The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini)

204 Pages·1991·61.7 MB·

by  P H Williams

Bull. Br. Mas.nat. //wf(Ent.)60(l):l-204 Issued25April1991
The bumble bees of the Kashmir Himalaya
(Hymenoptera: Apidae, Bombini)
PAUL H. WILLIAMS
do DepartmentofEntomology, British Museum (NaturalHistory), CromwellRoad, South Kensington,
London SW75BD
CONTENTS
Introduction 2
Materialexaminedanddepositories 8
DiagnosisofHimalayanBombini 10
Charactersstudied 11
Relationshipsamongspeciesbyancestry 13
GeneraofBombini 14
Discriminationofspeciesandintraspecificvariation 16
The'biological'speciesandtheRecognitionConcept 17
Mate-searchingbehaviourofmalebumblebeesinKashmir 18
Relationshipbetweenmalemate-searchingbehaviourandhabitatstructure 25
Inferenceofallopatric,conspecifictaxa 27
Inferenceofallopatric,separatespecies 27
Charactersofspeciesnotrelatedtomalemate-searchingbehaviour 28
Thedescriptionofintraspecificvariation 29
Nomenclaturalsummary 29
TaxonomyoftheKashmirBombini 31
Keytospecies(females:queensandworkers) 31
Keytospecies(males) 35
GenusBombusLatreille 38
Distributionofthefauna 107
Distributionwithaltitude 107
DistributionacrosstheHimalaya Ill
DistributionoutsideKashmir 112
Relationshipsbetweenpatternsatdifferentspatialscales 113
ColourpatternsoftheKashmirfauna 115
VariationofB.asiaticusinKashmir 115
Convergentcolourpatterns 118
Acknowledgements 120
References 121
Index 202
Synopsis. Bumble bees are extraordinarily variable in the colour patterns of their
pubescence.Thishascontributedtothelackofagreementamongauthorsconcerningthe
discrimination of species. Material from Kashmir has been listed previously under 80
names, butforthebroadestinterpretationsofthese species, morethan500nameshave
been published for differing individuals from throughout their distributions. In this
review,alloftheavailablevouchermaterialfromKashmir(6312museumspecimensfrom
78sites)isassignedtojust29species.Thereductioninthenumberofnominaltaxathatare
inferredtorepresentseparatespeciesdependsontwofactors.First,variationisexamined
amongthelargersamplesofnewmaterial.Second,currentspeciesconceptsrequireare-
assessmentofthisevidenceforthelimitsofreproductivecohesioninspaceandtime. A
particular study is made of the variation in male mate-searching behaviour and of its
relationship to the different kinds ofhabitat. For Bombusasiaticus, patterns ofcolour
variation across the Great Himalaya are compared with those expected from simple
PAULH.WILLIAMS
geneticmodels.Typematerialisexaminedfor103ofthenominalspecificandsubspecific
taxa. Seventeen lectotypes are designated and 80 new synonyms and provisional new
synonymsareestablished. Keystothespeciesareaccompaniedbydiagramsofthecolour
patternsandbydistributionmaps.
