Bumble Bees

Z k/ /4.evQ4h,"./Lt:ffNORSK ENTOMOLOGISK TIDSSKRIFTVOLUME 20, NO. I, T973Studies on ScandinaYuafiBumble Bees\ (Hymenoptera, Apidae)BYASTRID LAKENUNIVERSITETS FO RLAGET

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In memory ofOve Meidell (1 903- 1942)

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Studies on Scandinayran Bumble Bees(Hymenoptetl, Apidae)ASTRID LOKENLoken, A. 1973. Studies on Scandinavian Bumble Bees (Hymenoptera, Apidae).Norsk ent. Tidsskr. 20, l-218.A synopsis of the genus Bombus Latreille in Norway and Sweden is presented.Although primarily a taxonomic, phenotypic, and faunistic study, bionomic informationis incorporated and post-glacial immigration routes are stressed.A total of 29 species, divided into 34 subspecies, has been recognized. One newsubspecies, -8. muscorum liepetterseni (: B. m. smithianus auctt. nec White, inpart), is described, and holotype ? is designated. The following types are alsoselected: B. mastrucatus Gerstaecker lectotype d; B. martes Gerstaecker lectotype?; B. humilis ha/sahlianus Yogt lectotype H; B. pascuorum sparreqnus nov.nom. (: .8. agrorum bicolor Sparre Schneider, 1909 nec Friese, 1903) hototype g.Astrid Loken, Zoological Museum, (Jniversity of Bergen, lr{-5014 Bergen, NorwayPREFACEThe world-wide genus Bombus Latreille occurs inall zoogeographic regions with a great number ofendemic species represented in holarctic, neotropical,and oriental regions (Skorikov lgzzb).one species was recently recognized in the Ethiopianregion (Tkalcu 1966) and four Europeanspecies introduced into the Australian region, i. e.New zealand (Gurr 1964). The species in thisgenus, i. e. the true bumble bees, are beneficialinsects, pollinating a wide range of plants includingthose of great economic importance. Theyate, moreover, social bees and, being stronglypolymorphic, they have been intensively studiedfrom genetic, taxonomic, and zoogeographicalpoints of view (Vogt 1909, lgll; Friese &Wagner 1909, 1912, etc.; Krtiger 1928, 1931,1940, 1950, etc.; R"einig 1936, 1939, 1965, etc.;etc.) besides being subject to basic and appliedresearch devoted to their biology and ecology(references in Free & Butler 1959).Floral ecological observations and preliminarypollination research (Loken 1949, 1950, 1958a,l96lb, 1962) emphasized the need for a taxonomicand faunistic study of Norwegian bumble bees.A revision of the collections was therefore startedin the year 1962. To secure information for abetter understanding of the species, the scope ofthe study was later widened to present a mono-graph of Bombus in Scandinavia, i.e. Norway andsweden, the entire peninsula being a more welldefinedunit. Though taxonomy, phenotypic variations,and distribution are the topics of thestudy, bionomic information has been added andthe opportunity taken to refer to the world distributionof the species involved. To limit thesubject, the work does not, however, deal withplant preference, which in fact was an essentialaim in the author's field observations.INTRODUCTIONHistorical reviewLinnaeus (1758) listed a total of seven Bombusspecies occurring in his home country Sweden,six of which are recognized as species, viz. thosetermed Apis terrestris, A. lapidaria, A. muscorunt,A. hypnorum, A. subterranea and A. alpina,whereasthe seventh designation, A. aceruoruffi, is asynonym of A. subterrenea. All the species are infact mentioned by Linnaeus in his Acta (Jpsaliensrs(1736). Four more Swedish species were addedby Linnaeus (1761) and named Apis hortorum,A. pratorum, A. sy,lvarum and A. lucorum. Asmany as 32 Bombus species are ascribed to Scandinaviaby Dahlbom (1832), twenty of which arerecognized as species. The remainder are synonyms,subspecies, or infrasubspecific forms ex-

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Studies on Scandinavian Bumble Beeswooded passes at approximately 63" 30' N:(1) A southern range occupying most of SouthernNorway and from 61" N northwards also theadjacent border area of Sweden, the isolatedmountains in Dalarne being included. (As to thelocation of counties mentioned in the text, videFigs. 97-99.) The watershed divide is situatedrather far west. The range consequently slopesmuch more gradually towards east than towardswest, where the mountains reach the coast fromwhere the fjords with very steep sides cut moreor less far into the country. (2) A northern rangeextending in varying width along the Norwegian-Swedish border, r,vith a shift of maximum elevationinto Sweden between 67o and 68o 30' N, andfurther north continuing to the northern coast ofNorway. The heights of the ranges, being 900-1200 m with peaks rising to a maximum altitudeof 2468 m in the mountainous massif of Jotunheimenin Southern Norway, are in general loweron going north.Norway is a pronounced mountainous country.3 ' r t ' ' r yt5 23.-- l '| \\ r\ rt'f-i.^-^ \. r?-\ - \\ltI,\t rii\ \ r\ tr+'$I--J-7TT-.v''< $ao( lIlII\\i \ \\\\\-\\\\bFig. l. The mean annual precipitation in Fennoscandia. Black area : > 2000 mm. (From Lindroth 1949,p. a86.)-iF+

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Studies on Scandinavian Bumble Beesdrift of lowland species up to the alpine and subalpinebiotopes. The term regio quercina for thepart extending along the western coast (Fig. 2) istherefore somewhat misleading, but since thiszone depicts the northern border area for a numberof continental plants and animals, it is quitesatisfactory for the present purpose.MATERIALThe present work is based upon a study of approximately29,000 Norwegian and 27,700 Swedishspecimens. It comprises old and recent collectionspreserved in institutions, three small personalcollections, and material presented to theFig. 2. A simple regional division of Fennoscandia. Obliquely hatched : regio fagina; vertically hatched :regio quercina; white : regio coniferina (and regio subalpinalsubarctica); stippled' : regio alpinalarctica. Thenorthern coniferous limit drawn in Eastern Fennoscandia only. The isolated northernmost coniferous forestsblack. (From Lindroth 1949, p.437, somewhat remodelled.)

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Studies on Scandinavian Bumble BeesNatural History, London. Scandinavian materialin general collections, in the Pittioni collection,and in small unarranged collections (Benson,Holgersen, Sparre Schneider, British expeditions,etc.).Zoologisch Museum, University, Amsterdam.Norwegian and Swedish specimens in the generalcollection (Eyndhoven, Kruselnan, Reusch, Toxopeus,etc.). Large Norwegian collection (Barca,Esmark, Hafsahl cf. below, Lie-Pettersen, Lyshoim,Sparre Schneider, Scho-ven, Storm, etc.),and Swedish collection (Bengtsson, Klaue, Roman,etc.) in Vogt collection, which also includesKri.iger collection. Common Swedish specieswere not examined. Frofessor O. Vogt (1909,1911, 1947) communicated with a great manyentomologists and amateurs who fur"nished himwith bumble bees. Specimens not being labelledby the collector were provided by Vogt with smallred locality labels. The great number of individualscollected by the Norwegian medical doctorP. Hafsahl, who lived in Hamar about 100 kmnorth of Oslo, all have the locality F{amar printedon those red labels. However, according to hisnephew, Hafsahl collected bumble bees at variouslocalities, mostly in the lorvlands near by Hamar,but also in subalpine and alpine areas and evenon his travels as far as Germany (A. A. Hafsahlin litt.). Indeed, his travels are reflected in hisbumble bee collection, which represents taxa nototherwise knowu from Hamar, even species notreaching Norway. Owing to the erroneous labelling,the entire material of Hafsahl, part of whichis referred to by Vogt (1909, 1911 , 1947) andKrtiger (1950, 1954, 1956) must unfortunately beconsidered unreliable and is therefore omitted.Zoologisk Museum, University, Copenhagen.Small Norwegian collection (mainly labelled Mus.Drewsen and Mus. Schiodte).American institutions. Cornell University, Dept.of Entomology, Ithaca, New York. Srnall Norwegiancollection (Raffensperger).U.S.D.A. Entomological Research Div., Log&n,Utah. Small Norwegian collection (tsohart).U. S. National Museum, Washington D. C.Small Norwegian collection (SabroskY, SparreSchneider).Personal collectionsLarsson, FI., Uppsala, Sweden. Small collectionfrom Norway: Sor-Trondelag and Sweden:Skflne.Semb-Johansson, A., Oslo, Norway. Small collectionfrom SE Norway.Tjeder, T., Riittvik, Sweden. Small collectionfrom Sweden: Dalarne.In addition to Scandinavian collections, severalFinnish specimens kept at Zoologiska Museet,Helsingfors and Museum Zoologicum, A,bo,were revised.METHCDSTaxonomyCoincident with identificationlrevision of Norwegian.collections,the data. for the individualspecimens were codified, punched on cards, andthe species then sorted geographically by computer,except a minor material identified andadded to the locality lists during the years 1968-1970. The specimens in the Swedish collectionswere catalogued by species and afterwards reaffanged geo graphically.Keys to the Scandinavian Bombus spp.are provided.In the subsequent treatment of the individualspecies they have not been fully described,but are given a fairly broad diagnosis. The s3xesare treated separately. The female characteristics,presented in three sections viz., morphologicalfeatures, colour pattern of the coat, and phenotypicvariations, are introduced flrst and followedby those of the male. To save space, the descriptionof the worker is included in that of thequeen even though some of the features have lessdiagnostic value in that caste, being too diffuse orvariable.The species are treated in alphabetical orderwithin the subgenera and the sequence of the subgeneraare affanged according to Richards (1968).If more species are involved, each sex is given asubgeneric description in order to avoid needlessrepetition in the subsequent description of thespecies. If there is only one species in the sub-

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Studies on Scandinavian Bumble BeesMeasurements. The following queen measurementswere taken: (a) right malar space (Fig.3c-d); (b) 'radial length', i.e. the distance fromthe 'veinlet' in the middle of first Rr cell (: firstsubmarginal cell) to the distal end of second Rr( - radial cell) (Fig. 5); this measurement not tobe confused with the length of radial cell; (c)interalar width (Fig. 6); (d) distance from rightocellus to preoccipital ridge (Fig. 4d); (e) distancefrom right ocellus to nearest compound eye(Fig. 4e). The measurements a-c were taken forthe queen of each species or subspecies, whereasthose marked d-e apply to a few species only,the ratio d:e being used as a diagnostic character.Besides the diagnostic value, malar spacecan roughly be correlated with the tongue lengthwhich makes this measurement useful in the studyof bumble bees and their plant preference.Measurement of interalar widthwas chosenas an indicator of the body size. The featuresa-c were moreover selected for triangulargraphing, cf. below, and in one subgenusthose marked d-e were used for the samepurpose.Least reliable measurements are those of thetotal length, viz. the distance from face of headto tip of abrJomen when head is in vertical positionand venter of abdomen in horizontal position.This character is subject to considerablevariation, above all because it is dependent uponthe degree of extension of the gastral segmentswhich is difficult to measure accurately. Nonethelesseach species or subspecies gives the impressionof having a characteristic length whichat any rate can be expressed in relative terms.Three categories are used (the correspondingaverage measurements of interalar width followin brackets): Body of large size, the length exceeding20 mm (interalar width exceeding 5.00mm) ; body of medium size, the length 17 -20 mm(interalar width 4.50-5.00 mm); body of smallsize, the length less than 17 mm (interalar widthless than 4.50 mm).As corresponding measurements of males ateless meaningful in a broad diagnosis and are notneeded for triangular graphing, they have beenomitted. The male body size is, however, indicatedby the following three size categories:Body of large size, length exceeding 18 mm; bodyof medium size, the length 14-18 mm; body ofsmall size, the length less than 14 mm.There is a distinct difference in size betweenalive and dried bumble bees, and very soon afterdeath the shrinking of abdomen is particularlynoticeable. As museum collections were the onlyavailable material in some of the species, allmeasurements were taken on dried specimens"The measurements were made witha Leitz-Wetzlar binocular, ocular 12.5 x , objective 4 x ,i. e. 40 x magnification, and a micrometer eyepiecewith each unit equal to 0.05 ffiffi, exceptthat of interalar width, which was measured byvernier calliper and read to an accuracy of0.05 mm. The features were measured once oneach specimen except interalar width, which wasmeasured twice and the mean value used. Themeasurements are presented for the queen of eachspecies or subspecies by the number (N), area,mean, standard deviation, mean error, and range.Example: N - 15 ; SE Norway; malar space:0.08 mm (+O.O: +0.01) range: 0.70-0.80 mm.Needless to s&y, the area representing the measuredpopulations cannot all be the same owingto varying distribution of the taxa involved and/orthe available queens as well.Triangular graphing. Specific and subspecificposition of certain closely related taxaare indicatedby triangular graphing (Mayr 1969) based on themeasurements of the features marked a-c above,or those marked b, d-e. tsy this method areplotted not the actual values, but their percentagecontribr"rtion to the Sum ( - 100 per cent) of thecharacters which indicate the body proportionsrather than the absolute size. Specific differencebetween two or more populations in generalaccords with difference in body proportion. Thisis illustrated by the measured populations occupyingdifferent areas on the triangular graph(Figs. 52, 53, 59, 66,72), whereas conformity inbody proportion corresponds to congruentspreading of the markings and favours conspecificity(Figs. 66, 81).Melanism, geographical variations. Variabilityof colour is somewhat arbitrary and not suitablefor statistical treatment. The rather large materialat my disposal has nevertheless stimulated me tostudy melanism and clinal variations by a broadgrouping of certain colour features. By dividing

erueu osor{/h 's1run o^rleJlsrullupe uo possq uorsr^-!p pcqderEooE e Eursn go eEelue^pesrp eql JooJe.&\e sr roqtne oql '(96 ,1,6.sEIC) 696I ol pel€puooq oler{ s}1urll pue seru€u ]€rI] ldecxe (wADpusJls s^{olloJ serlrc pue s}3rr}srp prnJ Jo }€rllpue sorlunoc eql Jo ecuenbes er{I .JolcelloJ eq]ot sroJer sesoqtuered ur eru€u or{I .TCC[ (ptggs-ueso'l{) ZtgI ,(lnl SI ,p ul 0001 letgerzro6l :ernoq: (uO ) pun1ddg : szlrolloJ se pelueserd era eqppelrelop 'serceds erer Jer,IleJ Jo sls11 erll ur puesorlrl€col leJeqdued euros JoC .ul\ou{ JI sell[ecoleuldle JoJ uen6 sr epnlrll€ eql .Jle .ouelgol'UINZ g.WZ ur 0€6 enls)loc 'owz flwz suNutrl 0001 gefgerno6l :erloq : (uO) puolddg:s^\olloJse (.re{t,{J,) serlunoc lueregrp eq} up{tll\ .(}lcpue (.perreq,) lJrJlsrp IeJnJ ,{q ,{gecrleqegdlepe8uerru ote sorueu eql serlrlucol ue6ezluoN Jols11 eqt u1 '1de{ ore sprocoJ oqt eJoq^\ uor}nlr}sureql pue f,14ecol or{} ol polcrr}sor preueE ur oroJ-eJoql sr seluscol Jo lsll oqroceds e^rssorxe e)I?lplno^\ suerurceds 11e tuo4 ercp po1relop ]s11 oI'lseJolur Jelncrlrcd go Eureg JIpels[ .{po oJe qcrqzn 'surrrol eErel Jo ser]rc unllr^\esoqt ldocxo serqecol Jo lsll et{t ur peuor}ueuete seureu lecryder8odot ilV.serlrleool ,(greeule epetu suorlcelloc IeJeAes Jo flrlecol leql tepelce[oc sueunceds eJour Jo euo s]ueseJder 1opqceg 'tu{ 0I o} spuodserroc }op e ur snrp"J eql'sdeur eql uO 'peppe ueeq eleq ,eur ,{q peuruexe'sprocel qsruurg ,!\oJ V.suorleclpqeJnleJelrlol Eurprocce serJlunoc EurrnoqqEreu Eurureur-er eqt ur pue (Sq01 Eu1,gp) puelurd ur eEue.ru^\ou>I erlt lcrdep osl€ rlclr{A\ sdeur uo pettoldueeq on€r{ spJocoJ dlqecol .serceds uerl€urp-ueJs qc€o Jo uorlnqrJlslp oql ol osuereJer rocuounquts!(['sed,(1 se tueql Eurlels lnoqll^t lnq .uodn pelueu-utrroc'pue peuorlueu ore reded lueserd eql urpeleeJl 'dds snqwog ueeeuulr eql o1 EurueJeJ uo[]-celloJ ueoeuulT eql ur suerurcedg .GgAy r,(e61)petqnop olorl^\ eql uo sr uorlJelloc slql Jo enle^.(reluewncop eql'Jelel peppe lou pue uorlcellocueeeuurT eql ur penreserd .(lleurErro sueurrcedsol srSger esec sFIl ur 1eu6uo'ed.(1 se uourrcecls^(ue peleuElsep releu sneeuulf sV .(SEOI spreqc-ID sleurErro ete uopuoT ur uorlce[oc sneeu-ulT eqt w sueturceds snqwog rer,Ileql\ elqeleqepsr l! 'rener!\oq 'sedr(1 uBeeuurT etll EururecuoJ'suorlsuErsep eqt Jo qcee ol ecueJeJeJ eql s/KolloJed,tl etll lnoqe uorletuJoJul elqelrelv .sadr(1'uorssncsp cglcedsqns eql ur Jos>lJerueJ leJrruouoxe] Jepun peuorlue{u ueeq eA€r,I''cle'suorleJJeqe'sruro;cgrcedsqnse4ur petueuol secuereJe1 '(SqO I '0961 Eu4g1g) sreded luecerol pelclJlseJ ele 'dds snqwoBr r1sruulC eql o} socue-reJo1 'glesfw ro .O 11epletre]el eq] ,(q pesrlarsuorlcefioc uo pessq uorleuJoJur cr]srune3r lueserdf,..{l JI pe}sl1 ,{po ere qcreesor perldde ul po^lolursorceds ol secrreJoJol{ 'poppe uoeq os1e oluq euneJuerluurpuecs oql ol socuoroJor cqderEolqlg.uoxelorl] Jo snlels lueserd eql elec1pu1 ol uolrE sl urel-Wd Jnoloc eql q suorlelrel l€erE ,{eldsrp r{crr{a\serceds ureuec JoJ secueJeJeJ Jo Jequnu eErele elrnb lnq 'pepnlcur ueeq eêq ,(gOgt) errol€lleq ,{q pe}s11 lereueE ur ,seuleu plo ile ]oN'elelduroc eq ol puelerd tou op 'lueurleeJl qcseJo EuruulEeq eqt W ue^nrE'sur^,(uour{s o} socueJe-JeU'suorsnJuoc leJnlulcuoruou teetf posneo el"qEurztrttoufs snoeuoJJe pue suorleuErsep prlelul'sruJoJ cgrcedsqnseJJul pue,sarcedsqns,sru^{uou,(sol se pesrloJ ueeq ol€r{ qcFIA\ yo ,(ueru .suol}€u-6rsep cgrcedsJo Jequnu lsel € ol osrJ uonrE seq serc-eds oeq olqtunq len]ce oq] Jo oruos ur urelledrnoloceql q ,ttryqulJel l€eJC 'sa)ua"tata"t,ttur(uou,ts'erleurpuecs lnoqEnoJql suorlerJel crqdurE-oeE eleJrpur ,(gEnoJ esec sFIt ur seop serlunoc^(g suorlelndod Jo uors4lp e ,serpnls leJrqder8oeE-ooz ur pelceler eq ptnoqs s]run elr]eJ]srurupeqEnoql ueÊ Joererll lcrJlslp ro ,(1unoc B urr1lrl\lerJoletu elq€lrele eql ol sprrodserroc eloqe pesnse uortelndod lerol rure] egl.(7S-ES,09 ,gL,LL's8lg) suorlerrel leurlJ elgrssod elerlsnp ol surerE-elp ol peJroJsven ueeq wq,{celq ecue}sul JoJosernleeJoql Jo ouo Jo uollnqrrlsrp eEeluecred eq1'ep€r1s Jo uorsseJEord ur pue sdcls ur peEuerreseJnleeJ Jnoloo urepeJ of Eulprocce serceds eJo suorlelndod [eco[ eq] Eu,dnorE ,(q pe1pnls eJo&\suorlerrel lecqderEoeE Jeqlo .dcuenberg lecolet{t slcouoJ ,{1}cexe }l ueqt tusruelotu o} .{cuep-uel eqt eJoru 'esrnoc g:o 'selerlsuoruep uorl€nl-BAe eql'(SS 'VL'69'sElg)sgeqc-erd ol porreJ-sueJl pue 6peleFqel 'pelelnclec se.r\ suourrcedscruelotu Jo e8eluecred oqt 'slenprrrlpul Jrueletupue 1ecrd.'(1 olq serceds e Jo uorlelndod Iecot eqlua4o7 'v 0t

Studies on Scandinavian Bumble Bees 11and limits can be altered. However, until another of counties, rural districts, cities etc. have beendivision has been generally accepted by entomol- written in their native language. Most Finnishogists I hesitate to leave such a well established names arranged by counties (Fig. 99) have beensystem as that of Strand's (1943). It should be listed by Elfving (1968). Otherwise geographicalnoticed that old specimens labelled Dowe are in names outside Scandinavia follow essentiallythe lists mentioned as Oppland (On): Dowe: Grosvenor (1963).Dovrefjell 1000 m, as those specimens obviously * The Norwegian letter o (:ct) has been used inrefer to the mountainous plateau and not to On: topographical names between pp. 205-209 and inDovre: Dowe, i.e. the administrative centre of personalnamesthroughoutthetext.the rural district situated in the valley about Abbreviations500 m s.m.In locality lists of the Swedish collections theA' Museumstopographical names were arranged alphabeti- ,C.fA : AOo Akademi Museum Zoologicum,cally within the individual county ('landskap').FinlandHowever, the sequence of the letters follows that BCL : Bank collection, London (BML)of the Norwegian alphabet in which the letter a BML : British Museum, Natural History,is placed at the end. The sequence of the counties,Londonrangingfromsouthtonorth,followsthatpractised CCC : Copenhagen collection, Copenhagenin the Catalogus Insectorum Sueciae (publ. Opus-(ZMC)culaEntomologicaforinst. 1955, 1960).Thecoun- CIN - Cornell University, div. of Entomoltiesare presented in the geographical division ofogy, Ithaca, New YorkFennoscandia (Fig. 99).* CNO : Canadian National collection, OttawaLocalities of unrevised records, being published DCL : Dahlbom collection, Lund (ZML)or received by personal communications, succeed GNM : Giiteborg Naturhistoriska Museet, Gdtheindividual list of localities. However they haveteborgbeen added only if they contribute especially to M : Institution fdr Viixtpatologi, Uppsalathe fauna and are considered reliable as well. IZB : Institute f. spezielle Zoologie u. Zoo-The world distribution of the species and thelogisches Museum, Berlintotal distribution of Scandinavian subspecies JCL : Jansson collection, Lund (ZML)have been indicated in separate sections with the KCC : Kiel collection, Copenhagen (ZMC)references gathered at the end of the actual sec- KCL : Kirby collection, London (BML)tions to save space-taking repetitions. Reference KMT : Kongelige Yidenskabers Selskab, MutoYarrow in litt. concerns his unpublished col-seet, Trondheimlections, mainly from Portugal, Spain, Egypt, LSL : Linnaean SocietS LondonIrak and Iran, kept in the British Museum, MAL : Zoological Museum, Academy ofLondon, which he kindly allowed me to include.Science, LeningradThe Norwegian locality names refer to those MNP : Mus6um National d'Histoire Natuontopographical maps edited by Norges Geo-relle, Parisgrafiske Oppmiling, Oslo as far as they cover the NMV : Naturhistorisches Museum, Viennalocality in question. A great many names Eue NMW : U.S. National Museum, Washington,otherwise listed in the index to the map attached D. C.to the Time-table for Norwegian Communica- NRS : Naturhistoriska Riksmuseum, Stocktions('Rutebok for Norge'). The Swedish namesholmare mostly listed in Svensk Ortsfcirteckning, ed. PCL : Pittioni collection, London (BML)Kungl. Generalpoststyrelsen, Kungl. Telestyrel- SMS : Stavanger Museum, Stavangersen & Kungl. Jiirnviigsstyrelsen, Orebro, 1965 SCT : Spinola collection, Museo Zoolog1ca,and/or in Svenska Turistfcireningens Atlas

