of Kashmir. White and
yellow females also resemble some of the B.
himalayanus (Figs 247-249), B. novus males (Fig.
262), B. kashmirensis (Fig. 203) and B. rufofasci-
atus (Figs 435 & 436). B. lepidus can be recognised
by the characters given in the key (couplets 19 &
21). The yellow males with red hairs on the apical
terga may be similar in colour pattern (Figs 329 &
332) to the males of B. pyrosoma (Fig. 430), which
occur with them at some localities. B. lepidus
males can be recognised by their small size and by
their rounded gonostyli (Figs 188 & 189).
FOOD PLANTS. (Ranunculaceae) Aconitum laeve
Royle, A. heterophyllum Wallich ex Royle;
(Balsaminaceae) Impatiens glandulifera Royle;
(Scrophulariaceae) Scrophularia pauciflora Benth..,
Digitalis lanata Ehrh. [introduced].
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Bombus (Pyrobombus) biroi Vogt
(Figs 42, 70, 71, 110, 111, 150, 151, 190, 191, 214,
217, 335-346, Maps 41 & 42)
Bombus (Pratobombus) biroi Vogt, 1911: 51.
Lectotype queen by designation of Tkalci
(1969a: 195), U.S.S.R.: Kirgiziya S.S.R., Alai
mountains, 1905 (TM) [examined].
Bombus (Pratobombus) biroi [var.] flavobistriatus
Vogt, 1911: 52 (see Note 1 below). LECTO-
TYPE queen by present designation (see Note 2
below), U.S.S.R.: Kirgiziya S.S.R., Naryn,
5.vi.1904 (ITZ) [examined]. Syn. n.
[Bombus (Pratobombus) biroi flavostriatus Vogt,
1911: 52 [examined]. Infrasubspecific (see Note
1 below) (Art. 45g(ii)(1)), unavailable name. |
[Bombus (Pratobombus) biroi flavofasciatus Vogt,
1911: 52 [examined]. Infrasubspecific (see Note
1 below) (Art. 45g(ii)(1)), unavailable name. ]
Bombus nursei Friese, 1918: 84. Holotype queen
by monotypy (see Note 3 below), INDIA:
Kashmir, 8000-9000 ft [2400-2700 ml], vi.1901
(Nurse) (MNHU) [examined]. Syn. n.
Bombus abbotti Cockerell, 1922: 2. Holotype
queen by original designation [p. 3], CHINA:
Xinjiang, Taghdumbash, 13000 ft [4000 ml],
11.vii.1894 (Abbott) (USNM) [examined]. Syn-
onymised with Pyrobombus biroi (Vogt) by
Tkalct (1969a).
Bombus agnatus Skorikov, 1933b: 248. Lectotype
queen by designation of Podbolotskaya (in
press), INDIA: Kashmir, Ladakh, Khardung
La, 2-4.vii.1912 (Jacobson) (ZI) [examined].
Junior primary homonym of Bombus lapponi-
cus var. agnatus Skorikov, 1912a [not infrasub-
specific after Tkalci, 1969a (Art. 45g(ii)(1))].
Provisional synonym.
Bombus (Pratobombus) kotzschi Reinig, 1940:
227. Holotype male by monotypy, presumed
lost (see Note 4 below), AFGHANISTAN:
Baghlan, Andarab, 4000-4500 m, viii.1936
(Kotzsch) [not seen]. Synonymised with Bombus
agnatus Skorikov by Tkalct, 1969a. Provisional
synonym.
NOMENCLATURE. Note 1. Vogt states (1911:50,
footnote 1) that he intends ‘Varietas geographica’
to be equivalent to subspecies in rank and
“Aberratio extrema’ to be of infrasubspecific rank.
He uses ‘Forma’ for taxa that he could not assign
to either rank with certainty. In the second
paragraph on page 52 he describes ‘Die Varietat
von Narynj und dem Boro-Chorogebirge . . .’ and
the first name used in this paragraph is ‘(flavobis-
triatus)’. This is deemed to be the subspecific
name that was intended for individuals from this
area with this colour pattern, which Vogt charac-
terised by a greater tendency towards a yellowish
TW
colour for the pale pubescence. The four names
that follow in the same paragraph are all applied
to what can be regarded as variants of this local
colour pattern, even though only the first two
(dyspostzonatus and nigrozonatus) have their
infrasubspecific status specified by the use of the
term ‘ab.’. Therefore the taxon named flavobistri-
atus is deemed to be of subspecific rank, whereas
flavostriatus and flavofasciatus are deemed to be
of infrasubspecific rank (Art. 45g(1i)(1)).
TYPE MATERIAL. Note 2. Vogt’s description of B.
biroi var. flavobistriatus does not specify the
number of specimens examined. A single queen in
the ITZ collection agrees with the original des-
cription and carries a label (1) ‘Narynj / 5.6.04’;
(2) a red, handwritten label ‘flavobistriatus /
Type’; (3) ‘Collectie /C. et O. Vogt / Acq. 1960”.
This specimen, which lacks only the left antenna,
is designated as lectotype (Art. 74a).
Note 3. Friese’s description of B. nursei speci-
fies a single queen. A single queen in the MNHU
collection agrees with the original description and
carries labels with the data quoted, together with a
red, printed label “Type’ and one of Friese’s
determination labels ‘Bombus / nursei’, dated
1909. I believe that this is the single specimen on
which the original description is based and regard
it as the holotype (Art. 73a(ii)).
Note 4. Reinig’s description of B. kotzschi
specifies a single male from Andarab. This speci-
men could not be found either in the ITZ collec-
tion (W. Hogenes in litt.) or in the ZS collection
(E. Diller in litt.), which are the collections with
many of Reinig’s bumble bee type- specimens.
According to P. Rasmont (in /itt.), any of Reinig’s
pre-war material not in the ITZ collection must be
considered to have been destroyed by soldiers
during the winter of 1945. The types of P. rupestris
subsp. elisabethae Reinig, 1940, from Afghanistan
in the ZS collection may have survived because
they were on loan to a colleague.
AFFINITIES. Among the Old World species of the
subgenus Pyrobombus (see the comments on B.
hypnorum), the male genitalia of B. biroi are very
distinctive, because the sickle-shaped, recurved
hook of the penis valve head is strongly broadened
and apically rounded (Figs 70 & 71; Kriiger, 1943:
p. 530 figs 30-45). The other species that share this
character state belong to the North American
flavifrons-group (?synapomorphy of the broad
hook of the penis valve head) (the ‘Flavifrons
group’ of Thorp, 1969). This includes at least
B. flavifrons Cresson, B. centralis Cresson, B.
vandykei (Frison), B. caliginosus (Frison) and
B. vagans Smith. B. sandersoni Franklin and B.
78
frigidus Smith are similar but have the hook of the
penis valve head less strongly broadened.
The closest relatives of B. biroi in the Old
World may include the species of the pratorum-
group, which are primarily Eurosiberian (see e.g.
Lgken, 1973; BMNH). Among the species of the
subgenus Pyrobombus, they have relatively undif-
ferentiated male genitalia. But for B. jonellus
(Kirby), B. pyrenaeus Pérez and B. brodmannicus
Vogt in particular, not only is the ventro-lateral
angle of the penis valve just nearer to the base
than to the head of the penis valve (as for species
of the flavifrons-group and B. biroi, Figs 110 &
111), but the recurved hook of the penis valve
head is also weakly broadened (see Kriiger, 1943:
530). B. jonellus is widespread in the northern
parts of Europe and of the U.S.S.R. (e.g. Panfilov,
1982: map 150) and reaches eastwards to western
Canada (where it occurs together with B. flavi-
frons; PW). B. pyrenaeus is known from the
mountains of southern Europe, including the
Pyrenees, the Alps, the Carpathians and the
Balkans (e.g. Tkalci, 19695; BMNH, PW). B.
brodmannicus is known from both the western
Alps and from Turkey and the Caucasus (e.g.
Tkalct, 1973; BMNH, PW). The females of B.
biroi can be distinguished by their few, scattered,
small punctures among the large punctures of the
post ocello-ocular areas of the head, where the
other species have these areas closely and evenly
covered with small punctures.
The lectotype queen of B. biroi from the Alai
mountains has the pale bands on the thorax and on
terga I-II grey-white, whereas the pubescence of
terga I-II is yellow for the queen described by
Vogt (1911) under the name B. biroi var. flavobis-
triatus. Queens from the Himalaya are closely
similar in morphology, but the grey-white pubes-
cence is often extensively replaced by black,
especially on the thorax and on tergum I (des-
cribed under the name B. nursei by Friese, 1918).
Although the single male described under the
name B. kotzschiis apparently no longer available
(see Note 4), Reinig’s description of the broader
recurved hook of the penis valve head (Reinig,
1940: 228 ‘ist der Uncus etwas kraftige entwickelt
als bei B. pratorum.’) and of the unbanded yellow
colour pattern, similar to B. semenovianus, agrees
most closely with two males from the Zanskar
mountains that are otherwise similar to B. biroi
(Figs 70 & 71). Variation in the male genitalia of
B. biroi is greatest in the breadth of the recurved
head of the penis valve, but this is not associated
with the variation in the colour pattern of the
pubescence. It appears to have no more than in-
dividual significance and the males from Zanskar
are within the range of variation for banded B.
PAULH. WILLIAMS
biroi (e.g. Kriiger, 1943: p. 530 figs 30-45).
Females with a similar, unbanded yellow colour
pattern were described from Ladakh by Skorikov
(1933b) under the name B. agnatus. More
recently, Tkalct (1969a) synonymised B. kotzschi
with B. agnatus, but listed them as a separate
species from Pyrobombus biroi without further
comment. From an examination of the available
material, the unbanded yellow females of B.
agnatus seem to be closely similar in morphology
to the banded yellow and white females of B. biroi
from Kashmir, as well as to the banded yellow
females of B. biroi from further north in central
Asia. There is some variation in the extent of the
fine punctures in the ocello-ocular area and in the
breadth of the labral lamella of the females,
especially among individuals of B. nursei. These
characters also vary considerably within each
nominal taxon so that they do not appear to be
diagnostic. Thus B. biroi and B. kotzschi (as the
oldest available name for B. agnatus) may be two
separate species, but until there is better evidence
against interbreeding than just differences in
colour pattern, I consider that they are most likely
to be parts of the same species, B. biroi. More
material is needed from the possible hybrid zone
to the north of the Great Himalaya range in order
to resolve the problem.
DISTRIBUTION. B. biroi is a central Asian species
(Map 41). It is known from Mongolia (BMNH,
PW), the Tien Shan (Vogt, 1911; Skorikov, 1931;
Kriiger, 1943; BMNH, ITZ, TM), the Pamir
(Cockerell, 1922; Reinig, 1930, 1934; Skorikov,
1931; USNM), the Hindu Kush (Reinig, 1934,
1940; Tkalci, 19692; BMNH), Kashmir and
Himachal Pradesh (BMNH, PW).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. biroiis recorded from the Hindu Raj, Ladakh,
Zanskar, Great Himalaya and Pir Panjal ranges,
in subalpine scrub (Map 42, Fig. 6): 14 queens, 38
workers, 12 males, from 13 localities 2300-4600 m
(BMNH, MNHU, NR, PW, ZI).
During July 1986, I saw queens investigate gaps
among the large boulders that were lying in gullies
in the subalpine zone of Mt Apharwat. However,
I could not find any colonies there in August or
September.
VARIATION WITHIN KASHMIR. Females from the
Hindu Raj range have the thoracic dorsum uni-
formly yellow and terga I-III black (5/S queens,
1/1 workers, Fig. 335), like the lectotype queen of
B. agnatus from Ladakh. A single worker from
Rumbak in the Zanskar ranges has yellow hairs
intermixed on tergum I (Fig. 336). Two males
from the Zanskar ranges at Panichar near Suru
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
and at Tungri (Fig. 337) have the pubescence of
the thorax, of tergum I and of the basal three-
quarters of tergum II uniformly yellow, with the
apical and lateral margins of tergum II, all of
tergum III and for the male from Suru, the basal
margin of tergum IV black. This colour pattern is
otherwise known for this species only from the
Hindu Kush ranges (BMNH).
The species may not vary much in colour pat-
tern on the southern side of the Great Himalaya
range, where light individuals occur (Figs 338-
340), but it is very variable in the Pir Panjal range,
where dark individuals (Figs 341-346), similar to
those from Himachal Pradesh, also occur. On Mt
Apharwat the variation is chiefly in the form of
(a) a reduction of the pale thoracic bands to an
almost completely black thorax, with only a few
pale hairs intermixed anterior to the wing bases;
(b) replacement of the red with black progres-
sively across terga III-VI, the darkest queen
retaining red only on the posterior median part of
tergum IV and on the anterior median part of
tergum V; and (c) replacement of the yellow with
white on tergum II. Reduction of the pale thoracic
pubescence is pronounced for all 4 queens (Figs
341, 344, 346), but less common among workers
(2/18 workers, Fig. 345). Complete replacement
of yellow with white on tergum II may be common
among queens (2/4 queens, Figs 341, 346) and
among workers with broad bands of pale pubes-
cence on the thorax (6/16 workers, Fig. 342).
The unbanded yellow individuals of this species
(Figs 335-337) are closely similar to some B.
subtypicus (Figs 319-324), although they have not
been found together. They are also similar to
some B. avinoviellus (Fig. 232), B. marussinus
(Figs 254-257), B. branickii (Fig. 265), B. asiati-
cus (Figs 371-376) and B. semenovianus (Figs
397-402) that may occur with them. Unbanded
yellow B. biroi can be recognised by the charac-
ters given in the key (couplets 19 & 20). The light,
white and yellow individuals (Figs 338 & 339) are
closely similar in colour pattern to the B. lemnis-
catus (Figs 325 & 326) and B. lepidus (Figs 330,
331, 333, 334) that occur with them at some
localities. They also resemble some of the B.
himalayanus (Figs 247-252), B. novus males (Figs
260 & 262), B. kashmirensis (Figs 302-304) and B.
rufofasciatus (Figs 435 & 437) that occur with
them. B. biroi can be recognised by the characters
given in the key (couplets 19 & 20). The queens
with a dark thorax and a white tergum II (Figs 341
& 346) resemble some of the B. himalayanus (Fig.
250) and B. kashmirensis (Figs 308 & 309) that
occur in the same area. B. biroi can be distin-
guished from these by its combination of a shiny
outer surface to the hind tibia and by its simple
79
mandibles, which lack the many, pronounced
teeth of B. kashmirensis. Some of the darkest
individuals of B. biroi (Figs 344 & 345) show some
similarity in colour pattern to some individuals of
B. haemorrhoidalis or B. trifasciatus but have
the oculo-malar distance much shorter. These
species are completely segregated by altitude. The
banded yellow males (Figs 340 & 343) are similar
in colour pattern to the males of B. keriensis (Figs
410 & 416) that occur with them, but can be
recognised by the broader recurved hooks of the
head of the penis valve (Figs 70 & 71).
FOOD PLANTS. Unidentified dandelion-like com-
posites; (Gentianaceae) Swertia petiolataD. Don;
(Scrophulariaceae) Scrophularia pauciflora Benth..,
Picrorhiza kurrooa Royle ex Benth.
Subgenus BOMBUS Latreille
Leucobombus Dalla Torre, 1880: 40 (as a sub-
genus of Bombus Latreille). Type species: Apis
terrestris Linnaeus, 1758: 578 = Bombus ter-
restris (Linnaeus), by subsequent designation
of Sandhouse (1943: 564). Synonymised with
Bombus Latreille s.str. by Milliron (1961).
Terrestribombus Vogt, 1911: 55 (as a subgenus
of Bombus Latreille). Type species: Apis ter-
restris Linnaeus, 1758: 578 = Bombus terrestris
(Linnaeus), by subsequent designation of
Frison (1927: 67). Synonymised with Bombus
Latreille s.str. by Skorikov (1931).
Authors have disagreed concerning the correct
application of the name Apis terrestris Linnaeus
(see the comments on B. lucorum). This is
currently under consideration by the ICZN but
does not affect the application of the name
Bombus s.str.
For a general description of both sexes see
Richards (1968).
Bombus (Bombus) tunicatus Smith
(Figs 72, 73, 112, 113, 152, 153, 192, 193, 347-355,
Maps 43 & 44)
Bombus tunicatus Smith, 1852a: 43. Lectotype
queen by designation of Tkalci (1974b: 325),
INDIA: north, no further data (BMNH)
[examined].
Bombus gilgitensis Cockerell, 1905: 223. Holo-
type queen by monotypy (see Note below),
PAKISTAN: Gilgit (BMNH) [examined].
Change of status to Bombus tunicatus [Rasse]
gilgitensis Cockerell by Tkalct (1969a). Syn. n.
Bombus terrestris var. simlaénsis Friese, 1909:
80
674. Lectotype queen by designation of Tkalct
(1974b: 325), INDIA: Kashmir, ‘2—3000’ [?m],
vi.1904 (MNHU) [not seen]. Synonymised with
Bombus tunicatus subsp. tunicatus Smith by
Tkalct (19745).
Bombus terrestris var. fulvocinctus Friese, 1909:
674. Type worker not found (Tkalct, 1974b),
INDIA: Himachal Pradesh, Simla [not seen].
Synonymised with Bombus tunicatus Smith by
Tkalct (1974b).
TYPE MATERIAL. Note. Cockerell’s description
of B. gilgitensis specifies a single queen. A
single queen in the BMNH collection agrees
with the original description and carries a label
‘Gilgit’ and another label ‘Bombus / gilgitensis /
Ckll TYPE’, in handwriting identical to that of
Cockerell. I believe that this is the single specimen
on which the original description is based and
regard it as the holotype (Art. 73a(ii)).
AFFINITIES. The species of the subgenera Festivo-
bombus Tkalct,, Rufipedibombus Skorikov, Pressi-
bombus Frison and most Bombus s.str. share a
subapical constriction of the volsella (?synapo-
morphy). The species of the subgenera Rufipedi-
bombus, Pressibombus and Bombus share an
enlargement of the outer ridge of the penis valve
head and a reduction of the exterio-apical corner
of the gonostylus (synapomorphies). The species
of the subgenera Pressibombus (which includes
only the east Himalayan B. pressus (Frison)) and
Bombus share a ventrally and apically directed
extension of the outer ridge of the penis valve
head (synapomorphy). The species of the sub-
genus Bombus share a loss of the recurved hook of
the penis valve head and a great broadening of the
ventrally directed outer ridge, so that the penis
valve heads are flared outwards to form a funnel
(Figs 72-75, 112-115) (synapomorphy). The vol-
sella shows a secondary distal broadening by the
extension of its broad interio-apical process in-
wards towards the midline of the body (Figs 152—
155) (?synapomorphy of the subgenus Bombus).
Within the subgenus Bombus there are a very
large number of nominal taxa, although there
is relatively little variation in morphology in
comparison with some apparently monophyletic
subgenera with fewer species, such as Subterraneo-
bombus. The male genitalia of the Eurosiberian
B. sporadicus Nylander (distribution mapped by
Tkalci, 1967: 45) have broader and straighter
penis valves in lateral aspect than do those of B.
(Pressibombus) pressus (synapomorphy of the
subgenus Bombus). However, for B. sporadicus
the subapical inner margin of the penis valve is
nearly straight or convex in dorsal aspect, the
interio-apical corner of the gonostylus is produced
PAULH. WILLIAMS
beyond the exterio-apical corner by a distance
along the axis of the gonocoxite that is nearly
equal to the breadth of the volsella below its
interio-apical process, the interio-basal process of
the gonostylus is particularly weakly sclerotised
and the volsella projects beyond the gonostylus in
dorsal aspect by more than its own breadth below
its interio-apical process (?plesiomorphies within
the subgenus Bombus, showing some similarity to
B. pressus). The females of B. pressus and B.
sporadicus share a lack of large punctures in the
posterior part of the ocello-ocular area (for B.
sporadicus see Tkalcu, 1967: fig. 1).
The male genitalia of all species of the subgenus
Bombus except B. sporadicus have the subapical
inner margin of the penis valve concave in dorsal
aspect (usually with a small rounded projection
interio-basally, particularly Figs 72 & 73), the
gonostylus has the interio-apical corner produced
beyond the exterio-apical corner by a distance
that is much less than the breadth of the volsella
(Figs 192-195), the interio-basal projection of the
gonostylus is strongly sclerotised, and the exterio-
apical corner of the volsella projects beyond the
gonostylus in dorsal aspect usually by much less
than its own breadth (synapomorphies of B.
tunicatus + B. franklini + B. affinis + terrestris-
group). Species of the ferrestris-group (see the
comments on B. lucorum) have the ventro-basal
angle of the penis valve marked with a much more
pronounced right angle in lateral aspect (Figs 114
& 115) (synapomorphy).
B. tunicatus is unique among the Old-World
species of the subgenus Bombus for the combina-
tion of a reduced projection of the interio-apical
corner of the gonostylus (Figs 192 & 193) and a
broadly obtuse form of the ventro-basal angle of
the penis valve (Figs 112 & 113). The other species
with these character states, B. franklini (Frison)
and B. affinis Cresson, are restricted to a small
area near the west coast of North America (see
Thorp etal. , 1983) and to the eastern side of North
America (see Milliron, 1971) respectively. Males
of B. tunicatus are unique among all species of
the subgenus in that they have the gonostylus
extended by the development of a distal shelf, so
that the interio-apical corner is often particularly
broadly curved (Figs 192 & 193). The queens can
be distinguished by a distinct band of fine punc-
tures in a slight groove at the midline of the
clypeus.
Cockerell described a queen under the name
B. gilgitensis from north-western Kashmir that
differs from those from the Vale of Kashmir and
from those of the eastern Himalaya by an exten-
sive replacement of the pale pubescence by black
on the scutellum and on terga I-II. These dark
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
females from the lower Indus valley are otherwise
closely similar to B. tunicatus s.str. and are
considered to be part of the same species (see
below).
DISTRIBUTION. B. tunicatus is a Himalayan species
(Map 43). It is known from Sikkim (Bingham,
1897; Friese, 1918; Skorikov, 1933b), Nepal
(Tkalci, 1974b; BMNH, NMS), Uttar Pradesh
(Bingham, 1897; Frison, 1935; BMNH, MNHN,
UM), Himachal Pradesh (Friese, 1909; Cockerell,
1917; Frison, 1935; BMNH, MNHN, PW),
Kashmir, Pakistan (Frison, 1933, 1935; PW) and
the Hindu Kush (Tkalct, 1969a). It may also have
been genuinely recorded from the Indian plains,
at Calcutta (Dover, 1922).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. tunicatus is recorded from the Hindu Raj,
Karakoram, Ladakh, Great Himalaya and Pir
Panjal ranges, where it is widespread among
habitats, if most abundant around the lower
montane coniferous forest (Map 44, Fig. 6): 149
queens, 569 workers, 195 males, from 37 localities
1600-4100 m (AB, BMNH, BPBM, IZ, NR, PW,
RH, TL, ZM, ZS).
A colony was found in woodland near Harwan
(1700 m, 10.ix.1985). The nest entrance was a hole
3 cm wide in the soil amongst grass, 1m from the
foot of a tree.
VARIATION WITHIN KASHMIR. Queens from the
southern side of the Great Himalaya range and
from the Pir Panjal range have a lighter colour
pattern, with the pubescence of the scutellum,
tergum I and most of tergum II grey-white (Figs
350 & 353). The workers differ in that the black
band between the wing bases is more often
reduced or replaced by grey-white hairs, and the
pale pubescence of tergum II is usually chocolate-
brown (Figs 351 & 354). Males may have tergum
II either grey-white (Fig. 355) or chocolate-brown
(Fig. 352).
Queens from the northern side of the Great
Himalaya and from the Karakoram and Hindu
Raj ranges are darker, with the pale pubescence
of the scutellum and of terga I-II almost com-
pletely replaced by black (Fig. 347). The workers
and males associated with these queens retain a
pronounced pale band on the scutellum, but the
black band between the wing bases always extends
over at least one-third of the length of the thorax
(Figs 348 & 349). The chocolate-brown pubes-
cence is replaced by black on all but a narrow
crescent in the middle of the base of tergum II.
Most individuals from all localities can be
identified with one or other of these two extremes
of the variation within Kashmir, so that there
81
appear to be regional differences in colour pattern
without much overlap. For this species of the low
to mid montane zones, the segregation of the light
individuals in the Jhelum valley watershed (Vale
of Kashmir and the Kishanganga valley), from the
dark individuals of the Indus valley watershed
(Map 44), is consistent with regional colour
variation as parts of a single interbreeding popu-
lation and a single species. Opportunities for
dispersal of individuals over the intervening
barrier of high mountain ridges may perhaps have
been restricted, which could have resulted in the
apparently narrow colour cline (cf. the comments
on B. lucorum).
Light queens of this species (Figs 350 & 353) are
most similar in colour pattern to the B. avinoviel-
lus (Figs 238-241) and B. asiaticus (Figs 383-389)
that occur with them at some localities in the Vale
of Kashmir. They also resemble some B. kash-
mirensis (Figs 305-307), B. biroi (Figs 341 & 342)
and B. keriensis (Figs 408 & 409). Queens of B.
tunicatus can be recognised by their large size, by
the short oculo-malar distance and by the strongly
arched posterior margin of the hind basitarsus.
Workers that lack a black band on the thorax (Fig.
354) can be closely similar in colour pattern to
workers of B. simillimus (Figs 418 & 420), but can
usually be recognised easily because they have
tergum I white rather than chocolate-brown,
which is very rare for B. simillimus. Males can be
recognised by their outwardly flared heads of the
penis valves (Figs 72, 73, 112, 113).
FOOD PLANTS. (Ranunculaceae) Aconitum hetero-
phyllum Wallich ex Royle, A. hookeri Stapf;
(Balsaminaceae) Impatiens glandulifera Royle;
(Leguminosae) Lupinus sp. [introduced]; (Sam-
bucaceae) Lonicera quinquelocularis Hardw.;
(Compositae) Cirsium falconeri (Hook.f.) Petrak,
C. wallichii DC., Centaurea iberica Trevir ex
Spengel, unidentified yellow thistle-like compo-
sites, unidentified dandelion-like composites;
(Gentianaceae) Swertia petiolata D. Don;
(Scrophulariaceae) Digitalis lanata Ehrh. [intro-
duced], D. purpurea L. [introduced], Pedicularis
punctata Decne.; (Acanthaceae) Pteracanthus
urticifolius (Kuntze) Bremek.; (Labiateae) Stachys
sericea Wallich ex Benth., Prunella vulgaris L.
Bombus (Bombus) lucorum (Linnaeus)
(Figs 74, 75, 114, 115, 154, 155, 194, 195, 356-370,
Maps 45 & 46)
[Apis terrestris Linnaeus, 1758: 578. Lectotype
queen by designation of Day (1979: 74), ‘in
Europa’, no further data (LSL) [examined].
Application temporarily suspended (see Note 3
below) (Art. 80c).]
82
Apis lucorum Linnaeus, 1761: 425. Lectotype male
by designation of Day (1979: 66), SWEDEN:
no further data (LSL) [examined]. Synony-
mised with Bombus terrestris (Linnaeus) by
Day (1979).
Apis cryptarum Fabricius, 1775: 379. Lectotype
queen by designation of Lgken (1966: 199),
DENMARK: Copenhagen (Kiel) [not seen].
Change of status to Bombus terrestris var.
cryptarum (Fabricius) by Thomson (1872); syn-
onymised with Bombus lucorum (Linnaeus) by
Léken (1966). Provisional synonym.
Bombus lucorum (Linnaeus); Fabricius, 1804:
350.
Bombus modestus Cresson, 1863: 99. Holotype
queen by monotypy (see Note 4 below),
U.S.A./CANADA: Yukon river, no further
data (Kennicott) [not seen]. Junior primary
homonym of Bombus modestus Eversmann,
1852. Synonymised with Bombus lucorum
[subsp.] /ucorum (Linnaeus) by Milliron (1971).
Bombus moderatus Cresson, 1863: 109. Replace-
ment name for B. modestus Cresson. Synony-
mised with Bombus lucorum [subsp.] lucorum
(Linnaeus) by Milliron (1971).
[Bombus lucorum ab. mongolicus Vogt, 1909:
42. Infrasubspecific (see Note 1 below) (Art.
45f(iv)), unavailable name (Art. 45e).]
Bombus (Terrestribombus) lucorum forma magnus
Vogt, 1911: 56. Syntype queens, BRITAIN:
northern Scotland; Orkney Isles (ITZ) [not
seen]. Synonymised with Bombus lucorum
(Linnaeus) by Pekkarinen (1979). Provisional
synonym.
[Bombus (Terrestribombus) lucorum mongolicus
Vogt, 1911: 56. Published without description
or indication, unavailable name (Art. 12a).]
Bombus lucorum subsp. jacobsoni Skorikov,
1912b: 610. Lectotype queen by designation of
Podbolotskaya (in press), INDIA: Kashmir,
Maru, Fargabad, 19.v.1910 (Trubetskoy) (ZI)
[examined]. Syn. n.
Bombus terrestris var. lucocryptarum Ball, 1914:
82. Lectotype queen by designation of Rasmont
(1981: 151), BELGIUM: Brabant, Molenstede,
6.v.1913 (Brussels) [not seen]. Synonymised
with Bombus cryptarum (Fabricius) by Rasmont
(1983a). Provisional synonym.
Bombus pratorum var. longipennis Friese, 1918:
83. LECTOTYPE worker by present designa-
tion (see Note 5 below), INDIA: Sikkim, no
further data (MNHU) [examined]. Syn. n.
Bombus (Terrestribombus) lucorum var. alaiensis
Reinig, 1930: 107 (see Note 2 below). 3 syntype
queens, 7 syntype workers, U.S.S.R.: Kirgiziya
S.S.R.; Tadzhikistan S.S.R., 1950-3700 m,
21.vi-28.vii.1928 (ITZ) [not seen]. Synony-
PAULH. WILLIAMS
mised with Bombus lucorum subsp. jacobsoni
Skorikov by Tkalct (1969a). Syn. n.
[Bombus lucorum mongolicus Skorikov, 1933a:
57. Published without description or adequate
indication (only Vogt, 1911, cited), unavailable
name (Art. 12a).]
Bombus (Terrestribombus) magnus Rasse mon-
golicus Kriiger, 1954: 276 [by indication of Vogt,
1909: 42]. LECTOTYPE queen by present
designation (see Note 6 below), MONGOLIA:
north, no further data, 1892 (Leder) (ITZ)
[examined]. Change of status to Bombus
lucorum [subsp.] mongolicus Kriger [cited as
Vogt] by Tkalct (1974a). Syn. n.
Bombus reinigi Tkalct, 19746: 322. Holotype
queen by original designation, NEPAL:
Ringmo to Phoksumdo lake, 3700 m, 10.vi.1973
(Martens) (NM) [examined]. Syn. n.
NOMENCLATURE. Note 1. Vogt’s (1909) term ‘ab.’
must be taken to indicate infrasubspecific status
(Art. 45f(i1, iv)). For his subsequent paper, Vogt
states explicitly (1911: 50 footnote 1) that he
intends “Varietas geographica’ to be equivalent to
subspecies in rank and ‘Aberratio extrema’ to be
of infrasubspecific rank. He uses ‘Forma’ for taxa
that he could not assign to either rank with
certainty and these are treated as being of sub-
specific rank (Art. 45g(ii)).
Note 2. Reinig’s use (1930: 83 etc.) of the term
‘ssp.’ elsewhere in his paper is taken to indicate
that infrasubspecific rank is meant for B. lucorum
var. alaiensis. However, this name has subse-
quently been treated as a name in the species
group by Kriiger (1951), so it is deemed to be of
subspecific rank (Art. 45g(1i)(1)).
TYPE MATERIAL. Note 3. Both Lg@ken (1973) and
Day (1979) noted that the only admissible syntype
material of A. terrestris Linnaeus did not agree
with the traditional interpretation of the taxon
(which corresponds to Harris’s Apis audax, 1776:
130, not of page 137), but instead may be a queen
of the same species as B. lucorum. This syntype
queen has since been identified as conspecific with
B. cryptarum (Fabricius) by Rasmont (1988: 52).
If this is the only admissible syntype, then the
many citations of B. terrestris in its traditional
sense must refer to misidentifications of Linnaeus’s
taxon. This is unfortunate because the name B.
terrestris appears particularly widely in the litera-
ture on physiology, behaviour and ecology. To
minimise any possible confusion an application
has been submitted to the ICZN to use its plenary
powers for the conservation of the traditional
application of B. terrestris (L¢ken, Pekkarinen &
Rasmont, unpublished).
Note 4. Cresson’s description of B. modestus
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
(1863) specifies that a single female was examined.
If found, this specimen which was originally in the
‘Coll. Smith. Inst.’ should be regarded as the
holotype (Art. 73a(ii)).
Note 5. Friese’s description of B. pratorum var.
longipennis (1918) lists three queens and one
worker from Sikkim. I have seen three workers
from the MNHU collection that agree with the
original description and carry labels with the data
‘Sikkim / Coll. Bingham’ and Friese’s determina-
tion labels. Large workers of species of the
subgenus Bombus are often similar in size to small
queens of species of the subgenus Pyrobombus.
One of these workers in the MNHU collection
carries the labels (1) ‘Sikkim / Coll. Bingham’; (2)
‘B. prat./v. longipennis / [female] 1914 Friese det.
/n. n.’; (3) a handwritten label ‘selbstandige Art’;
(4) ‘LECTOTYPE / Bombus prato- / rum var.
longi / pennis Tkalcu det. / Friese’ [designation not
published, see Tkalct, 19746: 324, footnote 4]; (5)
‘Zool. Mus. / Berlin’. This specimen, which lacks
the left hind leg, is designated as lectotype (Art.