and in China, except for another corridor of
INTRODUCTION contactnearBeijing,whichisoccupiedbyrelatively
few species (Panfilov, 1957). Greater Kashmir
encompasses almost the entire mountain system
Bumble beesrank amongthe mostabundant and that links the high Tibetan (Xizang-Oinghai)
conspicuousofflowervisitorsinalpine,temperate plateau in the east with the Hindu Kush, Pamir
andarcticenvironmentsofthenortherncontinents. and Tien Shan mountain ranges to the west and
Inthesouthernhemispheretheyarenativeonlyin north. Hence Kashmir covers segments of the
the East Indies and South America (see Map 1), Pir Panjal, Great Himalaya, Zanskar, Ladakh,
where most species are associated with the high- Karakoram and Hindu Raj ranges (Map 2), and
lands. Adults feed mainly on nectar, which includes some of the highest peaks in the world
providesenergyforflight. Thelarvaearefedon a (altitudes range between about 400-8600 m; for
mixtureofpollenandhoney,whichprovidestheir anaccountofthegeologicalstructureseeSearleet
requirements for growth. Most species are social ai, 1988). Kashmir is slightly larger in area than
(reviewsofbehaviourbye.g. Alford, 1975;Morse, the European Alps and lies at the same latitude
1982; Plowright & Laverty, 1984). The colonies as Syria, Tunisia and Arizona. Since Indian
consist almost always of a single, mated queen, independencein1947,thesovereigntyofKashmir
which lays most of the eggs, and usually of no hasbeen amatterofdispute andsectionsarenow
morethanafewhundred, unmatedworkers. The administeredbyIndia(Jammu&KashmirState),
establishment and developmentofcoloniestakes Pakistan (Northern Areas) and China (aspart of
place each summer and may take less than two the Xinjiang Autonomous Region).
months (Richards, 1973), culminating with a The high relief of Kashmir provides a broad
switch to the production of males and young rangeofhabitatsforbumblebees. Patternsinthe
queens nearthe endofthe season. After mating, distributionsofbumblebeesinEuropehavebeen
young queens normally diapause through the linkedwithclimaticfactors(e.g.Pekkarinenetal.
,
winter,awayfromthenest,beforetheyattemptto 1981). The differing climates of Kashmir can be
foundcoloniesontheirowninthefollowingyear. summarised in three regions (e.g. Gurcharan
Only a few species from the tropics of South Singh & Kachroo, 1976). First is the subtropical
America (Sakagami, 1976) and South East Asia regionoftheJammufoothills, whichis subjectto
(Michener& Amir, 1977) mayhave coloniesthat the summer monsoon (Fig. 2). Second, beyond
persistformorethanoneyear. Bumblebeesshow the Pir Panjal range, is the more temperate Vale
unusuallywell-developedendothermy,whichcan ofKashmir, where most ofthe rain and snowfall
befacultativewhenforaging(reviewedbyHeinrich, in the winter months (Fig. 1). Third, in the rain
1979). Almost all species are generalists in their shadow ofthe Great Himalaya, is the arid alpine
choice of food-plants, visiting any remunerative region of the Zanskar, Ladakh and Karakoram
flowers.Theirforagingfollowsa'scramble'pattern, ranges (Figs 3 & 4). But of equal importance to
without either recruitment ofnest-mates to good climate, as an influence on whether or not a
food sources or defence of food sources. These speciesofbumblebeecouldpersistataparticular
characteristics of bumble bees may account for locality, maybethe natureofthelocalvegetation
theirabundance incool environmentsthathavea (e.g. Bowers, 1985; WiUiams, 1988, 1989), even
predictable season of adverse conditions, where though bumble bees are seldom dependent on
flowers are often fairly evenly dispersed. In particular species of food-plants. Map 3 shows a
contrast, the other social, long-proboscis bees summary ofthe vegetation in Kashmir, based on
(honeybeesandstinglessbees)havetheircentres the survey of the Himalaya by Schweinfurth
ofdiversityandabundanceinthetropicallowlands. (1957). Not only do the three principal climatic
The bumble bees of Kashmir are of particular regionswithin Kashmirdifferintheirflora, butin
interest because this narrow corridorofmountains combination with the influences oflocal altitudinal
is almost the only major, modern point ofcontact zonation and variation in local exposure (Troll,
between the large and divergent Oriental and 1972), this contributes towards a particularly
Palaearctic bumble bee faunas. These faunas are broadrangeofhabitats. Incomparisonwithsome
otherwise separated by deserts in central Asia partsofthe Himalaya, Kashmirretains relatively
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Fig.1SubalpinezoneofMtAphanvatinthePirPanjalrange,viewedfrom3000mtowardsthepeakat4143m(Map4
locality1:foregroundpasturewithEuphorbiawallichiana;lowerslopesdominatedbyBetulautilis,Salixsp.,Syringa
emodi,withafewAbiesspectabilis;middleslopesdominatedbyB.utilis.Rhododendroncampanulatum;higherslopes
dominatedbyR. anthopogon,Juniperusrecurva). Of17speciesofbumblebeesrecordedinthisimmediatearea,the
mostrestrictedtothiszoneisB. biroi, andthemost abundantisB. rufofasciatus.