-qns oaU ur snqulog peppp (tlAD reEnr;1'sernleeJ 1ec6o1olq pue cnuou-oxel Eurppe ,fq penordurr ,t1rpee1s sr qJrr{zlr Jouorlrugep oql 'uleueEqns peugep-lle^\ olq pep$lpsr snueE eErel eql 'serceds uregec ;o drqsuorleleJesolc eqt eleJlpul o1 EurdnorE cgrcedserdns e .en1l-cefqo eturrd eq] ot Eurdee>1 dg .(Sq6I) spreqcru,fq petnJeJ ulelsfs drolcegsllesun ue - sdnorEserceds Jo reqrunu e olq pepl^lp dllenpr^lpule.reueEqns ol\l rllll\ qJrqa Jo oAU .eleueE eeJrltolq plro^\ eql Jo seeq elqunq eql pogrsselr ( t qe t )uorrllrlN ,{1}ueceg 'uorleqordde lereueE ureE lorrplp snqLuog se qcns dnorE snoouoEouoq e dn Eull-ll1ds lnq leêl crroueE ol ereueEqns oql poleôlopue ror1pnJ dels e lue^\ (qZZeI ,qVI6I ,aVI6I)ô4rro{S '(Sq6IspreqclU i WAI osnoqpu€S)ereueEqns se peldope ero^{ (OSS1) errol e11eq Josuorsrllpqns JerlJeo esneceq lno clorp ol pepueloleq soureu plo IeJoAoS .eleueEqns eJoru dnllpq osle pu€ ereueEqns s.1Eon ur flreueEouloqEuorls oql pegrldue e^€q ('cle ie6E6I Iuolllld:ggOI 'LZ6I spreqcru itZel uosuC :OZ;I ,LT6lreEnry :qZZeI'7VI6I'eVI6l ô{rro>ls :VrctIIeg) sror{}ne Jo regtunu V'(tSSI) Hs.Âo{zsopel{,{q peruecuoc ,{lleurErJo seA\ qJIqA\ go 8urdnorE e'urlelrueE eleu egl q seJnleeJ uodn powq dlereulse^{\ lI 'srseq}odftl Eur4rom ]soporu e se (t tOt)FoA ,(q pecunouuu lsJU sel\ ereuoEqns olulellreJleT snquog snueE eErel eql Jo uorsrlrp VNOIIYCICISSV]CJorllne lue-sord eq1 dq uees ueulrceds - (r{sdolne) i(esues JeprdA € ul uoxel €JOAOC Ol pesn osle sr etueuoql JI) esues Je.&\oJJev E ul : (o1crr1s nsues) .s.s(esues JeA\oJJeu € uruoxel e Jeôc ol pesn osle sreureu eql JI) esuos reprl\ € ul --Sfeer(,qJg ue{} oJour rog ulr{uoudsJOTUOS e Se pesnun peureur-er seq l€ql uorleuElsepe'e'r oeureu uello8ro; e =_eJnlelouer,uou 1ec6o1ooz urEurpuels ou seq'e'r 'poqsll-qnd f1e1eu4lpe1 lou etueue _(otel nsues) '1'stunlrlqo ueurouunpnu ueruoueru€u peqslqelse ^{peerleue Eurcelder eureu ./y\eu e -uoxel u.{l'ou>I f,ueol elqecqdde lou erueu e -eJetu erdns seJleur -uorleuroJul peqs[qnd-uou -r,unôu ueruouunrqnp ueruou.UI'S UI(sue11q ul) 'tU ulsrurel ulleT 'clerredg ',[ 'Jeplour{Js - rIcS'O 'fieplontr - Ietr^t'Y 'ue4o1 - g-I'g 'uosspuelJg -,IJApuerlsEul1yg ')T 'd '>lcurrg - lA T gsJolcelloc uruueJ 'ceErreng '{l^l}g)il 'repef; 'I Ie{sgspueT -elesddn 'uossleT 'H 'pnls 'rEy -olso 'uossueqof-ques 'v 'Jp JosseJoJd -etrmaqorsuorlJelloc leuosJed'g'sellllBcol tlspe/y\S Jo sls{ eq} uI pellruo q -lWZesr^\e{r-I 'r{1uo ueEreg 'urnesnru {s}E olooz ruoJJspJoJeJ ,(q peluesordor sr ,(14ecol eql Jr pollnuosr 1EWZ ser]{ecol uet8ervuoN Jo }s11 eg] uJ {1srEo1o oZ :OWZpunT"{ltsren1un'ueuorlnlr}sul e4srEolooz -.-IV{Zsro;Eurs-leH '^{lrsren1un '1eesn141e>1srEo1oo/ -- HINZueEequed-oJ '.{ltsren1un 'urnesn141 4srEolo oZ : JWZueEreg'.(lrsrenlun'ulnesnntr {sFoIooZ -.BWZtuepJelsuIv'l1e1rsre^lun rep unesntr l seqcsrEolooz - vWZpuelod 'r!\9{-eJ) 6ueueqcsuessrd\ Jep ertuepe>lvueqcsrulod rep lnlrlsul seqcsEolooz _ >IIZfiWD punT 'uollcelloc tpelsrogoz - -IJZpu3[-ulC 'ueurlueurgH 'uorloelloc elre>lleA - CfA(VWD rueprelsulv 'uorlcelloc]Eon - VCAqeln 'ueEo1 ''llpqcreeseg ''V'C'S'n lec6oloruolug - nsnuaToT 'v 7I

Studies on Scandinavian Bumble Bees 13sections, viz. Odontobombus and Anodontobombus,based upon a few selective characters, viz.the structure of the distal edge of the intermediatebasitarsus in the female and the genitalia in themale. These sections and the actual subgenerahave been included in the keys to the Scandinavianspecies presented below.Biologically the subgenera of Bombus canbroadly be divided into two groups, viz.'pocketmakers'and 'pollen-storerso, according to theirmethod of feeding the larvae of second and succeedingbroods (Sladen l9l2). Generally speakingpocket-makers provide their larvae with solidpollen stuffed beneath them and stored in waxpockets built adjacent to the wax canopy surroundingthem. Lawae of pollen-storers are fedwith regurgitated suspension of nectar and pollen,the latter being stored in pollen-cylinders apaftfrom the brood mass. This classification breaksdown, however, in certain species (Hobbs 1964).KEYS TO THE SCANDINAVIANOF BOMBU,S LATREILLESPECIESIn bumble bees, variations in colour of the coatare remarkable, but this cannot be regarded as abasis for recognition of species. Some speciesdisplay an identical colour pattern while othersindividually exhibit a wide range in their colouring.Specific recognition must rely on other characters.The colouring is, however, a useful supplementin distinguishing the species and in particularin regarding species from a limited area as,for instance, Scandinavia, a fact which is takenadvant age of in the keys below.FemalesAntennae with twelve segments. Clypeus moderatelycovered with hairs which become sparse onthe disc. Labrum with median furrow and lateraltubercles. Robust mandibles with truncate distalmargin laterally toothed (more than laterallytoothed in -8. wurfleini Radoszkowski) and outersurface with distinct carinae. Hind tibia withcorbicle, i.e. with outer surface flattened, distallygrooved, and hairless except for anterior andposterior fringes. Mid- and hind basitarsi individuallystrong, flattened, about as long as remainingtarsi all together. Abdomen with sixvisible tergites, the hindmost pointed (truncate inB. Iapidarius (Linnaeus)). Well-defined coat invarious colour patterns.The workers are far more variable and oftenlack the diagnostic characters of the females.They can nevertheless in general be identified bythe key. Exceptions concern above all the subgenusBombu^s s. s., where the worker caste oftendisplays too diffuse characters for a reliable specificdetermination.1. Mandible without basal keel (Figs.78, 8A, 10A, 15, 164).Mid-basitarsuswith posterior distal angle obtuse,more or less rounded (Fig. lzc)(more or less acute in B. wurfleiniRadoszkowski (Fig. 7C)). . . section Anodontobombus 2Mandible with basal keel (Figs. 20,228). Mid-basitarsus with posteriordistal angle spined ' (Fig. 178) orpointed section OContobombus 182. Malar space about half the distalwidth (Fig. 7 A). Mandible stronglybent, distally strongly toothed (Fig.7B). Mid-basitarsus with distal posteriorangle acute (Fig. 7C)Alpigenobombus Skorikov1 specieS,B. wurfleini Radoszkowski (p. 25)Malar space markedly longer thanhalf the distal width. Mandible notstrongly bent, distally only laterallytoothed (Figs. 8A, 10A, 15, 164).Mid-basitarsus with distal posteriorangle right-rounded (Fig. lzc) 33. Malar space as long as distal widthor nearly. Mandible without sulcusobliquus, with indistinct incisura lateralis(Fig. 8A). Ocellar-orbital fieldas in Fig. 8B. Ocelli transected bysupra-orbital line (Figs. 8C-D). Hindbasitarsus almost parallel-sided (Fig.8E) ... Kallobombus DallaTorre1 species, B. soroeensis (Fabricius) (p. 31)2. Bumble Bees.

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Studies on Scandinavian Bumble Bees 15v,"il@12xd il,,"u,,.Figs. 7-16. Section Anodontobombas. Fig. 7: B. wurfleini Radoszkowski Q. A: Lateral view of lower part ofhead; B: Right mandible; C: Right mid-basitarsus. Fig.8: B. soroeensis (Fabricius) g. A: Rieht mandible;B: Ocellar-orbital field; C-D: Position of ocelli, I and g respectively; E: Right hind basitarsus. Fig. 9:ll.hyperboreus Schonherr ?. A:'Distal inner posterior process of hind tibia; B: Right hind basitarsus. Fig. l0:B. alpinus (Linnaeus)?. A: Right mandible; B: Distal inner posterior process of hind tibia. Fig. ll: B. iporadicusQ.ocellar-orbital field. Fig. 12:8. terrestris (Linnaeus) Q. A: I-abrum; B: Ocellar-orbitalfield; C: Righimidbasitarsus;D: Right hind basitarsus; E: Sting, a dorsal end ofsting; b inner thickening ofsting sheath. Fig. 13:B' lucorum (Linnaeus) 9. A: Ocellar-orbital field; B : Sting. Fig. 14: B. magnus Vogt g. Ocellbr-orbital field.Fig. 15: B. lapidarius (Linnaeus) 9. Rieht mandible. Fig. 16: B. lapponicus (Fabricius) Q. A: Right mandible;B: Distal inner posterior process of hind tibia.ocellus to eye. Yellow collar broader,extending part way down episternum.... ...B.magnusVogt(p. 46)12. Mandible with distinct sulcus obliquusand poorly to moderately developedincisura lateralis (Fig. 15).Hindmost tergites reddish or brightferruginous ....Mandible with inconspicuous sulqusobliquus and moderate to well developedincisura lateralis (Fig. 16,.{).l:' ::'::1 i'i: "'uz::;;ffi J+:lt13t413. Malar space about as long as distalwidth. Labral furrow well defined, inwidthless than I 13 labral width.Mandible as in Fig. 15. Tu with centraleminence in front of truncate apex(indistinct or lacking in workers)Melanobombus Dalla Torre1 species, B. Iapidarius (Linnaeus) (p. 89)Malar space shorter than distalwidth. Labral furrow moderate, inwidth about 113labral width. T6 withindication of impressed median longitudinalline, distally not truncate. .

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Studies on Scandinavian Bumble Bees 17aoooo r(17n ll.::: o D17a 18a 19@21Figs. 17-22. Section Odontobombus. Fig. 17: B. hortorum(Linnaeus)?. A: Position of ocelli; B: Rightmid-basitarsus. Fig. 18: ,8. consobrinus Dahlbom 9.A: Lateral view of lower part of head; B: Position ofocelli. Fig. 19: B. distinguendus Morawitz g. Lateralview of lower part of head. Fig. 20: B. veteranus(Fabricius)?. Right mandible. Fig. 2l: B. pascuorum(Scopoli)?. Labrum. Fig. 22: B. sylvarum (Linnaeus)9. A: Labrum; B: Right mandible.well-defined unpunctured areaand thepunctured band extremely narrow. . .. . Rhodobombus Dalla,TorreI species, B. pomorurn (Panzer) (p. 189)(8. pomorum has pile on Tr-u ferruginousand coat otherwise black).Malar space less than l/2 times thedistal width (nearly so in B. pascuorum).Ocellar-orbital field with welldefinedunpunctured area of moderatesize and the punctured bandrather wide. If hindmost tergites ferruginous-haired,this applies to T*-uor Tn-u not T, .22-nThoracobombus Dalla Torre 2424. Disc of clypeus distinctly flattened. :Mandible slender, elongated (Fig.t20).Coat pale greyish-yellow with exceptionof broad not parallel-sidedinteralar band and variable admixtureof black bristles on Tr-uB. t,eteranus (Fabricius) (p. 179)Disc of clypeus more or less convex.Mandible not elcifigated (Fig. 228).\U,wOther colour pattern. If with interalatband,Tn-uferruginous .... 2525. Malar space nearly l/z times distalwidth. Labral furrow widened towardsthe front (Fig. 2l). Coat shaggy.Dorsum of thorax and at leasthindmost tergites yellowish-brown ororange-brown .B.pascuorum (Scopoli) (p. 154)4 subspeciesMalar space shorter . Labral furrowwidened towards the base or almostparallel-sided (Fig. 224). Tu with ferruginousor black hairs 2626. Thorax dorsally black or with interalarband. Tn-u (or n-u) ferruginous. . 27Thorax dorsally yellowish-brownor orange-brown. T*-, not ferruginous,T6 black-haired..... . .. 2827 . Labral furrow as in Fig. 22A. Lamellaof labrum with knifelike edge, smooth,shiny. Mandible as in Fig. 22B. Surfaceof Tn-u smooth, shining, withsparse coarse puncturing. Broadinteralar band, thorax otherwisepale yellowish-grey (melanic individualswith pattern similar toB. ruderarius common). . . .8. sylvarum (Linnaeus) (p.172)Labral lamella with distal marginthickened, alutaceous, dull. Surfaceof Tn-u chagrinated, densely punctured.Thorax, Tr-u black. Corbicularfringes and T*- u bright or dull ferruginousB. ruderarius (Mtiller) (p. 169)28. Surface in front of lateral ocelli ratherdensely punctured. Distance fromlateral ocellus to preoccipital ridgemarkedly longer than distance betweenlateral ocellus and eye. Lateralhairs of TB arising from pustules.l: ':'1 ::lfH1,::"::::i'ii'I::[Jb Ia5)2 subspeciesSurface in front of lateral ocelliwith disperse, coarse punctures. Distancefrom lateral ocellus to preoccipitalridge barely or not longerthan distance between lateral ocellusand eye. Lateral hairs of T, arising

pueq fflerelul'J-A97, 'sEIC ur se€rlelruoE pue 8lS 'V9Z 'Elg ur se t-'V 'l(tOf 'd) (dqr121) sncp,to 'g' ' snour8nrroJpeounouord o1 eled Joqler seltErel ;oJepuretuoJ 'Icelg ,IJo yed, JorJoluelseel te pue 'I '(qSZ 'Elg) qleeueqruo4 uees elleslon Jo ged JorJelsod'J-11gZ 'sBIC ur se €rle}rueE pue 819'(VSZ 'EI{) q}pyvt 1etslp seull} zAE ueq}reEuol snsJelrseq pulH 'snoecelnle,{1Euor}s erqlt puFI Jo eceJrns relno-ur pecnpo"ld sn1,(lsouoE(96'd) (sneeuurl) snuldlo '9"'' ' ,g;EV,JZV ,W ,g;OV .q6g .sErg :CAV-CVVur se' ' ' {oelq esrl\Jeq1o 'snout8nr;eg t -, Leqelrueg'(V6V-yVV'yZV'V6t'sElg)'(qVZ '6lg) qleeueq tuo4 uees elleslo^' Jo l"led rorrelsod 'J-1VZ 'sErg ur seeqetrue8 pue tlS '(VVT,'E!i) ,(peeu roglpli!\ Ielslp uBrlt reEuol setull %tlnoqs snsrelrseq pulH 'snoecelnle louro ,{11qEr1s erqll puq Jo eceJrns re}no '9' ' ' snoutEnrre; ro qsrlrqa '{celq t -, L'snour8nrreJ lou*t '(VtZ 'fJ:gl'sEtg)pocnpord leqzlrouros urEreul lelsrp JooJluoc qlll\ xede le eleounrlqns 819'snourEnrreJ9,-*I '(ASZ-gVZ 'sElg) xede spreÂo1 Eurredel EIS 'Sg 's's .sr?qLuog r{sl}lqA\se1tEre1 lsorupun{ Jo esoq}'zlrogef, L Jo sJrer4 elqrsrl }seo[ le 're11o3'J6Z-)SZ .sEIC ur se erlelrue1l.qtpl,\\sI€lslp ueQl reuor{s snsre}rs?q purrl Joleql 'Qlptzn lelslp 3ql ecIAU ueql sselelqll puF{ go seEulrC 'q}pr^\ Ie}srp uurf}reEuol lou Jo .(1ereq eceds Jeleryi:llr"{S szswoqourdtv' gLZ 'J97,-JVT, 'sEIC ur see{etrueD'q}pl1Y\ Ie}slp errrlcedser seEuol se ecrlq lseel le snsrelrseq puF{pue elqll puFI go seEulrC 'qlpr^\ Ielsrpue{} reEuol ,{1pe>peul eceds mVW'V' ' 'xedeV1e pe{ooqlou Jo ,{lsnoncrdsuocur oAIeA srued'(ySZ 'yVT, 'sElg) eseq oqt spre^\olpe^\oJJeu qJnu lou snsJelrseq pulH(tE 'd) (snrcrrqeg) srsuaaotos 'g 'sarceds 1eJrol €lleQ snqwoqoilDx t{sllF{/y\seltErel lsorupuF{ 'A,\olled .-tI 'rego3'spJel\lno pe>looq ellel srued;o xede:C-C6Z 'sEIC ur s€ €rlelrueE pue 819'(A7EIDeseq eql sprelv\ol pe^\or-reu .(llcurlsrp snsrelrseq pulH '(VtZ'Etg) r{}pl,!\ Ie}slp rleq} ocrÛ }noqe,tllenprl1pq sr-sv 'Euo1 eeuuoluy 'g0I (gEE 'JZE 'JIe 'CIgg 'sElg) spre^\urpeurnl s>looq lelslp qtl,\\ o^[?n, slued'(CeZ'JgZ'gLZ'JTZ-JVT'qETE'sEtDsprel(tno peurnl $looq qll^\Jo xede le pe{ooq }ou eAIeA srued 'ZLI snqwoqotuopo uorlces' 'eulds Jo €lletuel eflml e olur spJel\os ^(1reeu Jo r{}pll\ Ie}slp Jreq} eJrAU,{llenplllpq 8r-ev 'Euo1 oeuue}uv7' "nT??::":?z uorrses'ssecoJd pedeqs-roEuq Jo pelurod-geuls e olur pecnp-ord lsotu le sprel\q sn1,{lsouoE :ggg,J7,8.J .CTO€,J6Z,J8Z,g.LZ,)92I €'yVZ'q.EZ'sEIC ur se eqelrue11'(V0€'y9Z'yEZ 'sEIC (fqrU) snununlprg',to4rro4g snqwoqouldlV'(snrcrrq€jl)swuaaotos 'g :suolldecxe) (VSE 'yVE,YT,E .VIT ,Y67 ,YBZ 'SETD q}PIA\1elslp Jleql ecr^\} ueql ssol ^,{1pe4reuldllenpl^lpul er-ev '1roqs eeuueluv'I'suJolled rnoloc snorJen rrr leoo peugep-lleA|pepunor,(lrorrelsocf lsorupuH eql'se1lEre1elqrsrl ueles qlll\ ueuopqv'rsJel Sulureuro.rue{l reEuol Jo Euol se'peleEuole 'peue1}noqe-leg ,{11enpr^lpq rsre}rseq puFI pue -pml 'srregluegtunJep ssel Jo eJoru qlll\ peJoloc dlelerepotuJo ssolJreq'xenuoJ Jo pouelleg eceJJns Jolno eql'seEurrg rorrelsod pue rorJolue glyvr elqll pulH'(r4srro>lzsopea rutagftnu 'g ul o]ecrnJrrl) elmJnJ-lq urEruur lelslp enbrlqo qll,!\ solqrpueu >leol$,'uelo runJqeT'dlletsry 'slr€r1 ldecxe r{lr1y\ poJoloc.(lesuep snedr(13 'slueurEes ueeglr{} Htyrr eeuueluvsalDw(0Et 'd) re8tnI sryfanq 'g' 'sellslrq {cBIq }noqll^\ ro rl}11tr o-'I'1se4rep eqt sr .I r{cpll\ Jo sepeqsSnoIJeA Jo uA\oJq-qstznolert n-tI'sernlcundEurcseleoc esJ€oJ ruo4uaToT'v8I

Studies on Scandinavian Bumble Bees t9(i r423cFies.23-27. Section Anodontobombzs. Fig. 23: B. soroeensr's (Fabricius)J. A: Antennal segments 3-8; B:Right hind basitarsus; C: Sternite 8; D: Genitalia. a penis valve; b volsella; c gonostylus; d gonocoxite,Fig. ?A: B. alpinus (Linnaeus)d. A: Right hind basitarsus; B: Right half of sternite 8; C: Right half of genitalia;D: Ventral view of posterior part of volsella. Fig.25: B. arcticus Kirbyd. A: Right hind basitarsus;B : Right half of sternite 8 ; C: Posterior right half of genitalia; D : Ventral view of posterior part of volsella.Fig.26: B. balteatus DahlbomJ. A: Antennal segments 3-8; B: Right half of sternite 8; C: Right half ofgenitalia. Fig. 27 t B. hyperboreus SchcinherrJ. A: Right half of sternite 8; B: Right half of genitalia.(except in melanic specimens). T u-,ferruginous, dull yellow or whitish. ......... .....8. balteatzs Dahlbom (p. 105)St, and genitalia as in Figs .27 A-B .Interalar band. Tr-, black . . . . . . . .. B. hyperboreus Schonherr (p. 114)8. A, slightly or not longer than Aa (Fig.28A). Wings elongated, strongly infuscate.St, and genitalia as in Figs.288-C; gonocoxite regularly convergingtowards apex; apex of volsellaas in Fig. 28D .8. sporadicus Nylander (p. 50)A, markedly longer than An (Fig.29A). Wings not elongated, moderatelyinfuscate. Gonocoxite not regularlyconverging towards apex, slightlyimpressed inside. . . . 99. Ar-a irS in Fig. 29A. Supra-orbitalline touching or just transecting theocelli dorsally. St r and genitaliaas in Figs. 29B,-C. Apex of volsellaas in Fig. 29D. Coat rather even.Colour pattern distinct. Yellow hairs,being deep yellow, restricted to collar:::i': *:::Tiii,i;,,i r,,;;";;,; 653)

'EIC) reEuol'ro nV ol lenbesy 'el,(p-uoc Jelnqrpueur rouelsod A\oleq Josprel\ol pelcerrp efe go eEpe Ieluo tC .Llsercedsqns Z(s|d) l:l,:iro*) .'.n.'::oddo1's'''znoled Jo snoulEnrre;t -, L'snourEnJJeJ t-.I .{JBIq xeJoq}Jo unsJop elorl^\ Jo pueg JeleJelurue reqll1'JLE 'Etg ur se e^lel srued'g,LE 'Etg ur se snlflsouog .y LE.Elgur se tls 'qlpp\ lseleer8 rreql ecrlUls€el te fllenpr^lpu! snsrelrseq puFIpue elqll puFI go seEur4 rorretsod(ZS'd) (sneeuufD wnnrutd 'g' 'snour8nJJeJ t -, L'rvro11ert,relloJ 'J9€ 'Elg ur se ollel sruod'g9E 'Elg ur se sn1,{lsouog .V9€ .Etgur se t lS 'qlpp\ lseleer8 Jror{} eJrA\}ueql ssel ,{11enpr^lpu} snsre}rseq puFIpu€ elqll pug Jo seEurrg rorrelsod .9I(tq'd) Ereqpe6 snrulnSmc'g'' . . . . . .' ' 'pueq relerelq ue Eunurog.(11euors€cco'cslp eql uo sJrer1 {celqJo eJnlxrupe elqerJe^ qlr^\ xeroql Jor,unsJoql '61St 'Elg ur se eAIeA srued'JSg 'Elg ur se sn1,{lsouog .ggt .EICur se tls 'luetuEes eql Jo qlpll\ lso-1eerE e{} Eurpeocxe seEurr; rouelsodJo srreq lse8uol qlr^t snsrelrseq pulH'(VSg 'Elg) nV ueql regor{s dlpe4reru'oV ueql re8uol lou ro ,t11qE[s rV(99 'd) wn"toudrhl 'g' ' ' ' ' '1t:n""r1) ' 'u^{\oJg-qsr^\oller{ ile xeJoq}Jo runsJoq 'gy1 'Elg ur se ellelsrued 'JVt, 'Elg ur se sn1.,(lsouoC'14VE'Elg ur se tls 'lueurEes eql Jor{lpll\ }se}eerE se Euol w }noqe seEuugJorJelsod Jo sJr€rl lseEuol qlllr snsJul-lseq pulH '(Vlg 'EIC) reEuol ro nVo1 'oV lenbe ue{l reEuol ,{1pe>1retu *v .S IS I pueq rcIerclur ue Eururrog z{11en1uele'cslp egl uo sJrer4 {celq Jo eJn}xru-pe elqerJsA rltla\ Jo rrA\oJq-qsrA\o11e,{11e xerog] Jo r,unsroq '(CSE 'qVE'sEIC) qleeueq pegtool e^1en srued(09'd) (,(qrry) snllauol' '€[' ' 'pueq releretqpepls-leller€d '(AEe 'EIC) qleeuoqr{}ool Jo uol}ecrptll lsoru 1B rI}rA\ eAIeA9tVT(os'd)Gz'd)7,7'sElD xede W pepurEreure Jo elecunJlelleslo^ 'eseq eql spre^\ol pea\oJJ?uqcntu tou snsrelrseq pulH -(VZt,.VIt's8l$reEuol Jo nV ol lenbe ty(gS'd) (,{qqX) snuownlpc .g osercods I€ IIIsrued 'J tt'Elg ur se snlztlsouog'g -Vtt 'sEIC ur se eqelrueE pue tls 'tl::: :::'::1':"'l:{ l: ^"'re't'-nrt{sl}lrl^\ ,-nI 'EI(sneeuurl) snrnpdol 'g 'serceds IOJJOI UIIeCI snqwoqouqaw''' .qsrppeJ t-rl'J-1ZE'sErg ul se e1el-rue8 pue 'lS'(VZE'El$ nV se Euol selnoqe 8y 'elecrnJlq elqpuel4tr .qlpl/KIelslp se Euol se lnoqe eceds relelNDISAlo{zsopeU rutagf"tnlr 'g 'selceds 1Ao>lrJo{S snqwoqoua&d1y. .snourEnr-roJ t-rl'C-flIt 'sEIC ur se e{e}ru-eE pue'lS '(VI€ 'EIc) nv ue{} re8uol'V 'elsrrnJlrl elqlpuel tr'Vlplzlr lelslp:::l :':::: jT:l:T eceds rcrerN' '(rztzr'JrEii:f.'.,.qT:?::?*,iln)' ' ''xsde 1e enbrlqoe11eslo^ : q-COg 'sElC ur se er1e1rueEpu€ tls '(gOE 'EIC) eseq oql spr€a\-ol pe^\oJJeu lsrl&\eIuos snsrclrseqpulH '(Vgt .EIC) nv uet{} reuorls rV .[ Iefiuc snquoqo*(4:1t:t ' "(gtg 'Elg) ssecord lueprle ouglll\ relnEueul sr r{cpll\ 'sn1.,t1souoEpuofeq Eurqceer ,{1preq €fleslo1...(CZE'JI€'CTOT'sElg) eseq sll le ssecord lu€reqnJl-ord ileuls e seq eprsur qcplr!\ .sn[-,(lsouoE puodeq JeJ Eurqceer ulleslo1 .0[' (tan"tocry g tuo4 peqsrnEullspeq ,(lurupoo gU/K louu€c snu&ow .g)(9V'd) lEoA snuSotu'g' "''''''''''''(O?'d)wnrc)ry'g''''''''' .. ..lt:r:""fD ' ' 'sJreq qsrderE go eJnlxnupe ueqlllr uetJo pue "I 'urn11n1ncs'runu-relsrde uo'ecug:goelld ur sJreq zlro11e.(Jo ernlxnupe elqerre^ Ieuorllppe qtl^\esnJlp Jo sl4sailat 'g Kq, s€ lcurlslproqlre urellud rnoloJ 'epeqs 11npeled e Jo Jo aollez( uoulol sJreg A\ol-le1 'dEEeqs ssel Jo eJoru leoJ .rllecoEurlcosueJl,(1ree1c eu11 Ie]rqJo-erdngtuilloT'Y oz

Studies on Scandinavian Bumble Bees2l33cftI'ilwHt'34ch35c37er37c - \--HH H3HH^H2Ba3on34n 35n30eFigs. 28-37. Section Anodontobombzs. Fig. 28:., B. sporadiczs NylanderJ. A: Antennal segments 3-8; B: Righthalf of sternite 8; C: Right half of genitalia; D: Ventral view of distal part of volsella. Fig. 29: B. terrestris(Linnaeus)J. A: Antennal segments 3-8; B: fught half of sternite 8; C: Right half of genitalia; D: Ventralview of distal part of volsella. Fig. 30: B. cullumanus (Kirby)6. A: Antennal segments 3-8; B: Right hindbasitarsus; C: Righthalf of sternite 8; D: Righthalf of genitalia. Fig.31: B.wurfleini RadoszkowskiJ. A:Antennal segments 3-8; B: Righthalf of sternite 8; C: Righthalf of genitalia. Fie.32:B.lapidarius(Lnnaeus)d.A: Antennal segments 3-8; B: Right half of sternite 8; C: Right half of genitalia. Fig. 33: B. jonellus (Kirby)$.A: Right half of sternite 8; B: Rieht half of genitalia; C: Rieht gonostylus, volsella; D: Lateral view of rightpenis valve. Fig. 34: B. hypnorum (Linnaeus)d. A: Antennal segments 3-8; B: Right half of sternite 8; C: Rightgonostylus, volsella; D: Lateral view of right penis valve. Fig. 35: B. cingulalas Wahlberg j. A: Antennalsegments 3-8; B: Right half of sternite 8; C: Right gonostylus, volsella; D: Lateral view of right penis valve.Fig. 36: B, pratorum (Linnaeus)d. A: Right half of sternite 8; B: Right gonostylus, volsella; C: Lateral viewof right penis valve. Fig. 37: B. Iapponicus (Fabricius) $. A: Right half of sternite 8; B: Right gonostylus,volsella; C: Lateral view of right penis valve.

