Bumblebees (Hymenoptera, Apidae, Bombus Latr.) of the thermal spring Pymvashor, north-east of European Russia

2019·0.3 MB·

by  Yulia Kolosova

©EntomologicaFennica.16December2016
Bumblebees (Hymenoptera, Apidae, Bombus Latr.)
of the thermal spring Pymvashor, north-east of European Russia
YuliaSergeevnaKolosova,GrigorySergeevichPotapov,NataliaGaryevnaSkyutte
&IvanNikovaevichBolotov
Kolosova,Yu.S.,Potapov,G.S.,Skyutte,N.G.&Bolotov,I.N.2016:Bumble-
bees (Hymenoptera, Apidae, Bombus Latr.) of the thermal spring Pymvashor,
north-eastofEuropeanRussia.—Entomol.Fennica27:190–196.
AbumblebeecommunitywasstudiedatPymvashor,theonlythermalspringin
NorthEuropeanRussia.Thebumblebeefaunacomprised12species,whichisa
largenumberwhencomparedtoothernativetundraecosystems.Mostofthespe-
ciesrecordedwereubiquitous,3wereforestspeciesand2weretypicaltundra
species.Thepresenceoftheubiquitousandforestspeciesinthebumblebeecom-
munityappearstobeduetothelandscapefeaturesandtheperennialimpactofthe
hotsprings,undertheinfluenceofwhichspecificextrazonalecosystemsarise
thataredifferentfromthosetypicalofthetundrazone.
Yu.S.Kolosova,G.S.Potapov,N.G.Skyutte&I.N.Bolotov,FederalCenterfor
Integrated Arctic Research, Russian Academy of Sciences, Naberezhnaya
Severnoy Dviny 23, 163000 Arkhangelsk, Russia; Corresponding authors’e-
mail:[email protected],[email protected]
Received23March2016,accepted5October2016
1.Introduction and fauna (Bolotov & Bogolitsyn 2011). Many
groups have not previously been studied in this
The study of invertebrates, including insects in area. As the object of our research, we chose
hydrothermal ecosystems, has always been of bumblebees, a taxonomic group with origins in
greatinterest.Thereisalargebodyofliterature the mountains of Central Asia (Hines 2008).
dedicated to research of different taxonomic Hence,bumblebeesarewelladaptedtolivingin
groups of insects at thermal springs in different theunfavourableclimaticconditionsofthenorth.
regions.Thisresearchhasconsideredtheimpact Thepresentarticledescribesthebumblebeecom-
ofhotspringsonthespeciesrichnessofinsects, munityatPymvashor.
adaptation to the specific conditions of thermal
waters,ecologyofdifferentspecies,etc.(Stock-
ner1971,Resh&Barnby1984,Glazier&Gooch 2.Materialsandmethods
1987,Pritchard1991,Pritchard&Kortello1997,
Lobkova&Lobkov2003,Garbuzetal.2008). ThethermalspringPymvashor(67°09’N,60°51’
The stimulus for our research at the thermal E)islocatedintheNenetsAutonomousRegion,
springofPymvashorwasthatthehydrothermal ontheeasternoutskirtsoftheBolshezemelskaya
systemislocatedwithinthesoutherntundra,and tundra,nearboththeAdzvaRiverandtheright
characterised by a unique composition of flora tributaryoftheUsaRiver.Itisinthesubzoneof
ENTOMOL.FENNICAVol.27 (cid:127) Bombusspp.ofathermalspringarea 191
Fig.1.Map
ofthePym-
vashorther-
malspring
areain
NorthEuro-
peanRus-
sia.
the dwarf-shrub tundra. Pymvashor is the only ronmentalProblemsoftheNorth(RussianAcad-
territoryofthemainlandnorthernEuropewhere emy of Sciences, Ural Division) in years 2009,
thermalspringsexist,andislocatedtothenorth 2010,and2012–14.Bumblebeeswerecollected
oftheArcticCircle(Bolotov&Bogolitsyn2011). usinganentomologicalnet,withintheareaofthe
Descriptions of the natural conditions, physical thermalspringinthevalleyandonthebanksof
and geographic characteristics of the study area thePymvashorstream.Atotalof233bumblebees
were published earlier (Bolotov & Bogolitsyn werecollected.