Fig. 2 Lower montane coniferous forest zone on the Patnitop ridge in the Jammu foothills, viewed from 2000 m
towardstheplainsofIndia(Map4locality11: Patnitopdominants, Cedrusdeodara[foreground],Pinuswallichiana,
P.roxburghii).Of3speciesofbumblebeesrecordedinthisarea,themostrestrictedtothiszoneisB.haemorrhoidalis
andthemostabundantisB. trifasciatus.
.
THEBUMBLEBEESOFTHEKASHMIRHIMALAYA
Fig.3High,aridalpinesteppezoneattheterminalmoraineofNimahngplainintheZanskarranges,viewedfrom4800
macrossthesouthernlateralmorainetowardsapeak(Kang-y-sey)at6400m(Map4locality51:foreground,shrubsof
Caraganaversicolor).Of9speciesofbumblebeesrecordedinthisimmediatearea,themostrestrictedtothiszoneare
B.personatus, B. obertiand B. ladakhensis,ofwhichthe mostabundantisB. oberti.
Fig.4SubalpinesemidesertzoneatLamayuruneartheIndusvalley,withaviewofvalleyterracesat3400m,againsta
backgroundofmountainridgesat4000-5000m(Map4locality42).Of3speciesofbumblebeesrecordedinthisarea,
themostrestrictedtothiszone,andthe mostabundant,isBombussemenovianus
1
8 PAULH.WILLIAMS
largeforests andyet access to some ofthe varied Table1 Localitiesthathavebeensampledforbumble
alpine areas is no longerdifficult. bees.Sitesincloseproximityaretreatedascombined
PreviousstudiesofthebumblebeesofKashmir andrepresentedbyasinglenumberonMap4.
havebeenbasedonatotalsampleofonlyacouple
ofhundredspecimens. Thefirst importantcollec- Localitiesfromwhichsamplesofbumblebeeshavebeen
tion of bumble bees from Kashmir was made by examined(numbersincircleson Map4).
Lt.-Col. C. G. Nurse in 1901. A complete inven- 1 (Mt) Apharwat 42Lamayuru
toryofthismaterialwasneverpublished, although 2Astor 43Leh
it provided the specimens that have since been 3AtroSar 44Lilam
described as the types of many nominal taxa 4Babusar 45Luskum
(Friese,1909,1918;Richards, 1928fl, 1928/),1930; 5 Baltal 46MahthantirGah
Tkalcu, 19746). Other collections were mostly 6Baltit Martselang = Chogdo
7Banidas 47Maru
small, but particularly important material was 8Banihal 48Matayan
obtainedby A. Jacobson inKashmirandLadakh 9BaraDeosai 49Muzaffarabad
during 1912 (Skorikov, 1914a) and by Col. R. 10Batakush Nagar = Holshai
MeinertzhageninLadakhduring 1925 (Richards, 1 Batote 50Naltar
19286). Skorikov(19336)compiledapreliminary 12Bulandi Nigagar = Baltal
Hst of the fauna of Kashmir and discussed the 13BurzilChauki 51 Nimaling
14Chakorkhand 52Nomal
faunaoftheentireHimalaya,thoughhisworkwas 15ChhantirGah Nowgam = Banihal
based on a different concept of the species from 16ChhotaDeosai 53Pahalgam
thatacceptedatpresent. Theonlyrecentrevision 17ChittakathaSah 54Parkutta
of any part of the large Himalayan fauna is ISChogdo Patnitop = Batote
Tkalcu's (19746) description ofa collection of73 19Chushul 55Rawat
bumble bees from Nepal. Although there are 20Dachhigam 56Rumbak
21 Daksum 57Saif-ul-MalukSar
manyelementsincommon betweenthefaunasof 22Dalti 58Sangisfaid