'ezrleir srued;o sseco.rd lecrdee;4 :c :sJlsllueE Jo Jpq 1q31.r rorrelsod:g '8 etlurels go ;1eq lq8rg :V '9 (sneeuurl) snauottaqqns 'g iEl '3rg 'e1e1rue6Jo JIeg fq8pl :3 !g elrurslsJo JI"q tqANIlI :g :8-€ slueuEes leuuetuv :y'Bzlrme.Io141snpuanSm$lp 'g :ZV'E!t'BIIelrueF Jo JI?r{ lrlSuJorrelsod 'P(rezued) utntoruod'g:IV'4C '€lleqo^ lqErr;o lred 1e1srp Jo ^\er^ I"sro([ :q lelrxocouo8 1q3rrJo U?d rolretsod Jo ,rer^ lerele-I:3 ie1e1rue8;o geq lq8rr rorrelsod :g !g e11ure1s go gleq fq8pl :V .Puoq-Ir4eqsnuqqosuoi'g :01 '8rC 'elleqo^ 1q3rr;o ged 1e1sp go ^\er^ I€sJoC :d :elxorouo8 lq8rr go ged rouelsodJo ^\erl lerepl:g :ell?tlueE go gyeq 1q8p1 :q !g e1u;e1s go geq rqElg :3 lerqr1 pulq 1q3p1 :g :g-g Eueur-Ees Ieuueluv :y 'B(sneeuurl) unto\.torl'g:6€ 'EIg 'elleslo^ tqEu Jo ged 1e1srp Jo ^\eF I"sroq :J :g elrurelsgo geq lqEpg :g ierqrl pulq lqSN :V 'P(srucuq?g) swotapnt'g :g€ '8lC 'snquoqoruopo uopres'et-g€ 'sEIdv68 a0? r68c88 s683ZV//v)'ool// //368o68ssel ro eroru elleslo^ Jo dll e{Il-}ooq:7 /44D:(EOS 'Elg) peugep ,(pee1c snff,ls'--ouoE Jo suotsserdep lerslel :61-J6E'sEIC uI se eIIelIueE pue 'tS 'g6E 'Elg6luI se €lgll pulH 'V6E 'EIC ul se '-'V '6IsI.. . ueqmIeletulJl'(g0E'Etg) sluetu-Ees eql Jo qlpp\ lseleer8 ue{} reEuol,(1pe4reur elqll puF{ Jo seEutrs iutE-J€ur luequnoep eql Eurpeecxe sellslJqqll^\ elqll puFI Jo u6retu IelslCI(SEI 'd) (snrcuqeg) snruDpn't'g ' 'qsltlt{^\, -oI 'pu€qJ€lerelul pepls-lellered0z8I'O8t 'Etg ol relltuls ssel ro orour elles-lo^ Jo dlt e{rl-]oofl 'g8t 'EIc uI se'lS '(VSg 'Elg) tueu8es eqt Jo qlpr^\lseleerE ueql reEuol lou Jo ,(1ereq€lqlt pulq Jo seEutr; lutEreur luoq-Iunoep eq] Eurpeecxe lou Jo ,(1ereq::l:T.:l,lputq ro setlslls lelstcl'8I(vev-vw 'YZv'sEtg) reUotIS ro nV o1 lenbetV 'el,(p-uoc Jelnqryueur rotrelsod Jo luo4ur pelceJrp de.,{e Jo e3pe Ie}uoJCeJJoI e11eq snqwoqn&ayTy''''''ggt 'CI6g 'sEI{ uI se e{e}IueD '(VAgua4o7 'v zz

Studies on Scandinavian Bumble Bees23similar to Fig. 39F. Interalar band(except in strongly melanic forms).Tn-u whitish . . . B.Itortorum(Linnaeus)St, and genitalia as in Figs. 40A-B ;lateral depressions of gono9tylte poorlydefined (Fig. 40C); boot-like tip ofvolsella more or less as in Fig. 40D.Whole dorsum of thorax yellowishbrown.Tn-u greyish with admixtureof black hairs. . .8. consobrinus Dahlbom20. AB as long as Au or nearly (exception:markedly shorter in B. pomorunt).Ar-r, individually parallel-sided ornearly (Fig. 424). Outer surface ofhind tibia sparsely to densely coveredwith short hairs. Genitalia as in Figs.42C,438As markedly shorter than Ar, atleast Ar-r, more or less swollen underneath(Figs. 441.-49,4'). Genitalia asin Figs. 44C-49CThoracobombus Dalla Torre21. Mandible without a beard. A, markedlyshorter than Au. Hind tibia withouter surface convex, densely coveredwith short hairs. Gonostylus transverse,outer edge curving obliquelyinwards and ending in a stout acuteprocess; volsella projecting well be-(p. 1le); ;'.y "l:';#::,""'3:J:;@ I 8e)Mandible with well-developedbeard. A, av long as Au or nearly(Fig. 42A). C ,: surface of hind tibiawith distal half flattened or slightlyconcave, rather sparsely covered withshort hairs. Genitalia as in Figs. 42C,438. Gonostylus transverse, outeredge not curving obliquely inwards@.12a)lateral swellings. St, and genitalia asin Figs . 428-C; preapical process ofpenis valve tridentate, usually onlyone tooth strongly projecting. Pare;ryr/-2and not ending in acute process; volsellamoderately projecting beyondgonostylus, broad, truncate distallywith a narrow process projecting inwards. . Subterraneobombus Vogt 2222. A"-s €rS in Fig. 42A. St6 with distal2lallel-sided interalar band. T ? black-haired, coat otherwise dorsally oliveyellow.. . . B. distinguendus Morawitz (p.182)Stu without lateral swellings. St,and genitalia as in Figs . 43A-B; preapicalprocess of penis valve tri- orquadridentate with lateral teethstrongly projecting (Fig. 43C) or bidentatewith one strongly projectingtooth. Interalar band posteriorly v-shaped. Other colour pattern. . . . .8 . subterraneus (Linnaeus) (p. 186)23. Subapical process of volsella almostparallel-sided, but suddenly dilatedand truncateat apex (Figs. 44C-46C).Dorsal of thorax all black or interalarband present 24Subapical process of volsella dif-23 ferently shaped, never parallel-sidedand truncate at apex. Dorsal of thoraxneither all black (except in rare melanicforms) nor with interalar band 2624. Ar-s &s in Fig. 441^. St, and genitaliaas in Figs. 448-C; subapical processof volsella markedly longer than distalwidth; penis valve toothed underneathyond gonostylus, moderately broad(Fig. 44D). Dorsal of thoraxexcept expanded considerably in-black, T n-, ferruginouswards at the end (Fig. 41). .8. ruderarius Miiller (p. 169)Genitalia as in Figs. 45C-46C;subapical process of volsella barely ornot longer than distal width 2525. As barely longer than An, Ar-r,swollen beneath (Fig. 45A). St' andgenitalia as in Figs. 45B-C; penisvalve without dlstinct tooth underneath(Fig. 45D). Broad interalarband i Tr black, Tn- u ferruginous,otherwise pale yellowish-grey (melanicform with pattern as in B. ruderariuslocally more common than the typicalform) B. sylvarum (Linnaeus) (p.172)Ar markedly longer than Ao, Au-',strongly swollen beneath (Fig. 464).St, genitaha as in Figs. 46F.-C. Inter-

GVt'd)LT,pu€ Eurpelord ,{1Euo4s e[[eslo^ JourEreur lelslp : 'sEIC J-gOt ur s? e\el-ruoE pue 8lS 'y6V'EIJ ul se t-*V(t".t:uulT) tanto)snta 'g' '' ' 'r\\ol1ef n-tl 'umolq11np-eEuero Jo ul\oJq-qsrzlroller( xeJoqlJo tunsJoq 'Egt 'Etg ur se elle^srued '(ASl 'Etg) qlool repEuerrllecrdeqns peorq e pue ssecord Eurlcel-ord qcntu lou peorq e qll^\ elleslo^Jo urEreru lels1p : 'sEICC-ggl ur se€rlelrueE pue'lS 'Vgt'8lC q se '-ry 'LZ' '(CeV-Cgt 'sEIC) rlleeuoqeleJJes 'xodelou le pe>looq eAIeAsruod '(VeV-Vgt 'sEl$ rll€eueq uel-[ol\s ,(lelerepotu o] ,(ltqEqs sr-svsercedsqns ?@St 'ol (tlldocg) tanton)sod 'g' ''ul\oJq-eEuero Jo ul\oJq-qsr-zno11e,( se1lEra1 lerlseE lsorupultl lseelle pue xeroql Jo runsroc'qleeueqeleJJes 'eldurs ellel sruod :I-Aw'sEIC ur se eqelrueE pue ']S '(Vw'Etg) qleeueq uego^rs fpuorls rr-s1.7 '92G4'd) (snrcrrqeg) snuo.rata^ '9" ",terE-rlsrly\Ol1er( eled leoc esr^\Jorllo' se11s rrqtcelq Jo ernlxrupe qll^\ ,{gensn t -, L'pecunouord ssel Jo oJour pueq tele'oAI€A srued lqErr ;o A\erA IEre]B'I : a : elleslo^ lJeI Jo .rerl l?rlue1 : q i e4zlrue8go geg lq8pl : j i g elu.re1s go geq 1g3p1 : g : g-€ slueu8es I"urroluy : y .B -re3ry1 s1lurnq .g : .Brg 'eqen67srued lqEu Jo ^\er^ Ierels'I :A :?lleqo^ tJel Jo ^rer^ IBrlueA :O :ellelrue8 go gleq fqElg :C :g etrurels Jo JIsgtqEpl :g ig-g slueur8es Ieuueluv :y 'p(sneeuur-I) unn)snu'g :gt .8lC .€JI€lrueAJo JIerI lgEI)I :J :g elrurelsJo JIeg rq8Jg :g !g-g slueu6es Ieuuetuv :y 'B(1odoc5) wntoncsod.g:LV-S:Hd.erl?true8 Jo JIer{ lq8r-a :3:8 elru.rots Jo Jleq tq8pl : g : g-€ sluew8es leuueluv : y .B(snrcuqe g) snuotalat ,g :gt .4C.e,r1e,r srued lg8lr Jo^\ehI?rolEf :q luqulrueEJoJIegfqEN:O:g elrurelsJoJleqfq6Ju:g ig-[ slueurEes I?uueluv:y.f,(sneeu-ur.1) wntod,1,(s'€.:St '8lJ 'eirlezr srued lqErr go aer^ IErel?-I :q !e11e1rue8 go geg 1qE1g :C :g elruJets Jo J:lEqfqEry :g :8-g slueruEes Ieuueluv :V 'P (rellnry) snttoopnt .g :W .EId,.snquoqoruopo uol1ces .6t-W .fitdavvo6?TY-reo8vgryv6?\'ili,l I i'ftV87 VLr S9VVI vvvvuaryoT'Y vz

with a bidentate subapical process(Fig. 49D); penis valve as in Fig. 498.Dorsum of thorax yellowish-brown,Tr-u yellowish of various shade of::::l l: :: 'T i:::;:'nu*;r,,i,i,!,, (p 13e)SUBGENUSSKORIKOVALPIGENOBOMBUSAlpigenobombus Skorikov, l9l4a, p. 128, typespeciesBombus mastrucatus Gerstaecker, 1869.(Mastrwcatobombus Krtiger, 1917, pp. 62, 66,type-species Bombus mastrucatus Gerstaecker.Monobasic.)Nomenclatural, taxonomic remarks.Sandhouse (1 943) stated that the type of Alpigenobombusis' Bombus lefebvrei Lepeletier, 1 836 "( :B. mnstrucatus auct.)." (Original designation).' ,8.lefebvrei Lepeletier, 1836, holotype ? MNP, iS,however, now revised to ^8. pomorum (Panzer),1802-05, by Tkalcu (1969) and this species belongsto the subgenus Rftodobombus Dalla Torre, 1880.In his meagre description, Skorikov (1914a) wrotethat he had gathered what he called all the endemicforms of '8. lefebvrei Lep. ( - B. mastrucatusauct.)' occurring in the mountain fauna of Eurasiainto one genus or subgenus, Alpigenobombus. AsSkorikov obviously meant B. mastrucatus, thistaxon is retained as type-species for Alpigeno'bombus in order to serve stability and uniformityof nomenclature.According to Tkalcu (1969), B. mastrucatus is,however, a European subspecies of B. wurfleiniRadoszkowski, 1859. Tkalcu examined the holotypeof the latter but did not trace the type seriesof .8. mastrucatus IZB, which unfortunately bythen was at my disposal. The series consists of29?,22??, 14 66 of which a male, selected as lectotypeby ffieo is labelled as follows: (1) Ober-Bayern, Kreuth Gerstaecker S (printed) ; (2) mastrucatusGerst. * d oper. Kreuth (handwritten?original); (3) red label marked parutype; (4) B.mastrucatus Gerst. d lectotype design. A. Laken1965. The specimen was chosen because of theasterisk in the text of the second label as 'DieseSternchen sind in unserer Sammlung zuverldssi-Studies on Scandinavian Bumble Bees 25gere Indikatoren ftir Typen als die roten Etiketten'(Konigsmann in litt. 1966). I have not seenthe holotype (a male) of -8. wurfleini, and havetherefore not been able to compare this specimenwith the type material of ,8. mastrucatus to confirmthe conspecificity of the taxa. Radoszkowski(1884, p. 66) emphasizes the structural similarityin the males of -8. wurfleini and B. mastrucatus,but maintains a specific difference in the females.However, he indicates that B. mastrucatus mightbe a variant of ,8. wurfleini - B. alpigenus Morawitz.Their conspecificity has been recognized inthe past (Morawitz 1881 , p. 71 ; Handlirsch 1888;Friese & Wagner 1909; Kruseman 1960; Reinig1965, p. ll7; etc.), but the priority of B. wurfleinihas been ignored and the species treated as B.mastrucatus. The nominate form of B. wurfleinihas a colouring of the coat completely differentfrom that of ,8. mastrucatus and the taxa have,moreover, an allopatric distribution. The statementof Tkalcu (1969) is no doubt correct, i.e.B. mastrucatus Gerstaecker - B. wurfleini mastrucatusGerstaecker.B. wurfleini mastrucatus is the only Scandinavianrepresentative of the subgenus.BOMBUS ( ALPIGENOBOMBUS )W U RFLEI N I RADOSZKOWSKIBombus wurfleini Radoszkowski, 1859, p. 482No. 4, holotype d ZIK, type area Caucasus(Tkalcu 1969).(Bombus levebvrei auctt. nec Lepeletier, 1836;Strand 1901 ; Brinck & Wingstrand 1949. Bombusmastrucatus Gerstaecker, 1869, p. 326 No. 19,lectotype d IZB! hereby selected cf. above. Typeloc. Germany: Ober-Bayern : Kreuth; SparreSchneider 1898, 1906, p. 183, 1909, 1918; Lie-Pettersen 1901, 1905, 1907; Friese 1902; Aurivillius1903; Bengtsson 1904, 1908; Ringdahl 1915;Gaunitz 1929; Meidell 1934a, 1946;Lsken 1949,1950, 1958a, 1960,1966c; Brinck 1951; Kruseman1959; Elfving 1960, 1968 ; Ander 1963, 1965.Bombus brevigenq Thomson, 1870, p.255 No. 12,lectotype ? TCL! selected by Ander (1967), typeloc. Sweden: Lapland (Lu. Lpm.): Kvikkjokk;Thomson 1872; Siebke 1880; Strand 1898a,1898b. Bombus alpigenus Morawitz, 1873, p. 132,type area USSR: Dagestan.)

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Studies on Scandinavian Bumble Bees 27rather long hairs except for a large distal barearea; anterior and posterior fringes of hind tibialong, the longest hairs in the latter about twicethe greatest width of the segment or longer. Longesthairs in posterior fringe of hind basitarsusabout twice the greatest width of the segment orjust shorter. T7 slightly dimpled in front of apex.St, and genitalia (Figs. 318-C), basic half ofgonocoxite distinctly caved inside; gonostyluswith well-defined finger-shaped process producedinwards; volsella reaching well beyond gopostylus,distally emarginated; penis valve distallybroad with dorso-ventral flattened, sickle-shapedhooks turned inwards. Coat shaggy. Body ofmedium to large size.Colour pattern Pile of face below antennalsockets and pile of vertex predominantly yellow,encroached by black hairs. Collar, including atleast adjacent margin of episternuffi, yellowhaired.Venter of thorax, dist al pafi of scutellumwith variable admixture of yellow hairs. Tn'r,Iateral part of fringes of Stn-u ferruginous. Trochanters,parts of all femora, the remainingsternites with pale yellow to greyish-white hairs.Fringes of hind tibia and hind basitarsus whitishtinged. Otherwise coat black.Variation. Yellow hairs on head and thoraxmore or less pronounced, occasionally T1 withyellow hairs. Ta distally with variable amount offerruginous hairs. Fringes of Stn- e entirely ferruginous,usually of paler shade than that of thetergites. Fringes of hind tibia and hind basitarsusmay be predominantly ferruginous.Distribution (Fig. 50)Ncrway. Widely distributed from the extremesouthern coast and north to nearly 68" N. In thesoutheasternmost lowlands, however, not reachingthe border area to Sweden, and only oncerecorded in Ostfold county.The frequency is locally high in woodland andsubalpine valleys, occasionally also in adjacentlower alpine zone. Along the western coast it issporadically recorded in the extreme archipelago.In Southern Norway recorded to 1300 m s.m.Biotopes: Meadows, leguminosae fields, gar-dens, roadsides, pastures, Aconitum biotopes,Vaccinium heaths. Barren land and Callunaheather are avoided.A total of about 2000 specimens was examined.List of localities. Ostfold; Ride: Tom H 1S July 1958(I. Meidell). Akershus.' Asker: BronnoY, Lushattdalen,Rustad; Brerum: Briten; Oslo : Bonntjern, Hellerud,Maridal, Slemdal TRM NRS, Tisen; Ullensaker:?loc. ZMO; Eidsvoll: S Eidsvoll, Feiring . Hedmark( HEs ).' Kongsvinger : W Eidsberg; A.snes : Asen;Hamar: Z}l4O, Helgoya ZMO. HEn: Trysil: Dammen,Trysil, S Sjoenden, Vestby; Amot: Glesubekken,Amot; Stor-Elvdal: Snodoldalen 1000 m; Tynset:Haugsvangen, Tyldal; Tolga: Hodalen; Engerdal:Torhus. Oppland (Os).' Gran: S Einavoll, Gjerdvika,Tingelstad; Ostre Toten: Stubdalen; Sondre Land:Fluberg; Nordre Land: Dokka USU ZIN4B, Torpa;Lillehammer: Jorstadmoen; Etnedalen: Bruflat; Sor-Aurdal: Ellingseter 820 m, Skard; Nord-Aurdal:Dalen, Fagernes, Flya, Hovda, Srebuoygard N Fagernes;Oyer: Aksjoseter 1000 m, Skii, Oyer, Astdalen900 m; Ringebu: ?loc. ZM.O, Flaksjoen. On: VestreSlidre: Kinnholt 1000 m; Vang: Eidsbugarden 1100-1200 m, Tyin 1100 m USU ZMB, N Oye USU ZMB;Sel: Heidal TRM, Klaksh&ug, Leirflata, Mysuseter900 m; Vigfl: Gjendesheim 1000 m, Klones, Leirungen1050 m, Randsverk TRM, Vigflmo; Lom: Bovertun900 m, Galdesand, Gjendebu 1000 m' Leirdalen900 m, Solell 1200 m; Skjak: Skj6ksetrene800 m; Dovre: Dovrefjell 1000 m KMT NRS TCLZ}rdB ZMC ZMO, Fokstua 930 m Z}l'dB ZMO, ToftemoZMO, Vfllflsjo ZMO. Buskerud (Bd): Hurum:Filtvedt TRM ZN.d.{B, Pinadalen; Drammen; Lier:Toverud; Ringerike: Hen; Krodsherad: BjorkerudoBrfltin, Fagerheim, Glesne, Veikflker; Modum: Dignes;Flesberg: Lampeland, Oydegarden; Kongsberg:Lauerud, Lintvedt, Skollenborg.Bv; Sigdal: Kopseng,Nedre Eggedal, Sarrdsbriten, Sigdal; Nore og Uvdal :Flitflseter 1000 m, Heggjeli 780 m, Norstebo, Rodberg,Uvdal stavkirke, Uvdal; Flfl: Grosland; Gol:Gol, Randalseter 940 m; Hemsedal: Holle bru, Kjolen1000 m, Lykkja 900 m, Muren 730 m; At: ?loc. ZMO,Bergsjostolen 1100 m, Tvist 1080 m, Vallehallell20 m, Vats 800 m; Hol: Geilo 800 m ZINI4A, Halfardokkistolen,Haugastol 1000 m VCA ZIM{.{B, NWHaugastol 1100-1200 m, Hovet, Kikut, Nygflrd1000 m, S Olsenvann 1100 m, Sundatrsfjord 750 m,Ustaoset 1000 m, Vikastolen 1000 m. Vestfold; Sande:S Sande; Botne: Angerskleiv. Telemark (TEy).' Skien:Kikut; Bamble : Trosby, AUv; Nome : Damtjern;Drangedal: Nordbo, Nos, Omnes, Tomyra; Kragero:Levang . TEi: Notodden: Gransherad; Bo : Sande,Vatnar; Seljord: Seljord, Svartdatr, Vedfallsgrend;Kviteseid: Brunkeberg, Mostol, Vridal; Nissedal:Kyrkjebygda, Lauviki, Lia, Tionnefoss, Treungen;Fyresdal: Moland; Tokke: Bandaksli, Holtet bro,Grimdalen, Mule g&rd, Amdal Verk; Vinje: Bossboen