2011,Bolotovetal.2012).Amapofthethermal Bumblebee identification followed Løken
springareaofPymvashorispresentedinFig.1. (1973,1984)andPanfilov(1978).Thesubgener-
Thezonalvegetationofthestudyareaisrep- icclassificationandthesynonymyofspeciesare
resented mainly by different variants of dwarf givenaccordingtoWilliams(2016).Ourdetermi-
birchandwillowtundra.However,underthein- nationswereverifiedusingthecollectionmateri-
fluenceofthewarmingimpactofthermalwaterit alsfromtheNaturalHistoryMuseum(London)
has formed an extrazonal ecosystem (Vinogra- andtheZoologicalInstituteoftheRussianAcad-
dova1962)whichischaracterisedbymoresouth- emyofSciences(SaintPetersburg).
ern types of vegetation, represented by sparse IdentificationofspeciesoftheBombusluco-
spruce-woods(Piceaabiesssp.obovata(Ledeb.) rum-complex followed the keys of Rasmont
Domin). These grow as a narrow band through (1984),RasmontandTerzo(2010),andwithout
thethermalvalley(Bolotov&Bogolitsyn2011, using DNA-barcoding. However, we have to
Bolotovetal.2012).Therearealargenumberof considerourspecimensasB.cf.cryptarum,be-
plant species of the taiga flora: Cirsium hetero- causemorphologicalidentificationofthesespe-
phyllum (L.) Hill, Filipendula ulmaria (L.) ciesisunreliable(Bossert2015).Itismostlikely
Maxim., Hieracium umbellatum L., Chamerion that B. cryptarum is represented at Pymvashor.
angustifolium(L.)Holub,VeronicalongifoliaL. RasmontandIserbyt(2010)notethat“according
andViciasepiumL.Thenomenclatureofplants toPamiloetal.(1997)andtomanypersonalob-
followsElven(2016). servations,B.cryptarummaytotallysupplantB.
Entomologicalstudieswereconductedaspart lucorumnorthoftheArcticCircle”.
ofthePolarExpeditionoftheInstituteofEnvi- The distributions of the bumblebee species
192 Kolosovaetal. (cid:127) ENTOMOL.FENNICAVol.27
Table1.Speciesofbumblebeesandtheirrelativeabundances(%)inPymvashor,NorthEuropeanRussia,indif-
ferentyears.
2010 2012 2013
B.(Megabombus)hortorum(Linnaeus,1761) 3.5 1.0 2.4
B.(Thoracobombus)pascuorum(Scopoli,1763) – – 7.1
B.(Th.)schrenckiMorawitz,1881 1.2 – –
B.(Psithyrus)flavidusEversmann1852 79.0 31.2 16.7
B.(Ps.)sylvestris(Lepeletier,1832) 1.2 – –
B.(Pyrobombus)lapponicus(Fabricius,1793) – 4.2 21.4
B.(Pr.)hypnorum(Linnaeus,1758) – 4.2 –
B.(Pr.)pratorum(Linnaeus,1761) 4.6 26.0 7.1
B.(Pr.)jonellus(Kirby,1802) 4.6 18.8 2.4
B.(Pr.)cingulatusWahlberg,1854 1.2 5.2 4.8
B.(Alpinobombus)balteatusDahlbom,1832 3.5 5.2 38.1
B.(Bombus)cf.cryptarum(Fabricius,1775) 1.2 4.2 –
Numberofspecimens 86 96 42
are based on the data of Pekkarinen and Teräs accountedfor25%,andtundraspecies17%ofthe
(1993), Byvaltsev (2009), Levchenko and total. As estimated from the proportion of each
Tomkovich (2014), and Williams (2016). The species caught, the relative abundance of each
main habitats of the bumblebee species are ac- bumblebeespeciesatPymvashorthermalspring
cording to Bäckman and Tiainen (2002), and isgiveninTable1.Themostabundantspeciesin
BolotovandKolosova(2006). the period from 28.VIII to 4.IX.2010 was B.