Nepal and Kashmir, 16 species that are known 23Darkot 59Shamran
fromKashmirarenotrepresentedinthiscollection. 24Dras 60Shardi
InthesamepaperTkalcureferstohis'Monographic Gamelti = Darkot 61 Shigar
der Unterfamilie Bombinae des Himalaya'' as in 25 Gilgit 62ShinghaiGah
26Gulmarg 63 Skardu
press. Unfortunatelythishasneverbeenpublished,
27Gulmiti 64Sonamarg
although recently he has described several new 28Gumri 65Srinagar
taxafromtheregion (Tkalcu, 1989).Theonlykey Harwan = Dachhigam 66Stakmo
that has been intended to cover any part of the 29Hemis 67Sumbal
Himalayan fauna is that published by Bingham 30Holojut 68Surgun
s(p1e8c97i)e.s nThoiwsrkeecyogarntiisfeicdiaflrlyomsuKbadsivhimdiers,fwohuerroefatshea 31 HHoulnshzaai= Baltit 69STuarnugmarg = Gulmarg
32Imit 70Taobat
further20species are not included at all. 33Janwai 71Teru
Forthisreview,muchoftheoldermaterialisre- 34Kangan Thajiwas = Sonamarg
examinedandcomparedwiththelargercollections 35 Kanur 72Thonde
ofnew material, in orderto describe some ofthe 36Karsa 73Tiggur
patterns of variation (and the discontinuities 37Kel 74TsoMorari
in these patterns) among the bumble bees of 38Khalsi Tungri = Karsa
39KhardungLa 75 Verinag
Kashmir. This evidence, together with informa- Khilanmarg = Apharwat76Wangat
tionfromthehabitatassociationsofthespeciesin 40Kishtwar 77Yasin
Kashmir and information from their broader 41 LaiPani 78Yusmarg
distributions beyond Kashmir, is used to discuss Zoji La — Gumri
the likely relationships of ancestry and inter-
breeding among the nominal taxa.
Localitiesatwhichnobumble beescouldbefound
duringbriefsurveysinAugust-September1980,1985or
1986(lettersintrianglesonMap4).
MATERIAL EXAMINED AND
DEPOSITORIES a Anantnag e Mulbekh
b Baramula f Ramban
c Chrar-i-Sharif g Udhampur
Large samples ofundescribed material are avail- d Jammu
able from western Kashmir in the collections
THEBUMBLEBEESOFTHEKASHMIRHIMALAYA
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madeduring1953and1954byF. Schmid(BMNH NMS Natur-Museum Senckenberg, Frankfurt,
accession 1962-457; account ofthe expedition in D.B.R.
Schmid, 1958). In addition, I was able to make NR Dr N. D. Rishi, University of Kashmir,
large, selective collections from southern and Srinagar, India.
eastern Kashmir during 1980, 1985 and 1986. PW Author'scollection, London, U.K.
Unfortunately, large areas near the borders, RH DrR. W.Husband,AdrianCollege,Adrian,
especiallytothenorthandeastofthecountry,are U.S.A.
prohibited to foreign visitors, so only a few SEMK Snow Entomological Museum, Lawrence,
specimensfrom oldercollections are available. Kansas, U.S.A.
The sites in Kashmir from which material has TL Dr T. M. Laverty, University of Western
Ontario, London, Canada.
beenexaminedarelistedinTable 1 andshownon TM Termeszettudomanyi Muzeum, Budapest,
Map 4. Geographical information is taken from Hungary.
the United States Survey ofIndiamaps (scale 1 : UM UniversityMuseum, Oxford, U.K.