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Studies on Scandinavian Bumble Bees 29Tuftenes; Marnadal: Ask, Finsland, Gretteland,Royseland, Solfls, Utkjrer, Oyslebo; Lindesnes: Ramsland,Reme, Spangereid; Lyngdal: Lyngdal TRM;Kvinesdal: Feda, Gjemlestad, Kvinesdal, Oy.; Farsund: Fjellesi, S Hanangervannet, Lodshavn; Flekkefjord:Abelsnes; Ersdal, Espetveit, Loga, NedreFjeldsfl, Nuland, NW Sandvann; Audnedal: Sveindal;Hregebostad: Skeie; Aseral: Espelid, Rosseland,Aseral; Sirdal: Nedre Sirdal ZMO. Rogaland (Ry):Lund: Eikjelandsdalen, Moi; Sokndal: Hauge i Dalane,Nesvflg, R.ekef.jord; Eigersund: Birkeland, Eigeroyeidet,NeCre Hetland; Bjerkreirn: Malmeim; H&:Ogna, Saite bro; Klepp : Jreren, Klepp; Gjesdal:AtgarO; Sandnes: Hana; Sola: R.joa; Stavanger:Lindoy; Strand: T'au; Rennesoy: Rennesoy; Vindafjord:Imsland; Tysvrer: Nedstiand; Karn-loy: Eide.Ri; Forsand: Lyse; F{jelmeland: Fister, Haugstri,Hjelmeiancl, Igland, Steinslandsvann, Ardal; Sutrdal :Br6tveit, Hf,vardstol, Jelsa, Mostol, Nesflaten, Sand,Suldalsosen, Svortebekk, Ullsneskvelven, Vetrhus,V≥ Sauda: Hellelandsbygd, Sauda, Saudasjoen.Hordaland ( HOy ); Bornlo : I\{osterhavn ; Sveio : NAustvik; Olen: Dommersnes, Innbjoa; Fana: Blomsterdalen,Dolvik VCA, Fjosanger VCA ZMB, Hop,Minde VCA, Skipanes VCA ZMB, Skjold, Scrfls,Titlestad, Tveiteris; Bergen: TRM VCA ZlldB ZMO;' Askoy: ? loc. VCA, Hegernes VCA, F{erdla; Osteroy:Kleppe; Meland: Brakstad, Io; Radoy: Rosnesset;Vaksdal: Eidslandet; Lindfls: Fosse, Herland, Srevrflsv6g.HOi: Etne : Frette, Skinevik; Kvinnherad :Lio, Varaldsoy; Odda: Austmannli 800 m, Dalengflrd 900 m, Midtleger .1 150 m, Roldal, Seljestad,Valldal 900 m; Ullensvang: Alvsaker, Djonno, Espe,Fossli 750 rn, Fresvik, Grythorgen 900 m, Hjoimo,Jistad, I-ofthus, Mabddalen, Sysendalen 750 m;Kvarn: Kvamskogen; Voss: Kinne, h/tjdlfjell; Granvin:Eide VCA; Ulvik: Finse 1250 r11, Gangdal1000 rn. Sogn og fiordane (SFy): Hyllestad: Botn,Hyllestad, Risnes, Skivenes, Yttre Dale; Gulen: Takle;Hoyanger: Lavik; Fjaler: Strandenes; Flora: Kinn;Naustdal: Framrnarsvik, Naustdal; Jolster: Ffamar,Vassenden; Gloppen: Sandane; Eid: Haugtrand,Kjolsdalen, Naustdal, Nordfjordeid; Selje: Ervik.,SFi; Vik: Vik; Aurland: Kvammadal 950 m, Myrdal870 m, Uppsete 850 m VCA ZMB, Vatnahalsen800 m, Ovstebo 800 m; Lnrdal: tsreistolen 1050 m,S Horgje; Sogndal: Slinde; Leikanger: Dalseter,Hermansverk; Balestrand : Brekka, Flesje, Kvan-lsoy;Luster: F&bergstolene 600 m, SW Hallbreim, Jostedal,Skjolden, Turtagro KMT NRS TRM VCA ZMB;Stryn: Flo, Hornindal, Skire, Stryn, Vevik, Videdalen,Videseter 600 m. Mrire og Romsdal ( MRy) :Sande: Gurskevig, Aram; Vannylven: Thue; Ulstein:Fl6, Vonheim; Orsta: Lyngstolvann, Oy.; Heroy:Runde; Alesund: VCA; Orskog: ZMO; Vestnes:Gjermundsnes, Vikebukt; Molde; Aukra: Aukrakirke, Falkehytta; Frrena: Bud; Eide: Vevang. MRi:Stranda : Kvanndalsretrene, Marik, Vollset; Norddal:Indreeidet, Valldal; Surnadal : Kvanne ZMO, Melhus,Stangvik, Ovre Surnadal; Rindal: Bolme. Sdr-Trdnde'3 Bumble Bees.Iag ( Sfy ).' Rissa : Gafsetisen. STi: Oppdal : Dalsbekk,Driva st., Drivstua 850 m ZMU, Gflv61i 975 m,Knutsho 1100-1300 m BML Z}ldB, Kongsvoll 900 mBML TRM ZN44r- ZMO ZMU, Skansen, Virstigen850 m; Rennebu: Rennebu; Alen: Reitan, IJrnholtet;Roros: N Adamsvoll 820 m, Evavollen 700 rn, Myrmoen700 m, Naustervoll, Storelvvoll 800 m; Tydal:Mo bru, Sakrismoen, Vekterstua 600 m; Selbu:Medbus; Trondheim: VCA ZMB. IVord-Trdndelag( NTy ).' Nreroy : Breiviken, Kvalviken, Teplingan.NTi: Merflker: Meriker, Tovmodal; Stjordal: Draveng,Hegra st., Tilleraunet;Levanger: Avdal; Verdal:Billingflaten, Sandvika, Sulstua, Stiklestad, Vuku;Inderoy: Kirkenesvflg, Skjelsvigen; Steinkjer: Sem,Snflsavann ZMA; Snflsa: Bronstad, Hegge, Leira bro;Lierne : Kvelia, Seterhaug, Smitjern; Hoylandet:Skilleberget; Narnskogan: Finnvollan; Royrvik:Bjorkhaug. Nordland ( Nsy).' Vega: TRM; Gildeskfll:Jelstad, Gildeskfil kirke; Bodo: Falkflaugdalen, Frostffio,Lopsviken. l/si; Grane: Bflfjellmo, Majavann,Skomstad; Vefsn: Luktvann; Hemnes: Finneidfjord,Strombotn; Rana: Randallsvollen, Rosvollheien;Beiarn: Ci6tflCalen; Satrtdal: ?loc. ZMO,Storjordet TRM, Solvigfjell 800 m; Fauske: Fauske,Kjeldvann BML, Sulitjelma 600 m BML. I,{nri: Sorfold:Bonnisjoen.Correction. Records from Buskerud (Bv): At(Strand 1898a) are revised to B. lapidarius (Linnaeus).Sweden Occurring from Dalsfand, about 58o30' N north to Virihaure atea in Lapland (Lu.Lpm.), nearly 67" 20' N and rnoreover sparselydistributed east to Gulf of Bothnia in Medelpadand ,4.ngermanland.A total of about 450 specirnens was examined.List of localities. Dalsland: Kroppefjiill NRS ZML,Kopmannebro, Rostock, Sk611erud NRS. Vrirmland:Arvika. Dalarne; Fulufjiill NRS, Slagufjiill NRS,Siirna ZMA. Medelpad: Liden NRS ZMB, Ange.Hcirjedalen: ?loc. NRS, Hamrafjail NRS ZML,KesusjonNRS, Tiinndalen NRS ZML. Jrimtland: BergeI)CL, Brattikkfjail DCL, Brunflo, Bydalen, FrostvikenNR.S, .Iormlien, Leipikvattnet NRS, Storlien,Svenstavik NRS, Undersflker NRS ZML, OstersundNR.S, Are TCL VCA Zl/'A ZML, Areskutan DCL.Angerrnanland: Bondsjo, Hdrnosand, Orsjon, SollefteflNRS, Stennds. Lapland (.4t. Lpm.).' S BorgafjiillNRS, Fatmo, Fatmomakke, Njakafjiill, Satsfjell NRS,Vilhelmina NRS. Ly. Lpm..' Ammarnds GNM NRSzMU, Bjorkfors, Tjultriisk BML GNM NRS ZMU,Tdtrna,Tiirnasjon, Vdstansjo, Ornds. P. Lpm.; Jeickvik,Peljekarse. Lu. Lpm.; Kvikkjokk TCL ZML, NjunjesVCA ZML, Sarek NRS ZML, Staloluokta, TarradalenNRS.

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Studies on Scandinavian Bumble Bees3lTable I. Data frorn a colony of .8.Stokke 8 June 1934 bv O. Meidellw.'mestrucatus Gerstaecker excavated Norway : Rogaland (RV) : Stavanger:and kept in observation box until 24 July 1934Queen, founder of theOffspring 8 June:lst brood: workers2nd brood: workerimaginestnestTotal number of progeny6Iworker cocoonsworker cocoon, 5 worker pupae3rd brood, lst batch: 5 worker pupae3rd brood, 2nd batch: 6 worker pupae4th brood, 1st batch: 9 worker larvae4th brood, 2nd batch: 14worker larvae5th brood, l st batch : eggs5th brood, 2nd batch: eggs6656914?,l46 + eggsI honey-potCocoons used as honey-potsOffspring produced 8 June-24 July: 11 batches of broodsImagines emerged 8-26 June: 35 workersImagines emerged 4-24 July;21 malesOffspring present in the colony 24 Jttly: 12 workers, 2 males, 0 pupae, a great number of moulded larvae,some healthy larvae and 2 batches of eggsworker broods. The first male was, moreover,eagerly incubating until the nest was left on10 Juiy. The remainder of the rnales left the nestafter a couple of days, several of which were alsoobserved incubating.The flight activity lasted during favourableweather conditions from about 4 a.m. to 10.30p.rn. which agrees with Laken (1949, p. 48), whomoreoversuggests that B. w. mqstrucatus in generalhas a longer working day than other Bombusspecies occurring in the same atea.Flisht season. From the beginning of May tothe middle of September. Queen: 5 May-6 Sept.;worker: 5 June-l3 Sept.; male: 14 June-l1 Sept.The males were only once observed earlier thanbeginning of July.SUBGENUS KALLOBOMBU,S DALLATORREKallobombus Dalla Torre, 1880, p. 40, typespeciesBombws soroeensis (Fabricius) by desi gnationof Sandhouse (1943).( Soroeensibombus Vogt, l9ll, p. 63, typespeciesBombus soroeensis (Fabricius) by monotvpv.)BOMBUS ( KALLOBOMBUS) SOROEET{SrS(FABRICIUS),Apis soroeensis Fabricius, 1776, p. 246; Zimsen1964, p.415 No. l0TT,lectotype ? CCC! selectedby Laken (1966a), type loc. Denmark: Sjelland.(Bombus soroeensis (Fabricius) ; Wahlberg1854; Thomson 1870, 1872; Siebke 1880; Strand1 898a, I 898b ; Friese 1902; Aurivillius 1903 ;Muchardt 1904; Bengtsson 1908 ; Sparre Schneider1918; Meidell 1934a, 1946; Wexelsen & Skflre1934; Lsken t949, 1950, 1958a, 1966c; Erlandsson1950, 1960; Ander 1953a, 1963, 1965; Kruseman1959; Elfving 1960, 1968; Frid6n, Eskilsson& Bingefors 1962; Hasselrot 1962; Frid6n 1967.Apis neutra Panzer, 1800-01, part 83 Teil 18,type not traced. Bombus virginalis: Dahlbom,1832,p. 35 nec Geoffroy in Fourcroy 1785 . Bombusproteus Gerstaecker, 1869, p. 325 No. 18, lectotype6 IZB! selected by Tkalcu (1969), type loc.Germany: Ober-Bayern: Kreuth. Bombus hortorumvar. soroeensr,r,' Siebke 1873.)First Scandinavian recordsIr{orway. Hedmark (HEs): Amot (Siebke 1873).Sweden Skine, Smflland, Vtistergotland (Dahlbom1832 cf. above).

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Studies on Scandinavian Bumble Bees 33There is a distinct difference in size betweenthe queen and the worker caste.Colour pattern Collar, extending to adjacentmargin of episternuffi, and visible hairs of T,lemon yellow. To-u, lateral part of fringes ofStn-u whitish haired. Otherwise coat black.Variation. T, with variable admixture of yellowhairs, occasionally entirely yellow-haired. Yellowband of T, more or less interrupted in the middle,i. e. yellow hairs replaced by a variable amountof black ones; additional black hairs often presentat extreme anterior and posterior margins. Whitishhairs on anterior paft of Tn to various degreesreplaced by black ones. Occasionally Tn-, clothedwith yellowish-white or pinkish hairs. SparreSchneider (1918, p. 13) indicates that only newlyemerged individuals clisplay a pale reddish topink tail, but this colouring does occasionallyoccur also in older specimens.f Among a total of 9 QQ and 2 A6 preserved froma colony excavated in Rogaland county by thi)late Meidell, two females display a pronouncedferruginous colouring of Tn- u, a pattern approachingthe yellow-banded variant of ,8. soroeensisproteus. A few queens, workers and males withcolouring as this subspecies, were also exarninedfrom southernmost Sweden: Sk6ne: Surroundingsof Hiilsingborg, where typical and melanicwhite-tailed individuals occur as well, indicatingthe closeness of the intergrading areain Denmark.MaleMorphological characters. Head about as iongas wide. Malar space about as long as distalwidth, as long as Ar*r.Mandible bifid with abroad ventral and a small dorsal tooth. Eyesdirected in front of posterior mandibular condyle.Antennae long, yet A, markedly shorter than Au,equal to An or barely longer I An barely longerthan distal width of the segment; Au-r, individuallyslightly bent and about twice as long asdistal width (Fig. 23A). Outer surface of hindtibia flattened or slightly convex, with cnly sparsedistribution of hairs of various length and withlongest hairs in fringes about twice the greatestwidth of the segment. Hind basitarsus stronglynarrowed towards the base (Fig. 238), longesthairs in posterior fringe exceeding the greatestwidth of the segment. St* and genitalia as inFigs. 23C-D. Gonostylus, seen from above,about twice as long as wide; penis valve withdistinct subapical tooth and with distal hooksturned outwards and produced into a point. Bodyof medium size.Colour pattern Pile of face black or with tracecf yellow hairs admixed. above and below theantennal sockets. Pile of vertex with variablemixture of yellow and black hairs. Collar, extendingdown entire episternuffi, Tr-, lemon yellow.Femora of all legs with variable mixture of yellowand black hairs. Posterior edge of T n, Tu-,whitish. Coat otherwise black.Variation. Yenter of head with trace of yellowhairs. Femur of hind leg clothed with black hairsonly. Whitish hairs of Tn ranging from coveringentire tergite to oniy the extrerne posterior edge,occasionally whitish hairs replaced by pink orferruginolls ones, rarely Tu entirely black. Tu-,with variable adn-lixture of black hairs, T 7 occasionallywith almost golden-brown hairs. Asingle black tailed specimen is examined. As mentionedabove, also males of the proteus formoccasionally occur in Sweden : Sk&ne: Hiilsingborgatea.Distribtttion (Fig. 5l )I{orway. Disperse, wide but local distributionthroughout the country north to 70o N. Occurringfrom sea level to subalpine valleys and sporadicallyrecorded in the lower alpine zone. InSouthern Norway recorded 1050 m s. rn.Biotopes : Meadows, pastures, roadsides.A total of about 1300 specimens was examined.List of localities. Ostfold.' Hvaler: Asmaloy, Kirkoy,Sondre Sando; Onsoy: Rorvik, Torp; Sarpsborg:ZMO; Halden: VCA, Asak, Berg kirke, Blakkmose,Hakelund, Nordbakke, Sponvika.Veggesdal ; Aremark :Gjeddetjern, Skodsberg, Stensbru; Marker: S Damholtet,Dypedal, Jival, Rodenes; Rakkestad: Levernes,Rorvik; Ride: Fuglevik; Moss: Jeloy VCA;Trogstad : Bingen, Monster bro, B6stad. Akershus:As: Vollebekk; Frogn: Degerud TRM, Drobak VCA;Cppegird : Svartskog ZMO; Asker: Rustad, Sem;Brerum: ? loc. ZMO, Hovik ZMA; Oslo: NRS TRMVCA ZN 4B ZMO; Aurskog-Holand: Bjorklangen,

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Studies on Scandinavian Bumble Bees35Nos, Omnes, Tomyra. TEi: Notodden: Gransherad;Bo: Sauda; Seljord: Seljord, Vedfallsgrend; Kviteseid: Kviteseid, Vrfldal; Nissedal: Treungen ZMO;Fyresdal: Hegglandsgrend, Moland; Vinje: Edland,Urdbo 700 m; Tinn: Vier, Vilen. Aust-Agder (AAy):Tvedestrand: Askeroy,Lyngor ZMO, Nes verk TRM;Tromoy: ZMB ZMO; Fjaere: Fjrere ZMO. AAi:,&rnli: Krossbekk; Valle: Kvestad, Setesdal TRM,Rygnestad, Valle TRM ZMB; Bykle: Hovden. Vest-Agder ( VAy ).' Marnadal: Finsland; Lyngdal: LyngdalTRM. VAi: Aseral: Aseral. Rogaland ( Ry):Soknedal: Hauge i Dalane, Vatland; Eigersund: Fotlandsvann;Bjerkreim: Ivesdal, Malmeim; Ha: Ogna,Fossfjellet; Klepp : Klepp; Gjesdal: Nese; Sandnes :Hommersflk, Todnheim, NW Ganddal; Sola: Solastrand;Stavanger: TRM ZMB, Lindoy; Finnoy:Sjernaroy; Rennesoy: Dale; Kalltveit; Karmoy:Vigsnes. Rl; Forsand: Lyse, Lysebotn, Meling; Hjelmeland:Fister, Hjelmeland, Ardal; Suldal: Bleskestadmoen875 m, Brirtveit, H6varstol, Jonstol 700 m,Kalltveit, Krokvassheia, Kyrkjesteindalen 800 m,Moen 600 m, Mostol 600 m, Nesflaten, Roaldkvam,Sand, Sandsa TRM, Svortebekk, Vandvik, Vige,Aimjo0lonuten 700 m; Sauda: Helielandsbygd, Saudasjoen. Hordaland ( HOy ); Fitjar : ?loc. ; Tysnes :Anuglo; Os: Lysekloster, Mobergslia, Moldeg6rd;Fusa: Ovre Hilandsdal; Samnanger: Hoyseter; Fana:Appeltun, Biol. st., Birkelundsbakken, Dolvik VCA,Fana kirke, Fantoft, Rdrdalen, Skjold, Steinsvik,Stend, Titlestad; Sund : Telavflg; Fjell: Selsto; Bergen:VCA ZN{B ZMC; Askoy: ?loc. VCA, Ask,Herdla; Asane: Golfbanen, Vinddalen; Osteroy:Greve, Kleppe; Vaksdal: Eidslandet. IIO|: Etne: Oslandsvig;Kvinnherad : Guddalsdalen, Prestvatn, Varaldsoy;Odda: Dalen g6rd, Odda, Roldal; Ullensvang:Aga, Borve, Djonno, Espe, Fossli 750 m,Fresvik, Grythorgi 900 m, Hjolrno, Jflstad, Kinsarvik,Kvanndal, Lofthus, Mebodalen, Nesheim, Sekse,tltne, O vre Eidfjord; Kvam: Kvamskogen, Nordheimsund,Valland, Vangdal, Vikoy, Oystese; Voss:Bomoen, Hangursfjellet 800 m, Voss; Granvin: EideVCA TRM, T16; Ulvik: R.ubbeliseter, Ulvik. Sogn ogFjordane ( SFy ); I{yllestad : Botn ; Gulen : Midt-Takle; Hoyanger : Lavik ; Gaular : Slotten ; Fjaler :Strandenes; Askvoll: Rauoy; Flora: Havrenesvig;Naustdal: Ovre Hella, Horstad; Forde: NE Flflten;Jolster: Hamar; Gloppen: Hope, Hyen, Lote, Lotsbergfjellet700 m; Eicl: Haugland, Heggjabygda,Kjolsdalen, Naustdal, Nordfjordeid, Vedvik ; VAgsoy :Degnepollen. SFi; Vik: Ovrisdal 960 m, Skjellingenseter 1000 m, Vik; Aurland: Aurlandsdalen, Berekvam,Fl6m, Gudvangon, Vassbygdi, Vatnahalsen800 m; Lnrdal: ? loc. ZMO; Sogndal: Kaupanger,Sogndal; Leikanger: Fosshagen, Hermansverk,Stavseter 780 m; Balestrand: Balholm NRS, Brekka,Flesje, Horpedalen, Kvarnsoy, Mel, Mundalsdalen,Tuftedalen; Ardal: Ardalstangen; Luster: Dalsdalen,Fortun, Fiberg, Hafslo, Jostedalen NRS ZMB ZMO,Okkleviki, Skjolden, Urnes; Stryn : Briksdal, Flo,Gillesdal, Hornindal, Hornindalsretra, Mindresundet,Skflre, Stryn, Stoverstein, Vanberg, Videdalen, OvreOldenvann. Mrire og Romsdal (MRy); Volda: Bjorkedalsvann,Folkestad, Straumshavn; Orsta: Lyngstolvann,Urke, Oy"; Ullstein: Vonheim; Alesund: VCA,Ase; Orskog: Skodje ZMO, Orskog ZMO; Vestnes:Vikebukt ; Molde : ZMO ; Aukra: ? loc. MRi.' Stranda :Bjordalsgirdene, Fivelstad, Geiranger NMW, Marik,Nordaldalen, Vollset; Norddal: Valldal, Yttredal;Nesset: Nauste; Sunndal: Branstad, Jordalsgrend;Surnadal: Ovre Surnadal. Sdr-Trdndelag ( STy):Hitra: Hamn; Agdenes: Lillevann SW Harnbora;Rissa: ? loc. VCA, Sdtvik; Orland: Hovde KMT.STi: Oppdal: Oppdal; Trondheim: KMT NRS ZN{4B.Nord-Trrindelag ( NTy ); Flatanger : Halmo TRM ;Nreroy: Skogenga. NTi; Meriker: Meriker; Stjordal:Hegra st. ; Verdal: Stiklestad, Sulstua, Verdal, Vuku;Snflsa : Bronstad, Hegge. Nordland ( Nsy ).' Somna :Sandvflg TRM; Bronnoy: Rommelsto TRM; Alstahaug:TRM; Nesna: Hamaroy; Meloy: Kunna,Reipfl; Gildeskil : Finnes, Gildeskfll, Storvika; Bodo:Lopsviken, Planteskolen, Skau. l/si.' Hemnes: Finneidfjord;Rana: Rovassdal TR.M; Saltdal: Drageid,Junkerdal; Fauske: Fauske, Kjeldvann BML, SulitjelmaBML. Nnii: Sorfold: Bonnisjoen; Hammaroy:Sandnes; Tysfjord: Tysfjord kirke ZMO; Ballangen:Hesjeli. Nnv: Vestvflgoy: Egguffi, Haukland, Kvalnes,Skulbru, Stamsund; Vflgan: Store Molla; Lodingen:Lodingen ZMO. Troms (TRy).' Tromso NRS TRM;Karlsoy: Hushattoy TRM. TRi: Balsfjord: ?loc.TRM ; Nordreisa : Sappen. Finnmark ( Fi ); Alta :Bossekop ? Juni 1924 (A. Strand) TRM, Kflfjord ?Juni 1924 (A. Strand) TRM.Correction. A worker specimen (Oppland (On):Dovre (Lysholm) NRS), labelled B. soroeensisvar. rarior b)' Friese (Erlandsson 1950) is revisedto B. balteatus Dahlbom. Moreover, no recordsconfirm a distribution 'nordwdrts bis zum Nordkap(Norwegen) . . .' (Friese & Wagner 1909,p. 28). Finnmark (Fv): Nordkapp is situated onthe northernmost coast of Norway.Sweden Widely distributed throughout thecountry, but so far there are only a few recordsof this locally-occurring species from Lapland, andonly a single specimen has in fact been collectedas far north as T. Lpm.: Abisko, apparcntlyowing to insufficient observations in forest biotopes.A total of about 800 specimens was examined.List of localities. Skdne: Barnakiilla GNM, Bjornstorp,Bokskogefl, Brosarp, Bdckaskog, Bokeberg,Dalby, Fjelkinge, Glimminge, Hjularod ZMB, HtigghultZN{B, Hiilsingborg NRS VCA ZN {.f.r_ ZML,Hiickberg&, Hdssleholm NRS, Ignaberga, Ilstorp,Klostersflgeno Kullaberg, Kungshult, Kflgerod, Lund,

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Studies on Scandinavian Bwmble Rees 37SUBGENUS BOMBU,S LATREILLE(s.s.)Bombus Latreille, 1802a, p. 385, 1802b, p. 437.Proposed as a genus. Type-species Apis terrestrisLinnaeus. Monobasic.(Bremus Jurine, 1801, p. 164. Proposed as agenus. Type-species Bremus terrestrzs (Linnaeus)- Bombus terrestris (Linnaeus) by designation ofMorice & Durrant, tr915. Generic name invalidated,Int. Com. Zool. Nomen., Opinion 135,1939. Leucobornbus Dalla Torre, 1880, p. 40, typespeciesBombus terrestrls (Linnaeus) by designationof Sandhouse (1943). Terrestribombzs Vogt,l9ll, p. 55, type-species Bremus terrestris (Linnaeus)- Bornbus terrestris (I-innaeus) by designationof Frison (1927).)Queen, worlcerHead slightly shorter than wide, occasionallyas long as wide. Malar space hardly to markedlyshorter than distal width, about as long as AB(longer in B. sporadicus Nylander, just shorterthan Ar+'). Disc of clypeus flattened anteriorlyand just shorter than distal width, with ratherdense punctures of various size throughout, withwell-defined impressions produced inwards andalmost meeting in the middle of anterior edge.Labral furrow well-defined, nearly one third aswide as labral width; labral tubercles deeply depressed(Fig. l2A), angled at inner end; labrallamella rather strongly curved (less so in B. sporadicus),slightly rvider than furrow. Mandible withoutbasal keel (indication of keel in B. sporadicus) ,with distinct sulcus obliquus and prorninent incisuralateralis, being almost semicircular in unwornspecimens. Eyes directed in front of posteriormandibular condyle. Supra-orbital line touchinglateral ocelli or just transecting them dorsally.Mid-basitarsus with alrnost right rounded distaldorsal angle (Fig. lzc). Hind tibia feebly or notalutaceous, dorsal inner distal angle slightly tomoderately produced; hind basitarsus abouttwice as long as greatest width of the segment(slightly longer in B. sporadicus), posterior edgemore or less curved (Fig. 12T)), longest hairs inposterior fringe markedly shorter than half thegreatest width of the segment.There is a distinct difference in size betweenthe queen and the worker caste (Lsken l96la).MaleHead slightly shorter than wide. Malar spacehardly longer than distal width or equal to thiswidth, longer than A, but shorter than Ar+r.Clypeus not longer than distal width. Mandiblebifid with ventral tooth broader than dorsal one.Eyes directed in front of posterior mandibularcondyle. Ocelli usually situated on a line. Antennaerather short, Au-r, individually l1/2 times thedistal width or hardly so, almost parallel-sided(Figs. 28A, 29A). Hind tibia slightly convexthough outer surface flattened or somewhat impressedtowards posterior margin, bare except.forat rnost proximal third being covered with sparsehairs of various length; posterior and anteriorfringes of the segment shorter than twice thedistal width; posterior fringe of hind basitarsusshorter than distal width. St' and genitalia (Figs.288-D , 298-D); gonostylus transverse with inwardlydirected spine, penis valve trvisted, distallyconvoluted, flaring.Scandinavian speciesThe subgenus is represented by a total of fourScandinavian species, viz. B. lucorum (Linnaeus),B. magnus Vogt, B. sporadicus Nylander andB. terrestris (Linnaeus).Taxonontical remarksThe four species present srnall sclerotic differencesand display a fairly uniform yellow-bandedcolour pattern which mainly differs in the yellowshade, in details in the extension of yellow hairsand in the length of the hairs (Kri.iger 1950, 1954,1956, 1958). Overlapping variations in diagnosticcharacters and fading of yellow hairs often preventreliable determinations of workers and males,An exception is B. sporadicLts) where sex andcaste are identified by several stablc features, cf.the description below.