The relative abundance of each species of flavidus (79%); other species in the community
bumblebeewastakentobeequaltothepropor- wererare.Theperiodfrom31.VIIto5.VIII.2012
tionofeachspeciescaught(Magurran2004).The wasdominatedB.flavidus(31.2%),B.pratorum
relativeabundancewascalculatedforeachyear (26%)andB.jonellus(18.8%).Bombusbalteatus
separately. The periods of collecting were from (38.1%), B. lapponicus(21.4%) andB. flavidus
28.VIIIto4.IX.2010,from31.VIIto5.VIII2012, (16.7%) were the most abundant species in the
and4–16.VII.2013.Thespecimensof2009and communityin4–16.VII.2013.
2014 were excluded from the calculations (and Inthelistoftheexaminedmaterialpresented
Table 1), because there were only few bumble- below, we note the numbers of individuals, the
bees(9individuals). datesoftherecords,thelocalitieswithinthestudy
The specimens of bumblebees from Pym- area,andthecollectornameinbrackets.Queens
vashoraredepositedintheRussianMuseumof ofbumblebeesaremarkedas$,workers(cid:1),and
theBiodiversityHotspots(RMBH),FederalCen- males#.
terforIntegratedArcticResearch,Arkhangelsk,
Russia.
3.1.Bombus(Megabombus)hortorum
(Linnaeus,1761)
3.Results
Examinedmaterial.1(cid:1)28.VIII.2010,bankofthe
The bumblebee fauna of the thermal spring of Pymvashorstream,[Bolotov];1#30.VIII.2010,
Pymvashor was represented by 12 species. The valley of the Pymvashor stream, [Bolotov]; 1(cid:1)
study area was dominated by Transpalaearctic 4.IX.2010,valleyofthePymvashorstream,near
species (58%), while Holarctic ones accounted tothehouse,[Bolotov];1(cid:1)3.VIII.2012,valleyof
for25%,andEurosiberianones17%.According the Pymvashor stream, near to the Iglovsky
totheirhabitatpreferences,mostspeciesarecon- snowfield,[Skyutte];1(cid:1)9.VII.2013,valleyofthe
sidered ubiquitous (58%), while forest species Pymvashorstream,neartothehouse,[Skyutte].
ENTOMOL.FENNICAVol.27 (cid:127) Bombusspp.ofathermalspringarea 193
Distribution.Transpalaearctic. 3.6.Bombus(Pyrobombus)lapponicus
Mainhabitatrequirement.Ubiquitous. (Fabricius,1793)
Examinedmaterial.1(cid:1)23.VII.2009,bankofthe
3.2.Bombus(Thoracobombus)pascuorum Pymvashor stream, [Bolotov]; 1(cid:1) 2# 31.VII.
(Scopoli,1763) 2012,1#1.VIII.2012,valley ofthePymvashor
stream, near to the house, [Skyutte]; 1#
Examinedmaterial.3(cid:1)14.VII.2013,neartothe 10.VII.2013,Goryachayagoraslope,[Skyutte];
thermalpond,[Skyutte]. 2(cid:1) 1# 14.VII.2013, valley of the Pymvashor
Distribution.Transpalaearctic. stream, near to thehouse, [Skyutte]; 4(cid:1) 15.VII.
Mainhabitatrequirement.Ubiquitous. 2013,valleyofthePymvashorstream,[Skyutte].
Distribution.Transpalaearctic.
Mainhabitatrequirement.Tundra.
3.3.Bombus(Thoracobombus)schrencki
Morawitz,1881
3.7.Bombus(Pyrobombus)hypnorum
Examinedmaterial.1#28.VIII.2010,bankofthe (Linnaeus,1758)
Pymvashorstream,[Bolotov].
Distribution.Transpalaearctic. Examined material. 2(cid:1) 1# 31.VII.2012, 1#
Mainhabitatrequirement.Forest. 1.VIII.2012, valley of the Pymvashor stream,
neartothehouse,[Skyutte].
Distribution.Transpalaearctic.