2(15908,70)0.0)Thaendwofrrlodm-wTihdeeTdiismtersibauttliaosnoofftshpeecwioerslids USNM DU.nCi.t,edUS.tSa.tAes.NationalMuseum,Washington
also shown by maps (see Map 1). These are ZI ZoologicalInstitute,Leningrad, U.S.S.R.
compiledwithapreferenceforrecordsfromthose ZM ZoologiskMuseum, Copenhagen,Denmark.
specimens that I have been able to examine, ZMMU ZoologicalMuseumofMoscowStateUniver-
supplementedbyalistofselectedreferences.The sity, Moscow, U.S.S.R.
source material is summarised briefly in the text ZS ZoologischeStaatssammlung,Munich,D.B.R.
either by political region (for the Himalayan/
Tibetan mountain system in India and China) or
bymajormountainsystem (forthemorerestricted
bumble bee faunas of central Asia), such as the DIAGNOSIS OF HIMALAYAN BOMBINI
Elburz, HinduKush, Pamir, Alai-Tien Shan (in
its broadest sense) and Altai mountains. More
precisedatacanbefoundintheoriginalreferences. AmongtheApidae,theBombinihavethelabrum
The specimens that have been examined are at least twice as broad as long. The labrum lacks
depositedincollectionsasshownbythereferences a longitudinal median ridge, although for the
to the followingabbreviations. females it has a strong transverse basal depres-
sion. The clypeus has a transverse subapical
depressionandtheapico-lateralcornersarecurved
AB DrA.A.Bhat,S.K.UniversityofAgricultural backtowards theocciput. A malararea (= malar
Sciences&Technology, Srinagar, India. space)separatesthecompoundeyefromthebase
BMNH British Museum (Natural History), London, ofthe mandible, often by a distance greaterthan
U.K. the breadth ofthe mandible at its base. The hind
BPBM BishopMuseum, Honolulu, U.S.A. wingslackajugallobe(= anallobe).Thevolsella
BT DrB. Tkalcu, Prague, Czechoslovakia. (=lacinia)ofthemalegenitaliaisgreatlyenlarged
FA FacultedesSciencesAgronomiquesdeI'Etat, andisproducedapicallybeyondthegonostylus(
Gembloux, Belgium. squama).
INHS Illinois Natural History Survey, Champaign, Bumble bees are large (body length 7-27 mm)
U.S.A. robustinsects.Theirbodieshaveadensecovering
rrZ Instituut voor Taxonomische Zoologie, ofvariously-colouredlongplumose hairs, although
Amsterdam, Netherlands. thesearefeworabsentonsomepartsoftheventral
LIZK DInrstiLt.utSe.ofKiZmosoelyo,gy,UnBieviejrisnigt,yCohfinCaa.lifornia, sounrfpaacretsofofthtehegaasntteerr,ioornfpaacretsooffgtahsetrpalrotpeordgeuummI,,
Davis, U.S.A.
LSL Linnean Society, London, U.K. andonpartsofthehead. Thescleritesareusually
MCSN Museo Civico di Storia Naturale, Genoa, black, or lighter brown on the distal parts ofthe
limbs, but are never marked with bright yellow,
Italy.
MI Dr M. Ito, Hokkaido University, Sapporo, red, or metallic (= interference) colours. The
Japan. wings maybe transparent (= hyaline) to strongly
MNHN MuseumNationald'HistoireNaturelle,Paris darkened(=infuscated),butrarelyshowstrongly
France. metallic reflections.
MNHU Museum fiir Naturkunde an der Humbolt- Femalebumblebeeshave12antennal'segments'
Universitat, Berlin, D.D.R. (= scape, pedicel and 10 flagellomeres) and six
NM NaturhistorischesMuseum,Vienna, Austria. visiblegastraltergaandsterna(abbreviatedtoTI-
NMP NarodniMuzeum,Prague, Czechoslovakia. VI,SI-VI).Maleshave13antennal'segments'(=