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Studies on Scandinavian Bumble Bees 39B. terrestns males are in general distinguishedfrom B. lucorum s.l. by several features, cf. thedescription below. In some specimens the diag-xtrstrP o\'k*",ft. xooaartclaa\ooIoof,oFig. 53. Triangular graphing of 'radial length' (B), distance from lateral ocellus to preoccipital ridge (D) anddistance from lateral ocellus to cornpound eye (E) of four speciesof Bombzs (s. s.).ments, together with those of 'radial length', givea better spreading of the rnarkings (Fig. 53).Though again there is overlapping between B.terrestris and R. magnus, the atea occupied byeach of the four taxa is well-defined. The separatedistribution of B. lucorum and B. magnus oneither graph (Figs. 52-53) is of vital irnportanceto indicate their specific difference, and thus confirmsthe statement of Krtiger (1954).As mentioned above, the queens of all fourspecies were recogn ized, whereas this was not thecase in the males. I am in fact at present not ableto distinguish B. magnus males. According toKrtiger (1954) they are separated from B. lucorummales by longer hairs and certain details in theextension of yellow hairs, but owing to variabilityin those characters more informatiorr is needed.The Scandinavian B. lucorum males have so farbeen broadly separated into a 'dark' and a 'blond'form. Recent chemic al analysis of odiferous compounds(Kullenberg et al . 1970) reveals howeverthat each of the two forms, presented as B. lucorum'dark' and R. lucorurn'blond', process different,characteristic secretion emanating fromglands in the heads. fnir difference may favoura theory of trvo species being involved, i.e. the'dark' form rnay refer to B. rnagnus.

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Studies on Scandinavian Bumble Bees4lmagnug usually with chagrinated microsculpturealso in the median posterior part of the disc. Innersting sheet (Fig. 13B) markedly narrower than inB. terrestris (Fig. lZE). Dorsum of thorax withmoderately long, somewhat uneven coat. Bodyless robust than in B. terrestris and B. magnus,wings rather infuscate.Queen measurements: N - 20; Hordaland(HOV); malar space: 0.59 mm (+O.OZ +0.01)range: 0.55-0.65 mm; 'radial length': 4.45 mm(+O.t+ +0.03) range: 4.10-4.70 mm; interalarwidth: 5.40 mm ( +0. tZ +0.04) range: 5.03-5.73 rnm; body of large size.Colour pattern Collar, extending to the extremeadjacent margin of episternum, and T, except atbase, lemon yellow. Hairs of distal half of Tn,entire Tr, vatriable part of Tu and at least lateralfringes of St^-u whitish. Otherwise coat black.Variation. Yellow collar varying in width andrarely reduced to an inconspicuous band. Yellowband on T, also varying in width according to avariable amount of black hairs at base anddistally. Occasionally the extreme posterior partof scutellum and rnedian part of Tt with a slightadmixture of yellow hairs. The shade of the colourfades and therefore ranges from bright lemonyellow to yellowish-white. Occasionally corbicularfringes tipped whitish. Cornpletely melanicspecimens rarely recorded. Workers occasionallywith black hairs of thorax whitish tipped.The diagnostic characters are in the workercaste often too diffuse for reliable identification,i. e. a number of specimens cannot be distinguishedfrom B. magnus or faded B. terrestris.MaleMorphological characters. Area between eyeand preoccipital ridge just outside genal furrowwith rather distinct uneven mainly coarse puncturing.Ocelli dorsally transected by supra-orbitalline. A, markedly longer than An, equal in lengthto Au or nearly so. T, usually not impressed distally.Str-, and genitalia difficult to distinguishfrom those of ,8. terrestris (Fig. 298-C); consideringthe variability in the shape of distal marginof St r-, and slight variations in details ofgonostylus, volsella, penis valve, etc. the specificdifferences in the features become indistinct,though genitalia of B. lucorum are usually lessstrongly chinitized than in B. terrestris. Coat ingeneral longer than by B. terrestrH, that of thoraxmore uneven. Body of medium to large size.Colour pattern The coat displays a colouringranging from distinct to pronounced diffuse pattern.When distinct, pile of face black or withonly singly yellow hairs admixed, T*-, whitishhaired, fringes of sternites mainly whitish, pile onfemora, tibia with a mixture of black and whitish;otherwise pattern as in the female. This odark'form is usually separated from B. terrestris bythe yellow band being of a paler shade, in newlyernerged specimens usuall.v lemon yellow, and bythe white hairs of Tn covering less than the posteriorhalf of the segment. In the 'blond' formpile of face below and above antennal sockets,pile of vertex pale to lemon yellow with variableadmixture of black; yellcw collar often extendingpart way down episternum; hairs of scutellumand Tr-, predominantly or entirely yellow; blackhairs on thorax and gastral tergites more or lesswhitish tipped ; coat of venter whitish. Specimensof the 'blond' form were traced in B. sporadicr,tscollections.Vuriation. trn addition to the variations mentionedabove, collar varies in width and maycontinue all way down episternum; only rarelyis collar reducod to a very narrow band. Thepattern in the 'blond' form beconaes particularlydifruse when black and yelloi.v hairs are more orless replaced by greyish-white ones. The 'blond'form usually has a more shaggy coat than the'dark' form.Remarks on the description of males. As mentionedabove more information must be gainedto distinguish B. magnus and B. lucorum and oneor several of the features above may be related toone of the species only.Distribution (Fig. 5a)Norway. Abundantlydistributed throughoutthe entire country, yet only occasionally observedin reg. alpina. In Southern Norway recorded to1300 m s.m.Biotopes: Salix biotopes, meadows, gardens,

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Studies on Scandinavian Bumble Bees43Orre, Orrevann, Orstad, Reve, Revtangoo, Vik, Otcsnevad;Sandnes: Brattebo, Brflstein, Dale, Floysvik,Ganddal, Helgeland, Hipet, Hoyland kirke, Riska;Sola: Gimre, Joa, Kolnes, Solastrand, Olbergstranda;Randaberg: Kverneviken SMS, Sande; Stavanger:Auklendhoyden, Hinna, Lindoy, Stavanger SMSTRM ZMB; Finnoy: Sjernaroy; Rennesoy: Dale;Kvitsoy; Karmoy: Brekkevann, Indre Eide, Skudesneshavn,Vigsnes; Haugesund: SMS. Ri.' Forsand:Forsand, Lysbotn, Lyse, Meling, Songedalen; Hjelmeland:Fister, Hjelmeland, Steinslandsvann, Ardal;Suldal: Bleskestadmoen, Britveit, Jelsa, Jonstol,Kyrkjesteindalen, kirdalen, Nesflaten, Mostol, Roaldkvam,Sand, Sandsa TRM, Suldalsosen, UllsneskvelenoAimjodlonuten ; Sauda : Birkeland, Hellelandsbygda,Sauda, Saudasjoen, Slettedalsvann, Ovre Sandvann1050 m. Hordaland (HOy).' Bomlo: Goddo,Rolfsnes, Spissoy, Sorstokka; Olen : Dommersnes ;Stord : Foynoy, Storsoy; Fitjar: Agoy, Brendoy,Fitjar, Koloyholmen, Risoy, Rubbestadnesset, Sl6tterriy;Tysnes: Anuglo; Austevoll: Ytteroy; Os: Forstrono,Gullholmen, Lysekloster, Midtsetra, Mobergslia,Nordstrono, Rod, Ulven; Samnanger: Hoyseter;Fana: Biol. st., Birkelundsbakken, Blomster-Fig' 54. B. lucorum (Linnaeus). Legend as in Fig. 50. Shaded areas refer to literatur€ indications and comprisesone or more subspecies,

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Studies on Scandinavian Bumble Bees 45Talvik; Kautokeino: Kautokeino, Karasjok. Fn: Kistrand:Hamnbukt, Skoganvarre; Vads6: Makkanes,Vads

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Studies on Scandinavian Bumble Bees 47Taxonomical remarksKrtiger (1954, p. 267) distinguishes B. m. latocinctusfrom the nominate subspecies'diebykurze greisgelbe Grundbehaarung der Area centralis,des caudalen wie des oralen Teiles der Areatriangulares. Zwar ist auch bei dem typischenmagnus der orale Teil mit kurzen greisen Haarenversehen, aber nicht in so starkem Grade wie beiseiner Rasse latocinctus. Ausserdem ist bei dieserdas Scutellum in grosserem oder geringerem Gradgelb bzw. weisslichgelb behaart.' In the examinedScandinavian material those characteristics ofpile of face and scutellum are not much pronounced,sometimes lacking. The present knowledgeof this critical species is in my opinion toomeagre for subspecific designation of the Scandinavianpopulation being treated as B. m. latocinctusby Kriiger (1954, p. 271).QueenMorphological characters. Malar space markedlyshorter than distal width. Ocellar-orbital fieldas in Fig. 14; the variations in the sculpturing tendtowards that of B. Iucorurn Distance from lateralocellus to preoccipital ridge (d) about 514 longerthan distance from ocellus to eye (e), mean valued: e - 1.24; distance between ocelli hardly tomarkedly longer than half the diameter of themedian ocellus, usually about % this diameter.As about I /2 times the distal width and about /,rthe length of Aaa u. Tz with distinct dense and,coarse puncturing, usually with chagrinated microsculpture;the unhaired distal margin in themedian part often narrower than in -8. lucorum.Length and evenness of coat moderate, i. e. ingeneral as for B. lucorum. Body more robust thanin ,8. lucorum. Wings rather infuscate.Queen measurements: N -- 20: Hordaland(HOV): malar space: 0.65 mm (+0.04 +0.01)range: 0.55-0.70mm1' 'radial length': 4.69 mm(+0.10 +0.02) range: 4.50-4.90 mm; interalarwidth: 5.48 mm (+0.14 +0.03) range: 5.10-5.73 mm. Body of large size.Colour pattern Broad yellow collar extendingabout l/a down the adjacent episternum and oftenprojecting backwards below base of wing. Usuallya few yellow hairs concealed by black ones at theextreme posterior part of scutellum. Median partof T, with variable amount of yellow hairs. T,entirely yellow-haired except perhaps black hairsat posterior margin concealed by yellow ones.Pile of almost distal half of Tn, entire Tu, variablepart of To tawny or yellowish-white with brownishbase. Fringes of sternites brownish, occasionallylateral fringes of Stn-u coloured as hindmosttergites. Corbicular fringes brownish or tippedso. Otherwise coat black. The yellow tone innewly emerged specimens is less bright than inB. lucorum, but not as deep as in ,8. teruestris,i. e. it is neither lemon-yellow nor sulphurousyellow.Faded specimens of B. magnus and B. lucorummay display the same yellow shade or rheformer is more dull.Variation. Pile of face with a more pronounceddark brown appearance compared with the blackpile in B. lucorum, and occasionally a minor partof the matted short pilosity is yellowish-grey.Yellow pile on scutellum is ranging from singlyposterior hairs to covering almost entire crescentshapedpart of the segment. Individuals with entirelyblack-haired scutellum were examined.Yellow pile on T, ranges from single hairs in thecentre to alanost all hairs on the tergite, the shadeis ranging from pronounced yellow to paleyellowish-grey. A few individuals have slightlynarrower yellow-haired bands. Tb may have anadmixture of black hairs, preferably in the median jpart. Tu is occasionally entirely black-haired.WorkerOnly individuals with a colour pattern as typicalfor the females were sorted out in B. lucorumcollections. Additionally about 200 workers froma colony revised from ,8. lucorum (Loken l96la)were studied, all of which display a yellow collarextending a little less than % down episternumand slightly varying in width. Pile on scutellumand T, was moreover entirely black except in avery few specimens presenting a slight admixtureof yellow hairs. As stated by Kriiger (1954, p. 266)the hairs on the hindmost tergites in this casteare more whitish than those of the queens.

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8" E.Grenwd 1d 12" 11' 16"Studies on Scandinavian Bumble BeesflFig. 55. B. magnus Vogt. Legends as in Figs. 50, 54.List of localities. Skdne: Ljunghusen, Nosaby.Halland.' Halmstad, Hasslov. Smdland: Jarstorp NRS,Jonkoping NRS, Kalmar NRS, Gasslanda, Markaryd,Ryssby Zli'4IJ, Vrirnamo NRS. Ostergtitland: KrokekNRS, Kvarsebo NRS, Norrkoping ZN,{A. Vristergcitland:Mosseberg in Falkoping NRS. Dalsland: BengtforsNRS, Kopmannebro, Steneby NRS, TingvallaNRS. Stidermanland: Uto NRS, Viksberg NRS, AttaNRS. Uppland.' ?loc. NRS. Viistmanland: Guldsmedshyttan.Ycirmland: Arvika NRS, Gylleby NRS,Lesjofors. Dalarne.' Transtrand V 10 Sept., 1955(Lundblad) NRS. Angermanland: Hdrnosand g 9 Aug.l93l (Fredlin), Siibnfl I 9 Sept. 1932 (Burrau). Lap-Iand (Ly.Lpm.).' Tiirna V 22-30 Sept. 1931 (?).Correction. Records from Lapland (Kriiger I1954, pp. 266, 271) refer to two specimens (Kolapen.: Umptek Mountains VCA) one of which isrevised to B. sporadicus (Kruseman in litt.) andthe other to B. lucorum by me.Finland. According to Krtiger (1958, p. 322)the species occurs in the southern part of the

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Studies on Scandinavian Bumble Bees5tQueen: Pile of face black, usually with admixtureof brownish in the short matted hairs. Yellowcollar just reaching margin of episternum, scutellumposteriorly with slight admixture of yellowhairs. T1 centrally with a variable amount ofyellow hairs; visible hairs of T, yellor,v, i. e. blackhairs on the extreme posterior margin usuallyconcealed by yellow hairs. Posterior half of Tn,entire Tu, often lateral fringes of Tu and at leastlateral fringes of Sto-u whitish. Otherwise coatblack. Worker: Crescent-shaped posterior part ofscutullum and T, yellow-haired; interalar band.Otherwise colouring as in the female.Variation. In the queens the amount of yellowhairs on scutellum and T1 ranges fronr coveringthe crescent-shaped posterior part of scutellum,and entire T1, to only single hairs in the centre ofT1. Otherwise colour-stable. Thus only a few ofthe examined queens display the prevailing patternin workers characterized by interalar band.MaleMorphological characters. Area between eyeand preoccipital ridge, outside genal furrow, withdistinct fine puncturing admixed with some coarseones. Ocelli dorsally just transected by supraorbitalline. A, just longer than A4 and justshorter than Au Gig. 28A). T, impressed in frontof distal margin. St, and genitalia (Figs. 28B-C);gonocoxite not grooved inside but regularly convergingtowards apex; gonostylus wider than inB. terrestris and B. lucorum,' apex of volsellausually as in Fig. 28D. Coat shaggy. Wings elongated,strongly infuscate. Body of large size.Colour pattern. As in the workers.Variation. Pile of face, pile of vertex with slightadmixture of yellow hairs. Fringes of hind tibiamay be whitish-tipped. The colouring is in generaldistinct, only very few specimens with slightlydiffuse colour pattern were examined.Distribution (Fig. 56)Norway. Occurring from inner district of Aust-Agder approximately 58" 30' N north to about66" N. A single queen from Troms, cf. the listbelow, is so far the only reliable record north ofthe Polar Circle. Another queen recorded in Hordaland,is the only observation outside the endemicspruce forest zone. In Southern Norwayrecorded to ll20 m s. m.Biotopes: Pastures, roadsides and meadows.A total of about 355 specimens was examined.List of localities. Ostfold; Eidsberg: Hole . Akershus:Brerum: Briten; Oslo: TRM; Aurskog-Holand: SotsjoenTRM; Fet: Bjorkflflten . Hednnrk ( HEs) :Hamar: ZMO. HEn: Trysil: Dammen, Enga, Sjoenden,Vestby; Amot: Glesubekken, Rena; Rendal:Solbakken SE Asheim; Tynset: Tynset; Engerdal:Bjorkoihl, Risbakken gird 750 m, Torhus. Oppland(Os).' Jevnaker: Randsfjord USU Z}ldB; Gran: Einavoll,Lander; Ostre Toten : Stubdalen; Sondre Land :Fluberg; Nordre Land : Dokka USU; Lillehammer:Jorstadmoen; Sor-Aurdal: Ellingseter 820 m, Skard700 m; Oyer: Aksjo 1000 m, Skei, Oyer, Astdalen900 m. On: Vang: Oye USU; Sel: Leirflata; Lom:Galdesand. Buskerud (Bd); Lier: Lokke gflrd; Krodsherad:Noresund; Kongsberg: Skollenborg. Bv; Sigdal:Kopseng, Nedre Eggedal; Nore og Uvdal: Gavlenseterll20 m, Rodberg, Uvdal, Uvdal stavkirke;Gol: Gol; At: ? loc. Z}.d:O. Telemark (fEy).. Drangedal:Grova, Naksjo, Omnes, Tomyra . TEi: Notodden:Gransherad, Tinnoset; Seljord : Svartdal; Kvitseid :Vrfldal; Nissedal: Lauvviki, Treungen ZMO; Fyresdal:Hegglandsgrend, Moland; Tokke: Holtet bro;Vinje: Krossen, Srerengrend, Amot; Tinn: Tinn,Atrstbygda. Aust-Agder (AAi); Evje og Hornes: Lauland;Bygland : Londal; Valle: Kvestad, Valle TRMZMB, Rygnestad. Hordaland ( HOi ).. Kvinnherad :Skeie ? ZZ July 1943 (Hans Tambs-Lyche). Miire ogRomsdal (MRy) ; Vestnes: Gjermundnes. Srir-Trtinde-Ias ( STy) ; Rissa: Gafsetflsen. STi: Roros: Myrmoen;Trondheim. Nord-Trrindelag ( Nfy).. Nreroy: Breiviken.NTi: Meriker: Tovmodal, Vardetun; Leksvik:Storvann; Verdal: Sandvika, Stiklestad; Inderoy:Kirkenesvflg; Overhalla: Gryten; Snisa: Bronstad,Telnes; Lierne: Kvelia, Seterhaug, Smfltjern; Hoylandet: Hoylandet; Namskogan : Brekkvasselv KMT,Finnvollan, Namskogan. Nordland ( Nsi).. Grane:Majavatn; Vefsn: Luktvann. Troms (TRi); Milselv:Olsborg ? 10 July 1955 (La).Corrections. Records from Aust-Agder (AAy):Tvedestrand: Skipvik (Strand 1898a), publishedas ,8. terrestrr,s var. d, were revised to B. lucorum.Sweden Wide, yot local distribution fromVdrmland, approximately 59o N, and northwardsthroughout the country. Three scattered recordsin Sodermanland and Uppsala indicate the extremesouth-eastern border area of the species.A total of about 550 specimens was examined,

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Studies on Scandinavtan Bumble Bees 53Erkheikki NRS, Haparunda ZMA ZML, HirvijiirviNRS, Kalix, Karungi NRS ZML, Kukkola NRS,Lulei NRS ZML, Lingtriisk, Mjolkudden NRS, NybyggetNRS, Pajala NRS ZML, Periijiivaara, Pite6,Pite havsbad ZMB, Sodra Sunderbyn, Overkalix NRS,Overtornefl, Ovre Svartlfl NRS. Lapland (As.Lpm.):Bingniis, Dorotea NRS, Risbiick NRS, VilhelminaNRS, Asele NRS ZCL. Ly. Lpm..' Ammarnds, LyckseleNRS, Malfl ZMU, Sickselberg NRS, SorseleZMU, Stensele, Tjultriisk BML ZML. Lu. Lpm.:Aktse, Kvikkjokk, Malmberget, Njunjes, SarekNational Park, Staloluokta, Ostansj o. T. Lpm.; AbiskoNRS, Bjorkliden NRS ZML.Unrevised records : Vcirmland; Rtimmen, Lesjofors(Wahlberg 1852) and Siiffle ZML (Ander in litt.).Correction. Records from Lapland (T. Lpm.):Abisko (Bengtsson l93l) were revised to B. lucorum(Linnaeus).Finland. Widespread throughoutthe countryexcept for Aland Isles and the extreme westernpart of the country (Elfving 1968).World distribution. Europe (Fennoscandia;Northern European USSR south to districts ofLeningrad, Kalinin, Ivanovo, Gorki, Kirov)Siberia Kamchatka Sakhalin Turkestan(Alma Ata) - Altai - Baykal - Northern Mongolia- Northern Manchuria - China (Koko Nor)(Panfilov 1957; Tkalcu 1967).BiologyFlight season. From the beginning of May tolast half of September. Queen : ?-25 Aug.;worker: 8 June-ZD Sept.: male:24 July-2O Sept.BOMBUS(s.s,) TERRESTRIS (LINNAEUS)Apis terrestns Linnaeus, 1758, p. 578 No. 30, typearea Sweden : Uppland. A queen LSL ! disagreeswith the usual interpretation of the taxon cf.below.(Bombus terrestris (Linnaeus); Dahlbom t832,p. 34 (partim) , 1837 (partim); Wahlberg 1854,1855 (partim); Thomson 1870 (partim), 1872(partim); Aurivillius 1903 (partim); Wahlgren1915 (partim); Ringdahl l92l (partim); Jansson1922 (partim) ; Krtiger 1956, 1958 ; Kruseman1959; Elfving 1960, 1968; Frid6n, Eskilsson &Bingefors 1962; Hasselrot 1962; Ander 1963,1965; Frid6n 1967. Apis virginalis Geoffroy inFourcroy, 1785, p. 450).Nomenclatural remarksA female in the collection of the LinnaeanSociety of London, labelled terrestris, has beenrevised to B. lucorum and the two subsequentspecimens in the same row, both of which arequeens with no labels, were revised to B. lucorumand B. terrestrr^r respectively. In referring to thedocumentary value of the Linnaean collectionp, 10, and to secure stability in nomenclature,B. terrestris, in sensu passed and present authors(Sladen l9l2; Schmiedeknecht 1930; Pittioni1939a; Kriiger 1950, 1954, 1956, 1958; Yarrow1959; etc.), is kept. The species is not rare inUppland county, the home area of Linnaeuswhich also is the type area.First Scandinavian recordsNorway. Hereby published for the first time.Sweden Uppland (Linnaeus 1758).Taxonomical remarksNorwegian records hitherto published as,B. terrestrisapply to B. lucorum (Loken 1960), cf. referencesto the synonymy of this taxon. Accordingto present revision, they may also include B. magnus.The mention of B. terrestris in Swedenconcerns ,8. terrestris, B" lucorum, possibly alsoB. magnus when referring to Southern Sweden,cf. authors (partim) in references to the synonymyabove, but only ,8. lucorum, possibly B. magnus,when referring to Northern Sweden, cf. referencesto fire synonymy of B. lucorum. Swedish B. terrestrisand B. lucorum were quite recently distinguished,cf. references above.Queen, workerMorphological characters. Malar space markedlyshorter than distal width. Labrum (Fig.l2A). Ocellar-orbital field (Fig. 12B). Distancefrom lateral ocellus to preoccipital ridge (d)about 514 longer than distance from ocellus toeye (e), mean d: f -_ 1.25; distance between ocelliI

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Studies on Scandinavian Bumble BeesitFig. 57. B. terrestrrs (Linnaeus). Legends as in Figs. 50, 54.lection of Hafsahl cf.not reliable.p. 7 , i. e. the labelling isSweden Widespread, local occurrence from theextreme southernmost coast northeast to about60'30' in Gtistrikland. It is uncertain whether theabsence inland in Southern Sweden, i. e. in Smfllandand in southern part of Viistergotland, is dueto insufficient investigations or indicates that B.terrestrus is conflned to the lowlands, areas below200 m s. m.Only recent finds in the northern border areasperhaps indicate a further moving of the northernlimit of the species.A total of about 850 specimens was examined.List of localities. Skdne: Alnarp, Arlov ZN{A, BokskogenZN{.{.[B, Bonderup ZM.B., Bromslla ZM.A,Brunnby, Bokeberg ZMB, Enslov ZMB, Falsterbro

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Studies on Scandinavian Bumble Beess7Taxonomical remarksThe complex cullumanus serrisquamaapollineus individually designated as species, hasrecently been considered conspecific (Kruseman1959; Tkalcu 1968; Yarrow in litt. l97l). Thethree taxa, having an allopatric distribution, areso far distinguished by the colour pattern onlyand the close relationship is emphasized bytreating them as subspecies.The Scandinavian population belongs to thenominate subspecies.DistributionScandinavia. Cf. the subspecies.World distributiorc. Europe (Southeastern England;Northern Spain; France; coast of theNetherlands, Germany; Denmark (Als Zll4.A,Bornholm VCA, Z]|dC); Southern Sweden;Poland; Hungary; in European USSR north todistricts of Vitebsk, Moscow, Ivanovsk, Molotovskand south to the northern limit of the aridstepped areas) Northern Caucasus Centraland Southern Ural Northern and EasternKasakhstan - southern part of Western SiberiaTurkestan (the mountains) Afghanistan(Yarrow 1954; Panfilov 1957; Tkalcu 1968).BOMBUS CULLUMANUS CULLUMAI\IUS(KIRBY)Queen, workerMorphological characters. Head slightly longerthan wide. Malar space about t/a the distal width,as long as A, or barely longer. Clypeus justtransverse, rather convex; disc of clypeus sparselypunctured in the centre, otherwise densely puncturedexcept for the impunctate anterior ateabetween well-defined coarsely punctured lateralimpressions. Labrum coarsely punctured ; labtalfurrow moderately defined, shallow, and inwidth about /s the labral width; labral tuberclesrather small, anteriorly flattened; labtal lamellaslightly to markedly wider than the furrow, themargin moderately curved. Mandible without ba-sal keel, with distinct sulcus obliquus, moderateincisur a lateralis, and with the bas al area stronglyalutaceous and dispersely punctured. Eyes directedin front of posterior mandibular condyle.Ocellar-orbital field with a broad band of coarseand fine punctures. Ocelli in front of supraorbitalline. A, nearly twice as long as distalwidth, markedly shorter thal1 Aa+ s. Aa about aslong as distal width, hardly shorter than A5.Mid-basitarsus with posterior distal angle roundedoblique, outside without tuft of long hairsnear the base. Hind tibia with outer surfaceinconspicuously alutaceous and with dorsal innerdistal angle distinct. Hind basitarsus with outersurface smooth, sparsely covered with shortbrownish pubescence, the posterior margin rathercurved. Surface of Tr-u smooth with ratherdense puncturing. T6 granulate with a more orless pronounced median longitudinal furrowcontinuing backwards in a keel, often beingconcealed by hairs; the distal edge thickened.Keel of St o rather feeble. Wings hyaline exceptthe infuscate distal ribbonless part. Coat rathereven, short.Queen measurements: N -6; Sweden: Skine;malar space: 0.59 mm ( +O.O: +0.01) range:0.55-0.63 mrn; 'radial length': 4.18 mrn (+0.t0+0.07) range: 3.95-4.40 mm; interalat width:5.04 rnm (+0.16+0.07) range: 4.78-5.28 mm.Body of medium to large size.Colour pattern Fringe of labrum and mandiblebrownish. Tn- u, Stn- u with reddish to brightferruginous hairs, those of the sternites usuallyof paler shade. Otherwise coat black.Variation. Extremely colour-stable. A singleworker with predominantly ferruginous corbicularfringes was examined.MaleMorphological characters. Head about as longas wide. Malar space about 415 shorter thandistal width and about as long as Ar. Mandiblebifid with a broad ventral and a small dorsaltooth. Eyes directed in front of posterior mandibularcondyle. Antennae rather long Fig. 30A;A, about 415 shorter than Aul Aaequal inlengthto the distal width and about 315 as long as Au.