74a).
Note 6. Kriiger (1954) elevates Vogt’s B.
lucorum ab. mongolicus to subspecific rank as part
of the nominal taxon B. magnus. Vogt’s original
description of B. lucorum ab. mongolicus refers to
more than one specimen. A queen in the ITZ
collection agrees with the original description and
carries a label (1) ‘mongolicus Vogt’ in hand-
writing identical to that of Vogt; (2) ‘terrestris /
det. Kohl.’; (3) ‘.gelb’; (4) a label with two
illegible lines written in pencil, possibly ‘B terres-
tris v. / or lucorum’; (5) ‘N. Mongolei / Leder 92’;
(6) ‘Coll. Zoologisch / Museum / Amsterdam’; (7)
ared printed label ‘TYPE’; (8) ‘LECTOTYPUS /
B.lucorum ab. / mongolicus Vogt / [female]
Tkalct’ det.’ [designation not published]. This
specimen, which lacks the distal joints of the right
hind tarsus, is designated as lectotype (Art. 74a).
AFFINITIES. B. lucorum is one of the least well
understood bumble bee taxa at present and appli-
cation of the name has been a matter of opinion.
There is both a biological problem in determining
how many species of the tferrestris-group are
present in Europe and Asia, and a nomenclatural
problem (see Note 3 above) that has arisen from
what appears to be the perpetuated misapplica-
tion of Linnaeus’s name Apis terrestris (currently
under consideration by the ICZN). The biological
limits of the species for which, at present, A.
lucorum is the oldest available name (Art. 80c)
remain very much a question of the interpretation
of indirect evidence.
Within the subgenus Bombus, species of the
terrestris-group (see the comments on B. tunica-
83
tus) have the ventro-basal angle of the penis valve
marked with a strongly pronounced right angle in
lateral aspect (Figs 114 & 115) (synapomorphy).
Only the most southerly species of the terrestris-
group in the lowlands of eastern Asia, B. ignitus
Smith (distribution mapped by Tkalci, 1962: 89;
Sakagami, 1975: fig. 13), can be distinguished
easily by the shape of the male genitalia. In
particular, the apex of the penis valve head of B.
ignitus is more elongated dorsally and finger-
like, whereas the ventro-apical corner is rounded
rather than acutely pointed. The remaining taxa
within the ferrestris-group are less strongly diver-
gent in their morphology. Those differences in the
form of the male genitalia that are known may not
always be reliably discrete, particularly among
samples from widely separated areas. Any cladis-
tic interpretations of relationship based on them
would be premature. In the northern part of its
range B. ignitus overlaps with the east Asian B.
hypocrita Pérez in northern China and Japan
(distribution mapped by Tkalcti, 1962: 89), and
with the Siberian and Chinese B. patagiatus
Nylander in northern China (distribution mapped
by Tkalci, 1967: 52). In North America the
terrestris-group is represented by the widespread
B. terricola Kirby (probably conspecific with B.
occidentalis Greene, see e.g. Milliron, 1971, for
distribution records) and by the north-western
B. moderatus Cresson (probably conspecific with
B. lucorum s.1.).
In Britain, at least two species of the terrestris-
group coexist over a large area: B. terrestris (of
most authors after Linnaeus, but not of Linnaeus,
see Note 3 above) and B. /ucorum. They can be
distinguished only by subtle differences in their
morphology, by the length of the pubescence and
by their colour patterns. For instance, British
queens of B. terrestris of authors can be recog-
nised by their larger size, by their shorter pubes-
cence, which has broader, more golden-yellow
bands (rather than lemon-yellow), and especially
by the brownish or ‘buff’- coloured pubescence of
terga IV-V. Throughout the rest of the species’
distribution at low altitudes in central and
southern Europe, the pubescence of the apical
terga is usually more nearly white, like that of B.
lucorum. There, B. terrestris of authors must be
identified using characters such as the number of
small punctures in an antero-lateral cluster in the
ocello-ocular area of the head (see Kriiger, 1951;
Loken, 1973; Pekkarinen, 1979; Rasmont, 1984).
B. terrestris of authors can probably be considered
conspecific with individuals of this group with
other colour patterns that are known from the
Canary Islands (described under the name B. ter-
restris var. canariensis Pérez) and Madeira
84
(described under the name B. maderensis
Erlandsson), because there is no reason to believe
that these very similar nominal taxa would not
interbreed if they were to occur together. B.
terrestris of authors is otherwise known from
Morocco (BMNH, PW), Algeria and Tunisia
(Kruger, 1956; L¢ken, 1973; BMNH) and from
across Europe, Turkey and the Caucasus (Kriiger,
1954, 1956; Lgken, 1973; Panfilov, 1981; BMNH)
to Syria (BMNH, PW), Lebanon (BMNH) and
the Elburz (Panfilov, 1981; BMNH); and from
across the central U.S.S.R. to as far as the Tien
Shan (Kruger, 1956; Panfilov, 1981) and Hindu
Kush ranges (BMNH).
It is perhaps because of the difficulty in distin-
guishing continental European specimens of B.
lucorum and B. terrestris of authors that a particu-
lar effort has been made to describe subtle
differences among individuals of the terrestris-
group. Even excluding B. terrestris of authors in
all of its colour patterns (i.e. considering only B.
lucorum in its broadest sense), at least 187 names
have been published, although the majority are
deemed to be of infrasubspecific rank (for a
discussion of the status of the major groups of
nominal taxa, see e.g. Kriiger, 1951, 1954, 1956,
1958; Léken, 1973; Pekkarinen, 1979; Erlandsson,
1979; Delmas, 1981; Obrecht & Scholl, 1984;
Rasmont, 1984; Rasmont et al., 1986 and refer-
ences therein). These samples of the terrestris-
group from Europe show colour variation that is
only relatively slight for females in comparison
with many other species. Colour variation is more
pronounced for males, with the frequencies of
light and dark individuals differing among local-
ities (e.g. Pekkarinen, 1979: fig. 7). This variation
in the colour patterns of the males shows some
correlation with variation in the compositions of
their volatile marking secretions (Bergstrom et
al., 1973) and with variation in the mobilities of
the enzymes phosphoglucomutase and esterase-1
under electrophoresis (e.g. Scholl & Obrecht,
1983; Pamilo et al., 1984).
In northern Europe, samples of the terrestris-
group that exclude B. terrestris of authors do show
correlated patterns of variation in the colour of
the pubescence and in the different phosphoglu-
comutase mobilities, although these combinations
of character states are not strictly segregated
between the two groups of individuals (Pekkarinen,
1979; Pamilo et al., 1984). This apparent recom-
bination of character states in some individuals
could arise by interbreeding between the different
parts of a population of a single species (B.
lucorum s.\.). Furthermore, males with both light
and dark colour patterns have been reared from
the same nest (Lgken, 1961).
PAULH. WILLIAMS
In central Europe, Scholl & Obrecht (1983)
found that individuals differed in the mobilities
of not only phosphoglucomutase but also of
esterase-1. As in northern Europe, the males
in the two groups also tended to differ in the
extent of the pale pubescence. Scholl & Obrecht
interpreted the lack of individuals with double
or hybrid enzyme bands expected from hetero-
zygotes, and the lack of the alternative banding
pattern between the two enzymes expected from
recombinants, as evidence against interbreeding
and concluded that two species were present.
However, the expected heterozygote individuals
with both phosphoglucomutase bands were also
absent from the samples from northern Europe
(Pamilo et al., 1984), where interbreeding was
believed to be occurring (Pekkarinen, 1979). In
fact phenotypes with two phosphoglucomutase
bands were found in samples from both the Alps
and from the Pyrenees, although at least one of
the bands had a lower mobility than expected fora
heterozygote. Furthermore, the close association
between the states of the two enzymes could
possibly result from close genetic linkage. But in
any case, a definitive interpretation of the differ-
ences between these electromorphs can only
emerge from studies of genetic inheritance.
Scholl & Obrecht (1983: fig. 2) went on to
describe differences in body shape between the
queens with the two enzyme phenotypes, al-
though there was considerable overlap in the
variation. Pekkarinen (1979, see fig. 12) con-
cluded that similar differences in shape among
individuals from northern Europe showed a con-
sistent allometric pattern as though they had all
been taken from a single population. Scholl &
Obrecht (1983; Obrecht & Scholl, 1984) also
associated one of the two enzyme phenotypes
(form A) particularly with highland samples, both
from the Alps (where it was identified as the
nominal taxon B. cryptarum) and from the
Pyrenees (where it was identified as the nominal
taxon B. magnus). However, covariation of size,
shape, colour and enzyme mobilities is also known
from across Europe within the single species of
western honey bee, Apis mellifera Linnaeus (see
Ruttner, 1988). I agree with Pamilo et al. (1984),
who concluded from a comparison of their results
with those of Scholl & Obrecht (1983), that a
closer study of the pattern of associations between
the colour characters, the enzyme mobilities and
the male pheromone compositions of B. lucorum
s.]. is still needed to solve the problem.
There is no direct evidence to say whether or
not males or young queens with any one of the
colour patterns of B. lucorums.1|. avoid those with
any other colour pattern as potential mates in
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
their natural habitats. Information on assortative
mating in the field is central to the concept of the
nature of species that is used for this review.
Unfortunately the results of experiments on
mating under laboratory conditions may not
reflect the discrimination by specific-mate recog-
nition systems under natural conditions (see the
introduction on discrimination of species). De
Jonghe (1982) showed this with bumble bees in
cages, where males of B. terrestris of authors
would even mate with young queens of B. luco-
cryptarum Ball, a nominal taxon that is more
closely similar to B. lucorum s.str. Therefore
under these artificial conditions neither positive
nor negative results necessarily have implications
for conspecific status. De Jonghe & Rasmont
(1983) attempted to test the fertility (i.e. post-
mating isolation) of five artificial crosses between
three of the nominal taxa by laboratory experi-
ment, using 11 queens that had survived the
winter, of which 6 laid eggs, although none
hatched. Control crosses were carried out within
each of two nominal taxa. A total of 5 of these
queens survived the winter, but only one queen
laid eggs, which hatched normally. As de Jonghe
& Rasmont stated in the discussion of their
results, it is perhaps premature to attribute the
failure to hatch of the eggs from the experimental
crosses to genetic incompatibility on the basis of
so few replicates. Nevertheless, more extensive
experiments may confirm that such incompati-
bility exists.
At present it can be concluded that there is a
broad set of individuals that appears to be discrete
(B. lucorum in its broadest sense). Within this set
there is variation in pubescence colour, morph-
ology and enzyme mobilities. What is not resolved
is whether the patterns of variation reveal entirely
discrete subsets or, if they do, whether such
patterns necessarily imply a lack of interbreeding
between separate populations. Therefore, until
more direct evidence against shared specific-mate
recognition systems and interbreeding is avail-
able, I shall follow many previous authors and
continue to treat these bees as parts of a single
species, B. lucorum s.1. (see the comments on B.
hypnorum). Of course this interpretation does not
preclude the idea of some restriction of gene flow
among some parts of the population (see Barton,
1988) of B. lucorum s.|., perhaps between the
lowland and highland bees in particular. Thus B.
lucorum is treated for the present as conspecific
with the European taxa described under the
names Apis cryptarum (infra-subspecific accord-
ing to Lgken, 1966, 1973; Tkalct, 1969b, 1975),
B. terrestris var. lucocryptarum and B. lucorum
f. magnus (conspecific according to Pekkarinen,
85
1979; Pekkarinen et al., 1981); and conspecific
with the Asian taxa described under the names
B. magnus Rasse mongolicus, B. lucorum var.
alaiensis, B. lucorum subsp. jacobsoni, B. reinigi
and B. pratorum var. longipennis.
DISTRIBUTION. The present broad concept of B.
lucorum leads to the conclusion that it is the most
widespread of all bumble bee species, even
though, unlike some (see Richards, 1973), it is
not quite circumpolar (Map 45). It may include
individuals recorded from Europe (e.g. Kriiger,
1951; Lgken, 1973; Alford, 1975; Rasmont, 1984;
BMNH, PW), Turkey and the Caucasus (Kriger,
1951; Panfilov, 1981; Rasmont, 1984; BMNH) to
the Elburz (Rasmont, 1984; BMNH); and from
across the northern U.S.S.R. (Kriger, 1951;
Panfilov, 1981; BMNH) to Mongolia (Morawitz,
1880; Vogt, 1909; Kriiger, 1954, 1958; Tkalci,
1974a; Panfilov, 1981; ITZ, PW), Inner Mongolia
(Reinig, 1936; Panfilov, 1981; Wang, 1982),
Heilongjiang (Kriger, 1954, 1958), North Korea
(Kim & Ito, 1987), Hokkaido in Japan (Tkalcu,
1962; Sakagami & Ishikawa, 1969) and Kamchatka
(Bischoff, 1930; Kriiger, 1951; Panfilov, 1981;
BMNH); and from Alaska almost to Hudson Bay
(Milliron, 1971; BMNH, PW). In central Asia
its distribution reaches southwards to the Tien
Shan (Morawitz, 1880; Skorikov, 1931; Kriiger,
1954; Panfilov, 1957, 1981), the Pamir (Reinig,
1930, 1934; Skorikov, 1931; Kriiger, 1951, 1958;
Panfilov, 1981), the Hindu Kush (Reinig, 1940),
Pakistan (BMNH, PW), Kashmir, and eastwards
to Himachal Pradesh (BMNH, PW), Uttar Pradesh
(BMNH), Nepal (Tkalct, 1974b; BMNH, NMS),
Sikkim (Friese, 1918; BMNH, MNHU), Tibet
(Kriiger, 1951, 1958; Panfilov, 1957; Wang,
1982, 1988; BMNH), northern Burma (BMNH),
Yunnan (Wang, 1987), Sichuan (Wang, 1982;
PW), Gansu (Bischoff, 1936; Kriiger, 1951, 1958)
and Shaanxi (BMNH). Part of this distribution is
mapped by Panfilov (1981: map 96).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. lucorum is recorded from the Hindu Raj and
Karakoram ranges, and from the southern side of
the Great Himalaya range and from the Pir Panjal
range, in the upper montane coniferous forest and
in subalpine scrub (Map 46, Fig. 6): 14 queens, 78
workers, 93 males, from 12 localities 2100-4600 m
(BMNH, NR, PW, ZI).
VARIATION WITHIN KASHMIR. As in Europe there is
some variation in the extent of the pale pubes-
cence and in its hue. Some of the females from
Kashmir (Figs 359, 363, 364, 368, 369) broadly
resemble British B. Jucorum in colour pattern, but
have the yellow band across the anterior of the
86
thorax extending down the sides of the thorax
from a third to nearly half of the distance to the
leg bases. The pubescence of the face, of the
scutellum, and of tergum I is almost completely
black and tergum II is often black in the apical
quarter. The pale pubescence of the anterior
thoracic dorsum of the queen may be lemon-
yellow (similar to the holotype of B. reinigi) or
grey-cream (lectotype of B. lucorum subsp. jacob-
soni and one paralectotype queen, BMNH: both
specimens show little wing wear, which is ex-
pected to be related to age and fading of yellow
pubescence). The possibility that these cream-
banded, dark individuals are semi-melanic B.
patagiatus has been considered, but the punctures
anterior to the lateral ocelli are more widely
spaced (cf. Tkalct, 1967: fig. 2). The dark indivi-
duals are rare in the Hindu Raj and Karakoram
ranges (1/4 queens, 0/26 workers, Fig. 359), but
are more common further south in the Great
Himalaya range (1/2 queens, 12/27 workers, Figs
363 & 364) and predominate in the Pir Panjal
range (7/7 queens, 23/24 workers, Figs 368 & 369).
Other females from Kashmir (Figs 356, 357,
360, 361, 366) are lighter in colour, sometimes
with pale pubescence on the face, but more
generally on the scutellum, tergum I and almost
all of tergum II (similar to B. magnus Rasse
mongolicus, but with more pale pubescence). The
possibility that these extensively pale individuals
are B. sporadicus has been considered, but they
have more of the large punctures in the ocello-
ocular area of the head, especially posteriorly (cf.
Tkalci, 1967: fig. 1). The pale hair of the thorax
and of tergum II is always lemon-yellow, although
it is susceptible to fading among older individuals.
Most individuals that have been examined from
the Hindu Raj and Karakoram ranges (2/4 queens,
26/26 workers, Figs 356, 357, 360) have this light
colour pattern, although they are rare in the Great
Himalaya range (0/2 queens, 0/27 workers, Fig.
361) and in the Pir Panjal range (0/7 queens, 1/24
workers collected, Fig. 366, — the frequency
is actually much lower: during more than two
months of field work on Mt Apharwat in each of
two years, many more of the dark workers were
seen, but no more of the light workers).
There is also variation in details of the morph-
ology among these females, though this appears
not to correlate closely with variation in colour.
For instance, there is subtle variation in the shape
and sculpturing of the labrum. These differences
allow individual specimens to be recognised, but I
found no discrete or reliable morphological differ-
ences in this character between individuals with
the two extreme colour patterns. Variation in the
sculpturing of the ocello-ocular areas also appears
PAULH. WILLIAMS
to be continuous rather than being ordered in
discrete states, and there is no strict association
between the states of these morphological and
colour characters among individuals. Generally
the light individuals have many fine punctures
antero-laterally in the ocello-ocular area of the
head, where they often form a cluster similar to
that of B. terrestris of authors (L@ken, 1973: fig.
12B; Rasmont, 1984: fig. 11). There are often few
or none of these fine punctures for the yellow-
banded dark individuals, although rather more for
the cream-banded dark individuals. But then the
four queens from the Hindu Raj range all have
these fine punctures, even though only two of
them have a light colour pattern (like most of the
workers), one of them can be described as inter-
mediate (only a very few pale hairs intermixed in
the black pubescence of the scutellum and tergum
I), and the other has a dark colour pattern (there
are slightly fewer punctures on the two darker
queens). In contrast, the single light worker
from Mt Apharwat in the Pir Panjal range has
none of these fine punctures, like most of the dark
workers. Much of the material from the Great
Himalaya range is more or less intermediate in
these characters. For instance, half of the material
from Lal Pani (0/1 queen, 11/21 workers) can be
described as intermediate in colour pattern and
has fine punctures on the head, whereas the
remainder (1/1 queen, 10/21 workers) has a dark
colour pattern and lacks these punctures.
Variation in the colour pattern of the males may
be related to the two extreme colour patterns
described from the females from Kashmir. The
majority of males from the Pir Panjal range
(6/9 males, Fig. 370) and some from the Great
Himalaya range (38/82 males, Fig. 365) are exten-
sively yellow. Other males from the Pir Panjal (3/9
males, Fig. 367), from the Great Himalaya (44/82
males, Fig. 362) and from the Hindu Raj and
Karakoram ranges (2/2 males, Fig. 358) are
darker, with a broad, black band on the thorax
and another on tergum III. Thus the dark males
are most frequent where the light or intermediate
females predominate in the Karakoram and Great
Himalaya ranges, whereas the light males are
restricted to localities with the dark females in the
Pir Panjal and Great Himalaya ranges. The
gonostylus of these males varies considerably in its
exterio-lateral length (Figs 194 & 195), but this
variation appears to be continuous and uncor-
related with the colour pattern of the pubescence
so that it is of only individual significance.
One interpretation is that the individuals from
Kashmir with the extreme light or dark colour
patterns of the pubescence could represent at least
two separate species, with some variation within
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
each (Rasmont et al., 1986:678). The bees of this
group from north-western Kashmir are more
similar to some of those from central Asia,
whereas those from south-eastern Kashmir re-
semble those from the eastern Himalaya. But my
present interpretation of the specimens available
from Kashmir is that a large proportion can be
classed as intermediates between the extremes of
variation (the intermediate female pattern is
similar to that described under the name B.
lucorum var. alaiensis) without any apparent
discontinuity in this pattern of variation. For these
bees of the subalpine and upper montane forest
zones, the tendency for the frequencies of light
and dark colour patterns to change between
north- western and south-eastern Kashmir (Map
46) is consistent with regional variation among
individuals from a single interbreeding population
and a single species. There is perhaps relatively
free movement of individuals along the mountain
chains, which has resulted in extensive hybridisa-
tion and a broad colour cline (cf. comments on B.
tunicatus). Some of the factors that may influence
the variation in the male colour pattern are
discussed in the section on the colour patterns of
the Kashmir fauna.
Females of B. lucorum are distinctive in their
appearance among the fauna of Kashmir. The
yellow males with white hairs on the apical terga
(Figs 365 & 370) may be similar in colour pattern
to the males of B. hypnorum (Figs 313 & 316),
which occur with them at some localities. B.
lucorum males can be recognised by their larger
size, by the numerous black hairs intermixed on
the thorax and on terga III-IV and by their
outwardly flared heads of the penis valves (Figs
74, 75, 114, 115).
FOOD PLANTS. (Leguminosae) Trifolium repens
L.; (Compositae) Cirsium falconeri (Hook.f.)
Petrak, C. wallichii DC., unidentified dandelion-
like composites; (Gentianaceae) Swertia petiolata
D. Don; (Scrophulariaceae) Digitalis lanata
Ehrh. [introduced], Pedicularis punctata Decne.;
(Labiateae) Prunella vulgaris L.
Subgenus SIBIRICOBOMBUS Vogt
Sibiricobombus Vogt, 1911: 60 (as a subgenus of
Bombus Latreille). Type species: Apis sibirica
Fabricius, 1781: 478 = Bombus sibiricus
(Fabricius), by subsequent designation of
Sandhouse (1943: 599).
Obertobombus Reinig, 1930: 107 (as a subgenus
of Bombus Latreille). Type species: Bombus
oberti Morawitz, 1883: 238, by monotypy.
87
Synonymised with Sibiricobombus Vogt by
Richards (1968).
Obertibombus Reinig; Reinig, 1934: 167 (as a
subgenus of Bombus Latreille). Unjustified
emendation (Art. 33b(i, iii)).
Sibiricibombus Vogt; Skorikov, 1938a: 145. Un-
justified emendation (Art. 33b(i, iii)).
For a general description of both sexes see
Richards (1968).
Bombus (Sibiricobombus) asiaticus
Morawitz
(Figs 19, 76-78, 116-118, 156-158, 196-198, 227,
371-391, Maps 47 & 48)
Bombus hortorum var. asiatica Morawitz in
Fedtschenko, 1875: 4. Incorrect original spell-
ing (Art. 32c(i)), without separate availability
in this form (Art. 32c). LECTOTYPE worker
by present designation (see Note 4 below),
U.S.S.R.: ?Tadzhikistan $.S.R., “‘Tschiburgan’,
4530-8650 ft [1400-2600 m], 26.vi.1871 (ZMMU)
[examined].
Bombus longiceps Smith, 1878a: 8. Syntype
workers and male, INDIA: Kashmir, Ladakh,
Dras; Kargil; Leh (Stoliczka) (Calcutta) [not
seen]. Synonymy with Bombus asiaticus
Morawitz suggested by Reinig (1940), con-
firmed here.
Bombus asiaticus Morawitz; Dalla Torre, 1896:
512. Justified emendation (Art. 33b(ii)).
Bombus Regeli Morawitz, 1880: 337. Lectotype
queen by designation of Podbolotskaya (in
press), CHINA: Xinjiang, Borohoro Shan,
near Yining [= Gulja] (Regel/) (ZI) [examined].
Synonymised with Sibiricobombus asiaticus
(Morawitz) by Skorikov (1922).
[Bombus regeli ab. miniatocaudatus Vogt, 1909:
50 [footnote 1], not of Vogt, 1909: 56 [examined].
Infrasubspecific (Art. 45f(iv)), unavailable name
(Art. 45e).]
Bombus (Sibiricobombus) regeli miniatocaudatus
Vogt, 1911: 61 (see Note 1 below). Holo-
type male by monotypy (see Note 5 below),
MONGOLIA: ‘Septentrionalis’, no further
data (ITZ) [examined]. Junior secondary
homonym in Bombus of B. soroeensis var.
miniatocaudatus Vogt, 1909. Synonymy with
Bombus asiaticus Morawitz suggested by Reinig
(1940), confirmed here.
[Bombus (Sibiricobombus) regeli form. fusco-
caudatus Vogt, 1911: 61 [examined]. Infrasub-
specific (see Note 2 below) (Art. 45g(ii)(1)),
unavailable name. |
[Bombus (Sibiricobombus) regeli form. albocauda-
tus Vogt, 1911: 61 [examined]. Infrasub-specific
88
(see Note 2 below) (Art. 45g(ii)(1)), unavail-
able name. |]
[Bombus (Sibiricobombus) regeli form. tenuifas-
ciatus Vogt, 1911: 61 [examined]. Infrasub-
specific (see Note 2 below) (Art. 45g(ii)(1)),
unavailable name. ]
Bombus (Sibiricobombus) miniatocaudatus race
falsificus Richards, 1930: 652 (see Note 3
below). Holotype queen by original designation,
CHINA: Xizang [= Tibet], Tingri, 14000 ft
[4300 ml], 4.vii.1924 (Hingston) (BMNH)
[examined]. Syn. n.
Sibiricobombus flavodorsalis Skorikov, 1933b:
248. Syntype queens, INDIA: Kashmir,
Ladakh, Suru valley (ZI) [not seen]. Junior
secondary homonym in Bombus of B. sonorus
subsp. flavodorsalis Franklin, 1913 [= B. pen-
sylvanicus (Degeer), the correct original spell-
ing of B. pennsylvanicus of authors]. Synonymy
with Bombus asiaticus Morawitz suggested by
Reinig (1940); synonymised with Pyrobombus
longiceps (Smith) by Tkalct (1969a).
Sibiricobombus oshanini Skorikov, 1933b: 248.
Syntype queens and male, INDIA: Kashmir,
Ladakh, Nubra valley (ZI) [not seen]. Syn-
onymy with Bombus asiaticus Morawitz sug-
gested by Reinig (1940); synonymised with
Pyrobombus longiceps (Smith) by Tkalcu
(1969a).
NOMENCLATURE. Note 1. Vogt states (1911: 50,
footnote 1) that he intends ‘Varietas geogra-
phica’ to be equivalent to subspecies in rank and
‘Aberratio extrema’ to be of infrasubspecific rank.
He uses ‘Forma’ for taxa that he could not assign
to either rank with certainty. No particular status
is specified for the trinomen B. regeli miniato-
caudatus. Therefore it is deemed to be of sub-
specific rank (Art. 45f(i)).
Note 2. The other forms of B. regeli described
by Vogt (1911) are all from the same series from
‘Siebenstrom’ and can be regarded as variants of
One principal local colour pattern. These indivi-
duals were collectively referred to as ‘typischen
regeli’. Therefore they are deemed to be of
infrasubspecific rank (Art. 45g(ii)(1)).
Note 3. Although Richards (1930: 634 etc.)
used the term ‘Subsp.’ as well as ‘var.’ and ‘race’,
he states that the ‘race’ falsificus is characteristic
of a particular geographical area (p. 652: ‘In the
Himalayas a much paler form is found, for which I
propose the name falsificus, race nov.’). This
nominal taxon is therefore deemed to be of
subspecific rank (Art. 45f(1i)).
TYPE MATERIAL. Note 4. Morawitz’s description of
B. hortorum var. asiatica lists several localities
and a range of body lengths so that he must have
PAULH. WILLIAMS
had a series of specimens. I have seen a series of
three queens, one large worker and three smaller
workers from the ZMMU collection that agree
with the original description (all have the pubes-
cence of tergum II entirely yellow). The largest of
these workers carries (1) a blue label ‘26’; (2)
[Chiburgan]; (3) ‘Bombus / hortorum L. / varietas
asiatica.” in handwriting identical to that of
Morawitz; (4) ‘F. Morawitz / det. 1875.’; (5)
‘26. VI.1871’; (6) [Moscow / Zoological / Museum
MGU]. This specimen, which bears data labels
consistent with the original description (and is the
only specimen to bear the full name on the third
label) and is the least abraded and faded, but
which lacks the tarsi of both mid legs and of the
right hind leg, is designated as lectotype (Art.
74a).
Note 5. Vogt’s description of B. regeli ab.
miniatocaudatus is rather oblique, but is adequate
to imply that it differs from B. regeli s.str. in that
the pubescence of the apical terga is red. A single
male in the ITZ collection agrees with this descrip-
tion (Vogt, 1909, specified that he had a single
male with this colour pattern from central Asia)
and carries a handwritten red label ‘regeli=
Gruppe / miniatocaudatus / Type’. I believe that
this is the single specimen on which the original
description is based and regard it as the holotype
(Art. 73a(ii)).
AFFINITIES. A reduction of the ventro-basal angle
of the penis valve is shared by a large group of the
bumble bees with sickle-shaped, recurved hooks
of the penis valve head (synapomorphy). In
Kashmir this includes the species of the subgenera
Sibiricobombus and Melanobombus (Figs 116—
125). The ventro-basal angle of the penis valve is
also reduced for the east Himalayan B. (Festivo-
bombus) festivus, although in this case it is likely
to be a convergent character state (as opposed to
the interpretation in Williams, 1985). In contrast
to the species of the subgenera Sibiricobombus
and Melanobombus, B. festivus shares a reduction
and ventral torsion of the interio-basal process of
the gonostylus with the species of the subgenus
Pyrobombus, and B. festivus also shares a sub-
apical constriction of the volsella with species of
the subgenus Rufipedibombus (?synapomorphies,
see the comments on B. hypnorum and B.
tunicatus).
Within the large group of bumble bees that have
a reduced ventro-basal angle of the penis valve,
the temperate steppe species of the subgenus
Cullumanobombus (see Panfilov, 1951) share a
reduced outer ridge of the penis valve head, a
reduced interio-basal process of the gonostylus, a
reduced interio-apical process of the volsella and
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
an extended outer apical corner of the volsella
(synapomorphies). Many of the remaining species
that have a reduced ventro-basal angle of the
penis valve appear to share a more pronounced
narrowing of the volsella towards the apex (?syna-
pomorphy, possibly secondarily expanded for B.
oberti and for species of the lapidarius-group).
These species include two species-groups.
Species of the first group share a concave
distal margin of the recurved hook of the penis
valve head (Figs 79-85) (?synapomorphy). This
group includes the oberti-group of the subgenus
Sibiricobombus and the species of the subgenus
Melanobombus. The centre of diversity for this
group is near Kashmir, with divergent species-
groups in the Oriental region and in Europe and
northern Asia (see the comments on B. oberti, B.
ladakhensis and B. simillimus).
Species of the second group share a thickening
of the volsella in its apical part, which is delimited
by a strongly raised ridge. This ridge rises below
the inner dorsal margin and curves back across the
ventral surface, often forming a pronounced and
coarsely-sculptured swelling or tubercle midway
along the volsella (Figs 156-158) (synapomorphy).
This second group includes the Mongolian B.
sibiricus and the central Asian montane asiaticus-
group of the subgenus Sibiricobombus, which
reach westwards in the Caenozoic mountains
to south-eastern Europe, as well as the large
fraternus-group (in its broadest sense) from North
America and the mountains of western South
America (see below).
If this interpretation of these character states is
accepted as the most likely estimate of relation-
ships available at present, then one part of the
subgenus Sibiricobombus would appear to be
paraphyletic with respect to one monophyletic
clade, whereas the other part would appear to be
paraphyletic with respect to another monophy-
letic clade. Consequently the subgenus Sibirico-
bombus may be a polyphyletic assemblage, which
would account for its apparent lack of exclusive
synapomorphies. Yet females of the species that
are currently placed in this subgenus do show
some phenetic similarity because they usually
have particularly many of the short, branched
hairs that continue onto the outer surface of
the hind basitarsus, below the postero-proximal
corner (Figs 227 & 228, see the key, couplet 11).
The use of the subgeneric name Sibiricobombus
for what appears to be an artificial group with both
B. asiaticus and B. oberti is continued here only
for the sake of stability. I consider it undesirable
to take any nomenclatural action until the males
of B. flaviventris and B. tanguticus are available
to clarify the relationships among the species-
89
groups (see the comments on B. oberti and B.
simillimus).
Within the group of species that have both a
reduced ventro-basal angle of the penis valve and
a thickened volsella, B. sibiricus has an expanded
gonostylus (plesiomorphic among the species of
Sibiricobombus) (see Skorikov, 1931: fig. 27;
Tkalct, 1974a: fig. 4) and the volsella is only
weakly thickened. B. sibiricus (of the ‘sibiricus-
Gruppe’ of Tkalci, 1974a) is known from the area
around Mongolia (Skorikov, 1931: fig. 7). The
eyes of the male are similar in relative size to those
of the female. Apart from B. sibiricus and B.
oberti, those males that are known from the other
species of the subgenus Sibiricobombus all have
very much enlarged eyes relative to those of the
females.
Also within the group of species that have both
a reduced ventro-basal angle of the penis valve
and a thickened volsella, the species of the New
World fraternus-group in its broadest sense share
a reduction of the inner apical corner of the
gonostylus and the thickened part of the volsella
is usually more pronounced (?synapomorphies).
These New World species are currently separ-
ated among what seems to be an unnecessarily
large number of subgenera, most of which have
very few species: Fraternobombus Skorikov,
Separatobombus Frison, Crotchiibombus Franklin,
Brachycephalibombus Williams, Robustobombus
Skorikov and Rubicundobombus Skorikov; the
volsella is further modified by a pronounced
reduction of the exterio-apical corner for the
other species of this group, all from Central
America and the mountains of western South
America, that are currently placed in the sub-
genera Dasybombus Labougle & Ayala (1985: fig.
5), Funebribombus Skorikov and Coccineobombus
Skorikov (which includes B. handlirschi Friese)
(?synapomorphy).