3.4.Bombus(Psithyrus)flavidus Mainhabitatrequirement.Ubiquitous.
Eversmann1852
Examined material. 31# 28.VIII.2010, bank of 3.8.Bombus(Pyrobombus)pratorum
the Pymvashor stream, [Bolotov]; 8# (Linnaeus,1761)
28.VIII.2010, 7# 30.VIII.2010, valley of the
Pymvashorstream,[Bolotov];7#30.VIII.2010, Examinedmaterial.1#28.VIII.2010,bankofthe
1# 4.IX.2010, valley ofthePymvashor stream, Pymvashor stream, [Bolotov]; 1$ 1(cid:1) 30.VIII.
[Bolotov]; 13# 28.VIII.2010, Goryachaya gora 2010,valleyofthePymvashorstream,[Bolotov];
slope, [Bolotov]; 2# 1.VIII.2012, near to the 1(cid:1) 4.IX.2010, valley of the Pymvashor stream,
Pityevoy stream, [Skyutte]; 13# 31.VII.2012, neartothehouse,[Bolotov];1(cid:1)1.VIII.2012,300
9#1$1.VIII.2012,5#5.VIII.2012,valleyofthe mfromthe house, near to the karst zone (kIII),
Pymvashorstream,neartothehouse,[Skyutte]; [Skyutte]; 1(cid:1) 1# 1.VIII.2012, near to the
2# 14.VII.2013, near to the thermal pond, Pityevoystream,[Skyutte];2#1.VIII.2012,near
[Skyutte]; 5# 15.VII.2013, valley of the Pym- tothekarstzone(kII);1(cid:1)7#31.VII.2012,2(cid:1)6#
vashorstream,[Skyutte]. 1.VIII.2012, 1(cid:1) 3# 5.VIII.2012, valley of the
Distribution.Holarctic. Pymvashorstream,neartothehouse,[Skyutte];
Mainhabitatrequirement.Ubiquitous. 3(cid:1) 14.VII.2013, near to the thermal pond,
[Skyutte].
Distribution.Eurosiberian.
3.5.Bombus(Psithyrus)sylvestris Mainhabitatrequirement.Forest.
(Lepeletier,1832)
Examined material. 1# 30.VIII.2010, valley of 3.9.Bombus(Pyrobombus)jonellus
thePymvashorstream,[Bolotov]. (Kirby,1802)
Distribution.Transpalaearctic.
Mainhabitatrequirement.Ubiquitous. Examinedmaterial.1(cid:1)28.VIII.2010,2(cid:1)30.VIII.
2010,valleyofthePymvashorstream,[Bolotov];
194 Kolosovaetal. (cid:127) ENTOMOL.FENNICAVol.27
1(cid:1) 30.VIII.2010, valley of the Pymvashor Distribution.Holarctic.
stream,[Bolotov];2(cid:1)4#31.VII.2012,1$1(cid:1)7# Mainhabitatrequirement.Ubiquitous.
1.VIII.2012, 2# 5.VIII.2012, valley of the
Pymvashorstream,neartothehouse,[Skyutte];
1(cid:1) VIII.2012, valley of the Pymvashor stream, 4.Discussion
[Skyutte].
Distribution.Holarctic. Therearetwomainfactorsthatappeartobere-
Mainhabitatrequirement.Ubiquitous. sponsibleforthespeciescompositionandabun-
danceofbumblebees in thePymvashor thermal
springarea.Thefirstoneisthegeothermalheat-
3.10.Bombus(Pyrobombus)cingulatus
ing,i.e.therearethespecificclimaticconditions,
Wahlberg,1854
differentfromthezonaltundrahabitats.Thesec-
ond factor is that the formation of bumblebee
Examined material. 1(cid:1) 30.VIII.2010, valley of
communitiesisinfluencedbyextrazonalecosys-
the Pymvashor stream, [Bolotov]; 1# 31.VII.
temsinthethermalarea.