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iStudies on Scandinavian Bumble Bees 59l''I r_I \ )/-(\,Pi )1--1"Ar:'.1' i, af(', )r ) ')iGtt"ry!v{, t-*$axf-\J\ - . t \t$--*$*oi-\tt \* 1N(Sf.rv/r\ r..((vlFig. 58. ,8. cullumanus (Kirby). Legends as in Figs. 50, 54.defined, coarsely punctured. Labraltuberclesanteriorly flattened or feebly depressed; labrallamella semicircular, not much broader thanlabral furrow. Mandible without basal keel, withno sulcus obliquus or this being indistinct andwith moderate to well-defined incisura lateralis(Fig. 16A). Eyes directed in front of posteriormandibular condyle. Supra-orbital line touchingor just above lateral ocelli (often transectinglateral ocelli in workers). A' twice the distalwidth or hardly so, about % as long as Aa+ s.Distal margin of mid-basitarsus posteriorly rounded.Hind tibia slightly or not alutaceous, dorsalinner distal angle feebly or not produced (Fig.16B); hind basitarsus with posterior marginstraight or slightly curved. Stu with short indistinctkeel or no keel. Wings evenly, ratherfeebly infuscate.There is a distinct difference in size betweenthe queen and the worker caste.

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Studies on Scandinavian Bumble Bees61treated as being conspecific, the latter consideredas a subspecies of the former, cf. p. 69. Therather srnall, well-defined area on the chartoccupied by B. cingulatus is moreover noteworthyand emphasizes the specific position ofthis taxon. B. cingulatus was, with exception ofthe designator and Boheman (1857), treated asa variety of B. hypnorum until Reinig (1936)re-established its specific rank.BOMBUS (PYROBOMBUS) CIIIGULATUSWAHLBERGBombus cingulatus Wahlborg, 1854, pp. 207 ,208-209, lectotype ? TCL ! selected by Ander(1967). Type loc. Sweden: Norrbotten: Nederkalix: Yttermorajiirv.(Bombus cingulatus Wahlberg; Wahlberg 1855 ;Boheman 1857, p. 17; Reinig 1936; Kruseman1959; Elfving t960, 1968; Laken i960; Ander1965; Bombus hyphorutn var. c Thomson 1872;Strand 1898a. Bombus hypnorum yar. cingulatusWahlberg; Friese 1902; Aurivillius 1903 ; Bengtsson1903 , 1904; Sparre Schneider 1909, 1918 ;Friese & Wagner I9I2; R.ingdahl tr915; Lundblad1924; Soot-Ryen 1925; Meidell 1934a;Laken 1949, 1950.)First Scandinavian recordsNorway. Troms (TRi): Mfllselv: Nordmo(Sparre Schneider 1909).Sweden Norrbotten: Nederkalix: Yttermorajiirv(Wahlberg 1854).Queen, workerMorphological characters. Malar space as longas distal width or just So, exceeding the lengthof A, but shorter than Ar* r. Disc of clypeusalmost impunctate centrally, otherwise with singlycoarse punctures in between sorne very fine onesand with well-separated lateral impressions whichare broad and with sparse, coarse punctures.Labral furrow moderate, varying in width butnarrower than /s the labral width ; labral tuberc-les usually densely punctured at the base, themicrosculpture smooth or indistinctly alutaceous.Ao just shorter than distal width or equal to it.Longest hairs in posterior fringe of hind basitarsusexceeding half the greatest width of thesegment. Surface of Tn-u with more or lessalutaceous microsculpture and moderate puncturing.T6 strongly alutaceous, dull, the puncturingrather dense, even, often with indicationof longitudinal median furrow and distally moreor less dimpled. Coat shaggy, less dense than inB. hypnorum.Queen measurements: N - 15; SE Norway;malar space: 0.58 mm ( +O.OZ +0.01) range:0.53-0.60 mm:''radral length': 3.75 mm (+0.11+0.03) range: 3.50-3.90 mm; interalar width:4.48 mm (+0.14+0.04) range: 4.18-4.65 trnm.Body of small to medium size.The smaller specimens of the workers weredifficult to distinguish from ,8. hypnorum.Colour pattern Pile of vertex yellowish-brownencroached by black hairs; dorsum of thorax,including narrow adjacent edge of episternum,yellowish-brown except for a more or less pronouncedinteralar band not reaching the extremeposterior margin of mesonotum. Pile of Tr,anterior margin or anterior lunate part of T2yellowish-brown; about distal half of Tu, entireTu- u, at least lateral paft of fringes of Stn-u,Sta whitish; otherwise coat black.Variation. Pile of vertex predorninantly or entirelyblack. Black hairs on dorsum of thoraxranging frorn a few single hairs on the disc to awell-defined interalar band. Occasionally, workershave no black hairs on dorsurn thorax, and awell-defined interalar band is only observed inqueens. Hairs on T1 more or less replaced byblack ones, preferably in lateral tufts. Occasionallyhairs on T'-2 completely black. White hairson T4 ranging from covering about distal halfof the tergite to a few hairs along the distalmargin.MaleMorphological characters. Malar space about aslong as distal width and as long as Arn r. Antennaeshort (Fig. 35A) i A' about % the lengthof Au, equal to Ao or just longer I An as long as5. Bumble Bees.

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IlIlIStudies on Scandinavian Bumble Bees6I- -16''t8"T39" ,t0"\-\-.J o/. IFig. 60. B. cingulatzs Wahlberg. Legends as in Figs. 50, 54.World distribution. Europe (Fennoscandia;Northern part of European USSR) - Northern,i * *-n*, Urals - Northern Siberia - Kamchatka. (Reinig t1939, p. 182; Panfilov in litt.).Biologytlishtseason. Beginning of May to beginningof September. Queen : 14 May - 7 Sept. ; worker:?-13 Aug.; male 12 June-t2 Aug.BOMBUS (PYROBOMBUS) HYPNORUM(LINNAEUS)Apis hypnorum Linnaeus, 1758, p. 579 No. 33,type area Sweden: Uppland. A queen LSL, labelledhypnorum, disagrees with the usual interpretationof the taxon. It was recently revised to.B.lucorum (Yarrow in litt. 1967).(Bombus hypnorum (Linnaeus); Sommerfelt1824-27; Dahlbom 1832, p. 50; Boheman 1844,

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Studies on Scandinavian Bumble Bees 65Queen measurements: N _ 20; SE Norway;malar space: 0.63 mrn ( +O.O: +0.01) range:0.58-0.68 mm ; 'radial length' : 4.44 mm ( +0.13+0.03) range: 4.15-4.64 mm; interalar width:5.15 mm (+0.18+0.04) range: 4.75-5.45 mm.Body of large size.Colour pattern Pile of vertex predominantlyyellowish-brown. Dorsum of thorax and adjacentedge of episternum yellowish-brown. Hairs ofTr, anterior lunate part of T2 predominantlyyellowish-brown. At least posterior half of T n,entire Tu- u, and at least lateral fringes of St+-,(or n- u) whitish-haired. Otherwise coat black.Variation. Pile of T1 ranging from entirelyblack to entirely yellowish-brown. Yellowishbrownhairs on T, more or less replaced by blackones, occasionally hairs on T, entirely black. ,A. fewindividuals with the yellowish-brown part of coatreplaced by dull clayish-brown or even darkbrown hairs were examined.A tendency to geographical variation in Scandinaviawas not recognized. More queens withhairs of Tr-, entirely black were, however,exqmined from Northern Scandinavia than fromthe southern part of the peninsula.MaleMorphological characters. Malar space as longas distal width or just longer, longer than Aubut shorter than Ar* r. A, longer than A*, equalin length to Au or just longer (Fig. 34A)i An aslong as distal width or just longer. Longest hairsin posterior fringe of hind tibia varying fromlyz to almost twice the greatest width of thesegment. Hind basitarsus about 3 times longerthan the greatest width, longest hairs in posteriorfringe about as long as this width. St, (Fig. 3aB);gonostylus and volsella (Fig. 34C); penis valve(Fig. 34D). Coat shaggy. tsody of medium size.Colour pattern Beard brownish. Pile of facebelow antennal sockets with variable admixtureof dull yellow hairs. Pile of vertex entirely orpredominantly yellowish-brown. Dorsum of thoraxand at least adjacentpart of episternum, T, atleast anterior lunate part of T, yellowish-brown.Distal margin of To, Tu-, whitish-haired. Venterof head and thorax, trochanters, fernora, andfringes of gastral sternites with variable admixtureof whitish hairs. Otherwise coat blackVariation. Hairs on Tr-, ranging from entirelyblack to entirely yellowish-brown. Whitish hairson the tergites rnay be reduced to an admixturewith black hairs on To-z only.Distribution (Fig. 61)Norw,ay. Widely distributed throughout thecountry north up to about 70' N. In SouthernNorway occurring from sea level to regio subalpinaand sporadically also recorded in the lowerbelt of alpine zone. Further north confined tothe conifer forest and only once observed on theextreme coast north of the Folar Circle, viz. onthe small island of Hushattoy in the archipeiagojust north of 70' N. Also in Southern Norwaythe abundance is greater inland than along thecoast. Recorded to tr000 m s.m.Biotopes: Meadows, gardens, orchards, roadsides,pastures.A total of about 1520 specimens was examined.List of localities: Ostfotd.' Hvaler: Asmaloy, Kirkoy,Sondre Sando; Krikeroy: Tangen, Odegird;Onsoy: Dypeklo, R-orvik, Torp; Sarpsborg : ZMO;Halden: Blakmose, Kuletjern, Sponvika; Aremark:Aremark, Gjeddetjern ; Marker : Damholtet, Rodenes ;Rakkestad: Rorvik; R.6de: Fuglevik, Oven; Moss:VCA, Jeloy VCA; Eidsberg: Holm; Trogstad: Monsterbru. Akershus: Frogn: Drobak TRM VCA; Asker:Lushattdalen; Brerum: Lysaker Z}l..{O; Oslo: TCLTRM Z}l..Ir_ ZMO; Aurskog-Holand: Bjorklangen,Gangsnes, Mo, Stigen, Ostegd.rd; Fet: Bjorkffiten;Skedsmo: Strommen ZMO; Hurdal: Tomte ZMO;Nes: Grinkelsrud; Eidsvoll: Eidsvoll ZMO . Hedmark( HEs ); Eidskog : Bolfoss, Gjelarisen ; Kongsvinger :Eidsberg; Sor-Odal : ? loc. ZMO; Grue: Finnskog;Hamar: ZMts ZMO. HEn: Trysil: Damrnen, Enga,Sjoenden, Vestby; Amot: Glesubekken, Amot, AsetZMO; Rendal: Solbakken SW Asheim ZMO; Tynset:Tyldal, Tynset, Ulsberg; Engerdal: Bjorkoihl,Gutuliseter NE Drevsjo 700 m, Torhus . Oppland( Os ).' Jevnaker: Randsfjord USU; Gran: Gjervika,Granvoltren, Rognstad; Ostre Toten: Stubdalen;Nordre Land: Dokka USU ZM.B; Lillehammer: Stor-Hflve; Nord-Aurdal: Knutshaugen SE Fagernes,Ler:a, AUjor; Gausdal: G6soya 875 m; Oyer: Aksjo1000 m, Oyer, Astdalen 900 m; Ringebu: Ringebu.On: Vestre Slidre: Oken; Oystre Slidre: Skammestein700 m; Vang: Oye USU ZMB; Sel: Heidal TRMZMB, Leirflata, Mysuseter 900 m, Otta ZMO; Vflgi:

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Studies on Scandinavian Bumble Bees 67Moland; Tokke: Amdals verk; Vinje: Krossen 700 m,Amot; Tinn: Vilen, Vier W Rjukan . Aust-Agder( AAy ).' Gjerstad: Fiane; Vegflrdshei: Ljostad; Tvedestrand:Eidbu, Nes Verk TRM, Strengereid; Arendal:ZMO; Motrand: Dal, Holrnsund, Tverrdalsoy, Voje;Tromoy: Bjelland, Grimstad; Landvik: Horte, Molland,Tonnescl; I-illesand : Blikksund, Kjopmannsvik,Amli; Birkenes: Langemyr, Rosseland; IvelanC: Fosstvedt.AAi: Evje og Hornes: Hornes, Syrtveit; Gjovdal:Krossbekk, Saurbekkflflt, Arh; Bygland: Londal,Ose; Valle: Kvestad, Valle; Bykle: Bykle. Vest-Agder ( VAy ); Kristiansand : Holskogen ; Sogne :Langeneset, Sbgne, Aros; Mandal: Mandal, Sflnum,Tuftenes; Marnadal: Finsland, Royseland, Utkjrer;LyngCal: TRM; Kvinesdal: Feda, Gjemlestad, Solis,Oy.; Farsund: hlellesfl, Lista Fyr, Lodshavn; F'lekkefjord:Ersdal, Espetveit, Flikkeid SMS. VAi: Audnedal:Sveindal; Aseral: Espelid, Rosseland, ,&seral.Rogaland (Ry); Sokndal: Bu, Hauge i Dalane, Sogndal,Ana-Sira; Eigersund : Egersund, Eigeroyeidet,Skjerpe; Bjerkreim: Hovland, Malrneim; Hfl: Ogna;Klepp: Bcrsheim, Klepp, Orre, R.eve, Vik, Oksnevad;Tirne : Mossige; Gjesdal: Algard; Sandnes : Brattebo,Ffana, Myrland, Sandnes; Scla: Gimre, Sclastrand,Tannanger; Stavanger: Forus, I{inna, Lindoy, StavangerSMS TRM ZMB, Sunde, Vflland; Strand:Tau; Finnoy: Judaberget, Sjernaroy; Rennesoy:Datre; Kvitsoy; Karmoy: Sreveland. Ri; Forsand:Forsand, Meling; Hjelmeland : Hjetrmeland, Kvamrne,Vadda, Ardal; Suldal: Jelsa, Sand, AirnjoAtonuten700 m; Sauda: Sauda. Hordaland (HCy); Bomlo:Royksund; Olen: Dcmmersnes; Fitjar: Fitjar, Risoy;Austevoli: Karlsoy; Os: Gullhotrmen, Lepsoy, Lysekloster,Nordstrono, Ravneberghaugen; Samnanger:Hoyseter; Fana: Biol. st., Birkelundsbakken, Blomsterdalen,Engjavik, Espeland, Fantoft, Lonninghavn,Milde, Ridalen, Skeie, Skipanes, Skjold, Steinsvik,Stend, Storetveit, Srloy, Titlestad, O vre Krflkenes ;Sund: Bjelkaroy, Glesvrer, Steinstrand, Tyssoy; Fjell:Bjoroya, Eide, Sclsvik; Laksevig: Alvoy, Bjorndalspollen,Hilleren, Storingaviken; Bergen; Askoy: Ask,Davanger, Fferdla, Marikova; Asane: F{aukedal, Astveitskogen;Osterdy: Fitje, Hamre, F{avratun,Kleppe, I-onevig, Nj6stad; VaksCal: Eidslandet;Meiand: Dalst5; Radoy: Vflgsbctten; Lindfls: Lauvfls,Srvrflsvig, VelanC. HCi: Etne: Oslandsvflg; Kvinnherad: Lio, Ljosmyr, Seimsfoss, Skeie, Varaldsoy;Ullensvang: Alvsaker, Espe, Fresvik, Jflstad, Kinsarvik,Lofthus, Tveito, Ovre Eidljord; Kvam: Kvamskogen,Nordheimsund, Steine, Oystese, Alvik, Asheim;Voss : Bomcen, Hjelle, Rong, Tyrlingen; Granvin:Granvin VCA ZN{{ts, Trfl; Ulvik: Bergc. Sogn ogFjordane ( SFy ).' Gulen: Innre Takle; H3rllestad :Hyllestad, Lekva, Skivenes; Gaular: tsygstad, Vikum;Flora: Floro, I{inn, Reksta, Skorpeidet; Naustdal:Frammarsvik; Forde: Fl6ten, F[atrtrbreim; Jolster:Harnar; Gloppen : Hjcrtset, F{ope, Hyen; Eid: Heggjabygda,Kjolsdale:r, Naustdal, Nordfjordeid, Stflrheim,Vedvik; V5gsoy: Brygg.ja, M&ioy, Roysen; Selje:Leikanger. SFi.' Vik: Tistel, Vik; Aurland: Kvamma-dalen 950 m, Vatnahalsen 800 m ; Lnrdal: Bergstolen,Grote; Sogndal: Kaupanger, Slinde, Sogndal; Leikanger:Hermansverk, Leikanger; Balestrand: Balestrand,Brekka, Fjrerland, Flesje, Horpedal, Kvamsoy,Mundalsdalen; Lusrer: Jostedal, Skjolden; Stryn:Hornindal, Stryn, Videdalen. Mdre og Romsdal( MRy); Volda: Bjorkedalsvann; Orsta: Lyngsrolvann,Orstavik, Oye; Hareid: Hjorungavig; Ulstein:Ulsteinvik; Alesund : Ase; Vestnes : Gjermundsnes ;Molde; Aukra: Falkehytta; Averoy: Vevang. MRi:Stranda: Geiranger NMW ZMts, Vollset; Norddal:Valldal; Sunndal: Jordalsgrend; Surnadal: Stangvik,Ovre Surnadal. Srir-Trdndelag ( SZ.y).. Hitra: Hamn;Rissa: ? loc. VCA, Gafsetflsen, Hasselvika, Stadsbygd,Sotvik; Orland : Bakken ; tsjugn : Kotengsvann ;Afjord:A. Sn; Oppdal: Driva st., Oppdal; Rennebu:Sorflfl; Midtre Gauldal: Storen; Alen: Reitan; Roros:Myrvann 700 m; Selbu: Medbus; Trondheim: KMTVCA ZN{B. l{ord-Trdndelag (NTy).' Flatanger: Sor-Flatanger; Otteroy: Hovika I(MT; Namsos: TRM,Strandmoen; Nreroy: Breiviken, Saltbotn . NTi: Meriker:Merflker, Tovmodal; Stjordal: Draveng, Hegrast. ; Leksvik: Storvann; Verdal: Sandvika KMT ZMB,Stiklestad, Vuku; Inderoy: Kirkenesv6g, Skjelvflgen ;Steinkjer: Z}l.f.A, Sem; Namdalseid: Hovika KMT;Grong: Fjerdingen, Grong SMS, Trangen; Snisa:Brenstad, Hegge, Telnes; Lierne: Seterhaug; Hoylandet:Hoylandet, Skilleberget, Skogenga; Namskogan: Brekkvasselv KMT, Finnvollan, Namskogan.l,{ordlqnd ( Nsy ).' Bronnoy : Flatmo TRM ; Nesna :Hamaroy. l/si.. Grane: Bifjellmo, Majavatn, Rotstokkmobekken; Vefsn : Store Bjornflvann ; Sor-Rana:Bleikvassli; Rana: Aenget, Mo, Rovassdal TRM;Saltdal: ? loc. ZN[O, Drageid, Junkerdalsura, Solvflgfjell,Storjordet TRM . Nnc): Ankenes: Beisfjord ZMA.Troms ( TRy ).' Lenvik : Finnsnes ; Tromso : TRMVCA, tsreivik SMS; Karlsoy: Hushattoy 5 VH 25 July1935 (Soot-Ryen) TRM. TRi: Bardu: tsardu, Stromsmo;M6lselv: ? loc. VCA ZMB ZMO, Bjerkeng TRMZMB, Kirkesdal, Moen TRM, Nordrno TRM VCAZl4B, Olsbcrg NMW ZMB, Rundhaug SMS ZM-r_ZMC; Balsfjord: ?loc. TRM, Storsteinnes VCA;Nordreisa: Nedrefoss, Sappen . Finnmark ( Fi ) : Alta:Jotkajavrre TRM; Karasj ok. Fn ' Forsanger: Hamnbukt.Fri: S5r-Varanger: Kirkenes TRM, LangfjorddalenTRM.Unrevised records are not included here except forthe following : Troms (f Ri ).' Lyngen: NE Jegervann,NW Furuflaten (Douglas in litt.).Sweden Widely distributed throughout themainland except in regto alpinalarctica. Moreover,occurring in 6lanC but not recorded inGotland.A total of about 120C specimans was exarnined.List of locaiities. Skine: Barkflkra, tsjiirred, BonderupZlu4 B, BokebergZN.[B, Bf,stad, Falsterbo NR.S,

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Studies on Scandinavian Bumble Bees 69Scandinavian species of this subgenus (Bengtsson1903; Hasselrot 1960 Table 21, 1962).Flight season. From the end of March to theend of September. Queen: 25 March-z4 Sept.;worker : 22 April-lO28 Sept.Sept.; male: 22 June-BOMBUS (PYROBOMBUS) JONELLUS(KTRBY)Kolari. A queen IZBI, labelled Schweden:Malung 7.31 Pfeiffer, has a red type label attached.The locality (Dalarne: Malung) disagreeshowever with the type locality published togetherwith the designation. Bombus jonellus suecicusFriese; Richards I 933. Bombus jonellus subboreAlisRichards, 1933, p. 64 ( - Bombus jonellusatrocorbiculoszs Vogt, lgll (partim), holotype !(Smith coll. originally named nivalis Dahlbom)BML!, type area Norway; Kruseman 1959).Apis jonella Kirby, 1802, p. 338 No. 90, holotypeC KCL! labelled type by Milliron (1960) First Scandinavian recordsand holotype by Yarrow (1968), type loc. England:Norway; Finnmark (Fv): Alta: BossekopLondon ('prope Londinum').(Bombus jonellus (Kirby); Dahlbom 1832, p.(Dahlbom 1832, p. 39).Sweden: Sk6ne, SmAland, Vdstergcitland, Lapland43; Strand 1901, 1910; Friese 1902; Aurivillius(Dahlbom 1832, p. 39, 43); queen and1903; Muchardt 1904; Sparre Schneider 1906,1909, 1918, p. 15; Wahlgren 1908; Ringdahl1915, I92l; Lundblad 1924; Soot-Ryen 1925;Gaunitz 1929; Bengtsson l93l; Jansson & Sjoberg1932; Hellen 1933; Meidell 1934a, 1946; Wexelsen& Skire 1934; Barendrecht 1941; tsrinck &Wingstrand 1949 ; Laken 1949, 1950, 1954,, 1966c;worker are treated as ,8. scrimshiranus and maleas B. jonellus. Though voucher specimens so farhave been traced from Lapland only, records fromthe remaining counties might be correct, exceptperhaps those fromVristergotland, as this areawasnot represented in other collections being revisedeither.Brinck 1951 ; Forsslund 1951 ; Elfving 1960, 1968;Erlandsson 1960 ; Ander 1963, 1965; Friden 1967 .Apis scrirnshirana Kirby, 1802, p. 342 No. 92, PKCL ! selected as type by Milliron (1960), isdisputed by Yarrovv (1968). Bombu scrimshiranus(Kirby); Dahlbom 1832, p. 391' Boheman 1844,p. 95; Wahlberg 1852, 1854, 1855; Siebke 1870;Thomson 1870, IB72; Aurivillius 1887; Schoyen1887; Ner6n 1892; Sparre Schneider 1895b, 1898 ;Strand 1898a, 1898b ; Lie-Pettersen 1901, 1905,Taxonomical remarksThe Scandinavian population is recognized asa subspecies, B. jonellus subborealis Richards.B. I. var. flavicolor Friese, 1909, B. j. suecicusand the sparreschneideranius form of Vogt (1911),all of which designated to Scandinavian forms,are infrasubspecif;c. A number of colour formsI9A7; Bengtsson 19A4, 1908. Bombus autwmnalis: presented to Sweden: Lapland (Lu.Lpm.): VirihaureZetterstedt, 1838 nec Fabricius, 1793. Bombusmartes Gerstaecker, 1869, p. 137 No. 4, lectotype? IZB! hereby selected, the labels are as(Brinck 1951), reflect the vast variationswithin a limited area as otherwise is the casealmost everywhere throughout Scandinavia.follows : 1) Ober-B ayern, Kreuth. 4-8. GerstaeckerS (printed); 2) red, type label ; 3) martesGerst. a? (handwritten); 4) B. jonellus A. Lskendet. 1965; 5) B. martes Gerst. lectotype designatedA. I-aken 1965. Type loc. Germany:Ober-Bayern: Kreuth. Bomhus hortorwm: Siebke1880 (partim). Bombus pratorum var. .ionellusDistributionFennoscandia. Cf. the subspecies.World distribution. Europe (British Isles; Iceland;on the continent ranging from NW Spain,(Kirby); Friese & Wagner 1909 ; Alfken 1913.France (the mountains), Belgium (Ardennes),Bon:bus pratorum - jorcellus var. suecicws Friese, the Alps,'phe Balkansi and north to Arctic Fennoscandia;I9ll, p. 572, type loc. Sweden: Norrbotten:European USSR (most freqr.lent in the. t