Also within the group of species that have
both a reduced ventro-basal angle of the penis
valve and a thickened volsella, the species of the
asiaticus-group share a concavity of the outer
margin of the middle part of the volsella, and the
thickened part of the volsella is more pronounced
particularly at the basal end, where it forms
a rounded swelling or tubercle (Figs 156-158)
(2synapomorphies). This tubercle may increase
the contact area of the distal, coarsely-sculptured
area of the male volsella for gripping the female
sting base during copulation. This asiaticus-group
includes the species of the ‘niveatus-Gruppe’ of
Tkalcii (1974a), the species of the ‘P. asiaticus-
Gruppe’ of Tkalcti (1969a, or ‘longiceps-Gruppe’
of Tkalci, 1974a, 1974b) and B. obtusus of the
‘morawitzi-Gruppe’ of Tkalct (1969a, 1974a, but
90
not B. morawitzi Radoszkowski, see the com-
ments on B. oberti).
Among the nominal taxa of the asiaticus-group,
B. obtusus Richards from the Hindu Kush ranges
has a broadly rounded gonostylus with a broad
interio-basal process, and the inner margin of the
head of the penis valve has only a single, straight
dorsal ridge (plesiomorphic within Sibiricobom-
bus). Three worker paratypes (BMNH) that are
from the same locality as the holotype male
(Richards, 1951) have few of the short, branched
hairs arising from the outer surface of the hind
basitarsus, so that this area appears to be more
shining than for B. asiaticus, and the pubescence is
generally shorter. These characteristics of B.
obtusus remain segregated where it is known to
occur with B. asiaticus, without any evidence
of recombination in the small samples that are
available.
The males of the remaining species of the
asiaticus-group share a slight reduction of the outer
apical corner of the gonostylus, and the interio-
dorsal ridge at the base of the penis valve head
curves from its dorsal origin to the ventral margin
before it joins the recurved hook (synapomorphies
of B. asiaticus + B. niveatus [+ B. vorticosus] + B.
sulfureus). Males of the many nominal taxa that
are similar to B. asiaticus share a narrowing of the
volsella towards the apex from the inner margin,
so that it appears even more strongly inwardly-
curved (Figs 156-158) (apomorphy).
The males of B. niveatus Kriechbaumer, B.
vorticosus Gerstaecker and B. sulfureus Friese
share the ventrally-curved form of the interio-
dorsal ridge at the base of the penis valve head,
but also show an unusually pronounced develop-
ment of an interio-ventral ridge at the base of the
penis valve head (synapomorphy) (see Skorikov,
1931: fig. 29). B. niveatus, B. vorticosus and B.
sulfureus also retain the tubercle of the volsella
despite a subapical expansion of the volsella
(synapomorphy) (see Skorikov, 1931: fig. 29),
which is most pronounced for B. sulfureus (?auta-
pomorphy). The only known differences between
B. niveatus and B. vorticosus are in colour (Vogt,
1909; Pittioni, 1938). B. niveatus has the bands of
light pubescence grey-white, whereas they are
yellow for B. vorticosus. B. vorticosus is broadly
distributed between Krasnovodsk and the Elburz
mountains in the east to Yugoslavia in the west
(e.g. Vogt, 1909, 1911; Pittioni, 1938; Reinig,
1967, 1971, 1974; Tkalci, 1969b; BMNH, PW).
B. niveatus apparently occurs only within the
more central part of this area, where it is less
abundant (e.g. Vogt, 1909; Pittioni, 1938; Reinig,
1967, 1971, 1974; D. B. Baker, pers. comm.;
BMNH, PW). Reinig (1967) described some
PAULH. WILLIAMS
individuals as possible hybrids and it may be that
the banded white individuals and the banded
yellow individuals are both parts of a single
species, B. niveatus (cf. the comments on yellow
or white B. keriensis). B. sulfureus is a rare species
that occurs from the Elburz mountains to Turkey
(e.g. Reinig, 1971, 1974; BMNH).
Reinig (1940) suggested that the many similar
nominal taxa of the central Asian asiaticus-group
(apart from B. obtusus) might actually be indivi-
duals with different colour patterns within a single
species, B. asiaticus, but he lacked sufficient
material, especially of the males, to draw firm
conclusions. Individuals of the asiaticus-group
from Ladakh have an unbanded, yellow thorax
(Figs 371-376, described under the names B.
longiceps Smith; Sibiricobombus flavodorsalis
Skorikov; Sibiricobombus oshanini Skorikov).
Individuals from the the other side of the Great
Himalaya range in the Vale of Kashmir usually
have a grey-white thorax with a black band
between the wing bases (Figs 383-391, referred to
as B. callophenax Cockerell by Richards, 1930:
652, a misidentification, see the comments on B.
avinoviellus). There is also variation in the extent
of pale pubescence on terga I-II and in the colour
of the wings, which are usually more clouded with
brown (infuscated) among specimens from the
Vale of Kashmir. There is variation in the shape
of the male genitalia, especially as to whether
the gonostylus has the apical margin convex or
concave and whether or not the interio-basal
process is constricted near its base (Figs 196-198).
However, individuals with both principal colour
patterns show all forms of the gonostylus, so
the variation appears to be of only individual
significance.
The male mate-searching behaviour of the
unbanded yellow individuals and of the banded
white individuals could not be distinguished (see
the introduction on male mate-searching be-
haviour and on the inference of allopatric, con-
specific taxa). Individuals with both regional
colour patterns occupy similar open habitats. The
males all perch to watch for potential mates in a
similar way and race in pursuit of moving objects
(but without holding exclusive territories, cf. the
comments on B. rufofasciatus). At a few localities
between the two regions where individuals with
each of the principal colour patterns predominate,
other individuals have been collected that have
colour patterns with combinations of the charac-
ter states of both regional colour patterns (Figs
377-382, see the comments below on variation
within Kashmir). This pattern of variation is
consistent with hybridisation between individuals
with divergent colour patterns within a single
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
population (see the comments on variation below
and the discussion section on the variation of
B. asiaticus in Kashmir). Therefore these, and
probably the many other nominal taxa of the
central Asian asiaticus-group for which inter-
mediates are known (i.e. apart from B. obtusus),
including the banded and pale-yellow B. miniato-
caudatus race falsificus from southern Tibet, are
likely to be interbreeding as parts of a single
species. Three further nominal taxa (B. huangcens,
B. heicens and B. xionglaris), similar to B.
miniatocaudatus race falsificus, have been des-
cribed by Wang (1982) from Tibet (not seen).
DISTRIBUTION. B. asiaticus is a central Asian and
Tibetan species (Map 47). It is known from
Mongolia (Vogt, 1909; ITZ, PW), the Altai
(Skorikov, 1931), the Tien Shan (Morawitz, 1875,
1880; Vogt, 1911; Skorikov, 1931; Panfilov, 1957;
BMNH, ZI, ZMMU), the Pamir (Friese, 1913;
Reinig, 1930, 1934; Skorikov, 1931; BMNH),
the Hindu Kush (Reinig, 1940; Richards, 1951;
Tkalci, 1969a; BMNH), Pakistan (BMNH, PW),
Kashmir, Himachal Pradesh (BMNH, PW), Uttar
Pradesh (BMNH), Nepal (Tkalct, 19746; BMNH,
NMS), Tibet (Richards, 1930; Wang, 1982;
BMNH), Qinghai (Bischoff, 1936; Tkalct, 1961;
Wang, 1982; BMNH, ZS) and Gansu (Bischoff,
1936).
MATERIAL EXAMINED FROM KASHMIR. B. asiaticus
is widespread in Kashmir, recorded from the
Hindu Raj, Karakoram, Ladakh, Zanskar, Great
Himalaya and Pir Panjal ranges, down to the
margins of the valley floor in the Vale of Kashmir.
Although it is widespread among open habitats,
it is most abundant in large meadows in the
montane forest zone (Map 48, Fig. 6): 156 queens,
428 workers, 434 males, from 42 localities 1800—
4800 m (AB, BMNH, BPBM, FA, IZ, MI,
MNRHN, NR, PW, RH, SEMK, TL, ZM).
Two colonies were found at Leh (3500 m,
8.vii. 1980, 19.ix.1986). The nest entrances were
between rocks in dry-stone walls.
VARIATION WITHIN KASHMIR. Even within Kashmir
there is great variation in the colour pattern of this
species (Figs 371-391, Map 48), often even within
samples from a single locality. Only the principal
aspects of this variation are described here.
All of the individuals from the Hindu Raj,
Karakoram, Ladakh and Zanskar ranges, and
from across the Great Himalaya range from the
northern side to at least as far as Gumri, have the
pubescence of the thorax yellow without a black
band between the wing bases (Figs 371-376).
These individuals from the arid region show
relatively little variation, except in the replace-
91
ment of black pubescence with yellow on terga I-II
(Figs 374-376), which is especially pronounced for
all of the specimens from central Zanskar (2
queens, 1 worker, 3 males). Individuals with this
unbanded yellow colour pattern are also known
from the Hindu Kush ranges, but not from Tibet,
where individuals have the thoracic pubescence
yellow but with a black band between the wing
bases.
Most of the individuals from the Kishanganga
valley (16/16 queens, 80/83 workers, 105/114
males) and from around the Vale of Kashmir
(24/26 queens, 132/198 workers, 42/82 males)
have an obvious band of black hairs between the
wing bases (Figs 377-382, 386-391). Most of these
specimens have some pale pubescence on terga
I-II (Figs 377-389). Some of the individuals
from the Kishanganga valley (9/16 queens, 35/83
workers, 4/114 males, Fig. 381) and almost all of
those from around the Vale of Kashmir (24/26
queens, 197/198 workers, 37/82 males, Figs 383—
387, 389-391) have the pale pubescence grey-
white rather than yellow. Individuals from the
more mesic region of the western Himalaya to at
least as far east as Nepal also have the pale
pubescence grey-white with a black band between
the wing bases.
Individuals from a few localities in the higher
valleys of the Great Himalaya range show com-
plex variation with intermediates between the
unbanded yellow colour pattern that is common in
Ladakh, and the banded white colour pattern that
is common in the Vale of Kashmir. This is
especially evident among the material from Lal
Pani in the upper Kishanganga valley (Table 7,
e.g. Figs 380-382), and from Nigagar in the
upper Sind valley (e.g. Figs 377-379). These two
samples are also unusual because they include
specimens that have the red pubescence of the
apical terga largely or completely replaced by
black (Lal Pani: 2/5 queens, 4/17 workers, 48/92
males, Figs 380 & 381; Nigagar: 0/2 queens, 2/12
workers, 2/6 males, Fig. 379). Individuals with this
colour pattern are otherwise known only from the
Hindu Kush ranges (BMNH). For an analysis of
these data, see the discussion of the variation and
genetics of B. asiaticus in Kashmir.
The individuals with an unbanded yellow thor-
acic dorsum (Figs 371-376) are closely similar in
colour pattern to some of the B. avinoviellus (Fig.
232), B. marussinus (Figs 254-257), B. branickii
(Fig. 265), B. subtypicus (Figs 319-324), B. biroi
(Figs 335-337) and B. semenovianus (Figs 397—
402) that occur with them at some localities. The
banded white individuals of this species (Figs 381,
386, 387, 389-391) are closely similar in colour
pattern to some of the B. avinoviellus (Figs 238-
92
241), B. kashmirensis (Figs 305-307), B. biroi
(Fig. 342), B. keriensis (Figs 408 & 409) and to the
queens and some males of B. tunicatus (Figs 350,
353, 355) that occur with them at some localities in
the Vale of Kashmir. Females of B. asiaticus can
be recognised by their very long oculo-malar
distance, by the many fine punctures between the
compound eye and the base of the mandible and
by the many short hairs at the base of the hind
basitarsus (Fig. 227, see the key, couplet 11). The
males are easily recognised by their combination
of greatly enlarged eyes and very long antennae.
FOOD PLANTS. Kashmir: (Balsaminaceae) Impatiens
glandulifera Royle; (Leguminosae) Trifolium
repens L., Lupinus sp. [introduced], unidentified
pink clover-like legume; (Compositae) Cirsium
falconeri (Hook.f.) Petrak, C. wallichii DC..,
unidentified green-flowered thistle-like compo-
site; (Scrophulariaceae) Digitalis lanata Ehrh.
[introduced], D. purpurea L. [introduced], Pedi-
cularis punctata Decne.; (Labiateae) Prunella
vulgaris L.
Ladakh: (Leguminosae) Medicago falcata L.,
Trifolium repens L., Caragana versicolor (Wallich)
Benth.; (Compositae) Echinops cornigerus DC.;
(Scrophulariaceae) Verbascum thapsus L.; (Labia
teae) Stachys tibetica Vatke, Nepeta podostachys
Benth., Prunella vulgaris L.
Bombus (Sibiricobombus) oberti Morawitz
(Figs 20; 795 119° 1595 1995 228, 230, 392, 393;
Maps 49 & 50)
Bombus Oberti Morawitz, 1883: 238. LECTO-
TYPE male by present designation (see the
Note below), U.S.S.R.: Kazakhstan S.S.R.,
mountains near Alma Ata (Kuschakewitsch)
(ZI) [examined].
Bombus Semenovi Morawitz, 1886: 198. Lecto-
type queen by designation of Podbolotskaya
(in press), CHINA: Qinghai, valley of the
Huang He [= Yellow River], 13500 ft [4100 m]
(Prshewalski) (Z1) [examined]. Syn. n.
Bombus (Subterraneobombus) duanjiaoris Wang,
1982: 444. Holotype worker by original desig-
nation [not seen]. Paratype worker, CHINA:
Xizang [= Tibet], Rutog, 5100-5400 m, 23.viii.
1976 (Huang) (IZ) [examined]. Syn. n.
TYPE MATERIAL. Note. Morawitz described B.
oberti from males from [p. 240] ‘Im Hochgebirge bei
Wernoye’. At the end of this description he noted
that a queen, which he also described, was most
probably of the same species (p. 240: “Als Weib-
chen gehort h6échst wahrscheinlich zu vorstehend
beschriebenem Mannchen folgendes.’). How-
PAULH. WILLIAMS
ever, because this specimen was only doubtfully
included in the taxon, it cannot be considered
to be a syntype. This is unfortunate, because
Skorikov (1931) subsequently used the term
‘Typus’ in reference to a ‘defekte’ queen of
Morawitz’s B. oberti from ‘Vernyj’ [= Alma Ata,
in the Tien Shan region], which might otherwise
have been taken to be a valid lectotype designa-
tion (Art. 74b). Skorikov also mentions four
males with the same locality labels “Vernyj’, but
states that this species was never collected there
again. The existence of several syntype males is
indicated in the original description of B. oberti by
the range of size measurements. I have seen a
male from the ZI collection that agrees with the
original description, carries a label ‘Werkoye /
Oberti’ and lacks the right antennal flagellum,
which is designated as lectotype (Art. 74a).
AFFINITIES. Within the subgenus Sibiricobombus
(see the comments on B. asiaticus), B. oberti
shares with B. morawitzi (see Skorikov, 1931: fig.
28) a constriction of the recurved hook of the
penis valve head that is shown to a lesser extent by
many species of the subgenus Melanobombus
(Figs 79-85) (?synapomorphy of B. morawitzi +
B. oberti + Melanobombus). But only B. mora-
witzi and B. oberti share the strongly curved form
of this narrowed hook (Fig. 79) (?synapomorphy).
B. morawitzi (of the ‘morawitzi-Gruppe’ of
Tkalci, 1969a, 1974a, in part) is known from the
Tien Shan (Morawitz, 1883; Skorikov, 1931;
BMNH), the Pamir (Reinig, 1930, 1934; Skorikov,
1931; BMNH) and the Hindu Kush ranges (Reinig,
1940; Tkalct,, 1969a). Unlike B. oberti, the males
have greatly enlarged eyes relative to the females.
Another species that may be closely related has
been recorded from just across the Tibetan border
from Ladakh by Wang (1982: Shazia). Friese
(1905) described some females from Qinghai and
Gansu that have the pubescence of terga I-III
yellow and of terga IV-VI black, under the name
B. flaviventris. I have seen one of Friese’s queens
labelled ‘Kukunor’ (MNHU), but not the putative
male he described later (Friese, 1909) under
the same name. Richards (1930) described a
series of females from southern Tibet under the
name B. (Subterraneobombus) flaviventris subsp.
ochrobasis. Friese’s queen (MNHU) and all of
Richards’s material in the BMNH appears to
belong to the subgenus Sibiricobombus, although
no males are present from which to confirm this.
These females have the malar area extensively
punctured, the lateral ocellus is separated from
the dorsal margin of the compound eye by less
than two ocellar diameters, the outer surface of
the hind tibia is coarsely sculptured, the many
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
short hairs at the base of the hind basitarsus
extend onto the outer surface, which has a particu-
larly dense covering of short branched hairs, and
tergum VI has a shallow apical notch and a
subapical rounded boss. In all of these characters
they resemble females of B. morawitzi most
closely, but differ in that the clypeus is nearly
smooth, with only a few, very widely-spaced
punctures and the subapical boss of tergum VI is
rounded rather than V-shaped. B. flaviventris has
also been recorded from Tibet by Wang (1982)
and from Qinghai by Skorikov (1931) and Panfilov
(1957).
B. oberti is morphologically very distinct within
the subgenus Sibiricobombus, both in the charac-
teristics of the female (see the key) and in the form
of the male volsella, which has a produced exterio-
apical corner but is broadly rounded interio-
apically with a very reduced interio-apical process
(Fig. 159; Reinig, 1930: fig. 19; Skorikov, 1931:
fig. 30). Reinig (1930) separated this species from
the other species of the subgenus Sibiricobombus
in the monobasic subgenus Obertobombus (the
‘oberti-Gruppe’ of Tkalct, 1974a) (the identity of
Reinig’s material was discussed by Skorikov,
1931; Reinig, 1934), although this name has since
been regarded as a synonym of Sibiricobombus
(e.g. Richards, 1968; Ito, 1985). Any further
nomenclatural action is considered undesirable at
least until the males of B. flaviventris and B.
tanguticus (see the comments on B. simillimus)
are available to elucidate the relationships of
the oberti-group to the species of the subgenus
Melanobombus.
Skorikov (1931) found scarcely any difference
between morphometric characters of B. oberti
and B. semenovi. I can find no evidence that these
nominal taxa are not parts of the same species. A
worker from just across the Chinese border from
Nimaling in western Tibet (IZ), which was des-
cribed under the name B. duanjiaoris by Wang
(1982), has a broad black band on tergum III like
queens of B. oberti from Qinghai (ZI). The
Tibetan worker has a shorter antennal segment 3
than for these queens, and the punctures of the
malar area are weaker. In these characters, as
well as in the colour pattern of the pubescence,
it resembles the queens of B. tanguticus from
southern Tibet (see the comments on B. simil-
limus), although they have the clypeus much
more coarsely sculptured, the labral tubercles are
strongly pointed and only narrowly separated,
and the oculo-malar distance is distinctly shorter
than the breadth of the mandible at its base.
Otherwise B. duanjiaoris is closely similar to the
queens of B. oberti from Ladakh and Qinghai in
characters such as the pubescence of the hind
93
basitarsus. Thus from the evidence available at
present, B. duanjiaoris is most likely to be part of
the same species as B. oberti, despite the small
morphological differences, which may be related
allometrically to body size. Wang (1982: 447)
described another queen from western Tibet
under the name B. zhadaensis as ‘closely similar’
to the type specimens of B. duanjiaoris, but with
tergum III red (not seen). Individuals of this
nominal taxon may also be conspecific with B.
oberti.
DISTRIBUTION. B. oberti is a central Asian and
Tibetan species (Map 49). It is known from the
Tien Shan (Morawitz, 1883; Skorikov, 1931; ZI),
the Pamir (Reinig, 1930, 1934; Panfilov, 1957;
PW), Kashmir, Tibet (Wang, 1982; IZ) and
Qinghai (Morawitz, 1886; Panfilov, 1957; ZI).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. oberti is recorded only from the Zanskar
mountains, in the high, arid alpine steppe (Map
50): 18 queens, Nimaling plain terminal moraine,
4800 m, 15—24.vii.1980 (Williams) (BMNH, IZ,
PW).
VARIATION WITHIN KASHMIR. The queens from
Nimaling generally have the black pubescence of
tergum III largely replaced by orange-red, so that
only a very few black hairs remain at the sides
(Fig. 392). At most the black pubescence on
tergum III is intermixed with the orange-red,
except in a narrow, basal band (2/18 queens, Fig.
393).
This species is similar in colour pattern to the B.
himalayanus (Figs 242 & 243), B. kashmirensis
(Figs 295-300) and B. ladakhensis (Figs 394-396)
that are known to occur with it. It is easily
recognised by its large size and by the characters
given in the key.
FOOD PLANTS. (Leguminosae) Caragana_ versi-
color (Wallich) Benth.
Subgenus MELANOBOMBUS Dalla Torre
Melanobombus Dalla Torre, 1880: 40 (as a sub-
genus of Bombus Latreille). Type species: Apis
lapidaria Linnaeus, 1758: 579 = Bombus lapi-
darius (Linnaeus), by subsequent designation
of Sandhouse (1943: 569).
Lapidariobombus Vogt, 1911: 58 (as a subgenus
of Bombus Latreille). Type species: Apis lapi-
daria Linnaeus, 1758: 579 = Bombus lapidarius
(Linnaeus), by subsequent designation of
Sandhouse (1943: 562). Synonymised with
Melanobombus Dalla Torre by Milliron (1961).
Kozlovibombus Skorikov, 1922: 152. Type
94
species: Bombus kozlovi Skorikov, 1910b: 413
[= Bombus keriensis Morawitz] in the sense
of Skorikov, 1922 [based on misidentified males
= Bombus pyrosoma Morawitz, see Reinig
(1934: 169), requiring designation by the ICZN
(see Note below)], by subsequent fixation of
Sandhouse (1943: 561). Synonymised with
Lapidariobombus Dalla Torre by Bischoff
(1936).
Kozlowibombus Skorikov; Bischoff, 1936: 10 (as
a subgenus of Bombus Latreille). Unjustified
emendation (Art. 33b(i, iii).
Lapidariibombus Vogt; Skorikov, 1938a: 145.
Unjustified emendation (Art. 33b(i, i1i)).
Tanguticobombus Pittioni, 1939c: 201 (as a sub-
genus of Bombus Latreille). Type species:
Bombus tanguticus Morawitz, 1886: 200, by
original designation. Synonymised with Melano-
bombus Dalla Torre by Richards (1968).
Note. Application of Kozlovibombus is a matter
that should mandatorially be referred to the ICZN
(Art. 70b). Since this name is treated here as a
synonym of Melanobombus (a position that is not
affected by either interpretation of the type
species), I intend to take no action.
For a general description of both sexes see
Richards (1968).
Bombus (Melanobombus) ladakhensis
Richards
(Figs 80, 120, 160, 200, 394-396, Maps 51 & 52)
Bombus (Lapidariobombus) rufofasciatus var.
ladakhensis Richards, 1928b: 336 (see Note 1
below). Holotype queen [not a worker] by
monotypy (see Note 2 below), INDIA: Kashmir,
Ladakh, Chushul, vi.1925 (Meinertzhagen)
(BMNH) [examined].
Bombus (Lapidariobombus) rufofasciatus var.
phariensis Richards, 1930: 642 (see Note 1
below). Holotype queen by original designa-
tion, CHINA: Xizang [= Tibet], Phari to
Gyangze, 13000-15000 ft [4000-4600 ml, vi.
1904 (Walton) (BMNH) [examined]. Change
of status to Pyrobombus ladakhensis subsp.
phariensis (Richards) by Tkalct (19745). Syn. n.
Bombus variopictus Skorikov, 1933b: 248. Lecto-
type by designation of Podbolotskaya (in press)
[not seen]. Paralectotype queen, CHINA:
Qinghai, valley of the Huang He [= Yellow
River], 13500 ft [4100 m] (Prshewalski) (ZI)
[examined]. Synonymised with Pyrobombus
ladakhensis subsp. phariensis (Richards) by
Tkalct (19746). Syn. n.
Bombus variopictus subsp. bianchii Skorikov,
1933b: 248. Syntype queen and worker, INDIA:
PAULH. WILLIAMS
Kashmir, Ladakh, Rupshu (ZI) [not seen].
Synonymised with Pyrobombus ladakhensis
subsp. ladakhensis (Richards) by Tkalct (19745).
Bombus (Pratobombus) reticulatus Bischoff,
1936: 7. Lectotype queen by designation of
Tkalct (1974: 336) [not seen]. Two paralecto-
type workers, CHINA: Gansu, Min Shan,
‘Drakana’, 3100-4000 m, 27 & 29.vii.1930
(MNHU) [examined]. Synonymised with Pyro-
bombus ladakhensis subsp. phariensis (Richards)
by Tkalct (1974b). Syn. n.
Bombus (Lapidariobombus) rufo-fasciatus [subsp.]
ladakhensis Richards; Tkalcu, 1961: 353.
Pyrobombus (Melanobombus) ladakhensis
(Richards); Tkalct,, 1974b: 335.
NOMENCLATURE. Note 1. Richards’s use (19285:
333 etc.; 1930: 634 etc.) of the term ‘Subsp.’ in
both papers is taken to indicate that infrasub-
specific rank is meant for B. rufofasciatus var.
ladakhensis and for B. rufofasciatus var. pharien-
sis. However, these names have subsequently
been treated as names in the species group by
Tkalct’ (1961, 1974b), so Richards’s taxa are
therefore deemed to be of subspecific status (Art.
45g(ii)(1)).
TYPE MATERIAL. Note 2. Richards’s description of
B. rufofasciatus var. ladakhensis specifies that a
single female was examined. A single queen in the
BMNH collection agrees with the original descrip-
tion and carries the data quoted, together with a
label ‘B. rufofasciatus. Sm. / var. ladakhensis
Richards / female. type’ in handwriting identical
to that of Richards. I believe that this is the single
specimen on which the original description is
based and regard it as the holotype (Art. 73a(ii)).
AFFINITIES. The species of the subgenus Melano-
bombus share a reduction in the outer ridge of the
penis valve head (Figs 80-85) and a shortened
form of the gonostylus with a reduced interio-
basal process (Figs 200-205) (synapomorphies,
see the comments on B. asiaticus and B. oberti).
Females often have short, branched hairs below
the postero-proximal corner of the hind basitarsus
(e.g. Fig. 229), although these hairs are not as
long, erect or numerous as for species of the
subgenus Sibiricobombus (Figs 227 & 228, see the
key, couplet 11).
Within the subgenus Melanobombus, the
species of the /apidarius-group (the ‘lapidarius-
Gruppe’ of Tkalct, 1974b, 1989) share a further
reduction of the interio-basal process of the gono-
stylus to a small, rounded knob (Figs 200-202)
(?synapomorphy). In contrast, the species of the
rufofasciatus-group share a further reduction of
the apex of the gonostylus (Figs 203-205) (syna-
pomorphy) (see the key, couplet 24).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Within the /apidarius-group, all of the species
with the exception of B. ladakhensis share a much
narrowed form of the penis valve shaft, in lateral
aspect (Figs 120-122), and a narrower interio-
apical process of the volsella (Figs 160-162)
(synapomorphies). Males of B. ladakhensis differ
from those of all other species of the subgenus
Melanobombus by the recurved hook of the penis
valve head, which is broadly fused to the shaft
(Figs 80-82) (autapomorphy).
A queen of B. ladakhensis from Uttar Pradesh
(BMNH) shares the yellow colour pattern with
individuals from Kashmir and from neighbouring
parts of the far west of Tibet (Wang, 1982). I have
seen much more material from southern Tibet, for
which the yellow of the pubescence is usually
replaced by cream or grey-white (described
under the name B. rufofasciatus var. phariensis
Richards). This grey-white colour pattern re-
sembles B. rufofasciatus. Skorikov (1933b) in-
dependently redescribed individuals with the
yellow colour pattern under the name B. vario-
pictus subsp. bianchii and redescribed those with
the white colour pattern under the name B.
variopictus s.str. Individuals with the white colour
pattern were again redescribed shortly afterwards
under the name B. reticulatus by Bischoff (1936).
All of these females are closely similar in morph-
ology and are considered to be parts of the same
species.
DISTRIBUTION. B. ladakhensis is a Tibetan species
(Map 51). It is known from Kashmir, Uttar
Pradesh (BMNH), Nepal (Tkalct, 1974b; NMS),
Sikkim (BMNH), Tibet (Richards, 1930; Wang,
1982; BMNH, MNHU), Qinghai (Morawitz,
1886; Skorikov, 1933b; Wang, 1982; ZI) and
Gansu (Skorikov, 1933b; Bischoff, 1936; MNHU).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. ladakhensis is recorded from the Zanskar and
Ladakh ranges, in the high, arid alpine steppe
(Map 52): 3 queens, 1 male, from 3 localities
3600-4800 m (BMNH, PW).
VARIATION WITHIN KASHMIR. The queen from
Chushul has the pale bands of the thorax and of
terga I-II yellow (Fig. 394). The hairs of tergum
III are orange, whereas those of tergum IV are
orange with white tips and those of tergum V are
white. There are few black hairs on these terga
and those that are present are mainly on the
lateral parts of tergum III. The two queens from
Nimaling share a similar yellow pattern, but the
longer hairs of terga IV-V are pinkish brown to
black basally with white tips, intermixed with
many black hairs (Fig. 396). Black hairs are also
more numerous for these two individuals on the
95
apical part of tergum II and laterally on tergum
III. The male has orange-red pubescence on terga
III-V (Fig. 395).
This species is similar in colour pattern to some
of the B. himalayanus (Figs 242 & 243), B.
kashmirensis (Figs 295-300) and B. oberti (Figs
392 & 393) that are known to occur with it. It is
easily recognised by the mixture of grey-white and
black hairs, rather than orange-red hairs, on terga
IV-VI.
FOOD PLANTS. (Leguminosae) Caragana_ versi-
color (Wallich) Benth.; (Labiateae) Mentha
longifolia (L.) Hudson.
Bombus (Melanobombus) semenovianus
(Skorikov)
(Figs 81, 121, 161, 201, 229, 331, 397-402, Maps
53 & 54)
Lapidariobombus semenovianus Skorikov, 1914a:
127. Lectotype by designation of Podbolotskaya
(in press) [not seen]. Paralectotype queen,
INDIA: Kashmir, Ladakh, Stakmo pass, 13.vii.
1912 (Jacobson) (Z1) [examined].
Bombus (Lapidariobombus) lapidarius subsp.
problematicus Bischoff, 1935: 255. Holotype
worker by monotypy, INDIA: Kashmir,
Ladakh, Lamayuru Gompa, 3300 m, 1.viii.
1930 (7MNHUV) [not seen]. Synonymised with
Bombus semenovianus (Skorikov) by Reinig
(1940).
Bombus semenovianus (Skorikov); Reinig, 1935:
S22.
AFFINITIES. Within the subgenus Melanobombus,
B. semenovianus belongs to the lapidarius-group
of species (see the comments on B. ladakhensis).
Within the /apidarius-group, B. semenovianus
may be most closely related to B. ladakhensis (see
the comments on B. ladakhensis). Both of these
species have the hook of the penis valve head
elongated and recurved to the shaft at an angle of
less than 45° (Figs 80 & 81). All the other species
of the /apidarius-group have the penis valve head
slightly reduced in size (smallest for B. lapidarius)
with the recurved hook forming an angle with the
shaft of 45° or more (Fig. 82) (?synapomorphy).
Males of B. semenovianus can be distinguished
from males of all the other species of the
lapidarius-group, including B. ladakhensis, by
their enlarged eyes (relative to those of the
females) and by the shape of the gonocoxite,
which is nearly parallel-sided, rather than con-
stricted, just proximal to the apex. The females of
B. semenovianus have the ocello-ocular area more
densely punctured than for the other species of the
96
lapidarius-group, with a continuous, broad band
of fine punctures along the eye margin.
Bischoff (1935) apparently redescribed B. seme-
novianus under the name B. lapidarius subsp.
problematicus from a single worker from Ladakh.
DISTRIBUTION. B. semenovianus is a central Asian
species (Map 53). It is known from the Hindu
Kush (Reinig, 1940; Richards, 1951; Tkalci,
1969a; BMNH), Pakistan (Frison, 1935; BMNH,
PW) and Kashmir.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. semenovianus is recorded from the Hindu
Raj, Karakoram, Ladakh, Zanskar and Great
Himalaya ranges, in dry subalpine steppe and
in high, subtropical semidesert (Map 54): 72
queens, 202 workers, 368 males, from 21 localities
2400-3700 m (BMNH, NR, PW, ZI, ZS).
A colony was found near Dras (3200 m, 10.viii.
1986). The nest entrance was under the edge of
the corrugated iron roof of a hut.
VARIATION WITHIN KASHMIR. This species shows
obvious variation only in the extent of the replace-
ment of black by yellow in the pubescence of terga
I-II (Figs 397-402). Extensive yellow on these
terga is very common among males from all
localities (Figs 399 & 402), but among the females
it is only shown by a few queens from Gilgit (3/51
queens have an obvious yellow band on tergum I,
Fig. 397).
This species is similar in colour pattern to some
of the B. avinoviellus (Fig. 232), B. marussinus
(Figs 254-257), B. branickii (Fig. 265), B. sub-
typicus (Figs 319-324), B. biroi (Figs 335-337) and
B. asiaticus (Figs 371-376) that may occur with it
at some localities. It can be recognised by the
combination of the presence of many punctures in
the ocello-ocular area of the head, the shining
outer surface of the hind tibia and the lack of many
short hairs at the base of the hind basitarsus (Fig.
229, see the key, couplet 11). The males can be
recognised by their short antennae, by their
slightly enlarged eyes and by the recurved hooks
on the heads of the penis valves (Fig. 81).