2012,3#1.VIII.2012,1(cid:1)5.VIII.2012,valleyof
Duetothecombinedeffectsofthesetwofac-
the Pymvashor stream, near to the house,
tors,thebumblebeefaunainPymvashorhasquite
[Skyutte]; 2(cid:1) 14.VII.2013, near to the thermal
a high number ofubiquitous and forest species,
pond,[Skyutte].
which are not typical of zonal tundra habitats.
Distribution.Eurosiberian.
Furthermore,thenumberofrecordedspecies(12)
Mainhabitatrequirement.Forest.
ishigherthaninotherlocalitieswithinthetundra
zonal habitats of North European Russia (Pota-
povetal.2014b).
3.11.Bombus(Alpinobombus)balteatus
However,thefaunaofbumblebeesinthesur-
Dahlbom,1832
roundingareashasbeenpoorlystudied,because
they are hard-to-reach. The materials represent
Examined material. 1$ 28.VIII.2010, valley of
mainlytheearly20thcenturyinthecollectionsof
the Pymvashor stream, [Bolotov]; 1$ 30.VIII.
the Zoological Institute of Saint-Petersburg
2010,valleyofthePymvashorstream,[Bolotov];
1(cid:1) 28.VIII.2010, on the slope of Goryachaya (Panfilov 1981, 1982, 1984). According to old
gora, [Bolotov]; 1(cid:1) 2# 31.VII.2012, 1(cid:1) 1.VIII. collections,onlyfivespecieshavebeenfoundin
2012, 1(cid:1) 5.VIII.2012, 1(cid:1) 9.VII.2013, valley of theseareas:B.lapponicus,B.hypnorum,B.pra-
torum, B. jonellus, B. balteatus. During our re-
the Pymvashor stream, near to the house,
[Skyutte];3(cid:1)10.VII.2013,ontheslopeofGory- search, we did not study the territories near
achayagora,[Skyutte];2$4(cid:1)1#14.VII.2013, Pymvashor.
ThebumblebeefaunaofPymvashorincludes
near to thethermalpond, [Skyutte];1$ 16.VII.
2013,neartotherocky slope,[Skyutte];1$6(cid:1) species that are widely distributed in the Palae-
arctic,whichindicatesthelowspecificityofthe
1#25.VII.2014,valleyofthePymvashorstream,
community. It contains two typical tundra spe-
neartothehouse,[Skyutte].
cies,B.balteatusandB.lapponicus.Theirabun-
Distribution.Transpalaearctic.
danceswerenothighinthelatesummer2010and
Mainhabitatrequirement.Tundra.
2012buttheydominatedinthecommunityinthe
middleJuly2013(Table1).
3.12.Bombus(Bombus)cf.cryptarum Theforestspeciesinthebumblebeefaunaof
(Fabricius,1775) thestudyareaincludeB.schrencki,B.pratorum
and B. cingulatus. These species are common
Examinedmaterial.1(cid:1)28.VIII.2010,onthebank within the taiga zone of the European North of
of the Pymvashor stream, [Bolotov]; 1(cid:1) Russia, but are not typical for tundra habitats
31.VII.2012,1(cid:1)1#1.VIII.2012,1(cid:1)5.VIII.2012, (Bolotov&Kolosova2006).
valley of the Pymvashor stream, near to the ThepresenceoftheubiquitousB.hortorumis
house,[Skyutte]. noteworthy, as it is usually characteristic of
ENTOMOL.FENNICAVol.27 (cid:127) Bombusspp.ofathermalspringarea 195
meadow communities (Bolotov & Kolosova Breeding of the Russian Academy of Agricultural Sci-
2006).Theotherubiquitousspecies,B.jonellus, ences, a member of Russian Botanical Society), R. I.