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Studies on Scandinat,ictn Bumble Bees7Ternerged individuals, or more or less replacedby black ones. In melanic specimens hairs ofventer and legs turn black and the amount ofyellow hairs is n lore or less reduced.Distribution (Fig. 62)Norway. Widely distributed throughout theentire country, from the extreme southern andwestern archipelagoes to the arctic coast. Ingeneral confined to heather and adjacent meadowsoccurring from sea level to regio alpinalarctica. Cultivated areas are avoided and thespecies. therefore only rarely recorded at thewestern side of Oslofjord. Sparre Schneider (1918,p. 15) and Meidell (1934&, p. 127) point out theabsence of the species in southeastern lowlands.F{owever, recent observations show frequentoccurrences also in this part of the country, btttrestricted to biotopes left between agriculturalareas. In Southern Norway recorded 1300 m s.{tl.tsiotopes: Calh.ma and Vaccinium heaths, rneadows,pastures, lnountains-meadows, etc.A total of about 2800 specimens was examined.List of localities. istfold.' Hvaler: Akeroy, Festningsoy,Herfol, Kirkoy, Sondre Sando; Onsoy: Dypeklo,Rarnseklo, Torp; Halden: Asak, Blakmose,Hakelund, Halden, Kuletjern, Nordbakke, Veggesdal;Aremark: Darnholtet, Gjeddetjern, Mymosetjern,Skodsberg; Marker: Damholtet, Dypedal,Flagghytta TRM , Jhval, R.odenes, C rje; R.akkestad :Levernes, Rorvik; Rflde: Tom; Eidsberg: Holrn.Akershus: Frogn: Drobak VCA, Scnderstoa TRM;Asker: Lushattdalen; Brerum: Britan; Oslo : ZMA,Strerndal TRI\4; Ralingen: Fjerdingsby; Aurskog-Holand: Bjorklangen, Loken, Mc, Skogen, Osterg6rd;Fet: Bjorkflflten; Nes: Grinkelsrud; Eidsvoll:Feiring. Hedmark (I{Es); Eidskog: Bolfoss, Gjelar-6sen; Kongsvinger: Eidsberg; Sor-Odal: ZMO; Hamar:ZMO. HEn: Trysil: Dammen, Enga, Sjoenden;Rendal: Solbakken SE Asheim ZMO; Folidal: Daien800 m, Krokvik 750 m; Tynset: Dallvang SMS, {Jlsberg;Tolga-Os: Hodalen 770 m; Engerdal: Bjorkdihl,Gutuliseter 700 m, Jyltingsvola 850 m, Torhus . Opp-Iand ( Os ).' Gran : Einavold, Gjervika, Rognstad ;Nordre Land: Dokka USU; Etnedal: Bruflat; Ncrd-Aurdal: Flya 850 m, Hovda 760 m, S:ebuoygard NFagernes, AUjor; Gausdal: Gisoya 875 n"r; Oyer:Aksjo 1000 nI, Oyer, Astdalen 900 m, Astclalsho1100 m; Ringebu: Flaksjoen 910 m, Ringebu, Venabyed.On: Vestre Slidre: Kinnholt 1000 m; OystreSlidre: Skammestein; Vang: Eidsbugarden 1100-1200m, Grunke 950 m, Oye USU, Oylo ZMO; Fron: ? loc.ZMO, Eldflseter 950 m; Sel: Klaksh&ug, Mysuseter900 m; Vigfl: Gjendesheim 1000 m BML ZMts,Hindseter 900 m, Klones, Leirungen 1050 m, Lemonsjoen860 m, Randsverk 800 rn, Sjodalsbu 950 m,Skjerva-Jondal 800 m, Storhaugseter 900 m, Valdresflya1150 m, Vflgflmo Z}dB Z}d}{, Ovre Sjodalsvann950 m; Lom: Galdesand, Kvan'lrne, Leirdalen 900 m;Skjak: Grotli 900 m, Skj6ksetrene 800 rn, Strynsfjellet1100 rn; Dovre: Domb6s ZMU, Dovrefjetrl 1000 mVCA ZMO, Fokstua 930 m Z N{4B_ ZMO, Vfllisjo950 m ZMO. Buslcerud ( Bd ).' Hurum: Pinadalen;Lier: Sylling; Modum: Snarum; Flesberg: HvilaNNW Kongsberg, Lampeland, Oydegarden NNWKongsberg; Kongsberg: Skollenborg. Bv: Sigdal :Haglebu, Kopseng, Nedre Eggedal, Nordbygda; Noreog Uvdal: Gavlenseter Il20 m, Norstebo, Solheiinstul1000 m, Sbnstervann 1050 m, Tunnhovd 850m,Ovre Hein 1150 m; Gol: Randalseter 940 m; Hemsedal:Kjolen 1000 m, Lykkja 900 m, Muren; Al: ? loc.ZMO, Bergsjo 1100 m, Laudalseter II50 m, Tvist1080 m, Vallehalle ll20 m; Hol: Dagali 900 m NRS,Geilo 800 m VCA Z}u.f'B, E Halne 1150 m, Haugastol1000 m VCA Zl,{4B, Kvasshogda 1200 ffi, N l-illevann1200 rn, Nygfird 1000 rn, Vikastolen 1100 rn, Hovet,Nyestolen 1050 m, Olsenvann 1100 m, Ustaoset1000 m. Vestfold; Lardal: Styrvoll; Tjorne: Havna,Kjrere. Telemark (TEy).' Barnble: AUv; Drangedal:Omnes. TEi: Seljord: Norsteteig, Svartdal; I{viteseid:Eidstad, Vridal; Nissedal: Treungen ZMO; Fyresdal:Hegglandsgrend, Vfoland, Veum; Vinje : Bossboen950 m, tr(romvika 930 m, Krossen 700 in, R.aulandkirke 700 m, Urdbo 700 m; Tinn: Borsjoen1150 m, Mosvatndammen 900 rn, Sneiseli 960 m.Aust-Agder ( AAy ).' Gjerstad : Fiane ; Tvedestrand :Nes Verk TRM, Oster&; Arendal; Moland: Voje;Tromoy: tsjelland; Fjrere: Fjrere kirke ZMO; Grimstad;Landvik: Holvika; Liliesand: Kvise, Ulvoysund;Birkenes: Sagen; Iveland: Frikstad, Fcsstveidt,Ivedal. AAi: Evje og Hornes: Hornes, S),rtveit; Bygland:Londal, Ose, Setesdal TRM; Valie: Ryg;restad,Valle TRM; Bykle: Breive 700 m, Breivann 900 m,Bykle, Hovde bro 700 m, Flovden. Vest-Agder ( VAy) :Kristiansand : ZMO, Stromme; Vennesla: OvreboZMC; Sogne: Scgne; Mandal: TRM ZNddf.B., Hogganvik;Marnadal: Finsland, Gretteland, Utkjer; Lindesnes: Ramsland, Spangereid ; Kvinnesdal : Forland,Gjemlestad, Scl&s, Oy.; Farsund: Lodshavn; Flekkefjord:Espetveit, Flikkeid SMS, Langevatn, Sandvann,Tennmei:li. VAi : AudneCal: Sveindal; Hregebostad:Bryggesfrk; Aseral: Espelid, Rosselancl, Aseral; Sirdal:Dcrgeseter, Dyneefjell 900 m, Handeland, Sinnes,Adneram BOC m. Rogaland (Ry).' Lund: Eikjelandsdalen,Ljosvatn, Iv{oi; Sckndal: Nesvig, Sogndal,Amct; Eigersund: Fotlandsvann, Gydal, Klungland;Bjerkreim: Asseiinvann, Hovland, Ivedal, Malrneim;Hfl: Aniqdai, Ogna, Salte bro, Sirevflg, Stolen, Vatnafflo;Klepp : 13Srsheltn, Orre, Orrevann, Reve, Vik,Oksnevad; Time: IV1elsvann, Time; Gjesdal: Ateflrd;Sandnes: Brflstein, Flcysvik, Hommersfirk, Myrland;

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Studies on Scandinavian Bumble Bees 73Solsvik VCA; Laksevig: Bjorndalspollen, Storingaviken;Oppdal, Skansen; Atren: Reitan; Roros: AdamsvollBergen: TRM Zh{4B; Askoy: ?loc. VCA, 820 m, Evavoll 700 m, Myrmoen 700 m, NaustervollDavanger, Erdal, Hanoytangen, Hegernes VCA, 700 m; Trondheim VCA ZMB; Skaun: SornypvannHerdla, Jacobsoy, Kleppesto, Asebo; Asane: Golfbanen,KMT; Orkdal: Hemnekjolen KMT, Scgnlia KMT,Haukedal, Jordalen, Steinesto, Tellevik; Oster-Sorvann KMT. Nord-Trrindelag ( NTy).' Flatanger:oy: Fitje, F{amre, Kleppe, Kvamffre, Nj6stad, Rakaneset;Bjornoyv&r KMT; Namsos: Strandmoen; Nreroy:Vaksdal: Eidslandet; Meland: Io, Kjeken;Oygarden: Buneset, Straumoy, Turoy; LindAs: Fosse,Breiviken, Dalene, Garmannsviken, Kvalviken, Saltbotn.NTi: Stjordal: Hegra st.; Leksvik: Storvann;Fyllingsfles, Kolis, Lauvis, Mykevoll, Srevrisv&g, Verdal: Billingflaten, Sandviken, Vergien KMT;Vollom . HOi: Kvinnherad: Gjermundshaffin, Jemtelandsvann,Steinkjer: Seim; Grong: Ekkerseter, Fjerdingon,I-josmyr; Odda: Austmannli 800 m, Dalen,Horra 1000 m, Mittleger 1150 m, Seljestad, ValldalTrangen; Snflsa: Bronstad, Hegge; Lierne: Kvelia,Seterhaug, Smfltjern; Hoylandet: Hoylandet, Skilleberget,900 m; Ullensvang: Bjoreidalshytta II00 m, Fresvik,Skogenga; Namskogan: Finnvollan, Nam-Fossli 750 m, Gryteskard 850 m, Grythorgen 900 m, skogan ; Royrvik : Bjorkhaug. Nordland ( I{sy ).' Heroy:Hallaskard 1100 m, Hjolmodalen, trsdalen 850 m,Syd-Heroy TRM VCA; Meloy: Kunna, Reip6,Kvanndal, Mebcdalen, Stavali 900 m, Storliseter Storglomvann TRIvI ; Gildeskfll : Finnes, Gildesk6l,900 m, Sysendalen 750 m, Tindholen n5A rn, Veivann Gilset, Jelstad, Skauvoll, Storvika, Sor-Fugloy; Bodo :1150 rn, Viveti 820 m, Ovre Eidfjord; Kvam: Djonne, Bodo BML, Falkflaugdalen, Karlsoyv&r, Lopsviken,Kleppfjell ZMO, Kvamskogen, Sjuseter, Alvik USU, Skau. l/si; Hatfjelldal: Krutvann; Grane: Bflfjellmo,Asheim; Voss : Armot, Bomoen, Hamlagro 600 m, Grane, Majavatn, Rotstokkmobekken, Skomstad;Hangurfjellet 800 m, Mjolfjell, Nesheim, Rong, OVseter;Vefsn: Ravassbakken, Store Bjornflvann; Hemnes:Granvin: ?loc. VCA; Ulvik: Finse 1200 m Finneidf.jord, Krokselvrncen; Rana: Bjellflnes ZMO,VCA, Loyningseter, Ulvik. Sogn og fiordane (SFy): Krokstrand, Randallsvollen, R.ovassdal TRM, Umskarstjern,Gulen: Takle; Solund: Faroy; Hyllestad : Botn, Eide,Aenget; Beiarn: Gr&tfldaien TRM; Salt-Hatleim, Hyllestad, Skivenes, Skor, Yttre Dale, Atefjell;Hoyanger: Lavik ZMts ZNd4O, Stoldalen; Gaular:Bygstad, Sande, SXotten, Vikum; Fjaler: Strandenes;dal : ?loc. ZMO, Junkerdal, Jr-rnkerdalsura, Lonsdal,Semska, Storjordet TR.M; Fauske: Fauske, KjeldvannBML, Sulitjelma BML . l{nd: Sorfold: Bonn&, Tennrova,Askvoll: Aralden, Raudoy, Rauoy; Flora: Askvann;Hamaroy: Brennvik, Fjerdevann, Hamnes,Grindholmen, Havrenesvig, Kinn, Nekkoyna, Hansbakk, Hauk6s, Innhavet, Oppeid, Sandnes,Rognaldsvig, Skorpeidet, Trollstua; Naustdal: Furehaugstolen,Skutevik, Strinda, Tranoy; Tysfjord : ?loc. ZN ddf.O;Horstad, Naustdal; Forde: Flflten, Hall-breim; Jolster: Flamar, Vassenden; Gtroppen: Hjortset,Hyen, Lote, Sandane; Eid: Nordfjordeid, Vedvik;Ballangen: Ballangen, Dyrhaug,Heggemoen, Hesjeli, Myrbakk,nes : tseisfjord ZNd.A, Bjerkvik,Elvesletten, Fors&,Skarneselva; Anke-Seterfjell; Evenes:Vigsoy: Hagen, Krflkenes, Kvalheim, Langenes, Maloy,Totland; Selje: Dalsto, Ervik, Lekanger, Sandvikseidet.Jansbakk. IYnv: Vestvflgoy: Egguffi, Knutstad, Skulbru,Starnsund, Siorhella, Torvclalsvann, Ure, Valvangen,SFi; Vik: Hflvfls, Vik; Aurland: Aurlandsberg;Vflgan: Store Molla, Svolvrer Z\AA; Andoy:Flf,m, Steinbergdalhytta 1000 m, Vassbygdi, Andenes SMS TRh4, Bjornskinn, Dverberg, Myre,Vatnahalsen 800 m, Ovstebo 800 m ; Lnrdal: BergstolenRisoyhavn SM S ZMB. Trcms ( TRy ); Kvrefjord :650 m, Breistolen 1050 m, Horgje, Lrerdal Bcrkenes, Vik; tsjarkoy: ? troc. TR.iVX; tr-envik: Finns-ZMO, Maristova 800 m; Sogndal: Sogndal; Leikanger:nes; Tromso: Bctnhamn SVtrS, Rarnfjord TRM,Hermansverk; Balestrand : Brekka, Flesje, Tromsdal VCA TRM ZMO, Tromso BML TRMTuftedalen; Luster: Fflberg, Fflbergstolene 600 m, VCA ZMO; Karlsoy: Hushattoy TRM, MflkeskjarTurtagro 900 m TRM; Stryn: Hornindal, Hornindalseter,TRM, Vanno TRM; Skjervoy: Arnoy, R.avelseidet,Stryn, Stoverstein, Videdalen, Videseter 600 m. Uloybukt. TR.i: Skflnland: Boltfls, Lavangseid; Gra-Mrire og Romsdsl ( MRy ).' Sande: Gurskevig, Kobbervik,tangen: Gratangen ZMA; Bardu: Bardu, Dorum,Larsnes, Aram; Volda: Bjorkedalsvann, Fol-Dypdal; Mfllselv: ? loc. TRM VCA ZMO, Andselv,kestad; Hareid: Hjorungavhg, Mork; Ulstein: Ulsteinsvik,Bjerkeng TRM Z}v4B., Frihetsli TRM, Kirkesdal,Vonheim; Heroy: Djupsvik, Runde, Stokk-sund; Alesund: Alesund, Ase; Orskog: ? loc. ZMO;Maukeil Zl.d O, Neset, Nordmo TRM ZFu .{O, Olsborg,Rundhaug SMS Z}.dF_ ZMO, Sorli, Trosdal; Ealsfjord:Vestnes: GjermunCnes; Molde; Aukra: Aukra kirke,? loc. TRM, tsl&fjell TRM, Lanes TRVI, Stor-Falkehytta, Lovik, Kolholmen, Rindarcy; Frrena: steinnes TRM, Takvann VCA ZldH_; Nordreisa:Gjendem, Hollinholm; Snnola: Andholmen. MRi:Stranda: Bjordal; Norddal: Valldal; Sunndal: Jordalsgrend;Surnadal: 6vre Surnadal. Srir-TrdndelagBakkeby, Gappruselv, Gapprusfjeilet 700 m, Nedrefoss,Sappen; Kvrenangen: Burfjord. Finnmark (Fv):Kvalsund : Cint

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Studies on Scandinavian Bumble Bees 75grading zone and the territorial linemits of B. j.subborealis and the nominate subspecies (Fig. 62),the latter reaching north to Denmark and BalticSea, cannot be indicated.BiologyI{est. Pollen-storer (Meidell 1934b, pp. 86, 1 13).It establishes nests in and below the surfaceof the ground and at various depths. Also recordedin abandoned bird nests built in trees.The size of the colonies is in general small.Meidell (unpublished) estirnated the maximumnumber of workers produced in a colony asabout 30 specimens. He states moreover thatthe species produces two generations in a season(Meidell 1968).Flight season. From last half of April to theend of September. Queen : 19 April-28Sept.;worker: 4 May-25 Sept.; male: 7 June-26 Sept.BOMBUS (PYROBOMBUS) LAPPONICUS(FABRTCTUS)Apis lapponica Fabricius, 1793, p. 318 No. 13;Zimsen 1964, p. 415 No. 1078, lectotype ? CCC!selected by Lsken (1966a). Type area Sweden:Lapland ('tr-apponia Dom.de Paykull').(Apis lapponica: Quensel in Acerbi 1802.Bombus lapponicus (Fabricius); Dahlbom 1832,p. 4l; Zetterstedt 1838; Boheman 1844, p. 103;Wahlberg 1854, 1855 ; Thomson 1870, 1872;Siebke 1880; Sparre Schneider I889, 1895a,1895b, 1898, 1909 ; Strand 1898b, 1901 , 1904,1910; Lie-Fettersen 1901 , 1907; Aurivillius 1903;Bengtsson 19A4, 1908, 1931; Friese & Wagner1912; Ringdahl 1915 ; I-undblad 1924; Soot-Ryen 1925; Forsslund 1929, 1951 ; Gaunrtz 1929;Hell6n 1933 ; Meidell 1934a; Barendrecht 1941;Pittioni 1942; Brinck & Wingstrand 1949;Laken1949, 1950; tsrinck 1951 ; Krusen'lan 1959; Elfving196A, 1968; Ander 1965. Bombus alpinus: Dahlbom1832, p. 32 (partim); Siebke 1880 (partim).Bombus lapponicus var. lugubris Friese, 1902,p. 495 nec Morawitz 1880, holotype ? TRM!,type areaNorway : Troms (TRi): M6lselv; SparreSchneider 1906, 1909, 1910. Bombus lapponicusvar. scandinavicus Friese, l9lI, p. 684, type -holotype of Bornbus lapponicus var. lugubris;Friese &. Wagner 1912; Meidell 1934a; Laken1949, 1950. Bombus lapponicus scandinavicusFriese; Fittioni 1942, 1943; Brinck 1951; Kruseman1959; Elfving 1960; Reinig 1965; Laken1966c. A bibliographic reference to the speciesis compiied by Pittioni 1942.)First Scandinavian recordsNorway. Troms (TRV) : Skjervoy; Finnmark(Fi) : Alta: Bossekop (Dahlbom 1832).Sweden. Lapland (Fabricius 1793).Taxonomical remarksApis lapponica mentioned by Quensel inAcerbi (1802) presurnably refers to this species.A coloured drawing applies to the nominate form,but the hindmost gastral segments are describedas 'apice albido'.The great variability in the colour pattern ofB. lapponicus throughout its range has beensubject for studies from a genetical and zoogeographicalpoint of view (Vogt 1909, 1911; Skorikov1912; Kcirner & Zarapkin 1938; Fittioni1943; Reinig 1955; etc.). A vast number ofcolour variants have been named and severalsubspecies recognized (Pittioni 1942, 1943), sevenof which refer to Europe (Reinig 1965, p. 118).Two Scandinavian subspecies were recognized,viz. B. l. lapponicus and B. I. scandinavicus.They are readily separated in their typical forms,cf. the description below, but there is a greatrange of intermediates and aberrants resultingin a nurnber of designations. B. l. var. praticolaSparre Schneider, 1909 nec Kirby 1837, B. l.var. ornatwlus Friese, l9Il, B. l. var. dissidensFriese, l91l , B. l. var. helveticws Friese, t9ll ,B. l. var. norvegicus Friese, 1911, B. l. var.noricus Skorikov, 1912, B. l. forrnolosr,rs Skorikov,1912, B. t. var. collaris Hell6n, 1933 , B. l. uutmelanarius F{ell6n, 1933, one or several of whichmentioned as occurring in Scandinavia by thedesignators and/or Friese & Wagner (1912),

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Studies on Scandinavian Bumble Bees 77Fie. 63. B. lapponicrzs (Fabricius). Oblique up to the left hatching : B. l. lapponicus (Fabricius); oblique upto the right hatching : B. l. scandinavicus Friese. Legend as in Fig. 50.Hjerkinn 950 mZMO, Toftemo,ZMO, Vilisjo 950 mZiMO. Buskerud (Bd).' Flesberg: Hvila; Kongsberg:ZM.O.,Bv; Sigdal: Haglebu, Kopseng, Nedre Eggedal,Nordbygda; Nore og Uvdal: Bjorkeflfltfl 600 m, Gavlenseterll20 m, Norstebo 700 m, Rodberg, Solheimstul1000 m, Sonstevann 1050 m, Tunnhovd 850 m;Ovre Hein ll20 m; Nes: Buglebjork 900 m; Gol:Gol; Hemsedal: Kjolen 1000 m, Lykkja 900 m,Stoytestolen 950 m; Al: Bergsjo 1100 m, Laudalseter1100 m, Levell 700 m, Tvist 1080 m, Vallehalle 1120 m;Hol: Fagerheim 1150 m, Geilo 800 m VCA ZIN4B.,6. Bumble Bees.Geitryggen l22O m, E Halne 1150 m, Haugastol1000 m VCA ZN{{B ZMO, Kvasshogda l20[ m, Lillevann1150 m, Nygflrd 1000 m, Vikastolen 1000 m,Hovet, Nyestolen 1050 m, Olsenvann 1100 m, Seim,Ustaoset 1000 m. Telemark (fEy).' Drangedal: Omnes.TEi: Seljord: Svartdal; Fyresdal: Moland; Mo:Grimdalen; Vinje: Edland, Kromvika 930 m, Krossen700 m, Rauland kirke 700 m, Srerensgrend, Urdbo700 m, Vfl 700 m; Tinn: SE Borsjoen 1150 m, Mosvatndammen900 m, Sneiseli 960 m. Aust-Agder( AAi) ; Amli: Smedland; Bygland: Londal, Ose;