FOOD PLANTS. (Leguminosae) Melilotus officinalis
(L.) Pallas; (Compositae) Echinops cornigerus
DC.; (Labiateae) Stachys tibetica Vatke, Mentha
longifolia (L.) Hudson, Perovskia abrotanoides
Karelin.
Bombus (Melanobombus) keriensis
Morawitz
(Figs 82, 122, 162, 202, 403-416, Maps 55 & 56)
Bombus keriensis Morawitz, 1886: 199. Lectotype
queen by designation of Podbolotskaya (in
PAULH. WILLIAMS
press), CHINA: Xinjiang, Kunlun Shan,
mountains near Yutian [= Keriya], 9000 ft
[2700 m] (Prshewalski) (Z1) [examined].
Bombus separandus Vogt, 1909: 58, 61 [footnote
1]. LECTOTYPE queen by present designa-
tion (see Note 3 below), CHINA: Xinjiang,
Borohoro Shan (ITZ) [examined]. Change of
status to Bombus keriensis f.g. separandus Vogt
by Reinig (1935). Syn. n.
Bombus kohli Vogt, 1909: 41, 61 [footnote
2]. Syntype worker (see Note 4 below),
MONGOLIA: north, no further data, 1892
(Leder) (ITZ) [examined]. Junior primary
homonym of Bombus kohli Cockerell, 1906 [=
B. morio (Swederus)]. Syn. n.
Bombus kohli var. postzonatus Vogt, 1909: 61
[footnote 2]. LECTOTYPE queen by present
designation (see Note 5 below), MONGOLIA:
north, no further data, 1892 (Leder) (ITZ)
[examined]. Syn. n.
Bombus kozlovi Skorikov, 1910b: 413. Replace-
ment name for B. kohli Vogt. Change of status
to Bombus keriensis f.g. kozlovi Skorikov by
Reinig (1935). Syn. n.
Bombus (Lapidariobombus) separandus incer-
toides Vogt, 1911: 58 [by indication of Vogt,
1909: 61 (footnote 1)] (see Note 1 below).
Holotype queen by monotypy (see Note 6
below), MONGOLIA: north, no further data,
1892 (Leder) (ITZ) [examined]. Syn. n.
Bombus lapidarius var. tenellus Friese, 1913: 86. 2
syntype females and 1 syntype male, U.S.S.R.:
?Zapadnyy Sayan, ‘Arasagun-gol’ (Staudinger)
(MNHVU) [not seen]. Synonymised with Lapid-
ariobombus incertoides (Vogt) by Skorikov
(1931). Syn. n.
Lapidariobombus separandus subsp. meridialis
Skorikov, 1914a: 127. Holotype queen by
monotypy (see Note 7 below), INDIA: Kash-
mir, Sind valley above Sonamarg, 2400-3000 m,
9-10.vi.1912 (Jacobson) (ZI) [not seen]. Syn.
n.
Bombus (Lapidariobombus) tenellus var. alpiva-
gus Richards, 1930: 639 (see Note 2 below).
Holotype queen by original designation,
CHINA: Xinjiang, Taghdumbash, 14000 ft
[4300 ml], 18.vi.1913 (Hingston) (BMNH)
[examined]. Syn. n.
Bombus (Lapidariobombus) kozlowi Skorikov;
Bischoff, 1936: 9. Unjustified emendation (Art.
33b(i, iii)).
Pyrobombus (Melanobombus) keriensis subsp.
karakorumensis Tkalct, 1989: 57. Holotype
queen by original designation, PAKISTAN:
Gilgit, Banidas 36°11’N 74°33’E, 2600 m,
30.vi-2.vii.1959 (Lobbichler) (ZS) [examined].
Syn. n.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
NOMENCLATURE. Note 1. Vogt’s (1911: 58) use of
the name incertoides under ‘Meine Separandus-
tiere ...’, for a female that he had described
previously (1909: 61 [footnote 1]), is deemed to be
of subspecific rank even though he did not state its
rank explicitly (Art. 45f(i)). Skorikov subse-
quently (1922) treated incertoides as an available
name and adopted it as the name of a species (Art.
45g(i1)(1)).
Note 2. Richards’s use (1930: 634) of the term
‘Subsp.’ elsewhere in his paper could be taken to
indicate that infrasubspecific rank is meant for B.
tenellus var. alpivagus (despite the comment
‘probably a geographical race’). However, this
has subsequently been treated as a name in the
species group by Reinig (1935: 333 [footnote 1:
‘f.g. means forma geographica (=subsp.)’ ]), so
Richards’s taxon is deemed to be of subspecific
status (Art. 45f(ii)).
TYPE MATERIAL. Note 3. Vogt’s original descrip-
tion of B. separandus mentions at least one female
and one male from the Alai Mountains and from
‘Siebenstromgebiet’. Vogt subsequently (1911:
58) refers to the same material ‘aus der Umgebung
des Festungswerks Narynj und dem Boro-
Chorogebirge [= Borohoro Shan] im Sieben-
stromgebiet’. A single queen in the ITZ collection
agrees with the original description and carries a
label (1) ‘Boro-Chorogeb / Tisilikau’; (2) a white,
handwritten label ‘separandus O.V.’; (3) a red
printed label ‘Type’. This specimen is designated
as lectotype (Art. 74a).
Note 4. Vogt’s description of B. kohli mentions
two queens (one of B. kohli var. postzonatus) and
a number of workers. However, the description is
specified to have been made primarily from a
queen that had the pubescence of tergum III
black. I have seen a single worker from the ITZ
collection that agrees with this description and
carries a label ‘N. Mongolei/ Leder 92’, and a red
printed label “Type’. I believe that this is one of
the workers referred to at the end of footnote 2
and regard it as a syntype. It would be preferable
to designate as lectotype the queen on which Vogt
concentrated for the original description, when
this queen is found.
Note 5. Vogt described B. kohli var. postzona-
tus as like the type, the queen of B. kohlis.str., so
he probably based his description on the second of
the two queens that he mentions at the end of
footnote 2, rather than on any of the workers. A
single queen in the ITZ collection agrees with
the original description and carries a label (1)
‘N. Mongolei/ Leder 92’; (2) a white, handwritten
label ‘kohli, ab postzonatus’; (3) a white hand-
written label ‘13’; (4) a red printed label ‘Type’.
97
This specimen is designated as lectotype (Art.
74a).
Note 6. Vogt’s description of B. separandus
incertoides specifies that only one specimen (refer-
ring to a second queen described separately under
the name B. separandus in 1909: 61 [footnote 1])
was available. A single queen in the ITZ collec-
tion agrees with the original description and
carries a white, handwritten label ‘separandus- /
incertoides-’ and a red, printed label ‘Type’. I
believe that this is the single specimen on which
the original description is based and regard it as
the holotype (Art. 73a(ii)).
Note 7. Skorikov’s description of Lapidario-
bombus separandus subsp. meridialis specifies
that only a single queen was available. If a single
queen with the appropriate data can be found then
this should be regarded as the holotype (Art.
73a(ii)).
AFFINITIES. Within the subgenus Melanobombus,
B. keriensis belongs to the lapidarius-group of
species (see the comments on B. ladakhensis).
Within the /apidarius-group, B. keriensis shares
with B. incertus Morawitz, B. lapidarius (Lin-
naeus) and B. sichelii Radoszkowski a slight
reduction of the recurved head of the penis valve
(Fig. 82) (?synapomorphy, see comments on B.
semenovianus). All of these species, except B.
incertus, have the volsella narrowed near its mid
point in ventral aspect (Fig. 162) (synapomorphy
of B. lapidarius + B. keriensis + B. sichelii). B.
incertus and B. lapidarius apparently do not occur
east of the Caspian Sea (distributions summarised
by Reinig, 1935: chart 3, 1939: fig. 15), except for
a couple of records of B. lapidarius from the
northern U.S.S.R. (Panfilov, 1981: map 92).
The closest relative of B. keriensis is probably
B. sichelii Radoszkowski [the correct original
spelling of B. sicheli of authors]. Unlike B.
lapidarius, females of these two species share an
apical notch on tergum VI and the males appear to
share a slight displacement of the weak ventro-
basal angle of the penis valve nearer to the mid-
point of the shaft (Fig. 122). B. sichelii has a broad
Eurosiberian distribution, although this is frag-
mented in Europe among the southern mountains
(mapped in Reinig, 1935: charts 2 & 5, 1939: fig.
4). The distributions of the two species overlap
in the Altai (Reinig, 1935; BMNH), Mongolia
(Reinig, 1935; Tkalct, 1974a; BMNH, PW) and
Gansu (Bischoff, 1936). Unlike B. sichelii, the
males of B. keriensis have the apex of the penis
valve head slightly less acutely pointed (Fig. 82)
and the interio-apical process of the volsella is
narrower, with a less prominent interio-basal
corner (Fig. 162). There is considerable variation
98
in the sculpturing of the ocello-ocular area of the
females, but females of B. keriensis often differ
from those of B. sichelii in that the band of fine
punctures is more distinctly interrupted, so that
there is a narrow shining area with only large
punctures adjacent to the eye margin.
Skorikov (1931) and Reinig (1935) both sug-
gested that the cream individuals that were des-
cribed under the name B. separandus Vogt,
which often also have pale hairs on the face and a
pale fringe in the pubescence of tergum III
(=‘ciliated’, e.g. B. kohli var. postzonatus Vogt
and Lapidariobombus separandus subsp. meri-
dialis Skorikov), are actually conspecific with the
more distinctly yellow and often unciliated B.
keriensis (also described under the names B. kohli
Vogt, B. lapidarius var. tenellus Friese, B. tenellus
var. alpivagus Richards and Pyrobombus kerien-
sis subsp. karakorumensis Tkalct). Reinig (1935)
also included the unciliated and white queen
described under the name B. separandus incer-
toides Vogt in this group. Individuals with the very
pale and ciliated colour patterns occur throughout
much of the total distribution of this group of
nominal taxa, but are most frequent in the region
of the Pamir (Reinig, 1935: chart 4, 1939: fig. 23).
The inference that they are all likely to be
interbreeding as parts of a single population and a
single species is supported by the apparently
continuous variation between the two extreme
colour patterns among the material from the
Hindu Raj range (BMNH).
A nest from the Pamir was described by
Bischoff (1931).
DISTRIBUTION. B. keriensis is a widespread, but
primarily central Asian and Tibetan species (Map
55). It is known from Mongolia (Vogt, 1909,
1911; Tkalct, 1974a; BMNH, ITZ, PW), the Altai
(Morawitz, 1880; Friese, 1913; Skorikov, 1931),
the Tien Shan (Morawitz, 1880; Vogt, 1909,
1911; Skorikov, 1931; Panfilov, 1957; BMNH,
ITZ, TM), the Pamir (Richards, 1930; Reinig,
1930, 1934, 1935; Bischoff, 1931; Skorikov,
1931; BMNH), the Hindu Kush (Reinig, 1940;
Richards, 1951; Tkalct, 1969a; BMNH), Pakistan
(BMNH, PW), Kashmir, Xinjiang [Kunlun Shan]
(Morawitz, 1886; TM, ZI), Tibet (Richards, 1930;
Wang, 1982; BMNH), Qinghai (Morawitz, 1886;
Panfilov, 1957; Tkalct, 1961; Wang, 1982; ZS)
and Gansu (Bischoff, 1936). In the west it is
also known from the Elburz (Skorikov, 1931;
BMNRH), Turkey and the Caucasus (Reinig, 1935;
BMNRH). Reinig (1935: chart 2, 1939: figs 4 & 23)
mapped a summary of this distribution. A similar
disjunct distribution between the Hindu Kush
ranges on the one hand and the Elburz and
PAULH. WILLIAMS
Armenian highlands on the other is shown by B.
melanurus.
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. keriensis is recorded from the Hindu Raj,
Zanskar, Great Himalaya and Pir Panjal ranges,
in alpine scrub and steppe (Map 56, Fig. 6): 73
queens, 155 workers, 38 males, from 18 localities
2600-5100 m (BMNH, FA, NR, PW, ZS).
VARIATION WITHIN KASHMIR. When these bees are
seen in flight, the two most obvious characters of the
colour pattern to vary are (a) the yellow or white colour
of the pale pubescence on the thorax and on terga I-
II; and (b) the presence or absence of a fringe of
pale hairs at the posterior and lateral margins of
tergum III (present in the ‘ciliated’ state).
Females from the Hindu Raj range generally
have the pale pubescence nearly white if queens,
but yellow if workers, although both are strongly
ciliated (cream-white: 49/50 queens, 0/68 workers,
no males available; strongly ciliated: 47/50 queens,
43/68 workers; Figs 403-407). In the Great
Himalaya range, workers are usually paler but
unciliated, both around the Kishanganga valley
(cream-white: 2/7 queens, 39/49 workers, 0/33
males; strongly ciliated: 0/7 queens, 0/49 workers,
0/33 males; Figs 408-410), and at the head of the
Sind valley (cream-white: no queens available,
32/33 workers, 0/3 males; strongly ciliated: 0/32
workers, 1/3 males).
Individuals from the Zanskar ranges usually
have the pale pubescence yellow and are uncili-
ated (cream-white: 1/5 queens, 1/2 workers, no
males available; strongly ciliated: 1/5 queens, 0/2
workers [more yellow and unciliated queens were
seen but not collected]; Figs 411-413) like those
from Tibet. Individuals from the Pir Panjal range
are similar, but with slightly more extensive
black pubescence on the thorax (cream-white: 0/8
queens, 0/5 workers, 0/2 males; strongly ciliated:
0/8 queens, 1/5 workers, 0/2 males; Figs 414-416).
This species is similar in colour pattern to some
of the B. himalayanus (Fig. 242), B. kashmirensis
(Figs 298-300, 305-307), B. subtypicus (Figs 317
& 318), B. biroi (Figs 340, 342, 343), B. tunicatus
queens (Fig. 350), B. asiaticus (Figs 386-388) and
B. oberti (Fig. 393) that occur with it at some
localities. Females can usually be recognised by
their combination of an apical notch on tergum VI
with a lack of many short hairs at the base of the
hind basitarsus (see the key, couplet 11). The
males are closely similar in colour pattern (Figs
410 & 416) to some of those of B. subtypicus (Fig.
318) and of B. biroi (Figs 340 & 343), which may
occur with them at some localities, but can be
recognised by the narrow recurved hooks of the
head of the penis valve (Fig. 82).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
FOOD PLANTS. Kashmir: (Ranunculaceae) Aconi-
tum hookeri Stapf; (Balsaminaceae) Impatiens
glandulifera Royle; (Leguminosae) Trifolium
repens L.; (Compositae) Cirsium falconeri
(Hook.f.) Petrak; (Gentianaceae) Swertia petio-
lata D. Don; (Scrophulariaceae) Picrorhiza
kurrooa Royle ex Benth.
Ladakh: (Leguminosae) Caragana versicolor
(Wallich) Benth.; (Labiateae) Stachys tibetica
Vatke.
Bombus (Melanobombus) simillimus Smith
(Plate 1, Figs 83, 123, 163, 203, 226, 417-421,
Maps 57 & 58)
Bombus_simillimis Smith, 18526: 48. LECTO-
TYPE queen by present designation (see Note
3 below), INDIA: north, no further data
(BMNH) [examined]. Incorrect original spell-
ing (see Note 1 below) (Art. 32c(i)), without
separate availability in this form (Art. 32d).
[Bombus similis Smith; Smith, 1854: 403. Incor-
rect subsequent spelling, unavailable name
(Art. 33c). Junior primary homonym of Bombus
similis Fabricius, 1804 [= Centris similis (F.)].]
Bombus simillimus Smith; Dalla Torre, 1896: 548.
Justified emendation (see Note 2 below) (Art.
33b(ii)).
[Bombus tonsus Skorikov, 1922: 160. Published
without description or indication, unavailable
name (Art. 12a).]
[Bombus tonsus Skorikov, 1931: 202. Published
without description or indication, unavailable
name (Art. 12a).]
Bombus terrestris var. grossiventris Friese, 1931:
303. LECTOTYPE worker by present designa-
tion (see Note 4 below), INDIA: Kashmir,
Srinagar, Shalimar, 1800 m, 7.x.1923 (Fletcher)
(MNHU) [examined]. Syn. n.
Bremus (Sibiricobombus) oculatus Frison, 1933:
335. Holotype male by original designation
[p. 338], INDIA: Himachal Pradesh, Baghi,
8800 ft [2700 m], 7-8.x.1921 (Kemp) (Calcutta)
[not seen]. Syn. n.
Sibiricobombus tonsus Skorikov, 1933b: 248.
LECTOTYPE queen by present designation
(see Note 5 below), INDIA: Kashmir, Kishtwar,
Datgash, 12.vi.1910 (Trubetskoy) (ZI) [exam-
ined]. Syn. n.
Bombus (Lapidariobombus) oculatus var. haemor-
rhous Richards, 1934: 87. Holotype worker by
original designation [p. 88], INDIA: Himachal
Pradesh, Dalhousie, 7.vii.1906 (Barrow)
(BMNH) [examined]. Syn. n.
NOMENCLATURE. Note 1. Smith’s original spelling
(1852b) is ‘simillimis’, which may have been
99
intended as the superlative of similis (comment
after the original description: “This species is very
like the Lapidarius of Linn.;’). There is no evi-
dence that this spelling is incorrect from the
original publication, except (Art. 32c(i)) in its
termination (Art. 31b). Therefore the subsequent
spelling by Smith (1854), which is not an emenda-
tion (Art. 33b), is deemed to be incorrect (Art.
33c), although the termination of simillimis must
still be corrected (Art. 32d(ii)) to simillimus.
Note 2. Dalla Torre’s change in the subsequent
spelling of ‘simillimis’ can be accepted as inten-
tional (Art. 33b(i)), even though he only quotes
‘simillimus’, because of his introduction (1896:
v): ‘So ist es z.B. wissenschaftlich gewiss nicht
gerechtfertigt, wenn jeder Elementarlateiner
befahigt und — berechtigt ist, WOrter, wie laevis,
coelestis, sylvaticus zu corrigiren, ohne dass man
in wissenschaftlichen Kreisen je davon Notiz
genommen hat, und ahnlich verhalt es sich auch
mit den Geschlechtsendungen, wo auch alle
erdenklichen grammatikalischen Fehler gemacht
und fortwahrend colportirt werden.’
TYPE MATERIAL. Note 3. Smith’s description of B.
simillimus is of a queen. A queen in the BMNH
collection agrees with the original description and
bears a purple-edged label (1) ‘Lecto- / type’; (2)
‘N. / India’, reverse side ‘48 / 132’ (this accession
number refers to 73 Hymenoptera from the Boyes
collection received in 1848); (3) ‘N. China / Baily
11/7/55’; (4) ‘60-15 / E.I.C.’ (see Note 2 on B.
haemorrhoidalis); (5) ‘similis / Type Sm.’; (6)
‘Bombus / similis / Smith Trans. / Ent. Soc. 1852’;
(7) a red-edged label ‘Type’; (8) ‘B.M. TYPE /
HYM. / 17B.981’. Other Smith syntypes might
have existed and may since have become dispersed
to other collections. Therefore this queen, which
lacks both fore tarsi, the left front basitarsus and
both hind tarsi, is designated as lectotype (Art.
74a).
Note 4. Friese described B. terrestris var. grossi-
ventris from three workers. I have seen one of
these from the MNHU collection that agrees with
the original description and carries a label (1)
‘Kashmir 200ft / Shalimar / Srinagar / 7 Oct. 1923 /
Fletcher coll’; (2) a handwritten label ‘Bombus /
grossiventris / Fr. / [worker] Friese det. 25’; (3) a
printed label “Zool.Mus. / Berlin’. This specimen,
which lacks the distal joints of the left hind tarsus,
most of the flagellum of the right antenna and the
distal joints of the left antenna, is designated as
lectotype (Art. 74a).
Note 5. Skorikov’s description of Sibiricobom-
bus tonsus is of a queen. A queen in the ZI
collection from near the village of Datgash, with a
label ‘B. tonsus type’, agrees with the original
100
description. Because other Skorikov syntypes
may exist, this specimen, which has two lateral
patches of orange-brown pubescence on the
anterior part of the scutum, is designated as
lectotype (Art. 74a).
AFFINITIES. Within the subgenus Melanobombus
(see the comments on B. ladakhensis), B.
simillimus belongs to a group of species, the
rufofasciatus-group (the ‘flavothoracicus-Gruppe’
+ ‘rufofasciatus-Gruppe’ of Tkalct, 1974b), for
which the male gonostylus is shortened with dis-
tinct inner and outer apical corners (Figs 203-205)
(synapomorphy). Skorikov (1922) recognised a
subgenus Kozlovibombus on the basis of the male
genitalia of what was probably a specimen of B.
pyrosoma s.str. (see Reinig, 1934, 1935). Females
of all of the species in this group apparently vary
in colour with size (see the comments on B.
pyrosoma) in a manner resembling that described
for the Central American B. (Pyrobombus)
ephippiatus Say by Owen & Plowright (1980) and
for the Himalayan B. (Festivobombus) festivus by
Ito et al. (1984). This variation is not only in the
extent of the pattern of replacement of black by
pale hairs in the pubescence, but smaller indivi-
duals often have at least one additional colour.
This is usually yellow or brown in the pubescence
on tergum II. Hence queens, workers and males
may appear strikingly different (see Plate 1).
Within the rufofasciatus-group, queens of B.
simillimus share with those of B. tanguticus
Morawitz a distinct and complete sulcus obliquus
of the mandible. They also share the fine,
branched hairs on the outer surface of the hind
tibia (Fig. 226) and the dense (often black) short
pubescence on the hind basitarsus. B. tanguticus is
known from Sikkim and Tibet (Richards, 1930;
BMNH) and Qinghai (Morawitz, 1886). It could
also be present in the high mountain ranges of
north-eastern Kashmir (unconfirmed record from
Kashmir by Skorikov, 19335). Queens of B.
tanguticus are very easily distinguished from those
of B. simillimus by their acutely and deeply
notched tergum VI, which is straight or only
shallowly indented for B. simillimus; by their
labral furrow, which is only one-quarter of the
total breadth of the labrum, as opposed to nearly
half of the breadth for B. simillimus; by their
oculo-malar distance, which is about equal to the
breadth of the mandible at the base, whereas it is
much longer for B. simillimus; by their ocello-
ocular area, which is much less densely punctured;
by their clypeus, whichis less strongly swollen, but
more coarsely punctured; and by their nearly clear
(subhyaline) wings. B. tanguticus was considered
sufficiently distinctive by Pittioni (1939c) to
PAULH. WILLIAMS
warrant the description of a monobasic subgenus,
Tanguticobombus. The male remains undescribed
and apparently unknown, so that the precise
relationships of this species are difficult to resolve
at present.
Within the rufofasciatus-group, some females
have a straight or only shallowly indented apex to
tergum VI as well as a distinct sulcus obliquus of
the mandible. Queens and workers with these
characters from the western Himalaya were des-
cribed under the names B. simillimus and B.
terrestris var. grossiventris respectively. They have
not been associated as castes of the same species
because of their very different colour patterns.
Queens of B. terrestris var. grossiventris and
workers and males of B. simillimus were un-
known. But within the rufofasciatus-group from
the Himalaya, only these nominal taxa share the
uniformly red pubescence of terga IV-V and the
intensively darkened (infuscated) wings. One of
the older queens (BMNH) has the pubescence of
terga I-II distinctly lighter than on tergum III, so
that it is dark brown rather than black (although
the brown is still not as light as for B. terrestris var.
grossiventris). The queens have particularly many
of the fine, short, branched hairs on the outer
(corbicular) surface of the hind tibia (Fig. 226),
although these hairs are also present for some of
the workers. These hairs are seldom numerous in
this position among bumble bees, except for
species of the subgenus Psithyrus. Among the
social species, the presence of these hairs may
be associated with particularly large body size
because they are otherwise very numerous for
B. (Melanobombus) tanguticus Morawitz, B.
(Rufipedibombus) rufipes Lepeletier and B.
(Rufipedibombus) eximius Smith, although they
are also present for some queens of the European
B. lapidarius. Almost certain confirmation that B.
simillimus (queens, Fig. 417), B. terrestris var.
grossiventris (workers, Fig. 418) and males (Fig.
419) described under the name Bremus oculatus
by Frison (1933) are the castes and sexes of a
single species comes from a colony discovered
near Harwan, in the Vale of Kashmir (see Plate 1
and the description of material examined from
Kashmir). The alternative interpretation is that a
colony of one species, for which the queen must
be unknown, had been usurped by a queen of
another species, which is closely similar in morph-
ology to the workers of the first queen.
Males of B. simillimus can be distinguished
from those of the other taxa of the rufofasciatus-
group by their greatly enlarged eyes; by the large,
sub-rectangular gonostylus, which is scarcely
reduced and retains a narrow interio-basal process
(Fig. 203) (plesiomorphic within the rufofasciatus-
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
group); and by the twisted interio-apical process
of the volsella (Fig. 163). The form of the
gonostylus is most similar to that of B. richardsiel-
lus (Tkalct, 1968a: figs 84-86; see the comments
on B. pyrosoma).
A queen with two light patches on the anterior
dorsum of the thorax was described under the
name Sibiricobombus tonsus by Skorikov (19335).
Two workers were redescribed under the name B.
oculatus var. haemorrhous by Richards (1934).
These individuals are closely similar in morph-
ology to those described under the names B.
simillimus and B. terrestris var. grossiventris
respectively and are considered to be part of the
same species. Females (probably workers) of
another nominal taxon from Tibet (B. trilineatus),
which is closely similar to B. simillimus, have been
described by Wang (1982) from slight differences
in colour pattern (not seen).
DISTRIBUTION. B. simillimus is a west Himalayan
species (Map 57). It is known only from Himachal
Pradesh (Frison, 1933; Richards, 1934; BMNH),
Kashmir and Pakistan (BMNH).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. simillimus is recorded from the lower slopes in
the Kishanganga valley, in the Vale of Kashmir
and along tributaries to the Chenab valley, in
open scrub around the lower montane coniferous
forest (Map 58): 9 queens, 102 workers, 8 males,
from 9 localities 1600-3000 m (AB, BMNH,
MNHU, IZ, NR, PW, ZI).
I found a colony near the edge of open wood-
land near Harwan (1700 m, 10.ix.1985). A fresh
hole, 50 cm deep by 30 cm wide, in the bank of an
irrigation channel contained 6 queens (including
an old and very abraded individual), 39 workers
and 5 males. Returning workers found their way
into the cavity via a separate tunnel, nearly
2 m long. The remains of only 4 cocoons were
present in the cavity. I had seen bears (probably
Solenarctos thibetanus (Cuvier)) in the vicinity
earlier on the same day.
VARIATION WITHIN KASHMIR. There is little varia-
tion among the queens (Plate 1 and Fig. 417). The
number of black hairs on the thoracic dorsum of
workers does vary, but none of them has even a
weak black band between the wing bases (Plate 1
and Fig. 418). One large worker (severely abraded)
from the colony near Harwan has the hairs of
tergum I white with dark bases, while tergum II is
very dark brown, with a posterior fringe of white
hairs (Fig. 420). The males vary from having terga
I-II entirely nearly white (5/5 males, Harwan,
Plate 1 and Fig. 419), to having tergum I white and
tergum II chocolate-brown, with the exception of
101
a posterior fringe of white hairs (3/3 males,
Banihal, Fig. 421).
Queens of this species are distinctive in their
appearance. Workers are similar in colour pattern
to some of those of B. tunicatus (Figs 351 & 354),
which occur with them. Workers of B. simillimus
can usually be recognised by their chocolate-
brown pubescence of tergum I (Fig. 418), rather
than white as for B. tunicatus (Fig. 354). Males are
also similar in colour pattern to some of those of
B. tunicatus (Figs 352 & 355), or even some B.
asiaticus (Fig. 385) that occur with them. They
can be distinguished from B. tunicatus by their
enlarged eyes and from B. asiaticus by their short
antennae.
FOOD PLANTS. (Balsaminaceae) Jmpatiens glandu-
lifera Royle; (Leguminosae) unidentified pink
clover-like legume; (Compositae) Cirsium fal-
coneri (Hook.f.) Petrak, C. wallichii DC.,
Centaurea iberica Trevir ex Spengel, unidentified
yellow thistle-like composite; (Gentianaceae)
Swertia petiolata D. Don.
Bombus (Melanobombus) pyrosoma
Morawitz
(Plate 1, Figs 84, 124, 164, 204, 422-430, Maps 59
& 60)
Bombus pyrosoma Morawitz, 1890: 349. Lecto-
type queen by designation of Podbolotskaya
(in press), CHINA: ‘Gansu’, ‘Utai’, 8900 ft
[2700 m], vi.1884 (Potanin) (ZI) [examined].
Bombus pyrrhosoma Morawitz; Dalla Torre,
1896: 544. Unjustified emendation (Art.
33b(i, iii)).
Bombus flavothoracicus Bingham, 1897: 552.
Lectotype queen by designation of Tkalcu
(19746: 338), INDIA: Sikkim, Lintu, 12500 ft
[3800 m], v.1894 (Bingham) (BMNH) [exam-
ined]. Junior secondary homonym in Bombus
of Psithyrus campestris var. flavothoracicus
Hoffer, 1889 [= B. campestris (Panzer)]. Pro-
visional synonym.
Bombus miniatus Bingham, 1897: 553. Holotype
male by original designation, INDIA: Sikkim,
Lintu, 12500 ft [3800 m], v[!].1894 (Bingham)
(BMNH) [examined]. Synonymised with Pyro-
bombus flavothoracicus (Bingham) by Tkalci
(19745). Provisional synonym.
[Bombus pyrrhosoma var. canosocollaris Skorikov,
1912b: 608 [not seen]. Infrasubspecific (Art.
45g(ii)(1)), unavailable name (Art. 45e).]
Bombus friseanus Skorikov, 1933a: 62. Holotype
queen by monotypy (see Note 1 below),
CHINA: Sichuan, Songpan, above 9500 ft
102
[2900 m], 1894 (Beresovski) (ZI) [examined].
Syn. n.
Bremus (Lapidariobombus) formosellus Frison,
1934: 163. Holotype male by original designa-
tion [p. 166], TAIWAN: ‘Roeichi’, 15.ix.1924
(Shiraki & Sonan) (INHS) [examined]. Pro-
visional synonym.
Bombus (Lapidariobombus) pyrrhosoma subsp.
hénei Bischoff, 1936: 10. LECTOTYPE queen
by present designation (see Note 2 below),
CHINA: Yunnan, Lijiang, 23.v.1934 (Hone)
(MNHU) [examined]. Syn. n.
[Bombus (Lapidariobombus) pyrrhosoma f. flavo-
corbicularis Tkalct, 1961: 353 [not seen]. Infra-
subspecific (Art. 45g(1i)(1)), unavailable name
(Art. 45e).]
Pyrobombus (Lapidariobombus) wutaishanensis
Tkalct,, 1968a: 39. Holotype queen by original
designation [p. 41], CHINA: Shanxi, Wutai
Shan, 3200 m, 15.vii.1936 (Héne) (MNHU)
[examined]. Syn. n.
TYPE MATERIAL. Note 1. Skorikov’s description of
B. friseanus specifies that only a single queen was
examined. A single queen in the ZI collection
agrees with the original description and bears a
label with the data quoted. I believe that this is the
single specimen on which the original description
is based and regard it as the holotype (Art.
73a(ii)).
Note 2. Bischoff’s description of B. pyrrhosoma
subsp. hoenei shows that several females, col-
lected ‘20.4’ and ‘1.5.—23.6.34’, were examined. A
queen in the MNHU collection bears the label (1)
‘Li-kiang. (China). / Provins Nord-Yuennan. /
23.5 1934.H.Ho6ne.’; (2) ‘pyrrhosoma / hoenei n.
sp. / [female] / det. Bischoff’; (3) a red printed
label ‘Typus’; (4) ‘LECTOTYPE / Bombus /
pyrrhosoma / Tkalci det. / hénei Bischoff /
[female]’ [designation not published]; (5) “Zool.
Mus. / Berlin’. This specimen, which is complete,
is designated as lectotype (Art. 74a).
AFFINITIES. Within the subgenus Melanobombus
(see the comments on B. ladakhensis), B. pyro-
soma belongs to the rufofasciatus-group of species
(see the comments on B. simillimus).
Within the rufofasciatus-group, the known
males, other than those of B. simillimus, are
characterised by pronounced reductions (shorten-
ing) of the gonostylus and a broadening of its
interio-basal process (Figs 204 & 205) (synapo-
morphies). Apart from B. simillimus and B.
rufofasciatus (see the comments on B. rufofasci-
atus), four principal nominal taxa have been
recognised in this group. These were described
under the names Lapidariobombus richardsiellus,
B. pyrosoma, B. friseanus and B. flavothoracicus.
PAULH. WILLIAMS
Of these, B. pyrosoma, B. friseanus and B.
flavothoracicus are morphologically closely simi-
lar and differ from each other primarily in colour
pattern.
B. richardsiellus was described (Tkalci, 1968;
BMNH) from 2 queens, 3 workers and 1 male
from northern Burma and from the neighbouring
parts of Tibet. The queens are similar to the B.
friseanus from the same region, but have the
clypeus slightly more coarsely punctured. Their
white-banded colour pattern has a much more
restricted distribution of pale pubescence. The
‘allotype’ male has the exterio-apical corner of the
gonostylus more strongly reduced than for B.
simillimus (synapomorphy of B. richardsiellus +
B. pyrosoma [+ B. friseanus]| + B. rufofasciatus),
so that it is intermediate in form between the
gonostyli of B. simillimus and of B. friseanus.