Gutiev(seniorengineeroftheLimitedliabilitycompany
isatypicalrepresentativeofthebumblebeefauna
“Design and development of the expedition”), S. O.
at high latitudes (Chernov 1966, Bolotov &
Yaylyan(GeneralDirectoroftheNon-CommercialPart-
Podbolotskaya 2003). The presence of B. flavi- nershipOff-roadClub“Bear”)forassistanceincollecting
dus,whichisasocialparasiteofB.jonellus,was bumblebeesandforcomprehensiveassistanceinthisre-
expected(Løken1984).Itsrelativeabundancein search.WearegratefulalsotothestaffoftheNaturalHis-
toryMuseum,LondonandtheZoologicalInstituteofRus-
the community was very high, 79% in 2010
sianAcademyofSciences,SaintPetersburgforproviding
(Table1).Suchhighvaluesofabundancemaybe
us with the opportunity to examine collections. Special
explainedby thefactthatwecollectedbumble- thanksareduetoDr.MatthewCopleyforimprovingthe
beesinthelatesummerof2010,whichistheperi- languageofthepaper.ThisstudywasfundedbytheRus-
od for the flight activity of B. flavidus males sianFoundationforBasicResearch,accordingtothere-
searchprojectNo.16-34-60035mol_a_dk.Thisstudywas
(Løken1984).In2012and2013,whenthecol-
supportedalsobytheFederalAgencyforScientificOr-
lectingperiodwasinthemiddlesummerorclose
ganisations(No.0410-2015-0034,No.0410-2014-0028,
to it, the relative abundance of B. flavidus was No.0410-2014-0025),theUralBranchofRussianAcad-
lower. emyofSciences(No.15-12-5-24,No.0410-2015-0032)
Unfortunately, we have no possibilities to and grants from the President of Russia (MD-7660.
2016.5),theRussianFoundationforBasicResearch(No.
compare the relative abundance of species be-
16-05-00854_a).
tweenPymvashorandthesurroundingareas,be-
causethereisfewinformationfromthere.Wecan
assume that the bumblebees develop more rap- References
idlyinPymvashorthermalspringthaninterrito-
Bäckman,J.P.&Tiainen,J.2002:Habitatqualityoffield
riesofzonalhabitats.Forexample,similarpro-
margins in a Finnish farmland area for bumblebees
cesses were observed in Kunashir Island (Rus-
(Hymenoptera:BombusandPsithyrus).—Agricultu-
sianFarEast)(Potapovetal.2014a). re,EcosystemsandEnvironment89:53–68.
OurconclusionisthattheareaofPymvashor Bolotov,I.N.&Bogolitsyn,K.G.(ed.)2011:(Operation
thermalspringssupportsalargernumberofbum- ofsubarctichydrothermalsystemsinthewinterperi-
od.).—UralBranchofRussianAcademyofSciences,
blebeespeciesthanistypicalofthesurrounding
Ekaterinburg.252pp.[InRussian.]
tundrahabitat,andthatreflectsthegreaterphysi-
Bolotov,I.N.&Kolosova,Yu.S.2006:Trendsinthefor-
calandfloristicdiversityofthethermalarea.The mationofbiotopiccomplexesofbumblebees(Hyme-
ecosystemofPymvashor,whichisformedunder noptera, Apidae: Bombini) in northern taiga karst
theinfluenceofthermalexposure,isakindof“re- landscapesoftheWesternRussianPlain.—Russian
JournalofEcology37(3):156–166.
servehabitat”(Lobkova&Lobkov2003)forthe
Bolotov,I.N.&Podbolotskaya,M.V.2003:Localfauna
typicalzonalspeciesofinsectsandfortheinhab-
of bumblebees (Hymenoptera: Apidae, Bombini) in
itantsfromthemoresouthernareas. theEuropeinNorthofRussia:theSolovetskyIslands.
We can assume that bumblebee species, — Vestnik Pomorskogo Universiteta, Seriya estes-
whicharecommonwithinthetaigazoneandnot tvennyeitochnyenauki1(3):74–87.[InRussianwith
summaryinEnglish.]
typical for tundra habitats, may represent relict
Bolotov,I.N.,Surso,M.V.,Filippov,B.Yu.,Gofarov,M.
populations, from a time when the climate was
Yu.&Tarakanov,A.M.2012:Changesintreestands
generallywarmerinthisarea.DuringtheHolo- onisolatedforestislandsintheEastofBolshezemel-
cenethermalmaximumontheeasternoutskirtsof skayaTundraforthelast100yearsinachangingcli-
themodernBolshezemelskayatundrawasrepre- mate.—LesnoyZhurnal5:30–37.[InRussianwith
summaryinEnglish.]
sentedbythetaigazone(Velichko2002).Hence,
Bossert,S.2015.Recognitionandidentificationofbum-
the Pymvashor thermal area became “a reserve
blebeespeciesintheBombuslucorum-complex(Hy-
habitat” for the bumblebee species of the taiga menoptera,Apidae)–Areviewandoutlook.—Deut-
zonelaterwhentheclimatebecamecooler. scheEntomologischeZeitschrift62(1):19–28.