Another 4 males from near the Burmese border of
Tibet (BMNH) may belong to B. richardsiellus,
but show a broad range of forms of the gonostylus,
from similar to that of the ‘allotype’, to more like
that of B. friseanus.
At a few localities in Kashmir (e.g. Mt
Apharwat [1986], Daksum), B. simillimus and B.
rufofasciatus not only occasionally occur together,
but also with another taxon of the rufofasciatus-
group. Males of B. simillimus and B. rufofasciatus
have colour patterns similar to their respective
workers, although the males have much larger
eyes. The males of the third taxon are often
predominantly yellow (Figs 424, 427, 430), unlike
the females (even when reared from the same
maternal colony, see Plate 1 and the comments on
the material examined from Kashmir), and have
eyes that are not enlarged relative to those of the
females. These yellow males differ in their mate-
searching behaviour from males of B. rufofasci-
atus and presumably from those of B. simillimus
(see the introduction on male mate-searching
behaviour, and the comments on B. rufofasci-
atus). There are no specimens with intermediate
colour patterns or morphology as evidence that
they interbreed with either B. rufofasciatus or B.
simillimus. Therefore the three taxa are believed
to represent three separate species.
B. flavothoracicus is the name that has been
used most recently for the third Himalayan taxon
of the rufofasciatus-group that is found in Kashmir
(e.g. Tkalci, 1974b). This name was used by
Bingham for a queen, which has light olive-yellow
pubescence at the front and rear of the thorax and
on tergum I. Gastral terga IV-VI have the pubes-
cence extensively white. The predominantly
yellow males were also described by Bingham,
but under the name B. miniatus. The workers
differ from the queens in that they have most of
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
tergum II dull olive-yellow or brown and most
of the pubescence of terga III-V red, which
replaces most of the white and even some of the
black hairs. This caste dimorphism may be size-
dependent, because the red pubescence is less
extensive in large workers (decreasing with in-
creasing body size in Figs 426, 428, 429) and red
may occasionally be intermixed on terga III-IV of
small queens. This nominal taxon is known from
as far east in the Himalaya as the area of the
Sikkim/Tibet/Bhutan border (BMNH), to as far
west as Kashmir. Queens of two further nominal
taxa from Tibet (B. eurythorax and B. steno-
thorax), both very similar to B. flavothoracicus,
have been described by Wang (1982) from slight
differences in colour pattern (not seen).
Material from further to the east in Yunnan and
Sichuan was described under the names B. pyr-
rhosoma subsp. hoenei by Bischoff (1936) and B.
friseanus Skorikov (1933a) respectively. Both the
queens and the workers from this region are
similar in colour pattern to the workers of B.
flavothoracicus from the Himalaya, although the
pale pubescence (including tergum II) is usually a
deeper orange-yellow and there are fewer black
hairs intermixed with it on the thorax. Thus both
castes have the pubescence of terga III-VI red,
although without any distinctly white hairs.
Workers of B. friseanus are known to occur as far
west as the lower Zangbo [= Tsangpo] valley of
Tibet and the Char Chu valley at the extreme
eastern end of the Himalaya, adjacent to Arunachal
Pradesh (BMNH, no queens available).
Material from further to the north in Gansu was
originally described under the name B. pyrosoma
s.str. by Morawitz (1890). The queens have the
thorax and tergum I black with just a very
few grey-white hairs intermixed. The pubescence
of gastral terga II-VI is predominantly red.
Morawitz also described the workers from Gansu
(listed as Var. b and Var. c), which have the
pubescence of the thoracic dorsum grey-white
with a black band between the wing bases, with
tergum I grey-white, tergum II brown and terga
III-VI red. Similar material is known from the
hills that reach eastwards to as far as Beijing and
Hebei. Tkalci (1968a) described two queens
under the name Pyrobombus wutaishanensis that
have distinctly fewer fine punctures in the ocello-
ocular area than B. pyrosomas.str. At least one of
these queens is from Shanxi. I have seen another
queen of B. pyrosoma s.str. from neighbouring
Beijing (PW) that also shows some reduction in
the number and extent of these fine punctures, so
that it is intermediate in this character between
Pyrobombus wutaishanensis and, for example, B.
friseanus. These specimens are otherwise similar
103
in morphology and in colour pattern to the
lectotype of B. pyrosoma. Until more evidence to
the contrary is available, I consider that they are
most likely to be part of the same species.
From the available material, each of the three
principal colour patterns is relatively constant
among the individuals of each caste from across
extensive regions, with transitions in colour
pattern apparently occurring over short distances.
Yet from Qinghai, between the regions occupied
by B. friseanus and B. pyrosoma s.str., Tkalci
(1961) described a worker (B. pyrrhosoma f.
flavocorbicularis, infrasubspecific under Art.
45¢(i)) that has both the yellow hairs on the head
and legs, as for B. friseanus, and the pale pubes-
cence of the dorsum grey-white (not seen), as for
workers of B. pyrosoma s.str. Skorikov (1912b)
even described banded grey queens from the area
of Gansu and Inner Mongolia, similar in colour
pattern to most of the workers rather than to
the queens with a predominantly black thoracic
dorsum from north-eastern China, under the
name B. pyrrhosoma var. canosocollaris (not
seen). I consider it likely that these intermediate
colour patterns are evidence of clines from hybrid
zones between the more widespread regional
colour patterns (see the comments on B. trifasci-
atus). This has also been recognised previously for
the Chinese nominal taxa by Bischoff (1936) and
Tkalct (1961).
Evidence of intermediates between B. flavo-
thoracicus and B. friseanus is not as strong, at least
partly because so little material is available from
where these nominal taxa occur in close proximity
in the eastern Himalaya. However, the few
workers and males that are available from the far
east of the Himalaya (BMNH) are difficult to
assign to either nominal taxon with any confi-
dence. The only available queens are two speci-
mens from between Gyangtse [=Gyangzé] and
Phari [=Pagri] in Tibet (BMNH). These large
individuals share the white-‘tailed’ colour pattern
of the west Himalayan B. flavothoracicus queens,
although they also have red hair extensively
intermixed in the pubescence of terga III-IV.
Overall, from west to east there may be a trend
towards an extension of the red pubescence on the
gaster, especially for queens, in which it com-
pletely replaces the white hair. There is also a
parallel decrease in the extent of the pale pubes-
cence on the thorax and on terga I-II, which is
accompanied by a change in colour from light
yellow to orange-yellow to grey-white.
The males that are associated with the three
principal nominal taxa, B. flavothoracicus, B.
friseanus and B. pyrosoma s.str., all have closely
similar genitalia, but vary in the form of the
104
interio-apical corner of the gonostylus. This is
narrowly bilobed or spinose (similar to some
putative B. richardsiellus) for B. miniatus (the
male associated with B. flavothoracicus, Fig. 204)
and also for some B. pyrosoma s.str., but may be
broadly bilobed (similar to B. rufofasciatus, Fig.
205) for other B. pyrosoma s.str. All of these
males have the pubescence predominantly yellow
and their eyes are not enlarged relative to those of
the females. Both of these states are characteristic
of males that patrol circuits of scent-marked sites
in the upper montane forest habitats to find mates
(see the introduction on male mate-searching
behaviour and the discussion of the colour pat-
terns of the Kashmir fauna). Therefore from the
evidence available at present I consider it likely
that the three principal nominal taxa are based on
the regional colour patterns of the females from a
single interbreeding population. If this is the case,
they would be considered to be parts of a single
species. More definite conclusions concerning the
status of these nominal taxa must await more
direct information on their mate-searching be-
haviour and on interbreeding between them,
particularly from the eastern Himalaya.
The name Bremus formosellus was applied by
Frison (1934) to a population that is isolated from
other B. pyrosoma s.\. by sea on the island of
Taiwan. He distinguished this nominal taxon from
B. miniatus by colour pattern alone, although he
had apparently not seen any specimens from the
Himalaya (p. 166: ‘It [“This new species’] differs
from the male (type) and workers of miniatus as
described by RICHARDS (1930) in the more
broader and distinct black band on the thorax
between the wings and lack of bright yellow
pubescence on first and second dorsal tergites.’).
The grey females and the extensively yellow males
of Bremus formosellus actually resemble closely
some of the B. pyrosoma s.str. from Qinghai and
Gansu in both colour pattern and morphology. I
know of no evidence to suggest that individuals
from the disjunct population on Taiwan would not
interbreed with those from the continental popu-
lation if they were to occur together, although I
have not yet had the opportunity to examine any
queens. So Bremus formosellus may also be a part
of the species B. pyrosoma s.1.
DISTRIBUTION. B. pyrosoma is primarily a peri-
Tibetan species, although it also occurs in other
highland areas in northern and southern China
(Map 59). It is known from Inner Mongolia
(Skorikov, 1912b, 1933a; Tkalct, 1960), Hebei
(Yasumatsu, 1951; PW), Beijing (PW), Shanxi
(Yasumatsu, 1951; Tkalci, 1968a; MNHU),
Shaanxi (BMNH), Hubei (Sakagami, 1972),
PAULH. WILLIAMS
Gansu (Morawitz, 1890; Skorikov, 1912b; Bischoff,
1936; ZI), Qinghai (Skorikov, 1912b; Panfilov,
1957; Tkalci, 1961), Sichuan (Skorikov, 1933a;
Sakagami, 1972; Wang, 1982; BMNH, PW, ZI),
Yunnan (Bischoff, 1936; Wang, 1987; BMNH,
MNHU), Taiwan (Frison, 1934; Chiu, 1948;
INHS, PW), Tibet (Richards, 1930; Wang, 1982,
1988; BMNH, PW), Sikkim (Bingham, 1897;
Friese, 1918; Richards, 1930; BMNH, UM),
Nepal (Tkalci, 19746; BMNH, NMS, PW),
Uttar Pradesh (Frison, 1935), Himachal Pradesh
(BMNH, PW) and Kashmir. A record from
Guangdong [Leizhou peninsula] (Sakagami, 1972)
needs to be confirmed with more material. A
similar disjunct distribution between the moun-
tains of south-western China and Taiwan is known
for B. parthenius (synonym of B. sonani (Frison),
see the comments on B. lepidus) (Wang, 1982).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. pyrosomais recorded from the southern side of
the Great Himalaya range and from the Pir Panjal
range, in montane coniferous forest (Map 60, Fig.
6): 56 queens, 172 workers, 62 males, from 10
localities 1900-4000 m (AB, BMNH, IZ, MNHN,
NR, PW, RH).
A colony was found on Mt Apharwat, in the
forest below Gulmarg (2500 m, 2.ix.1985). The
entrance tunnel was in the bank of a stream and
lead 40 cm to the nest cavity, 80 cm vertically
below the surface of the bank. The nest contained
478 small cocoons (33 occupied, 7 that were
opened all contained males), 126 large cocoons
(106 occupied), 20 queens, including one old and
very abraded individual, 36 workers and 31 males.
No younger brood was present.
VARIATION WITHIN KASHMIR. This species is very
variable in colour pattern, but among the females
from Kashmir the trends in variation appear to be
associated more with body size than with locality
(e.g. increasing body size in Figs 426, 428, 429,
425). In particular, small queens show the partial
replacement of black or white pubescence by red
on terga III-IV, so that they appear similar to the
largest workers (Fig. 429). Most workers have, in
addition, yellow (Fig. 426) to chocolate-brown
(Figs 428 & 429) hair basally on tergum II, so that
black hair remains only apically on tergum II and
basally on tergum III (Plate 1). Some of the
smallest workers have most of terga I-II dull olive-
yellow and most of terga III-V orange-red (Fig.
426).
The male colour pattern is predominantly
yellow (Plate 1 and Figs 424, 427, 430). The
pubescence of terga III-VII varies from entirely
yellow (Figs 424 & 427) to largely black on tergum
III and orange-red on terga IV-VII (Fig. 430). The
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
most frequent pattern is for the pubescence of
tergum III to be yellow with a band of orange-red
hairs and black hairs intermixed basally, whereas
terga IV-VII are orange-red with a few black hairs
laterally and a few yellow hairs apically and
laterally. Only one male (from Daksum) has a
well-defined black band between the wing bases.
The only pronounced geographical variation is
that queens from Kel, in the Kishanganga valley,
have unusually extensive patches of yellow hair on
the basal half of tergum II (10/10 queens, Fig.
422). Workers from Kel (26 workers) have few
black hairs on terga II-III, irrespective of size
(Fig. 423). Another queen with the same data is
severely abraded, so this series may represent the
members of a single colony that had been col-
lected by the Schmid expedition. There are also
very few black hairs on the gasters of workers
from 4 other sites in the Kishanganga valley (5/5
medium-sized workers, no queens available).
Queens and workers of this species are distinc-
tive in their appearance. The yellow males with
red hairs on the apical terga may be similar in
colour pattern to the males of B. lepidus (Figs 329
& 332), which occur with them at some localities.
Males of B. pyrosoma can be recognised by their
larger size and by the short gonostylus with an
interio-apical spine (Fig. 204).
FOOD PLANTS. (Balsaminaceae) /mpatiens glandu-
lifera Royle; (Leguminosae) Trifolium repens
L., Lupinus sp. [introduced]; (Sambucaceae)
Lonicera quinquelocularis Hardw.; (Compositae)
Cirsium falconeri (Hook.f.) Petrak, C. wallichii
DC., unidentified dandelion-like composite;
(Ericaceae) Rhododendron anthopogon D. Don;
(Gentianaceae) Swertia petiolata D. Don;
(Scrophulariaceae) Scrophularia pauciflora Benth..,
Digitalis lanata Ehrh. [introduced], D. purpurea
L. [introduced], Pedicularis rhinanthoides Schrenk,
P. punctata Decne.; (Labiateae) Prunella vulga-
ris L.
Bombus (Melanobombus) rufofasciatus Smith
(Plate 1, Figs 26, 30, 34, 38, 85, 125, 165, 205,
431-439, Maps 61 & 62)
Bombus rufo-fasciatus Smith, 1852b: 48. Lecto-
type queen by designation of Tkalct (1974b:
340), INDIA: north, no further data (BMNH)
[examined].
Bombus Prshewalskyi Morawitz, 1880: 342. Syn-
type worker [not a queen] and male, CHINA:
‘Gansu’, no further data (Prshewalski) (ZI) [not
seen]. Synonymised with Bombus rufofasciatus
var. championi Richards by Richards (1930);
105
synonymised with Pyrobombus_ rufofasciatus
(Smith) by Tkalct (19745).
Bombus rufocinctus Morawitz, 1880: 343. Lecto-
type queen by designation of Podbolotskaya
(in press), CHINA: ‘Gansu’ hills, viii.1871
(Prshewalski) (ZI) [examined]. Junior primary
homonym of Bombus rufocinctus Cresson, 1863.
Synonymised with Bombus rufofasciatus Smith
by Handlirsch (1888).
Bombus chinensis Dalla Torre, 1890[June 25]: 139.
Replacement name for B. rufocinctus Morawitz.
Junior secondary homonym in Bombus of
Apathus rupestris var. chinensis Morawitz, 1890
[April 30] [= B. chinensis (Morawitz)]. Synony-
mised with Bombus rufofasciatus Smith by
Richards (1930).
Bombus rufofasciatus var. championi Richards,
1928a: 107. Holotype queen by original desig-
nation, INDIA: Kashmir, 8000-9000 ft [2400-
2700 mJ], vi.1901 (Nurse) (BMNH) [examined].
Syn. n.
[Bombus (Lapidariobombus) rufofasciatus var.
rufior Richards, 1928b: 335 [examined]. Infra-
subspecific (Art. 45g(ii)(1)), unavailable name
(Art. 45e).]
[Bombus (Lapidariobombus) rufofasciatus var.
intermedius Richards, 1930: 643 [examined].
Infrasubspecific (Art. 45g(ii)(1)), unavailable
name (Art. 45e).]
Bombus (Lapidariobombus) waterstoni Richards,
1934: 88. Holotype queen by original designa-
tion [p. 89], ‘Himalayas’, no further data
(BMNH) [examined]. Synonymised with Pyro-
bombus rufofasciatus (Smith) by Tkalct (19745).
AFFINITIES. Within the subgenus Melanobombus
(see the comments on B. ladakhensis), B. rufo-
fasciatus belongs to the rufofasciatus-group (see
the comments on B. simillimus).
Within the rufofasciatus-group, B. rufofasciatus
can be distinguished from B. simillimus by its
strongly reduced male gonostylus (Fig. 205; see
the key, couplet 25). B. rufofasciatus is similar to
some B. pyrosoma s.\. in that it has a broadly
bidentate, interio-apical corner of the gonostylus,
although the apex of the interio-basal process
usually forms a right angle rather than being
distinctly acute (see the comments on B. pyro-
soma). The females of B. rufofasciatus can be
distinguished by their nearly clear (hyaline)
wings, which are more strongly darkened (infus-
cated) for the other taxa of the rufofasciatus-
group, and by their weaker punctures in the
ocello-ocular area of the head. The males can be
distinguished from males of B. richardsiellus or B.
pyrosoma by the size and shape of their eyes,
which are moderately enlarged relative to those of
106
the females (i.e. similar in shape to those of B.
simillimus).
The enlargement of the eyes of male B. rufo-
fasciatus in comparison with the females reflects
differences in their mate-searching behaviour that
may contribute to a failure to interbreed with
individuals of the other species where individuals
of B. rufofasciatus occur with them (see the
introduction on male mate-searching behaviour).
For instance, B. rufofasciatus and B. pyrosoma
overlap extensively in their foraging areas around
the upper limit of the montane coniferous forest
on Mt Apharwat (Fig. 6). However, workers and
queens of B. pyrosoma are very much in the
minority on the subalpine slopes above the forest,
whereas those of B. rufofasciatus are rare within
the forest. The males of B. pyrosoma, with eyes
similar in size to those of the females, patrol
circuits of scent-marked sites only within the
upper montane forest. The males of B. rufofasci-
atus, with enlarged eyes, watch for potential
mates from perches on open, subalpine hilltops,
where they usually establish exclusive territories.
So individuals from these two populations are
searching for mates in different ways and in
different kinds of habitat (cf. the comments on B.
asiaticus). No intermediates could be found in
samples containing both B. rufofasciatus and
B. pyrosoma s.\. [= B. flavothoracicus| from
Kashmir or Nepal (BMNH, PW); B. rufofasciatus
and B. richardsiellus from northern Burma
(BMNH); B. rufofasciatus and B. pyrosoma s.1.
[= B. friseanus] from Yunnan (BMNH) or
Sichuan (PW); and B. rufofasciatus and B. pyro-
soma s.str. from Gansu (Bischoff, 1936).
A queen from Kashmir that has a broad band of
yellow pubescence on tergum II was described
under the name B. rufofasciatus var. championi
by Richards (1928a). This individual is closely
similar in morphology to those queens of B.
rufofasciatus that have tergum II entirely black
and it is considered to be part of the same species.
DISTRIBUTION. B. rufofasciatus is a peri-Tibetan
species (Map 61). It is known from Gansu
(Morawitz, 1880; Bischoff, 1936; ZI, ZS), Qinghai
(Morawitz, 1886; Tkalct, 1961; Wang, 1982; PW,
ZS), Tibet (Friese, 1918; Richards, 1928a, 1928),
1930; Wang, 1982, 1988; BMNH), Sichuan
(Morawitz, 1890; Wang, 1982; PW), Yunnan and
northern Burma (BMNH), Arunachal Pradesh
(Friese, 1918), Sikkim (Bingham, 1897; Friese,
1918; BMNH, UM), Nepal (Tkalci, 1974);
BMNH, NMS, PW), Uttar Pradesh (Richards,
1928a, 1930; BMNH), Himachal Pradesh (BMNH,
PW) and Kashmir. A record from Inner Mongolia
[Ordos] (Morawitz, 1880) is dubious and needs
PAULH. WILLIAMS
to. be confirmed (cf. Skorikov, 1912b, on B.
pyrosoma).
MATERIAL EXAMINED FROM KASHMIR. In Kashmir,
B. rufofasciatus is recorded from the Hindu Raj,
Great Himalaya and Pir Panjal ranges, in mesic
alpine scrub (Map 62, Fig. 6): 58 queens,
351 workers, 260 males, from 28 localities 1900—
4600 m (AB, BMNH, BPBM, MI, MNHN, NM,
NR, PW, RH, SEMK, TL; 3 males photographed
in the upper Wangat valley, 1 worker photo-
graphed in the Sangisfaid valley, C. A. Chadwell).
A colony was found on Mt Apharwat, at the
lower edge of the subalpine pasture at Khilanmarg
(3000 m, 4.ix.1985). The nest entrance was among
boulders, mostly larger than 1 m across, that were
lying several deep in a gully. This nest could not be
excavated. I saw many queens investigate similar
sites in the subalpine zone during July 1986,
presumably in search of suitable nest sites.
VARIATION WITHIN KASHMIR. Most queens from
Mt Apharwat in the Pir Panjal range have a few
yellow hairs intermixed with the black near the
base of tergum II (Plate 1), but some individuals
(10/32 queens from Apharwat, also 2/2 queens
from around the Kishanganga valley) have an
obvious band of yellow pubescence on the basal
half of this tergum (Fig. 434). Queens from the
Hindu Raj range (12/12 queens), at the western
limit of the species’ distribution, are unusual in
that the pubescence of tergum II is predominantly
yellow, with at most only a narrow band of black
hairs apically (Fig. 431). Workers and males from
all parts of Kashmir generally have little black
pubescence on tergum II (Plate 1 and Figs 432,
433, 435, 436, 438, 439), especially among indivi-
duals from the Hindu Raj range (Figs 432 & 433).
At the opposite extreme, some of the largest
workers from Apharwat have only a narrow
yellow band on tergum II (Fig. 438), or even just a
few yellow hairs at the base of the tergum, so that
it appears generally black (2/92 workers). Queens
from further to the east in the Great Himalaya
range generally share the darker colour pattern
(Fig. 437), whereas workers generally retain
yellow pubescence on tergum II (Figs 435 & 438).
A male from Batakush has the pale bands on the
thorax and tergum I yellow (1/5 males).
The queens without obvious yellow pubescence
on tergum II are quite distinctive in their appear-
ance (Fig. 437). The yellow and white individuals
of both castes and both sexes (Figs 431-436, 438,
439) can be particularly closely similar in colour
pattern to the B. kashmirensis (Figs 302-304) that
occur with them at some localities. B. rufofasci-
atus can be recognised by the hairs of terga II-VI,
which are shorter and either red (most of terga
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
III-IV in workers and males; only tergum III in
queens) or white (most of terga IV-VI in queens,
terga V-VI in workers, terga V-VII in males),
rather than red with white tips. They also
resemble some of the B. himalayanus (Figs
247-249), B. novus males (Figs 260 & 262), B.
lemniscatus (Figs 325 & 326), B. lepidus (Figs 330,
331, 333, 334) and B. biroi (Figs 338 & 339) that
occur with them. Females of B. rufofasciatus can
be recognised easily because they have terga V-VI
with white hairs rather than with orange-red hairs.
FOOD PLANTS. (Ranunculaceae) Aconitum laeve
Royle, A. heterophyllum Wallich ex Royle, A.
violaceum Jacquem. ex Stapf; (Balsaminaceae)
Impatiens glandulifera Royle; (Leguminosae)
Trifolium repens L., Lupinus sp. [introduced];
(Sambucaceae) Lonicera quinquelocularis Hardw.;
(Compositae) Cirsium falconeri (Hook.f.) Petrak,
C. wallichii DC., Lactuca lessertiana (DC.) C. B.
Clarke, unidentified dandelion-like composites;
(Ericaceae) Rhododendron anthopogon D. Don;
(Gentianaceae) Swertia petiolata D. Don;
(Scrophulariaceae) Scrophularia pauciflora Benth. .,
Digitalis lanata Ehrh. [introduced], Pedicularis
oederi Vahl, P. roylei Maxim., P. pectinata
Wallich ex Benth., P. rhinanthoides Schrenk, P.
punctata Decne.; (Labiateae) Phlomis bracteosa
Royle ex Benth., Nepeta podostachys Benth.,
Prunella vulgaris L., Thymus linearis Benth. ex
Benth.
DISTRIBUTION OF THE FAUNA
Distribution with altitude
The slopes of Mt Apharwat in the Pir Panjal range
(Map 4 locality 1) show many of the more mesic
kinds of habitat in which bumble bees occur in
Kashmir. Differences in the distributions of the
bumble bee species with altitude can be described
from transect walks of the north-eastern flank
of Apharwat (see the introduction on mate-
searching behaviour of male bumble bees in
Kashmir). These transects ran from Tangmarg at
1800 m in the Vale of Kashmir, through Gulmarg
at 2700 m, to the peak of Apharwat above at
4143 m (13,592 ft). More than 2000 bumble bees
were examined in the field along these transects
during July, August and September 1985. Fig. 6
shows the records for each of the social species
between the 500 ft contours of the 1 : 63,360
British Survey of India map (Oxford University
Library). These patterns of distribution are simi-
lar to those found along the same transects in 1980
107
and 1986, and on other mountains around the
Vale of Kashmir in all three years.
Away from the foot of the mountain, the middle
of the Vale of Kashmir around Dal Lake (1500 m)
is marshy and much of this area is used for the
cultivation of rice. No bumble bees were found
there. As the land slopes gently towards the
mountains, terraces of rice give way on drier
ground to apple orchards and higher still, near
Tangmarg, to maize. Above Tangmarg, the lower
coniferous forest (1900-2700 m) is dominated by
Pinus wallichiana A.B. Jackson, with Cedrus
deodara (Roxb. ex D. Don) G. Don near the
lower margin. The forest is protected by law,
although its lower edge now largely coincides
with a pronounced increase in slope just above
Tangmarg. The understorey is very disturbed and
although few trees are felled, their branches
are still collected for firewood. Flowers used by
bumble bees are scarce in the lower forest (species
listed in Table 5) but are more abundant around
buildings and along roadsides. Bumble bee densi-
ties are very low, with B. tunicatus and B.
trifasciatus the most abundant species both within
the forest and around its lower edge.
Gulmarg (= ‘flower-meadow’, 2700 m) oc-
cupies a basin on a shoulder of the mountain. This
natural, marshy meadow is grazed intensively by
cattle, sheep and horses, yet the flowers from
which bumble bees feed (Table 5) remain abun-
dant. In the extensive wet areas of the meadow,
Prunella vulgaris is a major nectar source and
Pedicularis punctata a major pollen source for
much of the summer. Large stands of the thistle
Cirsium falconeri, which is often over 2 m tall,
occur on disturbed slopes around the edges of the
meadow. Impatiens glandulifera is abundant
around water outflows from houses. The density
of bumble bees is high (e.g. 89 bees/km, from
2.7 km transect, 14.ix.1986), with B. asiaticus and
B. tunicatus the most abundant species, although
there are fewer species than in the other zones
(Fig. 6).
The upper coniferous forest is dominated in
turn by Pinus wallichiana (2700 m), Picea smithi-
ana (Wallich) Boiss. (2700-2900 m) and Abies
spectabilis (D. Don) Mirbel (2900-3000 m).
Prominent members of the shrub layer are bushes
of Sambucus wightiana Wallich ex Wight & Arn.
and Euonymus hamiltonianus Wallich. Flowers
used by bumble bees (Table 5) are few, as in the
lower forest. Digitalis purpurea, and more com-
monly D. Janata, are naturalised just above
Gulmarg. Bumble bee density is low (23 bees/km,
from 4.2 km, 14.ix.1986), with B. pyrosoma and
B. tunicatus the most abundant species.
The lower slopes (3000-3200 m) of the sub-alpine
108
PAULH. WILLIAMS
Table 5 Lists of the plants that were visited most frequently by foraging bumble bees on Mt Apharwat during 1985
and 1986 in the different altitudinal zones (LF, lower forest; GM, Gulmarg meadow; UF, upper forest; SA,
subalpine; A, alpine; +, visited frequently; +, major food source).
Family Binomen
Altitudinal zone
LF GM UF SA A
Ranunculaceae
Balsaminaceae
Leguminosae
Caprifoliaceae
Compositae
Ericaceae
Oleaceae
Gentianaceae
Scrophulariaceae
Labiateae
Aconitum heterophyllum Wallich ex Royle
Aconitum hookeri Stapf
Impatiens glandulifera Royle =F
Lupinus sp.
Trifolium repens L.
Lonicera quinquelocularis Hardw.
Cirsium falconeri (Hook. f.) Petrak
Cirsium wallichii DC. *
Lactuca lessertiana (DC.) C.B. Clarke
yellow dandelion-like composites
Rhododendron anthopogon D. Don
Syringa emodi Wallich ex Royle
Swertia petiolata D. Don
Scrophularia pauciflora Benth.
Digitalis purpurea L.
Digitalis lanata Ehrh. +
Pedicularis oederi Vahl
Pedicularis roylei Maxim.
Pedicularis pectinata Wallich ex Benth.
Pedicularis punctata Decne. +
Phlomis bracteosa Royle ex Benth.
Stachys sericea Wallich ex Benth.
Prunella vulgaris L. +
Thymus linearis Benth. ex Benth.
++
*
*
++
*
++ ++
+++
Salvia hians Royle ex Benth.
Polygonaceae Bistorta vivipara (L.) Gray
zone (Fig. 1) are dominated by species of
Salix, Betula utilis D. Don and lilac, Syringa
emodi. Large areas are strewn with boulders.
Spaces between the rocks provide nest sites
for bumble bees and refuges from grazing for
Scrophularia pauciflora, which, despite its green
flowers, is very attractive to bumble bees. Much of
this zone is used as pasture for goats, sheep and
cattle, especially in the more gently sloping area,
known as Khilanmarg, that is immediately above
the forest. Pasture areas have many bushes of
Euphorbia wallichii Hook. f. and flowers of
Anemone obtusiloba D. Don. The steeper middle
slopes (3200-3400 m) are dominated by B. utilis
and Rhododendron campanulatum D. Don. The
upper slopes (3400-3800 m) are dominated by R.
anthopogon, Juniperus recurva Buch.-Ham. ex D.
Don and Bergenia stracheyi (Hook. f. & Thoms.)
Engl. Overall, the subalpine zone has the largest
number of bumble bee species (Fig. 6), although
their total density (32 bees/km, from 4.2 km,
++
12.ix.1986) is less than at Gulmarg. The most
abundant species throughout the subalpine zone is
B. rufofasciatus.
The alpine zone (3800-4143 m) is marked by an
absence of the low bushes of R. anthopogon
and by the presence, at least in exposed areas,
of a silver-grey mat vegetation with Anaphalis
triplinervis (Sims) C.B. Clarke and species of
Leontopodium (Edelweiss). Bistorta vivipara is
conspicuous, but the major food-plants for
bumble bees are species of Pedicularis, Swertia
petiolata etc. (Table 5). The most abundant
bumble bee species are B. rufofasciatus and
B. kashmirensis (density of all bumble bees 31
bees/km, from 2.3 km, 12.1x.1986). These two
species are closely similar in colour pattern and
general appearance, but all of the individuals with
this colour pattern that were caught visiting
flowers of Compositae were of B. rufofasciatus,
whereas those visiting Aconitum hookeri were all
of B. kashmirensis.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
The overall pattern in the distribution of the
species’ foraging activities with altitude on Mt
Apharwat (Fig. 6) is very similar to that found on
other mountains around the Vale of Kashmir (e.g.
near Pahalgam and near Daksum). The overlaps
in the species’ altitudinal ranges on these moun-
tains were more extensive than those measured
along the floor of the Sind valley (Fig. 12). For
instance, species such as B. trifasciatus and B.
tunicatus are restricted to the lower end of the
Sind valley (e.g. Kangan, 1900 m) and other
species such as B. rufofasciatus and B. keriensis
are restricted to the upper end (e.g. Sonamarg,
2700 m), although they have all been collected
from the same meadow on Apharwat (2700 m).
The Sind river runs from deep in the Great
Himalaya to the Vale of Kashmir, so the slope
of the valley floor is far less steep than the north-
eastern flank of Mt Apharwat. It is more likely
that individuals could wander or commute to
forage outside their optimal altitudinal zones on
Apharwat, simply because the zones are so much
closer together horizontally. B. simillimus was
common in the lower Sind valley (e.g. at Kangan,
1900 m) but was not found on Apharwat during
1980 or 1985. Three workers of this species were
found at 3000 m on Apharwat in 1986.
In general, the distributions of bumble bee
species are most likely to be governed by the
combined effects of climate and of local levels
of food resources (discussed in Williams, 1988,
1989). The environment in Kashmir is particularly
complex, because the regional climate may be
greatly modified locally by the effects of altitude
and exposure (e.g. Troll, 1972). This is likely to
have three principal direct effects on bumble bees.
First, lower average temperatures at higher alti-
tudes may increase thermoregulatory expenditure
of energy by bumble bees, especially for queens,
which must incubate their first brood alone in
the spring. Second, a shorter annual season of
temperatures above freezing point in the high
mountains constrains the time available for colony
development. However, daytime temperatures
even in the alpine zone of Kashmir may be higher
and more favourable than further north (e.g. in
the Alps), although the day lengths are shorter in
Kashmir. But third, at lower altitudes, bumble
bees may actually suffer from heat stress. For
instance, foraging activity by bumble bees (chiefly
B. trifasciatus and B. haemorrhoidalis) at Patnitop
(2000 m) in the Jammu foothills ceased during
the warmest part of the day, between 10 a.m.
and 5 p.m. (6-8.ix.1986), when the patches of
Impatiens glandulifera and Pteracanthus urtici-
folius from which they were foraging were no
longer in the shadows of the forest trees. The
109
bumble bees could not be found visiting any other
flowers elsewhere during these periods. A less
pronounced trend towards bimodality in diurnal
foraging patterns is well known for some temper-
ate bumble bees (e.g. Heinrich, 1979; Willmer,
1983) and is also reported for the large carpenter
bees of the genus Xylocopa at a subtropical
lowland site (Abrol, 1987). In both of those
situations, foraging activity by the bees was in-
versely related to air temperature and intensity of
direct solar radiation, but was positively associ-
ated with nectar availability. At Patnitop, small
and medium-sized anthophorine bees continued
to forage from the same flowers throughout the
day, so presumably at least some nectar was still
available. A restriction of diurnal foraging periods
for bumble bees is likely to reduce the daily rate of
food supply to colonies and so slow their develop-
ment within the limited foraging season. This has
at least the potential to limit a species’ distribution
at the southern edge of its range.
In common with many other organisms, bumble
bees tend to occur at higher altitudes in Kashmir
than they do further north, but in zones with
similar climate. B. lucorum is most abundant on
Mt Apharwat around the upper edge of the
montane forest zone (c. 3000 m) and is recorded
from 2100-4600 m at other sites in Kashmir.
Further north in the Alps, it is common on the
forested middle slopes (1300-2600 m recorded by
Pittioni, 1937). In cool temperate Britain, this
species is common in virtually all of the available
altitudinal zones down to sea level (0—-1200 m).
Climate may also affect bumble bees indirectly
through its effects on food resources. The lower
parts of the valleys in Kashmir often appear to
receive less rain or snow in summer than the
mountain ridges. In some of the drier areas the
availability of bumble bee food-plants in late
summer may be very restricted and bumble bees
are sometimes either absent or present at only
very low density (e.g. at Baramula and Anantnag
in the Vale of Kashmir). Food sources are also
scant in the shade of the evergreen, coniferous
forests on the hillsides. But higher in the mesic
subalpine zone, the density of nectar-rich food-
plants is often high and their diversity may be at
its highest. In the alpine zone, the frequent
freezing and overcast conditions throughout the
summer are likely to reduce plant productivity
and hence resources for bumble bees. The favour-
able balance of these climatic and food resource
factors in the subalpine zone may help to account
for its large number of species of bumble bees
(Fig. 6).
Although the meadow at Gulmarg has much the
highest density of bumble bees on Mt Apharwat,
110
@® Type ia] yellow banded/
@ Type 1B
@ Type al grey-white
© Type 2B
@ Type 3
HIGH oberti
ALPINE personatus
ladakhensis
ALPINE kashmirensis
himalayanus
keriensis =)
MESIC rufofasciatus (=) @
SUBALPINE _ biroi @ g
avinoviellus S
ARID subtypicus
SUBALPINE — semenovianus
STEPPE asiaticus on en ee)
melanurus @ @
UPPER lepidus g @
FOREST lucorum
hypnorum @
pyrosoma @ &
LOWER tunicatus iS) < (=) < S <
FOREST simillimus O O
trifasciatus g ® ®
haemorrhoidalis <
|
s
3
a
Predominant local <
colour patterns of workers: S
oD
Ks) a ‘
= iS oun,
unbanded thorax 7S i) & /
roy isi =I
eS & N ZANSKAR LADS
S RANGE
SI RANGE
banded thorax s nS GREAT HIMALAYA Z ee
PANJAL RANGE 3 $
black thorax RANGE 3 =
Sind R. Dras R. — y
PAULH. WILLIAMS
SES OES
Z @
oo
@
@<« Qs
@ @«
®9 8 S&S @ ; o ®@
Oo ® 6 ti)
: 3
AZ. /a\
Martselang
Tiggur
Fig. 12 Distribution of the social species of bumble bees (i.e. excluding species of the subgenus Psithyrus) and their
local colour patterns across the Himalaya. Only the locally predominant patterns among the workers are represented.
Triangles show the most abundant species at each locality. B. lemniscatus is not included for lack of information.
it has rather few species (Fig. 6). This may be
partly because the abundance of food-plants is
particularly uneven, although the relationship
between diversity of food-plants and diversity of
bees may not be straightforward (Williams, 1989).
Pedicularis punctata and Prunella vulgaris are by
far the most abundant of the food-plants used
by bumble bees in the meadow. The meadow is
often extensively flooded and some areas around
streams are permanently marshy, although these
plants are apparently tolerant of water-logged
soil. Most of the more abundant species of bumble
bees in the meadow, such as B. tunicatus and B.
pyrosoma, are also among the most common
species in the forest. Nests of both species have
been found in the forest, but not in the meadow,
so it may be that many of their foragers commute
from nests in the forest to the richer resources in
the meadow. B. asiaticus is unusual because it is
both abundant on Gulmarg meadow and gener-
ally associated with open habitats.
The locality at the highest altitude in Kashmir
from which I have collected a large sample of
bumble bees is on Nimaling plain (4800 m). This is
a high glacial valley, surrounded by large areas of
arid alpine steppe that have very few if any
bumble bees. Visits for 13 days in July 1980 and
for three days in September 1986 found nine
species of bumble bees (including the social para-
site B. (Psithyrus) branickii) visiting the flowers of
only a single plant species, Caragana versicolor.
This plant forms low, thorny bushes on the old
Altitude
6000
5000
4000
3000
2000
1000
0
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
lateral and terminal moraines (Fig. 3), from which
the glacier snout has retreated up the valley by
about 10 km to the south-east. If the foragers do
all genuinely use what appears to be a single food-
resource, and there were no other obvious poten-
tial food-plants to be seen, then this situation
deserves further study. It would follow from the
application of competition-based ideas of ‘com-
munity structure’ to these bumble bees (e.g.
Inouye, 1977) either that the flowers of Caragana
are very variable in the accessibility of their
rewards to the different bumble bee species, or
that other factors are preventing the local assem-
blage of bumble bee species from reaching equili-
brium by the local extinction of most of the species
(cf. Williams, 1989).
Fig. 6 shows the narrower range of altitudes and
habitats in which the males search for mates on Mt
Apharwat, in comparison with the range of habi-
tats in which the females forage. The possible
relationship between the different kinds of male
mate-searching behaviour and the different kinds
of habitat is summarised in the introduction.
Distribution across the Himalaya
The distribution of bumble bees across the
Kashmir Himalaya can be described from the
samples collected at selected sites in the Pir
Panjal, Great Himalaya, Zanskar and Ladakh
ranges during August and September 1986. These
data are supplemented with records from speci-
mens collected at Nimaling in 1980 and with
others examined in collections from Muzaffarabad
(BMNH), from the Khardung-La above Leh (FA,
ZI) and from Tiggur near the Nubra/Shyok
confluence (BMNH). Together this covers a hori-
zontal distance of about 430 km from west to east,
at about 34° latitude. Fig. 12 shows the distribu-
tions of the bumble bee species in these samples
and their locally predominant colour patterns.
The most widespread species in Kashmir are the
steppe species B. melanurus and B. asiaticus. B.
asiaticus shows a major change in colour pattern
between Nigagar and Gumri in the Great Himalaya
range, with some intermediate individuals present
at Nigagar (see the discussion of variation of B.
asiaticus in Kashmir). The alpine element in the
fauna is also widespread between the Vale of
Kashmir and Zanskar. Like B. asiaticus, B.
kashmirensis and B. himalayanus show a pro-
nounced change in colour pattern, but in this case
the major transition in the thoracic pattern is to
the east of Gumri (and to the west of Nimaling).
The principal boundary in the composition of
the fauna is associated with the Great Himalaya
range. This region of transition was studied in
111
more detail between 9-11.viii.1986 in samples
from Sonamarg, Nigagar, Baltal, Gumri, Matayan
and Dras (a gradient-directed transect, see Map 4
& Table 1). The forest element is represented at
Sonamarg, Nigagar and Baltal but, like the forest,
it is absent to the north-east, beyond the Zoji-La
(La = pass). There is also a large change in the
subalpine element across the Great Himalaya. Of
the subalpine species found on Mt Apharwat, B.
rufofasciatus reaches across the Great Himalaya
as far as Matayan, although B. biroi may be more
widely distributed in Zanskar and Ladakh (Map
42). At Matayan, B. rufofasciatus occurs together
with B. subtypicus, although B. rufofasciatus is no
longer the most abundant species, as it is in the
subalpine zones around the Vale of Kashmir. The
three high alpine species found at Nimaling are
distributed widely across the Tibetan massif but
are apparently absent from the mountains around
the Vale of Kashmir. This abrupt discontinuity in
the faunal composition reflects the transition in
climate and vegetation from the more mesic,
wooded environment of the Vale of Kashmir to
the arid (= xeric), steppe environment of the high
Tibetan massif (Map 3). The environmental dif-
ferences between these two regions in Kashmir
are more extreme than those across the eastern
Himalaya (Chang, 1981), where the Tibetan
plateau is generally lower, more humid and
dissected by wooded valleys. This is reflected in
the distribution of bumble bees such as B. rufo-
fasciatus, which penetrates much more deeply
into Tibet in the east (Wang, 1982; BMNH).
A second boundary in the composition of the
fauna is located on the outer slopes of the Pir
Panjal range. The low altitude fauna has not been
studied thoroughly here, largely for reasons of
inaccessibility (see the comments on material
examined). It is dominated by a lower montane
forest element (B. haemorrhoidalis and B. trifas-
ciatus). This includes many more species in the
more humid foothills of the central and eastern
Himalaya, where the effects of the monsoon are
more pronounced.
From the limited amount of material available
(BMNH, NMS, PW), the general pattern in the
distribution of bumble bees across the Himalaya
of Kashmir is similar to that found across the
Himalaya of western Nepal (Table 6). The major
difference is that the same arid subalpine element
is not known to be represented in Nepal or in the
adjacent parts of Tibet. The Kashmir-western
Nepal region of the Himalaya spans about
1000 km, but shares a broadly similar patterfiof
climate and vegetation (see the maps in
Schweinfurth, 1957). The pattern of vegetation
formations is highly convoluted locally because it
112
PAULH. WILLIAMS
Table 6 Distribution of the social bumble bees of Kashmir (i.e. excluding species of the subgenus Psithyrus) outside
Kashmir. X — present in the regional lists from world summary grid-squares (see Map 1): AP, western Alps; AR,
Armenia and the Caucasus; MG, northern Mongolia; TS, central Tien Shan; HK, Hindu Kush; NP, Nepal; GS,
southern Gansu; BM, central Burma; TW, Taiwan.
Region
Habitat in Kashmir: AP AR MG _ TS
High alpine x
Alpine
x x x
Mesic subalpine
x xX
Arid subalpine
x
Steppe x x
x ».4 x
Upper forest
x x x x
x x x
Lower forest
tends to follow contours, but throughout this
region it consists essentially of bands parallel to
the Great Himalaya (e.g. Map 3). So for bumble
bees, as for the vegetation and for many other
groups of organisms (e.g. Mani, 1986), the fauna
apparently changes to a greater extent in a small
distance across the Himalaya than it does along
them. This supports the familiar idea that distri-
butions are restricted less by the capacity of
Organisms to spread through a habitat than by
their capacity to colonise a different kind of
habitat (e.g. Fretwell & Lucas, 1970).
Distribution outside Kashmir
Only the broadest patterns in the distribution of
bumble bees outside Kashmir can be discussed with-
out an analysis of the distribution of all bumble bee
species (in prep.). Table 6 summarises the distribu-
tions of the social bumble bees of Kashmir (i.e.
excluding species of the subgenus Psithyrus) across
some of the mountainous areas of Europe and
Asia. The species are listed in a sequence to show
their altitudinal distribution and habitat associa-
tion within Kashmir (see Figs 6 & 12).
HK
mmx KM
g
Kashmir BM
oberti
personatus
ladakhensis
kashmirensis
himalayanus
keriensis
rufofasciatus
biroi
lemniscatus
avinoviellus
~~ Km Km KM RM
~ Km KM KR MR
marussinus
subtypicus
semenovianus
asiaticus
melanurus
lepidus
lucorum
hypnorum
pyrosoma
KKK MK mM
tunicatus
simillimus
trifasciatus
haemorrhoidalis
aM MRM KM
*
~*~
The alpine and high alpine species of Kashmir
are primarily associated with the Tibetan massif
(Table 6: reaching Nepal, Gansu). The distribu-
tions of B. keriensis and B. oberti also extend
westwards and northwards through the Pamir to
the Tien Shan ranges and, for B. keriensis, further
to Armenia in the west and to the mountains of
northern Mongolia in the north-east.
The subalpine and steppe species of Kashmir
are principally associated with the arid mountains
to the north in central Asia (Table 6: Hindu Kush,
Alai-Tien Shan s.1.). There are also a couple
of very widespread, cool temperate and upper
montane forest species (B. lucorum and B.
hypnorum). Species of the arid subalpine element
are at the south-eastern limits of their distribu-
tions in Kashmir. Species of the steppe element
are also more widespread in central Asia, but
reach beyond Kashmir eastwards into the Tibetan
massif. Together with the alpine species these
correspond to the ‘western hypsobiont element’
described from the Himalayan butterfly fauna by
Mani (1986).
The remaining species are found in and around
the forests at lower altitudes and are more
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
southern and eastern in their distribution outside
Kashmir. They are either restricted to the western
Himalaya (B. tunicatus and B. simillimus) or are
at the western limit of their distributions (Table 6:
reaching eastwards to Nepal and Taiwan). B.
haemorrhoidalis and B. trifasciatus reach east-
wards and southwards into the Oriental tropics in
the mountains of northern Thailand. Together
this group of eastern species corresponds to the
‘eastern forest element’ described from butterflies
by Mani (1986).
Four species are narrowly endemic to the west-
ern Himalaya, B. avinoviellus, B. himalayanus,
B. tunicatus and B. simillimus. It is possible that
B. himalayanus may prove to be conspecific with
the central Asian taxa B. turkestanicus and B.
marussinus (see the comments on B. marussinus
and B. himalayanus). The three remaining species
are among the more plesiomorphic species within
their respective species-groups. If they are relics
of a relatively ancient bumble bee fauna that was
once more widespread, then it now persists in a
region that has been subject to some of the most
active mountain building during the last few
million years (see Bally, 1983; Mani, 1986). But
although the western Himalaya may not appear to
have been a particularly stable refuge in this
period, the high relief could have provided some
buffering against any climatic fluctuations by
allowing altitudinal displacement of bumble bee
populations.
Overall, the bumble bee fauna of Kashmir
includes particularly few of the relatives of the
European B. hortorum (only B. trifasciatus) or B.
pascuorum (only B. haemorrhoidalis), but par-
ticularly many of the relatives of the European B.
lapidarius (8 species of the subgenera Sibirico-
bombus and Melanobombus). This pattern (9%
Megabombus + Diversobombus + Thoraco-
bombus + Orientalibombus : 35% Sibiricobombus
+ Melanobombus, Kashmir fauna excluding
Psithyrus) contrasts markedly with the repre-
sentation of these groups in the fauna of the
European Kalsbach Alps (31% : 6%, Pittioni,
1937), in the fauna of the whole of Europe (34% :
8%, Reinig, 1981) or in the fauna of the western
Palaearctic (32% : 10%, Rasmont, 1983).
Relationships between patterns at different
spatial scales
Among the social bumble bees, those species that
are more widespread world-wide are not generally
found at more localities within Kashmir (Fig.
13, which excludes the species of the subgenus
Psithyrus, Kendall coefficient of rank correlation
0.08, P > 0.5 n.s.). It is possible that these data
113
might not be representative because the sampling
effort was far from uniform at both spatial scales
and the sites in Kashmir were not randomly
located but selected by kind of habitat and for ease
of access. Nonetheless, there is pattern in the data
insofar as species with similar habitat associations
(Figs 6 & 12) do tend to occur closer together in
Fig. 13. One interpretation is that this pattern in
Fig. 13 may show a common limiting effect within
the groups of species with similar habitat require-
ments, which could be just a consequence of the
limited availability of the most suitable kinds of
habitat for these species at both spatial scales.
From a study of British bumble bees, patterns in
local distributions appear to be related to how
close the species are to the centres of their world-
wide distributions (Williams, 1988). Among the
Kashmir fauna, those species that are nearer to
the centres of their world-wide distributions (from
Maps 5-62: B. avinoviellus, B. himalayanus, B.
melanurus, B. tunicatus, B. asiaticus, B. semeno-
vianus, B. keriensis and B. simillimus) do also
tend to occur at more localities within the country
(Maps 5-62; Fisher-Pitman randomisation test,
W = 181, single-tailed P < 0.01, see Krauth,
1988). More precisely, those species that are
nearer to the centres of their distributions in
Britain have been found in a broader range of
kinds of habitat. For the fauna of Kashmir, this
greater breadth of habitat use for those species
that are nearer to the centres of their world-wide
distributions is apparent in the greater altitudinal
ranges of these species within the country (exclud-
ing single altitude records of B. oberti and B.
lemniscatus, Fisher-Pitman randomisation test,
W = 18400, single-tailed P < 0.05).
Fig. 13 also appears to contain information as to
how close the species are to the centres of their
world-wide distributions. For these data in which
the variance is similar in the log(world) and
log(kashmir) distribution values, the central/
marginal position of a species appears to be
related to log(kashmir)-log(world). Since all
species tend to occupy more localities nearer to
the centres of their distributions, then if the more
widespread species world-wide were generally
found at more localities within comparable parts
of their distributions, perhaps the deviation of the
Kashmir species from any underlying positive
relationship in Fig. 13 might show some influence
of how far Kashmir is from their centres of world-
wide distribution.
The relationship between distribution patterns
and patterns of local abundance found among
British bumble bees (Williams, 1988) is not consis-
tently evident in the samples from Kashmir.
Those species that were chosen from the maps to
114 PAULH. WILLIAMS
100
distribution centre WIDESPREAD
near Kashmir SPECIES
oO aSialticus
tunicalus e
rufofasciatus
A C] melanurus
EGIL OY
Bs kashmirensis (a keriensis
ow
28 i: A
ee biroi VIN
ie himalayanus 7 aris O lucorum
£5 10 avinoviellus
GI pyrosoma
aM ;
2s simillimus subtypicus
25
3 = wy rifasciatus
s 3
& marussinus L\ © hypnorum
ladakhensis we haemorrhoidalis
personatus Vv
RESTRICTED distribution margin
1-] SPECIES oberti OY e lemniscatus near Kashmir
1 10 100
distribution world-wide
(number of world grid-squares)
O Palaearctic & Oriental, cool temperate/upper montane forest species
ial Central Asian & Tibetan, temperate/montane steppe species
A Central Asian, montane steppe species
vV Tibetan, alpine steppe species
A Oriental (peri-Tibetan), alpine steppe species
@~ Oriental, upper montane forest species
W Oriental, lower montane forest species
p endemic West Himalayan species
Fig. 13 Plot of how widespread the species of social bumble bees (i.e. excluding species of the subgenus Psithyrus) are
within Kashmir against how widespread they are world-wide (logarithmic axes). Data are taken from occupancy of the
number of localities within Kashmir (see Map 4) and from the number of world summary grid-squares (see Map 1)
respectively. The distributions of B. tunicatus and B. lemniscatus are anomalous.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
be nearer to the centres of their world-wide
distributions in Kashmir do not appear to be
generally more abundant locally in these samples
(local abundance for a species estimated as the
total numbers of individuals examined from
Kashmir / numbers of localities in Kashmir,
Fisher-Pitman randomisation test, W = 145,
single-tailed P = 0.19 n.s.). However, there is still
a tendency for the more widespread species within
Kashmir to be more abundant locally (Kendall
coefficient of rank correlation 0.29, single-tailed
P < (00.025). It is possible that even if there were a
relationship with the world-wide pattern, it may
have been obscured in these data because the later
collections at least were strongly biassed against
including many specimens of the common species
from each site, whereas the rare species were
deliberately sought.
Relationships between the patterns of distribu-
tion at different spatial scales form the basis of a
simple model for the interaction of those ecologi-
cal factors that are most likely to govern bumble
bee distributions (discussed in Williams, 1988,
1989). It follows from this that where some
disturbance of sites causes a major reduction in
food availability, it is the species that are nearest
to the centres of their world-wide distributions
that are most likely to persist. This may provide
one explanation for the observation that B. tuni-
catus and B. asiaticus are the species that are often
found alone in severely over-grazed areas around
the edge of the Vale of Kashmir and in the Sind
valley.
COLOUR PATTERNS OF THE KASHMIR
FAUNA
The colour patterns of bumble bees often vary
greatly among individuals of one species (e.g. Fig.
11), but may be closely similar between indivi-
duals of unrelated species where these species
occur together (Fig. 12; see e.g. Vogt, 1909, 1911;
Sakagami & Yoshikawa, 1961; Tkalci, 1968),
1989; Pekkarinen, 1979; Plowright & Owen,
1980). Genetic determination of characteristics
of the colour pattern with simple Mendelian
inheritance has been demonstrated for the North
American B. melanopygus and B. rufocinctus
(Owen & Plowright, 1980; Plowright & Owen,
1980; Owen, 1986). But in the face of such marked
regional convergence, several authors have ques-
tioned whether variation among colour patterns
of individuals within a population is always deter-
mined genetically, or whether it is perhaps more
115
often determined directly by environmental
factors, such as diet, temperature or humidity
during development (e.g. Vogt, 1909, 1911;
Reinig, 1935; Pittioni, 1942, 1943).
The pubescence of bumble bees develops
during the pupal stage and the colours appear
soon after the adults emerge from their cocoons.
Adult queen and worker bees invest much time
and energy ensuring that the conditions in the nest
remain nearly constant, for instance by incubating
brood or by fanning air over the nest to regulate
temperature as necessary (e.g. Heinrich, 1979).
Hence the brood is shielded from many of the
possible environmental effects on colour pattern
during its development, so colonies usually pro-
duce individuals with relatively homogeneous
colour patterns. However, if a nest is exposed and
moved to a nest box, unusual colour patterns are
more often found among the adults that sub-
sequently emerge from pupation (pers. obs.).
The few examples of pronounced caste dimor-
phisms show that colour patterns are not solely
determined by genes, but must also be influenced
by the environment. Two examples have been
described previously from subtropical environ-
ments (B. ephippiatus from the mountains of
Costa Rica, see Owen & Plowright, 1980; B.
festivus from the eastern Himalaya, see Ito et al.,
1984), in which workers and males from the same
colonies share similar colour patterns that differ
strongly from those of the queens. Similar differ-
ences exist within species of the rufofasciatus-
group from Kashmir (see Plate 1). For B.
pyrosoma, males usually also differ in that they
have the pubescence predominantly yellow (Figs
424, 427, 430), a characteristic that can be
regarded as sex-linked (see Stiles, 1979). But
colour variation among the females from Kashmir
appears to be related to body size (which increases
through Figs 426, 428, 429, 425, see the comments
on B. pyrosoma) and so to larval food intake.
From the material available it seems most likely
that the white-tailed queens from Kashmir belong
to the same population as the nominal taxon B.
friseanus from Yunnan (synonymised with B.
pyrosoma s.1.). These queens from Yunnan have
the pubescence of terga IV-V red, so they closely
resemble their own workers from Yunnan and
also many of the workers from Kashmir. Thus
caste dimorphism in colour pattern may exist in
only part of the population of B. pyrosoma s.1.
Variation of B. asiaticus in Kashmir
The asiaticus-group is particularly well repre-
sented in collections for an investigation of the
variation in the colour pattern of the pubescence.
116
In Kashmir, two principal nominal taxa occur on
opposite sides of the Great Himalaya range (Map
48), one with an unbanded yellow thorax (Figs
371-376), the other usually with a black band
between the wing bases and an otherwise grey-
white thorax (Figs 383-391). These nominal taxa
are believed to be parts of a single species (see the
comments on B. asiaticus).
The basis of this colour variation can be exam-
ined where the regional colour patterns occur
together. At a few sites at the upper end of the
higher valleys in the Great Himalaya, the B.
asiaticus are not uniformly of one or other of the
two principal colour patterns, but instead many
individuals show a variety of combinations of their
differing character states (Figs 377-382). The
largest sample with this kind of variation was
collected by F. Schmid from Lal Pani in the
upper Kishanganga valley between 2-3.ix.1953
(BMNH: Table 7). This is likely to be a relatively
unbiassed sample insofar as he specialised in
collecting Trichoptera and not particular bumble
bee taxa. I have also collected a small sample that
shows similar variation, from Nigagar in the upper
Sind valley (9.viii & 17.ix.1986, PW). The occur-
rence of the black tail-colour state (Figs 379-381)
at both of these localities is curious. It is not
known for individuals from the adjacent regions
of Ladakh or the Vale of Kashmir, although it is
common in parts of the Hindu Kush ranges
(BMNH).
The ability of simple population genetic models
to account for the observed patterns of variation
Table 7 Numbers of individuals of B. asiaticus
collected by Schmid from Lal Pani 2-3.ix.1953 with
each combination of states of three characters of the
colour pattern of the pubescence. The scoring was
simplified prior to analysis in order to present discrete
states despite some limited intergradation. The black
thoracic band is deemed to be present if it appears well
defined from the dorsal aspect. The pale colour
‘white’ includes very pale cream. The ‘tail’ colour is
deemed to be red if the pubescence of at least two of
the apical terga is predominantly red.
Thoracic band Pale bands Tail colour Females Males
present white red 7 1*
absent white red - 1
present white black 2 1
absent white black 1 -
present yellow black 3 34
absent yellow black - 13
present yellow red 9 37
absent yellow red - Se
* B. callophenax of Richards, not of Cockerell
** B. longiceps Smith
PAULH. WILLIAMS
Table 8 Association between pairs of character states
of the colour pattern among the males of B. asiaticus
from Lal Pani (see Table 7). None of these
comparisons shows significant association (chi-square
two-sample test, all comparisons P>0.05).
Pale bands Tail colour
white yellow red black
Thoracic present 2 71 38 35
band absent 18 6 13
if
Tail red 2 42
colour black 1
among the specimens from these boundary sites
can be tested if certain assumptions are made.
Three aspects of the variation are chosen because
they appear to be among the most important
elements of the bees’ colour pattern as they
are viewed flying among flowers. It is assumed
initially that each character shows either of two
states and that these are each determined by one
of a pair of alleles at a single locus. In fact the three
characters selected do not show entirely discrete
states. However, the minority of intermediate
specimens from the possible hybrid zone can be
assigned to these states for a preliminary analysis
(Table 7), at least until further information on
modifier effects is available.
There is no association between the states of the
three colour characters among individuals from
the top of the Kishanganga valley (Table 8). This
apparent independent assortment of the charac-
ters shows an absence of the disequilibrium (e.g.
Mallet, 1986) that would be expected if gene flow
across the coincident character clines in the hybrid
zone were large. Hybrid zones are expected to be
captured by linear habitat features where popula-
tion density is particularly low (e.g. Mallet, 1986;
Hewitt, 1988). For B. asiaticus in Kashmir, this
role may be fulfilled by the high ridges and passes
of the Great Himalaya range. For instance, where
the upper Sind valley reaches the main ridge of the
Great Himalaya just above Nigagar, the valley
floor rises abruptly by about 700 m at the Zoji-La
pass. My small samples from either side of
Nigagar, at Sonamarg (2700 m) and above Gummi
(3700 m), each consist of material with a single
colour pattern (i.e. the alleles may be nearly fixed),
so the hybrid zone at the top of the Sind valley may
be less than 16 km wide. Such a narrow cline is
consistent with the interpretation that there is
little gene flow across the hybrid zone at the top of
the Kishanganga valley (Table 8). The hybrid
zone at the top of the Kishanganga may, nonethe-
less, be much broader (Map 48, see the comments
on variation within Kashmir for B. asiaticus).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
GLY?
Table 9 Observed numbers of individuals of B. asiaticus from Lal Pani with each state of three colour pattern
characters (see Table 7), together with the expected numbers from simple genetic models. Frequencies from a small
sample (n = 20) from Nigagar are included for comparison. Deviation from model significant at P<0.05 *,
P<0.001 ***, or not significant ns.
Observed:
females
(diploid)
Thoracic band present 21
absent 1
Frequency (absent) Lal Pani 0.05
[Frequency(absent) Nigagar 0.00
(Chi-square one-sample test, df = 1)
Expected:
males females females
(haploid) —nodominance -simple dominance
73 17 [est(p) = 73/92] 21
19 5 [est(q) = 19/92] 1
0.21
0.66]
(4.14) (0.00)
P>>0.05 ns
Probability that deviation of observation from expectation is due to chanceP<0.05 *
Pale bands white 10
yellow 12
Frequency (yellow) Lal Pani 0.55
[Frequency (yellow) Nigagar 0.50
(Chi-square one-sample test, df = 1)
3 1 [est(p) = 3/92] 1
89 21 [est(q) = 89/92] 21
0.97
0.66]
(84.86) (84.86)
P<<0.05 ***
Probability that deviation of observation from expectation is due to chanceP<<0.05 ***
Tail colour red 16
black
Frequency (black) Lal Pani 0.27
[Frequency (black) Nigagar 0.14
(Chi-square one-sample test, df = 1)
Probability that deviation of observation from expectation is due to chanceP<0.05 *
The frequency of alleles can be estimated and
the dominant allele can be determined from the
frequencies of the character states, because
although female bumble bees are diploid, the
males are usually haploid (see Plowright &
Pallett, 1979). The frequency of the character
states among males in the large sample from Lal
Pani is assumed to be representative of the allele
frequencies in that local part of the population
(which assumes that even if the material was
collected in the proximity of only a very few
colonies, it still represents a random sample of the
local gene pool). These frequencies may differ at
other localities in the Kishanganga valley or at
Nigagar, so these samples cannot be combined for
analysis. If there is simple, complete dominance
by one of the two alternative alleles at each locus,
then the frequencies of character states should
differ between males and females, with higher
frequencies of expression for the recessive allele
among males. The higher frequencies among
males of the lack of a black thoracic band, of
yellow pale bands and of black ‘tails’ at both Lal
Pani and Nigagar (Table 9) are consistent with
their determination by recessive alleles. Further-
more, the frequencies of characters among males
44 11 [est(p) = 44/92] 16
48 11 [est(q) = 48/92] 6
0.52
0.33]
(4.55) (0.00)
P>>0.05 ns
can be used to calculate the frequencies among
females that would be expected if there were
no dominance (which might result from Hardy-
Weinberg equilibrium with co-dominance of both
alleles, or from some mechanism other than
straightforward Mendelian genetics). These cal-
culated frequencies are found to differ signifi-
cantly from those observed for all three characters
(Table 9).
The observed frequencies of the states of the
black thoracic band and of the tail colour among
females are consistent with those expected from
the Hardy-Weinberg equilibrium model with
simple, complete dominance between two alleles
(Table 9). However, this is rejected for the pale
band colour character, the distribution of which
was not consistent with the dominance-free model
either. It has to be noted that the chi-square one-
sample test is not reliable when an expected
frequency is less than 5 (e.g. Siegel, 1956), but the
values of chi-square that result here are so large
(84.8, df = 1) that the conclusion that the model
should be rejected is unlikely to be altered.
The reason for the failure of the models must be
that their assumptions are violated. In addition
to those mentioned above, assumptions of the
118
equilibrium model include insignificant dispersal,
random mating and unselective predation.
Firm conclusions as to the genetic basis of the
patterns of colour variation among these bumble
bees cannot be drawn without breeding experi-
ments. This would allow ideas of the pattern of
inheritance to be tested, although this is not
particularly easy with bumble bees (but see Owen
& Plowright, 1980). However, the pattern of
variation for two out of three of the characters that
have been analysed is consistent with simple
genetic models. It is therefore also consistent with
the assumption that individuals with these colour
patterns are interbreeding as parts of a single
population and a single biological species. Thus
local climate or diet does not appear to be the
proximate factor governing the close convergence
in colour pattern between this species and the
regionally covarying groups of species on either
side of the Great Himalaya.
Convergent colour patterns
It is noticeable that only a few of the possible
colour patterns are actually found among bumble
bees world-wide, given the range of colours shown
by the pale pubescence of bumble bees and the
observed patterns of disposition of bands of pale
and black pubescence on their bodies (Plowright
& Owen, 1980; BMNH, PW). For instance, at
least half of all species of bumble bees have some
individuals with a ‘red-tailed’ colour pattern.
Other common elements in the colour pattern are
a band of black hairs on the thoracic dorsum
between the wing bases, and a black band on
tergum III. The most widespread colour pattern
of the dorsal pubescence among all bumble bees
of the world appears to be for the thorax to be
yellow with a band of black hairs between the wing
bases and for the gaster to be yellow — black — red,
from base to apex (cf. Figs 233-235, 242, 243,
298-300, 317, 318, 377, 378, 382, 388, 393, 407,
412-416, 426).
The three principal regions of Kashmir have
groups of bumble bee species with characteristic
colour patterns of the dorsal pubescence (Fig. 12):
(1A) In the high alpine zone of Ladakh, the most
common colour pattern of the pubescence is
for the thorax to have yellow transverse
bands anteriorly and posteriorly, with a
black band between the wing bases. The
gaster is yellow (terga I-II) and red (terga
III-V), although often with a narrow black
band between them (basal quarter or less of
tergum III).
(1B) At lower altitudes in the arid steppe of
PAULH. WILLIAMS
Ladakh, the common colour pattern is for
the pubescence of the thoracic dorsum to be
uniformly yellow and for the gaster to have
a yellow band (terga I-II yellow, or just
tergum I yellow, or terga I-II completely
black) — black (at least tergum III) — red
(usually terga IV-V).
(2A) Around the Vale of Kashmir, above the
forests, the most common pattern is for
the pubescence of the thoracic dorsum to
be grey-white with a black band between
the wing bases. The gaster is grey-white
(tergum I) — yellow (usually tergum II) —red
(terga III-I[V, sometimes with white on
terga IV-V).