Byvaltsev,A.M.2009:(Bumblebees(Hymenoptera:Api-
dae,Bombini)offorest-steppeandsteppeinthesout-
Acknowledgements.Theauthorsexpresstheirappreciation hernWest-SiberianPlain:faunaandcommunity.PhD
toA.E.Skopin(PhD,theleadresearcheroftheDepart- thesis.). — Institute of systematics and animal eco-
mentofAnimalEcology,InstituteofHuntingandAnimal logy,Novosibirsk.21pp.[InRussian.]
196 Kolosovaetal. (cid:127) ENTOMOL.FENNICAVol.27
Chernov,Yu.I.1966:(Complexofanthophilousinsectsin ra, Apoidea, Apidae. — In: Gorodkov, K. B. (ed.),
thetundrazone.).—Voprosygeografii69:76–97.[In ProvisionalAtlasoftheInsectsoftheEuropeanpartof
Russian.] USSR.Maps179–221:28–32.Nauka,Leningrad.61
Elven,R.2016:AnnotatedChecklistofthePanarcticFlora pp.[InRussian.]
(PAF).Vascularplants.URLhttp://nhm2.uio.no/paf/. Pekkarinen,A.&Teräs,I.1993:ZoogeographyofBombus
(Sitevisitedon31August,2016) andPsithyrusinnorth-westernEurope(Hymenoptera,
Garbuz,D.G.,Zatsepina,O.G,Przhiboro,A.A.,Yushe- Apidae).—AnnalesZoologiciFennici30:187–208.
nova,I.,Guzhova,I.V.&Evgen’ev,M.B.2008:Lar- Potapov,G.S.,Kolosova,Yu.S.&Bolotov,I.N.2014a:
vaeofrelatedDipteraspeciesfromthermallycontras- Thestructureofbumblebeecommunities(Hymenop-
tinghabitatsexhibitcontinuousup-regulationofheat tera,Apidae,Bombusspp.)insomeecosystemsofKu-
shockproteinsandhighthermotolerance.—Molecu- nashir Island and Southern Sakhalin (Russian Far
larEcology17:4763–4777. East).—RussianJournalofEcology45(4):304–307.
Glazier,D.S.&Gooch,J.L.1987:Macroinvertebrateas- Potapov,G.S.,Kolosova,Yu.S.&Gofarov,M.Yu.2014b:
semblagesinPennsylvania(USA)springs.—Hydro- Zonaldistributionofbumblebeespecies(Hymenopte-
biologia150:33–43. ra,Apidae)intheNorthofEuropeanRussia.—Ento-
Hines,H.M.2008:Historicalbiogeography,divergenceti- mologicalReview94(1):79–85.
mes,anddiversificationpatternsofbumblebees(Hy- Pritchard,G.1991:Insectsinthermalsprings.—Memoirs
menoptera:Apidae:Bombus).—SystematicBiology oftheEntomologySocietyofCanada155:89–106.
57:58–75. Pritchard,G.&Kortello,A.1997:Roosting,perching,and
Levchenko,T.V.&Tomkovich,K.P.2014:Contribution habitatselection in Argia vividaHagen and Amphi-
to the bee fauna (Hymenoptera: Apiformes) of the agrion abbreviatum (Selys) (Odonata: Coenagrioni-
Khanty-MansiAutonomousRegion,WesternSiberia, dae),twodamselfliesinhabitinggeothermalsprings.
Russia.—Entomofauna:ZeitschriftfürEntomologie —CanadianEntomologist129:733–743.