(2B) Inthe Vale of Kashmir and the surrounding
upper montane forest, the common colour
pattern is banded and white as for type 2A,
but the gaster lacks any yellow and is grey-
white (tergum I, sometimes tergum IT) —
black (at least tergum III) — red (usually
terga IV-V).
(3) Inthe lower montane forests of the Jammu
foothills, the common colour pattern is for
the pubescence of the thorax to be black and
for that of the gaster to be yellow (terga I-II)
and red (terga III-V), but without an
obvious black band.
The type 1A colour pattern deviates only slightly
from the widespread bumble bee pattern of three
yellow bands and a red ‘tail’ by the replacement of
much of the black on tergum III by red. This type
1A pattern is otherwise most predominant among
individuals from the Pyrenees in Europe.
The type 1B pattern differs from the wide-
spread pattern principally in the absence of a
black band on the thorax, although it retains an
extensive black area on the gaster. It is predomi-
nant in the Hindu Kush and occurs elsewhere in
the Caucasus (B. (Melanobombus) lapidarius)
and quite independently in South America (B.
(Fervidobombus) bellicosus Smith).
The type 2A pattern differs from the wide-
spread pattern by a change in the pale pubescence
from yellow to grey-white on all but tergum II.
The type 2A pattern is perhaps most predominant
in Kashmir, but is also found around the southern
and eastern margins of the Tibetan massif.
The type 2B pattern completes the transition of
all of the yellow pubescence of the widespread
pattern to grey-white. Individuals with this colour
pattern occur widely in the Himalaya, the central
Asian mountains and westwards in the Caenozoic
mountains of southern Europe to as far west as the
Alps (B. (Pyrobombus) brodmannicus) and the
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
Atlas mountains of North Africa (B. lapidarius).
Males from populations with the type 2B pattern
often retain the common banded yellow pattern
(e.g. B. wurflenii and B. lapidarius from the
Caucasus, B. brodmannicus from the Alps, B.
lapidarius from the Atlas, BMNH). A variant of
the type 2B pattern has tergum II or terga I-II dark
chocolate-brown. This is seen in the Himalaya
only among the males and workers of B. tunicatus
(Figs 348, 349, 351, 352, 354) and B. simillimus
(Figs 418, 420, 421), but it is also known from the
Caucasus mountains for B. (Thoracobombus)
mlokosievitzii Radoszkowski (BMNH, PW).
The type 3 pattern differs from the widespread
pattern by the replacement of the pale pubescence
on the thoracic dorsum by black and by the
reduction of the black band on the gaster. In the
western Himalaya individuals with this pattern are
restricted to the outer foothills. Further east in
Uttar Pradesh the same pattern is shared by B.
(Alpigenobombus) breviceps and B. (Pyrobom-
bus) rotundiceps.
So what is responsible for the apparent con-
straints on the range of bumble bee colour
patterns that are actually observed? Why do these
colour patterns appear to have converged es-
pecially closely among many species within each
geographical region? These species are often not
closely related, as shown by differences in their
male genitalia. Colour pattern had no effect on
acceptance of females by patrolling males of
British B. pratorum (L.) (Free, 1971). Stiles
(1979) suggested that colour affects radiative
properties that may be crucial to thermoregula-
tion in extreme climates. Sakagami & Ito (1981)
challenged this interpretation of sexual dimorph-
ism in colour pattern, because the two faunas
compared by Stiles were dominated by different
groups of species. Within each of these subgenera,
Fervidobombus and Pyrobombus, no regional
trends were found. However, there may be differ-
ences in the kind of habitat and in mate-searching
behaviour between the species of these two
groups that could also affect the relative advan-
tages of the two kinds of colour patterns. The
peculiar behaviour of males of some species of the
subgenus Fervidobombus has been described by
Lloyd (1981) and by Villalobos & Shelly (1987). It
could be that many of these differences between
the two groups, including differences in colour
pattern, are each related to an underlying differ-
ence in physiology that is linked to their regional
patterns of distribution more directly.
Many species of bumble bees commonly pro-
duce individuals that are entirely black, but these
are most often predominant in tropical environ-
ments (Franklin, 1913). These species include B.
119
(Rufipedibombus) rufipes, B. (Megabombus)
melanopoda Cockerell and workers and males of
B. (Senexibombus) senex Vollenhoven in the
East Indies (BMNH), and B. (Fervidobombus)
brevivillus Franklin, B. (Fervidobombus) niger
Franklin [= B. atratus Franklin, preoccupied
name], B. (Fervidobombus) pullatus Franklin and
B. (Fervidobombus) morio (Swederus) in South
America (BMNH, PW). An extensively black
body is likely to have a lower reflectance even in
the infra-red part of the spectrum and so should
have a higher rate of approach to thermal equi-
librium with the environment by an exchange of
long-wave radiation. This may be an advantage to
actively foraging bumble bees in a warm environ-
ment, because they produce a lot of excess heat
from the flight muscles and so might otherwise
suffer from heat stress. The subtropical black bees
listed above may often be forest-dwelling. In the
shade they can escape the additional problem of
rapid overheating that could arise from an equally
efficient absorption by black pigments of the
short-wave radiation in direct sunlight. So this
likely advantage of the black pubescence for bees
foraging in a shaded warm environment may
account for why the type 3 pattern, with a black
thorax, the source of the endogenous heat, pre-
dominates among bumble bees at the lowest
altitudes at which they occur in Kashmir (Fig. 12).
It is very noticeable that near their lower altitudi-
nal and latitudinal limits in Kashmir these bees (B.
trifasciatus and B. haemorrhoidalis) forage par-
ticularly both in the shade and during the cooler
periods of the day (see the discussion of the
distribution of the fauna with altitude). However,
further south in their distributions in South East
Asia, these same species have extensively orange-
brown colour patterns of the thoracic pubescence
(Fig. 11). More information is needed on the
foraging behaviour of these bees in South East
Asia, where there is a tendency among the few
large nocturnal bees and wasps to have an orange-
brown colour pattern (M. C. Day, pers. comm.;
Roubik, 1989). There is also clearly a need for
measurements of the radiative properties of
bumble bee colour patterns in general.
Male bumble bees that patrol in the higher
forest on Mt Apharwat tend to be almost uni-
formly bright yellow (Fig. 6 and see the comments
on B. pyrosoma, B. lucorum and B. hypnorum).
This may also be true of those males that patrol
in forests, but not in open habitats, at higher
latitudes (see Pekkarinen, 1979: fig. 7, cf. Loken,
1973: fig.2). In contrast to black, a pale, highly
reflective body colour should reduce the rate of
approach to thermal equilibrium with the environ-
ment by an exchange of long-wave radiation, so
120
that following an active warm-up, a sufficiently
high excess thoracic temperature for flight can be
reached quickly and maintained in a shaded cool
environment. In contrast, males of those species
that patrol in the forests at low altitudes, in open
areas outside the forest or in a broad range of
habitats in Kashmir tend to be darker and share
the colour pattern of the females (Fig. 6). Uni-
form bright yellow may be disadvantageous in
open areas beyond the forest, for instance in
alpine and arctic zones, because it is very con-
spicuous to predators. This could be a particular
problem for the more sedentary ‘perch and sprint’
males (e.g. B. asiaticus and B. rufofasciatus, see
the introduction on mate-searching behaviour of
male bumble bees in Kashmir) and those males
that aggregate at nest entrances (which includes
several species of Fervidobombus, e.g. review by
Schremmer, 1972; Lloyd, 1981; Villalobos &
Shelly, 1987). Yet the most uniformly light-
coloured female bumble bees in Kashmir, which
belong to the type 1A pattern-group, are found at
the highest altitudes sampled for bumble bees, in
Ladakh. This trend towards lighter colours in
cooler environments is the reverse of that des-
cribed by Pekkarinen (1979) from European
species.
Regional convergences in the finer details of the
colour pattern between bumble bee species are
likely to be the result of selective predation.
Predators with perception of colour and great
visual acuity, perhaps certain birds, may avoid the
familiar common colour patterns of these insects
because the females can inflict a painful sting, and
preferentially attack any individuals with unusual
colour patterns. This kind of selection could bring
about Millerian mimicry among the colour pat-
terns on the dorsa of the females (Plowright &
Owen, 1980) and Batesian automimicry by the
colour patterns on the dorsa of the stingless males
(Stiles, 1979). The putative models for this
process would be the colour patterns of the
females of the most abundant species. In Kashmir,
these species tend to be regionally restricted and
relatively invariant in their colour patterns (type
1A, B. oberti, Figs 392, 393; type 1B, B. semeno-
vianus, Figs 397, 398, 400, 401; type 2A, B.
rufofasciatus, Figs 431, 432, 434, 435, 438; type
2B, B. tunicatus, Figs 350, 353; type 3, B.
haemorrhoidalis, Figs 279, 280; see Fig. 12). Many
of the mimics occupy the middle classes of abun-
dance, but are often more widespread among the
regions of Kashmir. These species covary in their
colour patterns among the regions so that they
show the protective livery of the local model (e.g.
B. avinoviellus, Figs 232-241; B. himalayanus,
Figs 242-253; B. kashmirensis, Figs 295-310; B.
PAULH. WILLIAMS
lepidus, Figs 327, 328, 330, 331, 333, 334; B. biroi,
Figs 335-346; B. asiaticus, Figs 371-391).
A subtle difference between the colour patterns
of B. kashmirensis and B. rufofasciatus could be
interpreted as further circumstantial evidence in
support of the mimicry hypothesis for colour
convergence. Workers of these two species ap-
pear to be identical (type 2A colour pattern) when
they are seen manoeuvring slowly from flower to
flower in the alpine and subalpine zones of the
mountains around the Vale of Kashmir. Most
workers of B. rufofasciatus have the hairs of
gastral terga III-[V almost uniformly red and
those of tergum V uniformly white (Fig. 435). The
B. kashmirensis from the same region often have
all three terga (III-V) covered with hairs that are
red at the base, but which become more exten-
sively white-tipped towards the apex of the gaster
(Fig. 303). This achieves the same white-tailed
effect as for B. rufofasciatus, because the apex of
the gaster curves slightly towards the ventral for
bumble bees.
Colour variation within many of the bumble bee
species of Kashmir does appear to be much
greater than within the bumble bee species of
Britain, or even within the more variable species
of the Alps. This may be blamed on taxonomic
‘lumping’ in this review. But if bumble bee colour
patterns have converged into separate and distinc-
tive mimetic groups within each of the neighbour-
ing regions of Asia, then the position of Kashmir
at the frontier between the large and divergent
Oriental and Palaearctic faunas (which are still to
some extent separated even inside Kashmir by the
divide of the Great Himalaya) may account in part
for the great diversity of colour pattern within
some of the Kashmir species.
Acknowledgements. I am very ‘prateful to all of
those who gave advice, with special thanks to
B. Bolton and M. C. Day. D. Agosti, N. H. Barton,
R. L. Blackman, S. A. Corbet, A. O. Divers,
W. G. Eberhard, P. J. Eggleton, K. Gaston, C. J.
Humphries, L. S. Kimsey, R. P. Macfarlane, A.
Pekkarinen, R. C. Plowright, P. Rasmont, M. J.
Scoble and R. I. Wane-Wright commented on parts
of the manuscript. C. R. Betts, G. R. Else, N. D. M.
Fergusson, L. Ficken, M. G. Fitton, P. Gilbert,
P. M. Hammond, J. M. Harvey, B. R. Pitkin, A.
Polaszek, J. Spratt, N. D. Springate, D. A. Sutton,
B. C. Townsend, C. R. Vardy, S. L. Ward and
D. B. Williams helped with equipment, library
searches and translations. M. A. Mir (No. 1 Guide,
Gulmarg) provided support in the field and C. A.
Chadwell helped with certain plant determina-
tions. I am very grateful to the Keeper of
Entomology for providing facilities for this work.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
I am also much indebted to the following who
helped by lending material: A. V. Antropov
(ZMMU), K. Arakaki (BPBM), E. Diller (ZS), M.
Fischer (NM), W. Hogenes (ITZ), F. Koch and I.
Wegener (MNHU), J.-P. Kopelke (NMS), J. Macek
(NMP), K. C. McGiffen (INHS), R. J. McGinley and
B. B. Norden (USNM), C. D. Michener (SEMK), J.
Papp (T,), B. Petersen (ZM), V. Raineri (MCSN), P.
Rasmont (FA), B. Tkalct, S.-f. Wang (IZ), J. C.
Weulersse (MNHN) and especially Y. A. Pesenko
and M. V. Podbolotskaya (ZI). N. R. Bogatyrov,
W. Budenburg, C. A. Chadwell, C. S. Cockell, R.
Desmier de Chenon, A. O. Divers, R. W. Husband,
A. Hutchings, M. Ito, L. W. Macior, S. Martin, H.
Osmaston, B. Petersen, N. Saville, G. S. Vick and
S.-f. Wang kindly donated or exchanged other
Asian specimens used here.
The work was carried out during a Postdoctoral
Research Fellowship funded by the Natural
Environment Research Council, which also
awarded a grant towards publication costs. Addi-
tional assistance for the purchase of a microscope
was provided from the Entomological Fund, and
assistance for the purchase of computer hard-
ware for the automated mapping system was pro-
vided by the Systematics Association.
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THE BUMBLE BEES OF THE KASHMIR HIMALAYA
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Plate 1 Representatives of the rufofasciatus-group from the Vale of Kashmir. In this part of their distribution, all three
species show caste-dependent colour dimorphism among females. There is also gender-dependent colour variation
among Bombus pyrosoma and B. simillimus. Top row: B. rufofasciatus (left to right: queen, worker, male), from a nest
on Mt Apharwat, in subalpine pasture. Middle row: B. Pyrosoma (queen, worker, male), from a nest on Apharwat, in
the coniferous forest. Bottom row: B. simillimus (queen, worker, male), from a nest near Harwan, in valley scrub.
Photographed by Harry Taylor, BMNH photographic unit.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 129
ocellus
ocello-ocular area
compound eye
antenna
clypeus
oculo-malar distance
malar area
mandibular breadth
labrum
15
mandible
se 2 ocellar breadth
ocello-ocular distance
Figs 14-22 Morphology of the head: (14) anterior aspect of the head of a banded white queen B. avinoviellus,
Gulmarg; (15) left lateral aspect of a worker B. Jemniscatus, Apharwat, showing measurements of the malar area; (16)
dorsal aspect of a banded white queen B. avinoviellus, Gulmarg, showing measurements of the ocello-ocular area;
dorsal aspect of segments 3—5 of the left antenna of queens of (17) B. trifasciatus, Gulmarg; (18) B. haemorrhoidalis,
Muzaffarabad; (19) banded white B. asiaticus, Gulmarg; (20) B. oberti, Nimaling; the same, males of (21) B.
bohemicus, Lal Pani; (22) B. ferganicus, Gulmarg.
130
basal depression
transverse ridge
apical depression
PAULH. WILLIAMS
median furrow
lamella
lateral tubercle
23 24 25
a)
26
29 30
| | / { |
31 Ki 33 34
basal tooth | apical tooth
7
Yi
basal keel sulcus obliquus
incisura
novus
intercalary tooth
avinoviellus kashmirensis rufofasciatus
Figs 23-38 Morphology of the head: anterior aspect of the labrum of females of (23) banded white B. avinoviellus,
Gulmarg; (24) B. novus, Apharwat; (25) yellow and white B. kashmirensis, Apharwat; (26) B. rufofasciatus,
Apharwat; anterior aspect of the labrum of males of (27) banded white B. avinoviellus, Gulmarg; (28) B. novus,
Apharwat; (29) yellow and white B. kashmirensis, Apharwat; (30) B. rufofasciatus, Apharwat; outer lateral aspect of
the left mandible of females of (31) banded white B. avinoviellus, Gulmarg; (32) B. novus, Apharwat; (33) yellow and
white B. kashmirensis, Apharwat; (34) B. rufofasciatus, Apharwat; outer lateral aspect of the left mandible of males of
(35) banded white B. avinoviellus, Gulmarg; (36) B. novus, Apharwat; (37) yellow and white B. kashmirensis,
Apharwat; (38) B. rufofasciatus, Apharwat.
THE BUMBLE BEES OFTHE KASHMIR HIMALA YA 131
proximal margin
femur
i tibia
oD
S
E
i=] & corbicular area
50 =
I =
E Z
z 3
S a.
i=
2
i=
s
proximo-posterior
distal margin corner
basitarsus
43
gonobase
gonocoxite
spatha
penis valve
gonostylus
volsella
44
Figs 39-45 Morphology of the thorax and gaster: outer lateral aspect of the basitarsus of the left mid leg of workers of
(39) B. trifasciatus, Gulmarg; (40) B. haemorrhoidalis, Patnitop; outer lateral aspect of the basitarsus of the left hind
leg of queens of (41) unbanded B. subtypicus, Dras; (42) banded yellow and white B. biroi, Apharwat; (43) outer
lateral aspect of the left hind leg of a banded white female B. avinoviellus, Gulmarg; (44) dorsal aspect of the genital
capsule of a banded white male B. avinoviellus, Gulmarg; (45) ventral aspect of the same (weakly sclerotised areas
stippled).
132 PAULH. WILLIAMS
60
Se ee, 51
61
a ee)
62
63
Baa 55 Cre ia 65
Figs 46-65 Morphology of the male genitalia: postero-dorsal aspect of the right penis valve of (46) banded yellow B.
avinoviellus, Lal Pani; (47) banded white B. avinoviellus, Gulmarg; (48) yellow and white B. himalayanus, Apharwat;
(49) B. marussinus, Batakush; (50) B. bohemicus, Lal Pani; (51) B. novus, Apharwat; (52) B. branickii, Chogdo; (53)
B. ferganicus, Gulmarg; (54) yellow B. morawitzianus, Batakush; (55) B. skorikovi, Lal Pani; (56) B. trifasciatus,
Harwan; (57) B. haemorrhoidalis, Muzaffarabad; (58) unbanded B. melanurus, Chogdo; (59) unbanded B.
melanurus, Gulmarg; (60) B. personatus, Chogdo; (61) yellow B. kashmirensis, Chogdo; (62) yellow and white
B. kashmirensis, Apharwat; (63) white B. kashmirensis, Lal Pani; (64) B. hypnorum, Apharwat; (65) banded B.
subtypicus, Lal Pani.
THE BUMBLE BEES OF THE KASHMIR HIMALA YA 133
(eee l- —_ 7 ;
73
fo y ,
Figs 66-85 Morphology of the male genitalia: postero-dorsal aspect of the right penis valve of (66) unbanded B.
subtypicus, Lal Pani; (67) B. lemniscatus, Uttar Pradesh; (68) B. lepidus [with yellow females], Dalti; (69) B. lepidus
[with yellow and white females], Nigagar; (70) unbanded B. biroi, Suru; (71) banded B. biroi, Apharwat; (72) dark
B. tunicatus, Shigar; (73) light B. tunicatus, Gulmarg; (74) dark B. lucorum, Gilgit; (75) light B. lucorum, Apharwat;
(76) unbanded yellow [red-tailed] B. asiaticus, Leh; (77) banded yellow black-tailed B. asiaticus, Nigagar; (78) banded
white [red-tailed] B. asiaticus, Gulmarg; (79) B. oberti, Pamir; (80) B. ladakhensis, Hemis; (81) B. semenovianus,
Hemis; (82) B. keriensis [with yellow females], Apharwat; (83) B. simillimus, Banihal; (84) B. pyrosoma, Apharwat;
(85) B. rufofasciatus, Apharwat.
75
134
86
87
88
89
95
=
a
PAULH. WILLIAMS
96
97
99
100
101
102
104
105
Figs 86-105 Morphology of the male genitalia: outer lateral aspect of the right penis valve of (86) banded yellow B.
avinoviellus, Lal Pani; (87) banded white B. avinoviellus, Gulmarg; (88) yellow and white B. himalayanus, Apharwat;
(89) B. marussinus, Batakush; (90) B. bohemicus, Lal Pani; (91) B. novus, Apharwat; (92) B. branickii, Chogdo; (93)
B. ferganicus, Gulmarg; (94) yellow B. morawitzianus, Batakush; (95) B. skorikovi, Lal Pani; (96) B. trifasciatus,
Harwan; (97) B. haemorrhoidalis, Muzaffarabad; (98) unbanded B. melanurus, Chogdo; (99) unbanded B.
melanurus, Gulmarg; (100) B. personatus, Chogdo; (101) yellow B. kashmirensis, Chogdo; (102) yellow and white
B. kashmirensis, Apharwat; (103) white B. kashmirensis, Lal Pani; (104) B. hypnorum, Apharwat; (105) banded
B. subtypicus, Lal Pani.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 135
2 eee
a 106 —- 116
——
107 117
a 108 — 118
en
a= 112 122
yi 113 123
Figs 106-125 Morphology of the male genitalia: outer lateral aspect of the right penis valve of (106) unbanded B.
subtypicus, Lal Pani; (107) B. lemniscatus, Uttar Pradesh; (108) B. lepidu [with yellow females], Dalti; (109) B.
lepidus [with yellow and white females], Nigagar; (110) unbanded B. biroi, Suru; (111) banded B. biroi, Apharwat;
(112) dark B. tunicatus, Shigar; (113) light B. tunicatus, Gulmarg; (114) dark B. lucorum, Gilgit; (115) light B.
lucorum, Apharwat; (116) unbanded yellow [red-tailed] B. asiaticus, Leh; (117) banded yellow black-tailed
B. asiaticus, Nigagar; (118) banded white [red-tailed] B. asiaticus, Gulmarg; (119) B. oberti, Pamir; (120)
B. ladakhensis, Hemis; (121) B. semenovianus, Hemis; (122) B. keriensis [with yellow females], Apharwat; (123) B.
simillimus, Banihal; (124) B. pyrosoma, Apharwat; (125) B. rufofasciatus, Apharwat.
136 PAULH. WILLIAMS
126 Gee ae
128 cay 138
131 (5
132 142
133 143
134 144
135 145
Figs 126-145 Morphology of the male genitalia: interio-ventral aspect of the right volsella of (126) banded yellow B.
avinoviellus, Lal Pani; (127) banded white B. avinoviellus, Gulmarg; (128) yellow and white B. himalayanus,
Apharwat; (129) B. marussinus, Batakush; (130) B. bohemicus, Lal Pani; (131) B. novus, Apharwat; (132) B.
branickii, Chogdo; (133) B. ferganicus, Gulmarg; (134) yellow B. morawitzianus, Batakush; (135) B. skorikovi, Lal
Pani; (136) B. trifasciatus, Harwan; (137) B. haemorrhoidalis, Muzaffarabad; (138) unbanded B. melanurus, Chogdo;
(139) unbanded B. melanurus, Gulmarg; (140) B. personatus, Chogdo; (141) yellow B. kashmirensis, Chogdo; (142)
yellow and white B. kashmirensis, Apharwat; (143) white B. kashmirensis, Lal Pani; (144) B. hypnorum, Apharwat;
(145) banded B. subtypicus, Lal Pani.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 137
156
146
157
147
158
148
159
149
160
3 NS 150 161
v NS 151
162
152
163
153
164
¢ \ 154
165
J
Figs 146-165 Morphology of the male genitalia: interio-ventral aspect of the right volsella of (146) unbanded B.
subtypicus, Lal Pani; (147) B. lemniscatus, Uttar Pradesh; (148) B. lepidus [with yellow females], Dalti; (149)
B. lepidus [with yellow and white females], Nigagar; (150) unbanded B. biroi, Suru; (151) banded B. biroi, Apharwat;
(152) dark B. tunicatus, Shigar; (153) light B. tunicatus, Gulmarg; (154) dark B. lucorum, Gilgit; (155) light B.
lucorum, Apharwat; (156) unbanded yellow [red-tailed] B. asiaticus, Leh; (157) banded yellow black-tailed
B. asiaticus, Nigagar; (158) banded white [red-tailed] B. asiaticus, Gulmarg; (159) B. oberti, Pamir; (160) B.
ladakhensis, Hemis; (161) B. semenovianus, Hemis; (162) B. keriensis [with yellow females], Apharwat; (163)
B. simillimus, Banihal; (164) B. pyrosoma, Apharwat; (165) B. rufofasciatus, Apharwat.
j=
iS)
co
PAULH. WILLIAMS
/\ 166 C 4 176 = 186 (g 196
/\ 167 fA 177 a 187 & 197
Nw ON hn ee
178
(| 169 ae ee 189 (yh 199
179
(ee 170 CY 180 a 190 CS 200
Ou 2 181 EY 191 ee. 201
Nex 182 i) 192 a. 202
172 (A
(SS 183 193 a. 203
173
Figs 166-205 Morphology of the male genitalia: dorsal aspect of the right gonostylus of (166) banded yellow B.
avinoviellus, Lal Pani; (167) banded white B. avinoviellus, Gulmarg; (168) yellow and white B. himalayanus,
Apharwat; (169) B. marussinus, Batakush; (170) B. bohemicus, Lal Pani; (171) B. novus, Apharwat; (172) B.
branickii, Chogdo; (173) B. ferganicus, Gulmarg; (174) yellow B. morawitzianus, Batakush; (175) B. skorikovi, Lal
Pani; (176) B. trifasciatus, Harwan; (177) B. haemorrhoidalis, Muzaffarabad; (178) unbanded B. melanurus, Chogdo;
(179) unbanded B. melanurus, Gulmarg; (180) B. personatus, Chogdo; (181) yellow B. kashmirensis, Chogdo; (182)
yellow and white B. kashmirensis, Apharwat; (183) white B. kashmirensis, Lal Pani; (184) B. hypnorum, Apharwat;
(185) banded B. subtypicus, Lal Pani; (186) unbanded B. subtypicus, Lal Pani; (187) B. lemniscatus, Uttar Pradesh;
(188) B. lepidus [with yellow females], Dalti; (189) B. lepidus [with yellow and white females], Nigagar; (190)
unbanded B. biroi, Suru; (191) banded B. biroi, Apharwat; (192) dark B. tunicatus, Shigar; (193) light B. tunicatus,
Gulmarg; (194) dark B. lucorum, Gilgit; (195) light B. lucorum, Apharwat; (196) unbanded yellow [red-tailed] B.
asiaticus, Leh; (197) banded yellow black-tailed B. asiaticus, Nigagar; (198) banded white [red-tailed] B. asiaticus,
Gulmarg; (199) B. oberti, Pamir; (200) B. ladakhensis, Hemis; (201) B. semenovianus, Hemis; (202) B. keriensis [with
yellow females], Apharwat; (203) B. simillimus, Banihal; (204) B. pyrosoma, Apharwat; (205) B. rufofasciatus,
Apharwat.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 139
= 184 194
fh
204
195 205
| L 174
a 175
X VA 206
207 208
~ 7 209 210 211
212 213 214
Figs 206-214 Morphology of the gaster: ventral aspect of gastral sternum VI of females of (206) B. bohemicus, Britain;
(207) B. novus, Apharwat; (208) B. branickii, Nimaling; (209) B. ferganicus, Gulmarg; (210) B. morawitzianus, Lal
Pani; (211) B. skorikovi, Lal Pani; dorsal aspect of the apex of gastral tergum VI of queens of (212) B. lemniscatus,
Nepal; (213) yellow and white B. lepidus, Apharwat; (214) banded yellow and white B. biroi, Apharwat.
140 PAULH. WILLIAMS
eR nee
Figs 215-219 Morphology of the head (scanning electron micrographs, pubescence partially removed): antero-dorsal
aspect of the left ocello-ocular area of the head of a queen of (215) B. haemorrhoidalis, Muzaffarabad; dorsal aspect of
the left ocello-ocular area of the head of queens of (216) unbanded B. subtypicus, Afghanistan; (217) banded yellow
and white B. biroi, ‘Krishnye’ Kashmir; (218) B. lemniscatus, Nepal; (219) yellow and white B. lepidus, Apharwat.
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 141
Figs 220-224 Morphology of the head and thorax (scanning electron micrographs): antero-ventral aspect of the labrum
of queens of (220) banded yellow and white B. avinoviellus, Gulmarg; (221) yellow and white B. himalayanus, Atro
Sar; (222) B. marussinus, Chhantir Gah; postero-dorsal aspect of the middle of the scutum, pubescence removed, of
queens of (223) B. lemniscatus, Nepal; (224) yellow and white B. lepidus, Apharwat.
PAULH. WILLIAMS
Ws
Ors
\
q y
Figs 225-231 Morphology of the legs and gaster (scanning electron micrographs): outer lateral aspect of the corbicular
area of the hind tibia of queens of (225) banded white B. avinoviellus, Gulmarg; (226) B. simillimus, Srinagar [the
circular structures are attachment cement from the peri-anal glands of uropodid mites, det. D. Macfarlane]; outer
lateral aspect of the proximo-posterior corner of the hind basitarsus of queens of (227) unbanded yellow B. asiaticus,
Nimaling; (228) B. oberti, Nimaling; (229) B. semenovianus, Baltit; dorsal aspect of gastral tergum VI of queens of
(230) B. oberti, Nimaling; (231) B. semenovianus, Baltit.
a "T)
. \
Bombus (Md. ) avinoviellus ae
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 5 Summary of the world-wide distribution of B. avinoviellus (for an explanation of the grid see Map 1).
‘Specimen’ records are from material examined by the author and ‘Literature’ records are from published data.
‘Unconfirmed’ records include both dubious identifications in the literature and records without precise locality data.
The grid-square that includes Kashmir is shown with a solid outline.
Map 6 Distribution of B. avinoviellus in Kashmir (for a comparison with the full range of localities sampled see Map 4).
All records are from specimens that have been examined. The different symbols represent the principal colour
patterns that predominate among workers in each local sample (key to colour pattern symbols see Figs 232-241).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 145
QUEENS
@
WORKERS
MALES
Figs 232-241 Colour patterns of B. avinoviellus from localities in Kashmir: (232) Banidas 30.vi—2.vii.1959, (233)
Shardi 1-13.viii.1953, (234) Lal Pani 2-3.ix.1953, (235) Lal Pani 2-3.ix.1953, (236) Gulmarg vii.1986, (237) Gulmarg
viii.1985, (238) Gulmarg viii.1985, (239) Gulmarg vii.1986, (240) Gulmarg viii.1985, (241) Gulmarg viii.1985 (for the
colour key see Fig. 258).
146 PAULH. WILLIAMS
Bombus (Md. ) himalayanus
@ Specimen G@ Literature © Unconfirmed
Map 7 Summary of the world-wide distribution of B. himalayanus (for an explanation of the symbols see Map 5).
100km
Land above 3000m.
Map 8 Distribution of B. himalayanus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs
242-253).
THE BUMBLE BEES OF THE KASHMIR HIMALA YA 147
QUEENS
)
WORKERS
LEER
me)
ae
id
a
wy
253
MALES
Figs 242-253 Colour patterns of B. himalayanus from localities in Kashmir: (242) Nimaling 17.vii.1980, (243) Chhantir
Gah 5-7. viti.1954, (244) Atro Sar 11.viii.1954, (245) Atro Sar 11.viii.1954, (246) Gumri 10—11.viii.1986, (247) Atro
Sar 11 .viii.1954, (248) Holojut 13-14. viii.1954, (249) Apharwat 23.viii.1985, (250) Apharwat 30.viii.1985, (251)
Apharwat vili.1985, (252) Apharwat 15.viii.1980, (253) Sangisfaid 1985 (for the colour key see Fig. 258).
148 PAULH. WILLIAMS
Bombus (Mad. ) marussinus Che
@ Specimen G@ Literature © Unconfirmed Equal area grid
Map 9 Summary of the world-wide distribution of B. marussinus (for an explanation of the symbols see Map 5).
——— 1!
0 100km
Land above 3000m.
Map 10 Distribution of B. marussinus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 254-257).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 149
QUEEN
WORKERS
MALE
Figs 254-257 Colour patterns of B. marussinus from localities in Kashmir: (254) Chhantir Gah 5-7.viii.1954, (255)
Holojut 13-14. viii.1954, (256) Batakush 8.ix.1954, (257) Chhantir Gah 5—7.viii.1954 (for the colour key see Fig. 258).
150 PAULH. WILLIAMS
Bombus (Ps.) bohemicus
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 11 Summary of the world-wide distribution of B. bohemicus (for an explanation of the symbols see Map 5).
| ee |
0 100km
| Land above 3000m.
Map 12 Distribution of B. bohemicus in Kashmir (see Map 6).
THE BUMBLE BEES OF THE KASHMIR HIMALA YA 151
COLOUR KEY
grey-white/cream-white
LJ
LC
lemon yellow/golden yellow
light brown
chocolate brown
orange-red
dense mixture of black hairs
black
Biv
MALE
Fig. 258 Colour pattern of B. bohemicus from Kashmir: Lal Pani 2-3.ix.1953
152 PAULH. WILLIAMS
Bombus (Ps.) novus
@ Specimen @ Literature © Unconfirmed
Map 13 Summary of the world-wide distribution of B. novus (for an explanation of the symbols see Map 5).
| ee |
0 100km
Land above 3000m.
Map 14 Distribution of B. novus in Kashmir (see Map 6).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 153
CAD
|)
| |
7
a ae
259
FEMALES
&
MALES
Figs 259-262 Colour patterns of B. novus from localities in Kashmir: (259) Apharwat 25.vii.1986, (260) Apharwat
23.vili.1985, (261) Apharwat 25.vii.1986, (262) Apharwat 23.viii.1985 (for the colour key see Fig. 258).
154 PAULH. WILLIAMS
Bombus (Ps.) branickii iS
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 15 Summary of the world-wide distribution of B. branickii (for an explanation of the symbols see Map 5).
100km
Land above 3000m.
Map 16 Distribution of B. branickii in Kashmir (see Map 6).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 155
FEMALES
MALE
Figs 263-266 Colour patterns of B. branickii from localities in Kashmir: (263) Nimaling 19.vii.1980, (264) Chogdo
24.ix.1986, (265) Nimaling 17.vii.1980, (266) Atro Sar 11.viii.1954 (for the colour key see Fig. 258).
156 PAULH. WILLIAMS
Bombus (Ps.) ferganicus
@ Specimen G@ Literature © Unconfirmed Equal area grid
Map 17 Summary of the world-wide distribution of B. ferganicus (for an explanation of the symbols see Map 5).
m="
0 100km
Land above 3000m.
Map 18 Distribution of B. ferganicus in Kashmir (see Map 6).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA US i7/
FEMALE
268
MALE
Figs 267, 268 Colour patterns of B. ferganicus from localities in Kashmir: (267) Gulmarg viii.1985, (268) Gulmarg
viii.1985 (for the colour key see Fig. 258).
158 PAULH. WILLIAMS
Bombus (Ps.) morawitzianus ie
@ Specimen @ Literature © Unconfirmed Equal area
as
Map 19 Summary of the world-wide distribution of B. morawitzianus (for an explanation of the symbols see Map 5).
100km
Land above 3000m.
Map 20 Distribution of B. morawitzianus in Kashmir (see Map 6).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 159
c
CORREA
4
p—“
|
a
_/
W
269
FEMALE
MALE
Figs 269, 270 Colour patterns of B. morawitzianus from localities in Kashmir: (269) Lal Pani 2—-3.ix.1953, (270) Lal
Pani 2-3.ix.1953 (for the colour key see Fig. 258).
160 PAULH. WILLIAMS
Bombus (Ps.) skorikovi ss
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 21 Summary of the world-wide distribution of B. skorikovi (for an explanation of the symbols see Map 5).
| a |
0 100km
Land above 3000m.
Map 22 Distribution of B. skorikovi in Kashmir (see Map 6).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 161
273
FEMALES
ELEN
ey
, |
ay
274
MALES
Figs 271-274 Colour patterns of B. skorikovi from localities in Kashmir: (271) Lal Pani 2-3.ix.1953, (272) Lal Pani
2-3.ix.1953, (273) Apharwat viii.1985, (274) Lal Pani 2-3.ix.1953 (for the colour key see Fig. 258).
162 PAULH. WILLIAMS
Bombus (Dv.) trifasciatus o
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 23 Summary of the world-wide distribution of B. trifasciatus (for an explanation of the symbols see Map 5).
| ee |
0 100km
Land above 3000m.
Map 24 Distribution of B. trifasciatus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 275-278).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 163
QUEEN
WORKERS
MALE
Figs 275-278 Colour patterns of B. trifasciatus from localities in Kashmir: (275) Gulmarg 28.vii.1985, (276) Patnitop
6.ix.1986, (277) Harwan 24.viii.1980, (278) Gulmarg 1913 (for the colour key see Fig. 258).
164 PAULH. WILLIAMS
Bombus (Or.) haemorrhoidalis
@ Literature
@ Specimen © Unconfirmed Equal area grid
Map 25 Summary of the world-wide distribution of B. haemorrhoidalis (for an explanation of the symbols see Map 5).
| ee |
0 100km
Land above 3000m.
Map 26 Distribution of B. haemorrhoidalis in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 279—
281).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 165
QUEEN
WORKER
MALE
Figs 279-281 Colour patterns of B. haemorrhoidalis from localities in Kashmir: (279) Muzaffarabad 10-12.v.1954,
(280) Patnitop 6—8.ix.1986, (281) Muzaffarabad 15.x.1953 (for the colour key see Fig. 258).
166 PAULH. WILLIAMS
Bombus (St.) melanurus eo
@ Specimen @ Literature © Unconfirmed Equal area grid
100km
Land above 3000m.
Map 28 Distribution of B. melanurus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 282-290).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 167
QUEENS
@
WORKERS
MALES
Figs 282-290 Colour patterns of B. melanurus from localities in Kashmir: (282) Nimaling 18.vii.1980, (283) Leh
9.vii.1980, (284) Leh 3.vii.1980, (285) Chogdo 24.ix.1986, (286) Chittakatha Sar 19-24.viii.1953, (287) Gulmarg
vili. 1986, (288) Gulmarg 13.viii.1980, (289) Gulmarg 16.viii.1980, (290) Gulmarg viii.1985 (for the colour key see
Fig. 258).
168 PAULH. WILLIAMS
Bombus (St.) personatus es
@ Specimen G@ Literature © Unconfirmed Equal area grid
Map 29 Summary of the world-wide distribution of B. personatus (for an explanation of the symbols see Map 5).
|
0 100km
Land above 3000m.
Map 30 Distribution of B. personatus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 291-294).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 169
QUEEN
WORKER
MALES
Figs 291-294 Colour patterns of B. personatus from localities in Kashmir: (291) Nimaling 16.vii.1980, (292) Nimaling
21.vii.1980, (293) Chogdo 24.ix.1986, (294) Chogdo 24.ix.1986 (for the colour key see Fig. 258).
170 PAULH. WILLIAMS
Bombus (Ag.) kashmirensis aS
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 31 Summary of the world-wide distribution of B. kashmirensis (for an explanation of the symbols see Map 5).
100km
Land above 3000m.
Map 32 Distribution of B. kashmirensis in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs
295-310).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 171
&
Nas
fore
®
°.
Ty wh»n
My wy
vy 295 we 298
QUEENS
WORKERS
Pua
is ee
Bo 4 4
Tn
(CuI
SSE
307
MALES
Figs 295-310 Colour patterns of B. kashmirensis from localities in Kashmir: (295) Nimaling 20.vii.1980, (296)
Nimaling 15.vii.1980, (297) Chogdo 24.ix.1986, (298) Nimaling 20.vii.1980, (299) Nimaling 19.vii.1980, (300) Chogdo
24.ix.1986, (301) Gumri 10-11. viii.1986, (302) Apharwat 24.vii.1986, (303) Apharwat viii.1985, (304) Apharwat
30. viii.1986, (305) Apharwat 24.viii.1986, (306) Apharwat viii.1986, (307) Apharwat 23.viii.1985, (308) Nigagar
17.ix.1986, (309) Nigagar 17.ix.1986, (310) Lal Pani 2-3.ix.1953 (for the colour key see Fig. 258).
72} PAULH. WILLIAMS
Bombus (Pr.) hypnorum _—S
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 33 Summary of the world-wide distribution of B. hypnorum (for an explanation of the symbols see Map 5).
100km
Land above 3000m.
Map 34 Distribution of B. hypnorum in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 311-316).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 173
QUEENS
WORKERS
fo
Eee]
Caan,
EE /,
313
MALES
Figs 311-316 Colour patterns of B. hypnorum from localities in Kashmir: (311) Apharwat 25.vii.1986, (312) Apharwat
viii.1986, (313) Apharwat 16.viii.1980, (314) Apharwat 25.vii.1986, (315) Apharwat viii.1985, (316) Apharwat
vili. 1985 (for the colour key see Fig. 258).
174 PAULH. WILLIAMS
Bombus (Pr.) subtypicus ae
@ Specimen G@ Literature © Unconfirmed Equal area grid
Map 35 Summary of the world-wide distribution of B. subtypicus (for an explanation of the symbols see Map 5).
|
0 100km
Land above 3000m.
Map 36 Distribution of B. subtypicus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 317-324).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 175
ie
Ww
—_
v=)
QUEENS
| Oe
| ie.
®
Lo)
—
_
=)
12°)
Ne
—)
WORKERS
.-
S “
’
ie
hee
io)
—
foe)
vs)
i)
_
MALES
Figs 317-324 Colour patterns of B. subtypicus from localities in Kashmir: (317) Lal Pani 2-3.ix.1953, (318) Lal Pani
2-3.ix.1953, (319) Dras 10-11 .viii.1986, (320) Dras 10-11. viii. 1986, (321) Lal Pani 2—-3.ix.1953, (322) Astor 29.v.1954,
(323) Dras 10-11.viii.1986, (324) Rawat 20-21.viii.1954 (for the colour key see Fig. 258).
176 PAULH. WILLIAMS
Bombus (Pr.) lemniscatus eS
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 37 Summary of the world-wide distribution of B. lemniscatus (for an explanation of the symbols see Map S).
0 100km
Land above 3000m.
Map 38 Distribution of B. lemniscatus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 325 &
326).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 177
WORKERS
Figs 325, 326 Colour patterns of B. lemniscatus from Kashmir: (325) Apharwat viii-ix.1986, (326) Apharwat
viii-ix.1986 (for the colour key see Fig. 258).
178 PAULH. WILLIAMS
Bombus (Pr.) lepidus oe
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 39 Summary of the world-wide distribution of B. lepidus (for an explanation of the symbols see Map 5).
|__|
0 100km
Land above 3000m.
Map 40 Distribution of B. lepidus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 327-334).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
a
<°,%,
SV
Qtr,
ey cao rate ats
QUEENS
WORKERS
MALES
179
TW TOW
wy WH
WV WV
Figs 327-334 Colour patterns of B. lepidus from localities in Kashmir: (327) Banidas 30. vi-2. vii.1959, (328) Bulandtar
3.vili.1954, (329) Dalti 7.ix.1954, (330) Lilam 17.viii.1953, (331) Surgun 29-30.vii.1953, (332) Nigagar 17.ix.1986,
(333) Apharwat 25.vii.1986, (334) Apharwat viii.1986 (for the colour key see Fig. 258).
180 PAULH. WILLIAMS
Bombus (Pr.) biroi
@ Specimen G@ Literature © Unconfirmed
100km
Land above 3000m.
Map 42 Distribution of B. biroi in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 335-346).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 181
cy
nw
ee SB
i
CWE?
|
WOM
UU Wy
Vv Vv
335 338 341 344
QUEENS
Th
4
@ 336 vw]
WORKERS
Sw
A___A
|
A
MALES
Figs 335-346 Colour patterns of B. biroi from localities in Kashmir: (335) Chhantir Gah 5—7.viii.1954, (336) Rumbak
12.viii.1980, (337) Tungri, (338) Gumri 10-11. viii.1986, (339) Gumri 10-11. viii.1986, (340) Gumri 30.ix.1986, (341)
Apharwat 25. vii. 1986, (342) Apharwat viii-ix.1986, (343) Apharwat 23.viii.1985, (344) Apharwat 25.vii.1986, (345)
Apharwat 15.viii.1980, (346) Apharwat 29. vii.1986 (for the colour key see Fig. 258).
182 PAULH. WILLIAMS
Bombus (Bo.) tunicatus
@ Specimen @ Literature © Unconfirmed
100km
Land above 3000m.
Map 44 Distribution of B. tunicatus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 347-355).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 183
QUEENS
WORKERS
349
MALES
Figs 347-355 Colour patterns of B. tunicatus from localities in Kashmir: (347) Luskum 30.v.1954, (348) Gilgit vi.1954,
(349) Shigar 1-3.x.1953, (350) Gulmarg vii—viii.1986, (351) Gulmarg viii.1986, (352) Gulmarg viii.1985, (353)
Gulmarg vii—viii.1986, (354) Gulmarg 10.viii.1980, (355) Gulmarg viii-ix.1986 (for the colour key see Fig. 258).
184 PAULH. WILLIAMS
Bombus (Bo.) lucorum
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 45 Summary of the world-wide distribution of B. Jucorum (for an explanation of the symbols see Map 5).
| es |
0 100km
Land above 3000m.
Map 46 Distribution of B. lucorum in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 356-370).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 185
QUEENS
‘Wle> <1 3e»
Wie» «le»
|
- 361 @ 364 ) Y 369
| eee Lee
fe fo)
es. ee
ee
ez, ay:
eer / ag
362 365 7370
MALES
Figs 356-370 Colour patterns of B. lucorum from localities in Kashmir: (356) Baltit 14-17.vi.1954, (357) Baltit
14-17.vi.1954, (358) Baltit 15.viii.1913, (359) Baltit 14-17.vi.1954, (360) Atro Sar 11.viii.1954, (361) Chittakatha Sar
19-24. vili.1954, (362) Lal Pani 2-3.ix.1953, (363) Maru 19.v.1910, (364) Lal Pani 2-3.ix.1953, (365) Lal Pani
2-3.1x.1953, (366) Apharwat 30.viii.1986, (367) Apharwat viii.1985, (368) Apharwat 17.vii.1986, (369) Apharwat
16. viii. 1980, (370) Apharwat viii.1985 (for the colour key see Fig. 258).
186 PAULH. WILLIAMS
Bombus (Sb.) asiaticus a
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 47 Summary of the world-wide distribution of B. asiaticus (for an explanation of the symbols see Map 5).
100km
Land above 3000m.
Map 48 Distribution of B. asiaticus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 371-391).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
—
87
| oe
qq
a
a
es
|
—"
w
—
P=
we
sa
si
os)
oe
So
QUEENS
Oe
Io»
| |e
| OO
«f
<
<
©
we
a |
nN
©
17%)
~
nn
@
w
~~
oo
®
w
So
—)
WORKERS
p-)
es
oo
hc
qi ice
hor
v*)
—I
io)
ies)
XQ
nN
Lo)
|
=)
ae
&
=)
—
MALES
Figs 371-391 Colour patterns of B. asiaticus from localities in Kashmir: (371) Leh 19-27.ix.1986, (372) Leh 8.vii.1980,
(373) Leh 19-27.ix.1986, (374) Thonde 1980, (375) Leh 3. vii. 1980, (376) Thonde 1980, (377) Nigagar 17.ix.1986, (378)
Nigagar 9.viii.1986, (379) Nigagar 17.ix.1986, (380) Lal Pani 2-3.ix.1953, (381) Lal Pani 2-3.ix.1953, (382) Lal Pani
2-3.1x.1953, (383) Gulmarg viii.1985, (384) Gulmarg viii.1985, (385) Gulmarg 29.viii.1986, (386) Gulmarg viii.1985,
(387) Gulmarg viii.1985, (388) Gulmarg 12.viii.1980, (389) Yusmarg 8.ix.1985, (390) Gulmarg viii.1985, (391)
Gulmarg 12.viii.1980 (for the colour key see Fig. 258).
188 PAULH. WILLIAMS
eke Lg = okt
Bombus (Sb.) oberti |
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 49 Summary of the world-wide distribution of B. oberti (for an explanation of the symbols see Map 5).
| ee |
0 100km
| Land above 3000m.
Map 50 Distribution of B. oberti in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 392 & 393).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 189
&
=
im
TOW
WM
wv
392
QUEENS
@
@
Figs 392, 393 Colour patterns of B. oberti from Kashmir: (392) Nimaling 15.vii.1980, (393) Nimaling 20.vii.1980 (for
the colour key see Fig. 258).
190
PAULH. WILLIAMS
Bombus (MI.) ladakhensis
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 51 Summary of the world-wide distribution of B. ladakhensis (for an explanation of the symbols see Map 5).
EE —,
0 100km
Land above 3000m.
Map 52 Distribution of B. ladakhensis in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs
394-396).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 191
QUEENS
MALE
Figs 394-396 Colour patterns of B. ladakhensis from localities in Kashmir: (394) Chushul vi.1925, (395) Hemis
25.ix.1986, (396) Nimaling 23.vii.1980 (for the colour key see Fig. 258).
192 PAULH. WILLIAMS
Bombus (MI.) semenovianus es
@ Specimen G@ Literature © Unconfirmed Equal area grid
Map 53 Summary of the world-wide distribution of B. semenovianus (for an explanation of the symbols see Map 5).
| ee |
0 100km
Land above 3000m.
Map 54 Distribution of B. semenovianus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs
397-402).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 193
(p>
QUEENS
qo» dio
©
wu
oO
oO
©
PS
—)
—
| |e
WORKERS
he.
ac
io)
N=)
v=)
PS
i=)
N
MALES
Figs 397-402 Colour patterns of B. semenovianus from localities in Kashmir: (397) Baltit 14-17.vi.1954, (398) Baltit
14-17.vi.1954, (399) Martselang 25.ix.1986, (400) Baltit 14-17.vi.1954, (401) Dras 10~-11.viii.1986, (402) Hemis
25.ix.1986 (for the colour key see Fig. 258).
194 PAULH. WILLIAMS
Bombus (ML. ) keriensis
@ Specimen @ Literature © Unconfirmed
Land above 3000m.
Map 56 Distribution of B. keriensis in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 403-416).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 195
SX
“es
oC
= =
403 405
QUEENS
\wmm
Nem
@® ~ 404
WORKERS
fe
|
416
MALES
Figs 403-416 Colour patterns of B. keriensis from localities in Kashmir: (403) Atro Sar 11.viii.1954, (404) Atro Sar
11.vili.1954, (405) Atro Sar 11.viii.1954, (406) Atro Sar 11.viii.1954, (407) Rawat 20—21.viii.1954, (408) Lal Pani
2-3.ix.1953, (409) Lal Pani 2-3.ix.1953, (410) Lal Pani 2-3.ix.1953, (411) Nimaling 15.vii.1980, (412) Nimaling
15.vii.1980, (413) Nimaling 24.vii.1980, (414) Apharwat viii.1985, (415) Apharwat viii.1985, (416) Apharwat
23.vili.1985 (for the colour key see Fig. 258).
196 PAULH. WILLIAMS
Bombus (MI.) simillimus NA
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 57 Summary of the world-wide distribution of B. simillimus (for an explanation of the symbols see Map 5).
| ———— |
0 100km
Land above 3000m.
Map 58 Distribution of B. simillimus in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 417-421).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 197
QUEEN
WORKERS
MALES
Figs 417-421 Colour patterns of B. simillimus from localities in Kashmir: (417) Harwan 10.ix.1985, (418) Harwan
10.ix.1985, (419) Harwan 10.ix.1985, (420) Harwan 10.ix.1985, (421) Banihal 3.ix.1986 (for the colour key see Fig.
258).
198 PAULH. WILLIAMS
Bombus (MI.) pyrosoma Ae
@ Specimen @ Literature © Unconfirmed Equal area grid
Map 59 Summary of the world-wide distribution of B. pyrosoma (for an explanation of the symbols see Map 5).
——————————,
0 100km
Land above 3000m.
Map 60 Distribution of B. pyrosoma in Kashmir (see Map 6; for a key to the colour pattern symbols see Figs 422-430).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 199
QUEENS
WORKERS
LEE LEAN
fF) CO
| |
ee eee) CLA
EL EL CN
Gizy Gay YU
94 "7 430
MALES
Figs 422-430 Colour patterns of B. pyrosoma from localities in Kashmir: (422) Kel 16.viii.1953, (423) Kel 16. viii.1953,
(424) Kel 16. viii.1953, (425) Gulmarg 2.ix.1985, (426) Gulmarg viii.1985, (427) Apharwat 14.ix.1986, (428) Gulmarg
vili.1985, (429) Gulmarg viii.1985, (430) Apharwat 14.ix.1986 (for the colour key see Fig. 258).
200 PAULH. WILLIAMS
Bombus (ML) rufofasciatus
@ Specimen @ Literature © Unconfirmed
Map 61 Summary of the world-wide distribution of B. rufofasciatus (for an explanation of the symbols see Map 5).
aT
0 100km
Land above 3000m.
=
NI:
Map 62 Distribution of B. rufofasciatus in Kashmir (see Map 6; for a a key to the colour pattern symbols see Figs
431-439).
THE BUMBLE BEES OF THE KASHMIR HIMALAYA 201
QUEENS
WORKERS
MALES
Figs 431-439 Colour patterns of B. rufofasciatus from localities in Kashmir: (431) Atro Sar 11.viii.1954, (432) Atro Sar
11.viti.1954, (433) Rawat 20-21.viii.1954, (434) Apharwat 23.viii.1985, (435) Apharwat viii-ix. 1986, (436) Apharwat
viii. 1985, (437) Apharwat 24.vii.1986, (438) Apharwat viii.1985, (439) Apharwat viii.1985 (for the colour key see
Fig. 258).
202
PAUL H. WILLIAMS
INDEX
This index includes references to the names of bumble bees used in the text and figures, but not to those in the
nomenclatural summary (pages 29-31), in the keys to species (pages 31-38), or in the lists of similar species at the end
of the comments on each species. Invalid or unavailable names are in italics; principal references affecting the
application of names are in bold; references to the figures that follow the text are in (parentheses).
abbotti 77
abnormis 70-71, 74
affinis 80
afghanus 44
agnatus (Skorikov, 1912a) 77
agnatus (Skorikov, 1933b) 77-78
agrorum 38
alaiensis 82, 85, 87
albocaudatus 87
albohirtus 66
albolateralis 52-54
albopleuralis 52, 55, 59
Allopsithyrus 45
Alpigenobombus 36, 62, 65-69
Alpigenibombus 65
alpigenus 65
Alpinobombus 39, 46
alpivagus 96-98
altaicus 42
amurensis 62, 65
Anodontobombus 69-70
Apathus 44
arcticus 39
ashtoni 45-46
Ashtonipsithyrus 45-46
asiaticus 13, 19-27, 29, 49, 87-92, 107,
110-118, 120 (129, 133, 135, 137-138,
142, 186-187)
assamensis 58, 60-61
assellus 42
atratus (Friese) 14
atratus (Franklin) 14, 119
audax 82
avinoviellus 15, 19, 21, 24, 39-41, 72, 110,
112-114, 120 (129-132, 134, 136,
138, 141-144)
barbutellus 45
bellicosus 118
beresovskii 66, 68-69
bianchii 94-95
biroi 6, 19, 24, 74, 77-79, 110, 112, 114,
120 (131, 133, 135, 137, 138, 139, 140,
180-181)
bohemicus 45—46 (129, 132, 134, 136,
138-139, 150-151)
Bombias 39, 46
Bombini 10, 31
Bombus 14-16, 31, 38, 62, 67, 79-87
braccatus 60
Brachycephalibombus 89
branickii 47, 48-49, 110 (132, 134, 136,
138-139, 154-155)
Bremus 38
breviceps 54, 60, 67, 119
brevivillus 119
brodmannicus 78, 118-119
bryorum 70-71
buccinatoris 58
caliginosus 77
callophenax 39-40, 90
campestris 41, 45, 49-50, 101
canariensis 83
canosocollaris 101, 103
centralis 77
Ceratopsithyrus 45
championi 105-106
chayaensis 67
chinensis (Morawitz) 42, 47, 105
chinensis (Dalla Torre) 105
chinensis (Skorikov) 42
cinnameus 59-60
Citrinopsithyrus 45
citrinus 45
Coccineobombus 89
Confusibombus 39, 46
confusus 39
convexus 15, 42
coreanus 46
cornutus 45
Crotchiibombus 89
cryptarum 82, 84-85
Cullumanobombus 46, 88
Dasybombus 89
defector 15, 40-42
dentatus 67
difficillimus 62-65
Diversibombus 52
Diversobombus 52-60, 113
diversus 52, 54
duanjiaoris 92-93
dyspostzonatus 77
elisabethae 48, 77
Eopsithyrus 45
ephippiatus 100, 115
eriophoroides 48
eurythorax 103
Eversmannibombus 42, 46
exil 14-15, 46
exiln 46
Exilobombus 15, 46
eximius 100
falsificus 88, 91
fedtschenkoi 62, 64-65
ferganicus 21, 49-50 (129, 132, 134, 136,
138-139, 156-157)
fernaldae 45, 51
Fernaldaepsithyrus 45, 50-51
Fervidobombus 119-120
Festivobombus 70, 80, 88, 100
festivus 70-71, 88, 100
flavescens 55, 74
flavidus 51
flavifrons 77-78
flaviventris 62, 89, 92-93
flavobistriatus 77-78
flavocorbicularis 102-103
flavodorsalis (Franklin) 88
flavodorsalis (Skorikov) 88, 90
flavofasciatus 77
flavopilosus 73-74
flavostriatus 77
flavothoracicus (Hoffer) 101
flavothoracicus (Bingham) 100, 101-103,
106
fletcheri 70-71
formosellus 102, 104
fragrans 62, 65
franklini 80
Fraternobombus 89
fraternus 89
frigidus 78
friseanus 101-103, 106, 115
fulvocinctus 80
Funebribombus 89
funerarius 60
fuscocaudatus 87
gansuensis 50-51
gantokiensis 52-54
geminatus 52-54, 58
genalis 67
genitalis 75-76
gilgitensis 79-80
grahami 67
griseofasciatus 62-63
grossiventris 99-101
haematurus 71-72
haemorrhoidalis 6, 52, 54, 58-61, 67, 74,
109-114, 119-120 (129, 131-132, 134,
136, 138, 140, 164-165)
haemorrhous 99, 101
handlirschi 89
handlirschianus 15, 40-42
heicens 91
hilaris 75-76
himalayanus 15, 19, 24, 41-44, 110-114,
120 (132, 134, 136, 138, 141, 146-147)
hoenei (Bischoff, 1936:10) 102-103
hoenei (Bischoff, 1936:26) 47
Hortobombus 52
hortorum 13, 87, 113
huangcens 91
hummeli 54
hyperboreus 39
hypcorum 70
Hypnorobombus 70
Hypnorubombus 69
hypnorum 19, 24, 69, 70-72, 110, 112,
114, 119 (132, 134, 136, 138, 172-3)
hypocrita 83
ignitus 83
THE BUMBLE BEES OF THE KASHMIR HIMALAYA
incertoides 96-98
incertus 97
indicus 49
infirmus 74-75
infrequens 75
insidiosus 52, 54
intermedius 105
Jacobsoni 82, 85-86
jonellus 78
Kallobombus 39, 46
karakorumensis 96, 98
kashmirensis 19, 23-28, 49, 65, 66-69, 75,
108, 110-112, 114, 120 (130, 132, 134,
136, 138, 170-171)
keriensis 19, 24, 49, 94, 96-99, 109-110,
112-114 (133, 135, 137-138,
194-195)
khasianus 58, 60-61
klapperichi 45
kohistanensis 72
kohli (Cockerell) 96
kohli (Vogt) 96-98
kotzschi 77-78
kozlovi 94, 96
Kozlovibombus 93-94, 100
kozlowi 96
Kozlowibombus 94
kuani 51
Laboriopsithyrus 45
laboriosus 45
ladakhensis 7, 94-95, 110, 112, 114 (133,
135, 137-138, 190-191)
Lapidariibombus 94
Lapidariobombus 93-94
lapidarius 13, 89, 93-97, 99-100, 113,
118-119
Laponicobombus 70
Lapponicobombus 69
lapponicus 69, 77
lefebvrei 65-66
lemniscatus 72, 73-75, 112, 114 (129, 133,
135, 137-141, 176-177)
lepidus 19, 24, 74, 75-76, 110, 112, 114,
120 (133, 135, 137-141, 178-179)
Leucobombus 79
leucopygos 72
leucopygus (Illiger) 72
leucopygus (Morawitz) 72
leucurus 72
longiceps 87-90
longipennis 82-83, 85
longipes 54
lucocryptarum 82, 85
lucorum 13, 19, 24, 46, 75, 81-87,
109-110, 112, 114, 119 (133, 135,
137-138, 184-185)
lugubris (Kriechbaumer) 42
lugubris (Morawitz) 42
luteipes 75—76
maderensis 84
magnus 82, 84-86
magrettianus 52-53, 55
maidli 62-63
makarjini 15, 39-41, 43
malaisei 54
margreiteri 42
marussinus 15, 40-43, 44, 112-114 (132,
134, 136, 138, 141, 148-149)
Mastrucatobombus 65
mastrucatus 65-68
maxillosus 42
maxwelli 52-53, 55
Megabombus 53, 113
meinertzhageni 66-67
Melanobombus 62, 67, 88-89, 92, 93-107,
113
melanopoda 119
melanopygus 115
melanurus 19, 21, 24, 61-65, 110-114
(132, 134, 136, 138, 166-167)
Mendacibombus 14-16, 38-46
mendax 15, 38-44
meridialis 96-98
mesoxanthus 50
Metapsithyrus 45, 49-50
metcalfi 60
mimeticus 52, 54—55
miniatocaudatus (Vogt, 1909) 87
miniatocaudatus (Vogt, 1911) 87-88, 91
miniatus 101-102, 104
minshanicus 52-53, 55
mirus 73-74
mlokosievitzii 119
moderatus 82-83
modestus (Eversmann) 82
modestus (Cresson) 82
mongolicus 82-83, 85-86
montivagus 52, 54-55
montivolans 58-60
morawitzi (Radoszkowski) 89-90,
92-93
morawitzi (Friese) 47
morawitzianus 50 (132, 134, 136, 138-139,
158-159)
morawitziides 65-66, 67
morio 96, 119
Mucidobombus 46
mucidus 14
nemorum 45
nepalensis 47
ningpoensis 52, 55
nevadensis 14, 39
niger 14, 119
nigrozonatus 77
niveatus 39, 89, 90
Nobilibombus 65-66
nobilis 65—66, 67
novus 46-48 (130, 132, 134, 136, 138-139,
152-153)
nursei 73, 75, 77-78
oberti 7, 87, 89-90, 92-93, 110, 112, 114,
120 (129, 133, 135, 137-138, 142,
188-189)
Obertibombus 87
Obertobombus 87, 93
obtusus 89-91
occidentalis 83
ochrobasis 92
oculatus 99-100
orichalceus 67
Orientalibombus 46, 58-61, 113
Orientalobombus 58
orientalis 58-61
oshanini 88, 90
203
pamirus 65
parthenius 71, 74, 75-76
pascuorum 38, 113
patagiatus 83, 86
pectoralis 59
pennsylvanicus 88
pensylvanicus 88
peralpinus 73-74
perplexus 71
persicus 42
personatus 7, 62, 64-65, 110, 112, 114
(132, 134, 136, 138, 168-169)
Phariensis 94-95
pieli 49-50
Poecilobombus 69
pomorum 65
postzonatus 96-98
Pratibombus 70
Pratobombus 69
pratorum 69, 71, 78, 82, 119
Pressibombus 80
pressus 80
problematicus 95-96
prshewalskyi 105
Psithyrus 14-16, 39, 44-51
pulcherrimus 65, 66
pullatus 119
pyrenaeus 78
Pyrobombus 46, 62, 67, 69-79, 119
pyrosoma 19, 22, 24-27, 94, 100, 101-107,
110, 112, 114-115, 119 (128, 133, 135,
137-138, 198-199)
Pyrrhobombus 69
pyrrhosoma 101-102
quasibreviceps 52, 54
redikorzevi 50
regeli 87-88
reinigi 82, 85-86
reticulatus 94-95
Rhodobombus 39, 65
richardsiellus 101-102, 104-106
roborowskyi 64-65
Robustobombus 89
rotundiceps 54, 60, 74, 119
Rubicundobombus 89
Rufipedibombus 80, 88, 100
rufior 105
tufipes 100, 119
rufitarsus 42
rufocinctus (Cresson) 105, 115
rufocinctus (Morawitz) 105
rufofasciatus 6, 13, 19-27, 47, 94-95, 100,
102, 104, 105-112, 114-115, 120 (128,
130, 133, 135, 137-138, 200-201)
rufoflavus 55
rupestris 42, 44, 47, 48, 105
sandersoni 77
secundus 52
semenovi 92-93
semenovianus 7, 78, 95-96, 110,
112-114, 120 (133, 135, 137-138, 142,
192-193)
semibreviceps 59, 61
semicoloricontrarius 59, 61
senex 119
Senexibombus 53
separandus 96-98
204
Separatobombus 89
shaposhnikovi 15, 42
Sibiricibombus 87
Sibiricobombus
113
sibiricus 87, 89
sicheli 97
sichelii 97-98
signifer 75
similis 99
simillimis 99
simillimus 99-102, 105-106, 109-110,
112-114, 119 (128, 133, 135, 137-138,
142, 196-197)
simlaensis 79
sitkensis 69
skorikovi 50-51 (132, 134, 136, 138-139,
160-161)
sonani 75
sonorus 88
soroeensis 28, 39, 87
sporadicus 80, 86
stenothorax 103
stramineus 66
subdistinctus 62-63
Subterraneibombus 61
Subterraneobombus 61-65, 80
subterraneus 61-62, 64
subtunicatus 39
16, 62, 67, 87-94,
subtypicus 72-73, 110-112, 114 (131-138,
140, 174-175)
suckleyi 46
sulfureus 90
superbus 42
susteraianus 49
tajushanensis 49
Tanguticobombus 94, 100
tanguticus 89, 93-94, 100
tenellus 96, 98
tenuifasciatus 88
Terrestribombus 79
terrestris 38, 79-80, 81-86, 99
terricola 83
tetrachromus (Cockerell) 66, 75
tetrachromus (Friese) 75-76
Thoracobombus 60, 113
tibetanus (Morawitz) 45, 74
tibetanus (Friese) 74
tonsus 99, 101
trifasciatus 6, 19, 24, 52-60, 67, 74, 107,
109-114, 119 (129, 131-132, 134, 136,
138, 162-163)
trilineatus 101
tschitscherini 62-63
tunicatus 19-22, 24, 79-81, 107, 109-110,
112-115, 119-120 (133, 135, 137-138,
182-183)
PAUL H. WILLIAMS
turkestanicus 15, 41-44, 113
turneri (Richards, 1929a) 52
turneri (Richards, 1931) 52-54
Uncobombus 69-70
ussurensis 54
vagans 77
validus 66, 67
vandykei 77
variopictus 94-95
varius (Lepeletier) 41
varius (Skorikov) 15, 41-43
vestalis 46
vorticosus 90
waltoni 15, 40-41, 42
waterstoni 105
wilemani 52, 55
wurfleini 67
wurflenii 28, 65, 67-68, 119
wutaishanensis 102-103
xionglaris 91
xizangensis 67
yuennanicola 75-76
yunnanicola 75
zhadaensis 93
\
1 The bumble siete of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini).
Paul H. Williams
ENTOMOLOGY SERIES
Vol. 60, a. is April 1991
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