35(5):85–100. Rasmont,P.1984:LesbourdonsdugenreBombusLatreil-
Lobkova,L.E.&LobkovE.G.2003:(Roleofbiological lesensustrictoenEuropeoccidentaleetcentral.—
components in ecosystems of the thermal fields of Spixiana7:135–160.
UzonandValleyofGeysers,andsomeaspectsofpro- Rasmont,P.&Iserbyt,S.2010:GenusBombus.Atlasof
tections of thermal ecosystems.) – Conservation of theEuropeanBees.STEPProject.AtlasHymenopte-
biodiversityofKamchatkaandcoastalwaters.Materi- ra.URLhttp://www.atlashymenoptera.net.(Sitevisi-
alsofIIIscientificconference:258–262.[InRussian.] tedon31August,2016)
Løken, A. 1973: Studies on Scandinavian bumblebees Rasmont,P.&Terzo,M.2010:Catalogueetclédessous-
(Hymenoptera, Apidae). — Norsk Entomologisk genresetespècesdugenreBombusdeBelgiqueetdu
Tidsskrift20(1):1–218. norddelaFrance(Hymenoptera,Apoidea).—Uni-
Løken,A.1984:ScandinavianspeciesofthegenusPsithy- versitédeMons,LaboratoiredeZoologie,Mons.25
rusLepeletier(Hymenoptera,Apidae).—Entomolo- pp.
gicaScandinavica23:1–45. Resh,V.H.&Barnby,M.A.1984:Distributionofshore
Magurran,A.E.2004:Measuringbiologicaldiversity.— bugsandshorefliesatSylvansprings,Yellowstone
BlackwellPublishing,Oxford.215pp. NationalPark.—GreatBasinNaturalist44(1):99–
Pamilo,P.,Tengö,J.,Rasmont,P.,Pirhonen,K.,Pekkari- 103.
nen,A.&Kaarnama,E.1997:Pheromonalandenzy- Stockner,J.G.1971:EcologicalEnergeticsandNatural
megeneticcharacteristicsoftheBombuslucorumspe- History of Hedriodiscus truquii (Diptera) in Two
cies complex in northern Europe. — Entomologica ThermalSpringCommunities.—JournaloftheFishe-
Fennica7:187–194. riesResearchBoardofCanada28(1):73–94.
Panfilov,D.V.1978:(ThekeysforthespeciesofFamily Velichko,A.A.(ed.)2002:Dynamicsofterrestrialland-
Apidae–Bees.).—In:Medvedev,G.S.(ed.),(The scape components and inland and marginal seas of
keysforinsectsoftheEuropeanpartofUSSR)3(1): NorthernEurasiaduringthelast130,000years.Atlas-
508–519.Nauka,Leningrad.584pp.[InRussian.] monographEvolutionoflandscapesandclimatesof
Panfilov,D.V.1981:BombusLatreille,1802.Hymenopte- Northern Eurasia. Late Pleistocene-Holocene-ele-
ra, Apoidea, Apidae. — In: Gorodkov, K. B. (ed.), ments of prognosis. II. General paleogeography. —
ProvisionalAtlasoftheInsectsoftheEuropeanpartof GEOS,Moscow.296pp.[InRussian.]
USSR.Maps73–125:22–28.Nauka,Leningrad.57 Vinogradova,V.M.1962:(Floraofthethermalsprings
pp.[InRussian.] Pym-Va-ShorintheBolshezemelskayaTundra.).—
Panfilov,D.V.1982:BombusLatreille,1802.Hymenopte- VestnikLeningradskogoUniversiteta,Seriyabiologi-
ra, Apoidea, Apidae. — In: Gorodkov, K. B. (ed.), cheskaya9(2):22–34.[InRussian.]
ProvisionalAtlasoftheInsectsoftheEuropeanpartof Williams P. H. 2016: Bumblebees of the world. URL
USSR.Maps126–178:25–28.Nauka,Leningrad.77 http://www.nhm.ac.uk/research-curation/pro-
pp.[InRussian.] jects/bombus/index.html.(Sitevisitedon31August,
Panfilov,D.V.1984:BombusLatreille,1802.Hymenopte- 2016)