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Studies on Scandinavian Bumble Bees 79nes TRM, Elvenes TCtr-, Galgoaivve KMT TRM,Gjobukta in Ovre Pasvik ZMO, Grense-Jakobselv,Kirkenes IZts KMT TRM ZN4A ZN4B, KarpefjellZII'{O, Langfjorddal TRM VCA ZN{{B ZMO, NeidenTRM, Solheien TRM, Strand TRM, Svanvik, OvreNeiden TRM.Sweden Widely distributed from about 61" Nin Dalarne north throughout Lapland. Singlerecords in the eastern lowlands, from Uppland,and northwards, are noteworthy.A total of about 1000 specimens was examined.List of localities. Uppland.' Vassunda ? 28 June 1951(Torsell) IVU, Osterflker V t7 July 1908 (Wretlind)ZMU, Dalarne; FulufjAil NRS, Hundfjiill, Idre,Lima,Lingfjiillet NRS, Nipfjiillet NRS, Slagufjtill NRS,Storvdtteshigna NRS, Stora Tandfln NRS, Stddjan,Stilen Z} 4A, Siirna NRS, Tofsingdalen National ParkNRS. Hcilsingland: Delsbo Hd 1903 VCA. Hrirjedalen:Girtibauna, Harnrafjiill NRS ZML, Skatfjiill NRS,Sinfjtill NRS, Tdnndalen NRS ZML. Jcimtland: Brattikfjall,Enafors NRS ZML, Giiddede, Handol, LeipikvattnetNRS, Mullfjiill, Snasahogarna NRS, Stalltj.stugan, Storlien, Svenstavik NRS, UndersikerZNIL ZMIJ, Are IVU ZML ZM\J, Areskutan ZML.Angermanland: Hiirnosand C 1 July 1942 (Fredlin).Norrbotten: ? loc. ZMIJ. Lapland (,{r. Lpm.); Bingnds,Dorotea NRS, Fatmomakke, Njakafjtill, SatsfjeilZML NRS, Saxnds NRS ZML. Ly. Lpm..' AmmarndsGNM, Bjorkfors VCA ZML, Gargnds NRS,Grevattnet in Sorsele NRS, Grundforsen, HiiggflsGNM NRS, Iltebursfors NRS, Kviirnbiicken NRS,Laxfjiill, Lycksele VCA, Norra Storfjtillet NRS, Oltokjaure,Sorsele NRS, Tjultriisk, Tdrna NRS, Tiirnasjon,Viinniis NRS, Viistansjo, Orniis NRS. P. Lpm.:Arjeplog, Jtickvik, Peskehaure ZMU, Svalesjaure.Lu. Lpm.; Giillivare NRS, Kutjaure NRS, KvikkjokkNRS ZMA ZML, Malmberget, Markovaare NRS,Meatajaure NRS, Muddus National Park NRS,Njunjes ZN4A, Ruotjajaure NRS, Sarek NationalPark NRS ZML, Sjitnjajarnga NRS, Sjofallet, Tarra'dalen NRS, Vaisaluokta NRS, Virihaure area (BirnapakkZMU - Killanjaure ZN{{IIL). - Rapasdter ZN4U- Staloluokta NRS ZML Stora Titir ZMU T.Lpm.; Abisko NRS Z}l4A ZML ZMU, BjorklidenNRS ZMAZML, Jukkasjiirvi NRS ZML, KaisepakteNRS, Karesuando NR"S, Karsavaggejokk, KesajtirviNRS, Kiruna NRS ZMA, Kummavoipio NRS, KurravaaraNR.S, Nakerijiirvi NRS, Njuolja IVU, NjuorajaureNRS, Nuonjevare NRS, Piiltsa NRS, SarvajokkNR.S, Sinnukasjiirvi ZMU, Stenbacken NRSZML, Stordalen NRS, Tornetrd.sk area ZIll4A, VadvetjokkoNaticnal Park ZlvTA, Vassilaure NR"S ZMA,Vietovare NRS, Vittangi NRS.Finlond. Widely distributed from about 65'30' I{ north tc the arctic coast (Elfving 1 958).World distribution. Arcto-alpine.Europe (inBritish Isles restricted to Southwestern England,Wales, Northern England, Scotland; NW Portugal;NW Spain; The Pyrenees; The Appennines;The Alps; The Balkan mountains; Greece(Olympus); Fennoscandia; Northern EuropeanUSSR incl. Kola pen., Novaya Zemlya) -Northern Siberia - Kamchatka - Altai - Pamirs- Tien Shan (Pittioni 1942, pp. 194-198; Reinig1966; Yarrow 1959 and in litt.).Zoogeographical remark sB. lapponicus is divisible into two Fennoscandiansubspecies on the basis of the yellowpile extension of the vesture. The subspeciesdescribed below meet and intergrade in part ofNorthern Scandinavia in a zone tentatively illustratedon the map (Fig. 63). The distributionof the subspecies indicates immigration intoScandinavia from two directions, viz. the palercoloured B. l. lapponicus invading arctic areasfrom east and the darker B. l. scandinavicusimmigrating from south. Kruseman (1959) indicatesthat the latter might be an interglacialrelict. Reinig (1965, pp. 124-126) suggests thatthe strongly variable appearance in the areaof overlap is caused by intergrading of threepopulations, viz. (1) B. l. lapponicus immigratingfrom east ; (2) B. l. scandinavicus invading Norwayfrom south in interglacial period and developingstrongly melanic appearance while isolated inthe western refuges during Wiirm; (3) an additionalpost-glacial population immigrating fromsouth. However, the melanic appearance of B. l.scandinavicus does not necessarily favour a theoryof glacial survival and the recent distribution ofthe subspecies may be explained by post-glacialimmigration only.BiologyI{est. Pollen-storer. A number of nests havebeen recorded, all of which are situated on orjust below the surface of the ground, preferablyin vacated rodent nests. The colonies are rathersmall, the nurnber cf workers produced in a

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Studies on Scandinavtan Bumble Bees8lTable II. The frequency of B. l. lapponicus (Fabricius) in Fennoscandia-Petsamo. Areas suggested as beingoccupied by this subspecies in italics. Figures in O refer to queens only.* Mainly based on collection ZMHrevised by ElfvingNumber of specimensArea total B. l. lapponicusNorwayTroms (TRv)Troms (Tri)Finnmark (Fv)Finnmark (Fi): AltaFinnmark (Fi ) : Kautokeino-KarasjokFinnmark (Fn)Finnmark ( Fd )(-r!(7'+ Y O134188l95139t\286(?(5e(8e(7(32(4(27(32a ) x 4+ Y O2l43733317079(?(l3l(22(3(2s(4(24(29629 (250) 284 (120)B. l. Iapponicusa) x 1\,+ Y Ot6233765796392in per cent(?)\22)(2s)(43)(78)(100)(8e)(e 1)SwedenLapland (As. Lpm.)Lapland (Lv. Lpm.)Lapland (P. Lpm.)Lapland (Lu. Lpm.): VirihaureLapland (Lu. Lpm.): MalmbergetLapland (T. Lpm.): Tornetriisk area -Karesuando4l72647r11(41)(47)(3)(20)(44)J104l0103(3)(4)(3)(6)(41)230 (54) 113 (18)507 (20e) 213 (7 s)714672l9349(7)(e)(loo)(30)(e3)(33)Finland *-PetsamoLapponia (Le) : KilpisjiirviRemaining areas north of polar circleSouth of Polar circle122245many86230all7094100Finland. This subspecies apparently occupiesthe entire area of the distribution of the speciesexcept the westernnrost area, viz. Lapland (Le),cf. Table II.The total Cistribution of the subspecies, B. l.lapponicas (Fabricius). Northeastern Fennoscandia- Kola pen. - Northern Russia - NorthernSiberia to Anadyr Bay (Pittioni 1943, p.29).Faunistic remarksThe distribution of the nominate subspecies,reaching northeasternmost part of Norway,agrees with Sparre Schneider ( I 906, 1909) butdisagrees with Reinig (1965, p. 120, the map).The latter considers Kola pen. as the rvesternborder area of this subspecies and NortheasternNorway and Ncrthern Finland as an intergradingzane. Consequently, he includes SwedishLapland, the type area of the species, in thearea occupied by B" l. scandinavicus.The more westerly occurrence of the norninatesubspecies indicated herein is based on a phenotypicalstudy of the collections, and the frequencydistribution of this subspecies is presented inTable II. By keeping to the so-calle d 75 per centrule, the districts suggested as occupied by thenominate subspecies are indicated. The westernborder line (Fig. 63), based on the figures inTable II, is of course artificial and tentativelyindicated. tsy using the frequency presented inbrackets, referring to the mcre colour-stablequeens only, the western lirnit should be moveda bit further westo i.e. B. l. lapponicus wouldoccupy rnore of Finnmark. Larger material is,however, needed to adjust the indicated limitsof the transitional area.

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Studies on Scandinavian Bttmble Bees83(Bombus pratorum (Linnaeus); Dahlbom 1832,p. 36; Nylander 1848, 1852; Wahlberg 1854,1855 ; Siebke 1870, 1873, 1880; Thomson 1870,1872; Sparre Schneider 1895b, 1898, 1909, 1918 ;Ner6n 1892; Strand 1898a,1898b, 1901 ,1904; Lie-Pettersen 1901, 1905, 1907; Friese 1902; Aurivillius1903; Bengtsson 1904, 1908, l93l; Itingdahl1915; Wahlgren l9I7-18; Lundblad 1924;Gaunitz 1929; Soot-Ryen 1925 ; Jansson & Sjoberg1932; Hell6n 1933; Meidell 1934a, 1946; Wexelsen& SkAre 1934; Brinck & Wingstrand 1949;Laken 1949, 1950, 1958a, 1966c; Brinck l95l;Ander 1953a and b, 1963, 1965; Tjeder 1954;Kruseman 1959; Elfving 1960, 1968; Erlandsson1960; Frid6n, Eskilsson & Bingefors 1962; Hasselrot1962; Frid6n 1967. Apis fidus Harris, 1776,p. 134 No. 4. The description is provided witha colour-drawing of a queen which is regardedas the type. Type area Great Brittain. Apissubinterrupta Kirby, 1802, p. 356 No. 99, holotype? KCL! labelled by Yarrow (1968), typeloc. England: E. Suffolk: Barham. Apis donovanellaKirby, 1802, p. 357 No. 100, holotype QKCL! labelled by Yarrow (1968), type loc. England:E. Suffolk: Barham. Apis burrellanaKftby,First Scandinavian recordsNorway. Oppland (On) : Vang; Buskerud (Bo) :Ringerike; Krcidsherad (Siebke 1870).Sweden Uppland (Linnaeus 176l).Taxonomical remarksThe Scandinavian population belongs to thenominate form.The sparreschneideri form designated by Vogt(1909) as a long-haired form occurring in northeasternmostNorway, i.e. Finnmark (Fo): Scir-Varanger, cannot be reco gnized as a subspecies.The hair length was not measured but presentsapparently only a very slight clinal increasedirected from Southern to NorthernNorway.The difference in size between the southern andnorthern Norwegian population, characterizedby the difference in malar space, 'radial length',and interalar width in the female, is presented inTable III.It reveals no significant differencebetween the southern population and that restrictedto Finnmark (Fo): Sor-Varanger (Table III1802, p. 358 No. 101, holotype d KCL! labelled column A and B). Between the southernby Yarrow (1968), type loc. England: E. Suffolk:Barham. Bombus subinterruptus (Kirby); Dahlbom1832, p. 35. Bombus ephippium Dahlbom,population and that of northernmost Norwayin a wider sense (Table III column A and C)there is a significant difference in one feature, the1832, p. 37 No. 10, lectotype ? ZCL! designated interalar width. The populations display, moreover,by Ander (1967) type loc. Lapland (T. Lpm.):Jukkasjiirvi ; Zetterstedt 1838 . Bombus burrellanus(Kirby); Dahlbom 1832, p. 43. ? Bombus arbustorum:Zetterstedt 1838, p. 473 (partim) necFabricius 1776. Bombus lullianrzs Nylander, 1848,p. 236 No. 21, holotype C ZIV..{EI!, type areaFinland: Uleflborg. Bombus pratorum ephippiumDahlbom; Kruseman 1959.)identical colour pattern with the same vafia-tions; perhaps specimens with yellow hairs onT, more often occur in the northern population.Thus there is no basis for subspecific discrimination.The population in Swedish Lapland is treatedas B. p. ephippium wrthout comments by Kruseman(1959). It is, however, identical with theTableIIL B. pratorum (Linnaeus). Average measurements in mm of (A) 2099 from Southern Norway: AkershusandOppland(Os)60'61'15'N,(B)gggfromNorthernNorway:Finnmark(Fci):Scir-Varanger69'40'N,(C) 20 9? from Northern Norway: Finnmark and Troms 69"-69"40'N of which gg from B includedAt4-gtestCmean s mean s rnean st4-gtestMalar space'Radial length'Interalar width0.60 *0.02 0.593.62 +0.08 3.574.36 i_0.12 4.33+0.01 r.63+0.08 1.35+0.18 0.440.613.514.49+0.02 r.24+0.09 0.18+0.20 2.42

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Studies on Scandinavian Bumble Bees 85gf,eat in woodland and in subalpinelsubarcticzone, less so in the western archipelago and inalpine/arctic regions. fn Southern Norway recordedto 1200 m.Biotopes: Meadows, orchards, gardens, roadsides,pastures, Vaccinium heaths.A total of aboutzTW individuals was examined.List of localities. Ostfold.' Hvaler: Asmaloy, Kirkiiy,Sondre Sando; Krflkeroy: Tangen, Odegird;Onsoy: Dypeklo, Ellingflrd, Lervik, Ramseklo, Ror-vik, Torp; Sarpsborg: ZMO; Halden: ZMA, Asak,Berg, Blakmose, Hakelund, E Kuletjern, Nordbakke,Sponvika, Veggesdal; Aremark: Gjeddetjern, Mymosetjern,N Skodsberg; Marker: Damholtet, Dybedal,J6val, N Rodenes; Rakkestad : Levernes, Rtirvik;Ride: Fuglevik, Oven, Tom; Moss: Jeloy ZMAZMB; Eidsberg: Holm; Trogstad: V Bingon, Fransrud,Monster bro, Vassbotn. Akershus; As: ?loc.ZMO, Vollebekk; Frogn: Drobak TRM; Oppegird:Svartskog; Asker: Lushattdalen, Rustad; Barum: EBritan, Fornebo ZMO; Oslo: NMW TRM ZIvdAZ|JMB ZMO; Lorenskog: Elvflga ZM.B ZMO; Rel-Fig. &. B. pratorum (Linnaeus). Legends as in Figs. 50, 54,

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Studies on Scandinavian Bumble Bees 87Sreloy, Troldhaugen, Titlestad; Sund: Bokken, Telavhg;Fjeli: Solsvik; Laksevflg: Alvoen, Bjorndalspolien,Hetrleren, Storingaviken; Bergen: TRM ZMB;Askoy: Davanger, Hegernes VCA, Herdla, Jacobsoy;Asane: Eidsvflgsnesset, Golfbanen, Haukedal, Jordalen,Steinesto, Tellevik, Tuft, Astveit; Osteroy:Fitje, Hamre kirke, Kleppe, Njistad; Vaksdal: Eidslandet,Stamneshella; Meland: Brakstad, Io; Lindfls :Fosse, Fyllingsnes, Kolfls, Lauvfls, Seim, Srevrisv6g,Veland, Vollom . HOi: Kvinnherad: Dimmelsvik,Gjermundsharnn, Guddalsdalen, Hatlestrand, Holen,Lio, Ljosmyr, Lofallstrand, Rosendal, Seimsfoss;Odda: Dalen gflrd, Hordasvingene 1000 m, Roldal,Seljestad; Ullensvang: Aga, Alvsaker, Djonno, Fossli750 m, Fresvik, Grythorgi 900 m, Hesthammer, Kinsarvik,Kvalvik, Lofthus, Mibcdalen, Rjoto 1000 m,Stavali 900 m, Sysendalen 750 m, Ovre Eidfjord;Kvam: Kvamskogen, Nordheimsund, Sjuseter, OVstese,W Atvit; Voss: N Armot, Bulken, Bomoen,Hamlagro 600 m, Hangurfjellet 800 m, Mjolfjell,Nesheiin, Rong, Tyrlingen; Granvin: Eide TRM,Skjervet, Trfl; Ulvik: Bergo, Finse 1200 m ZM.A,Gangdal 1000 ffi, E Mjolfjell, W Rubbeliseter. Sognog Fjordane (SFy).'Gulen: Takle; Hyllestad: Botn,Yttre Dale, Hyllestad, Lekva, Skivenes, Skor; Hoyanger:Gare bro, Stoldalen; Gaular: Bugstad, Sande,Slotten, Vikum; Fjaler: Dale, Strandenes; Flora:Floro, NE Heia, Kinn, Sundaneset, Trollstua, Verpevik;Naustdal: Frammarsvik, Furehaugstoleno FIorstad,Naustdal, Ovre Hella; Forde: NE Flflten, FordeVCA, SW Hallbreim; Jolster: Vassenden, Ahus;Gloppen: Hjortset, Hope, Lote, Sandane; Eid: Kjolsdalen,Nordfjordeid, W Stflrheim, Vedvik; Vflgsoy:Bryggja, E Kjolstad, Mflloy, W Totland. SFi; Vik:Havis, Langedalen 850 m, Seljedal, Vik; Aurland:Flim, Vassbygda, Vatnahalsen 800 m, Ovstebo 800m; Lrerdal: Bergstolen 650 m, Horgje, Maristova800 m; Sogndal: Kaupanger; Leikanger: Hermansverk,Leikanger, Stavseter, Suphammer; Balestrand :Balholm NMW, Brekka, Flesje, Eitodn, Horpedalen,Kvamsoy, Mel, Suphellesetra; Ardal: Ovre Ardal;Luster: Dalsdalen, Fiberg, Fflbergstolen, Jostedal,Okkleviki, Turtagro 900 m; Stryn: Flo, Gillesdal,Hornindal, Hornindalseter, Skflre, Stryn, W Vanberg,Videdalen, Videseter 600 m. Mcire og Rontsdal ( MRy) :Vannylven: Thue; Volda: Bjorkedalsvann, SE Folkestad;Orsta: Lyngstolvann, Orstavik, Oy.; Hareid:Hjorungavhg, Mork; Ulstein: Flo; Heroy: Djupvik,Rodvik; Atresund: VCA; Orskog: Skodj e ZMO; Vestnes:Gjermundnes: Vikebukt; Molde: TRM ZMB;Aukra: Rindaroy; Frrena: Gjenden. MRi; Stranda:Bjordalsgflrdene, Fivelstad, Fredheim, NorangCalen,Vollset; Norddal : Indreeidet, Krike bru, Valldal,Ytredal; Nesset: Nauste; Sunndai: Dalen, Jordalsgrend;Surnadal: Lahjell, Melhus, Stangvik, OvreSurnadal; Rindal: tsolme. Srit'-Trdndelag (STy): Rissa:Gafset6sen, Nordseter gflrd, Rrssa VCA, Stadsbygd,Sotvik; Bjugn: Kotengsvann; Afjord: Molsletten,Morreaune, A. Sfi.. Oppdal: Dalsbekk, Driva st.,Drivstua 850 m ZMU, Knutsho 1200 m KMT ZN{B,Kongsvold BML ZN {B ZMO, Oppdal; Mittre Gauldal:Rogstadoy; Holtfllen: Holtdalen; Alen: Reitan;Roros: Evavoll 700 m, Naustervoll 700 m, N Storelvvoll800 m; Tydal: Mo bro, Sakrismoen; Trondheim:KMT VCA ZMB; Melhus: Benna; Skaun:Sornypvann KMT; Orkdal: Sognlia KMT. Nord-Trt)ndelag ( Nfy).' Namdalseid : Hovika KMT; Namsos:TRM; Nreroy: Breiviken, Dalene, Garmannsviken,Saltbotn, Teplingan. NTi: Merflker: GudienKMT TRM VCA, Tovmodal; Stjordal: Draveng;Leksvik: Storvannet ;Levanger: Avdal; Verdal : SandvikaKMT ZN4B., Stiklestad, Sulstua, Verdal KMT,S Vuku; Inderoy: Kirkenesv6g, Skjelvigen; Steinkjer:Sem ZMA ZMB; Overhalla: Gryten, W Sellotoren,Strandmoen; Grong: Ekkerseter, Fjerdingefl, Trangen;Snflsa: Bronstad, Hegge, Strindmoen, S Telnes;Lierne: Kvelia, Nordli, Seterhaug, Smfltjern; Hoylandet:Hoylandet, Skilleberget, Skogenga; Namskogan:Brekkvasselv ZN4A, Finnvollan, NamskoganVCA Z}.'4A ZMB, AktejAvre; Royrvik: Bjerkhaug,Namsvatn KMT. Nordland ( Nsy) ; Somna: SandvAgTRM, Somnes TRM; Bronnoy: Nevernes ZMA;Vevelstad: Aursletten TRM; Meloy: Dalen, Glomfjord,Reipi, Storglommvann TRM; Gildeskfll:Finnes, Gildeskfll, Gilset, Inndyr, Jelstad, Skauvoll,Storvika, Sor-Fugloy; Bodo: Bodo, Falkflaugdalen,Frostmo. ,A/si: Hattfjelldalen : Bjorkisen brygge ;Grane: S Bflfjellmo, Majavatn, Rotstokkmobekken;Vefsen: Luktvann, Ravasbakken, Store Bjornflvann;Hemnes: Bleikvassli, Finneidfjord, Krokselvmoen,Strombotn KMT; Rana: Bjellflnes ZMO, Dunderlandst., Krokstrand TRM Z]N{B., Randallsvollen,Ravni ZMO, Rosvollheien, Rovassdal TRM, StokkvikTRM; Beiarn: GrAtfldalen; Saltdal: ? loc. TRMZ}r'.4.O, Junkerdalsura, S Lonsdal, Solvflgfjell, StorjordetTRM, Vensrnoen; Fauske: Fauske, Vargflsen.Nnci: Sorfold: Bonnfl, Bonnflsjoen, Tennvann; Hamaroy:Brennvik, NE Fjerdevann, Innhavet, KrikmoSMS, Oppeid, Sandnes, N Strinda; Tysfjord: ? loc.ZMO; Ballangen: Bakkerud gflrd, Ballangen, Dyrhaug,Elvesletten, Forsfl, Hesjeli; Ankenes : BeisfjordenZMA, E Seterfjell, Trollviken; Narvik; Evenes:Dragvik, Jansbakk. Nnv: Vestvigoy: Eggum,Finstad, Knutstad, Skulbru; Vflgan: Store Molla,Svolvrer ZMA ZMO; Hadsel: Kongselv KMT. Troms(TRy ); Kvrefjord : Borkenes; Lenvik: Finnsnes;Tromso: TRM VCA, Breivik SMS, Tromsdal TRMZMB; Karlsoy: Hushattoy TRM, Karlsoy, NordfugloTRM, Vanno TRM; Skervoy: Ravelseidet,Vaddas . TRi: Skflnland : Boltis, Lavangseid ; Bardu :Bardu kirke, Rotvoll; M&lselv: ? loc. ZM.O, BjerkengTRM, Elvevoll, Frihetsli TR.M, Kirkesdal, KlettenZMO, Moen, Nordmo TRM VCA Z'MO, Olsborg,Rundhaug, S5rli; Balsfjord: I-aksvatn SMS, LanesTRM, Takvann TR.M; Storfjord: Signaldalen; Nordreisa:Bilto, Mollifossen; Kvrenangen: Burfjord,Kvaenangfjetrl. Finnmsrk ( Fv ).' Kvalsund: iinkasjavrre,unna l{atteras. Fi.' Alta: Bossekop TRM ZMB,Arones, Jotkajavrre TRM, Kvalfjord ZMO, KflfjordTRM: Kautokeino: Kautokeino TRh'I . Fn.' Porsan-

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Studies on Scandinavian Bumble Bees 89N{jolkudden NRS, Lulei NRS ZML, Pajala NRSZMI-, Periijiivaara, Pite havsbad ZMB, StromsundNRS, Tore ZMB, Overkalix NRS, OvertorneA NRSZML, Ovre Svartli NR.S, Alvsbyn NRS. Lapland( As. Lprn.): Biingnds, Dorotea NRS, Kittelfjdll, OrrnsjoNRS, Itisbiick NR.S, Vilhelmina NRS. Ly. Lprn.:Ammarnds NRS ZMU, Bjorkfors, Forsvik, LyckseleNRS, Sorsele, Stensele ZM{J, Stensundforsen NRS,Stromnris NRS, Tjultriisk BML, Tdrna NRS ZML,Tdrnasjo, {Jmfors. P. Lpm.: Skattrdsk. Lu. Lpm.:Giillivare NR.S, I(vikkjokk, Malmberget, MuddusNRS, Njr-rnjes ZMA ZML, Sarek, Tarradalen NRS.T. Lpm.; Abisko NR.S ZML, Bjorkliden, Jukkas.jtirviNRS ZML, Kaisepakte NR.S, Kauppinen, N,{asungbyn,Mell. Meriisjdrvi, Nuolja, Njuonjevare NRS,Svappavaara NRS, Tornehamn ZN{U, VietovareNRS, Vittangi NRS.Finland. Abundantlythe country (Elfving 1958).distributed throughout[Vorld distribution. Europe (British Isles; onthe continent ranging from NW Spain, ThePyrenees, ltaly, The Balkans and north to ArcticFennoscandia; entire European USSR) - Turkey- Northern Iran - Crimea - Caucasus - TranscaucasusUral Southern Siberia (Panfilov\957; Yarrow 1959 and in litt. ; Tkalcu 1960;Reinig 1958).is correspondingly completed during relativelyshort time. In England the colonies are finishedby mid-July or even earlier (Free & Butler 1959,p. 32; Alford 1969), yet according to Yarrow(in litt.) B. pratorum is sporadically recordedduring August-October, which indicates that theseason is occasionally lengthened. Alfken (1913)states that the season of this species ends nearthe middle of August in Northern Gerrnany:Brernen: 'Sie erscheint wie ihre Rasse B. jonellussehr zeitidg, verschwindet aber bedeutend frtiherals diese.' Workers and males were recorded inthe Austrian Alps at the end of August (Loken1954). In Norway, the frequency is stronglyreduced after middle of August, &S far as Icould observe. The local variations in seasonallength are noteworthy. Sex and caste occurringAugust-September may be the progeny ofl lateappearinghibernating females or perhaps offspringfrorn those emerging from coionies completedJune-July, i.e. B. pratorum in favourableseasons may produce two generations as statedfor B. jonellus (Meidell 1968. )BiologyIt{est. Follen-storer. Nests were recorded inand below the surface of the ground, in abandonedbird nests, in nest-boxes for birds set outat various heights above the ground, in betweenwalls in houses, etc. The nesting sites range fromhumid to dry biotopes and agree with the unconventionalchoice of dwellings generally knownfor the species (Free & Butler 1959, p. 7). Thesize of the colony is rather small. Meidell (unpublished)estimates the maximum number ofworkers produced in colonies excavated in Rogalandat about 40 individuals.Flight season. From the end of March to themiddle of September. Queen : 29 March-5 Sept.;worker: 25 April-25 Sept.; male: 29 May-15 Sept.Biological remarksThis early-appearing species produces sexualoffspring early in the summer and the life cycleSUtsGENUS A4ELAI{OBOMBUS DALLATCRRE,Melanobombus Dalla Torre, 1880, p. 40, typespeciesBombus lepidarius (Linnaeus) by designationof Sandhouse (1943).(Lapidariobombus Vogt, i 91 X , p" 58, typespeciesBornbus lapidarius (Linnaeus) by designationof Sandhouse (1943).)BOMBUS ( MELANOBO MBUS ) LAPI DARI US(LINNAEUS)Apis lapidaria Linnaeus, 1758, p. 579 No. 31,type area Sweden: Uppland. A small ? LSL !labelled lapidaria agrees with the usual interpretationof the taxon.(Bombus lapidarius (Linnaeus); Dahlbom 1832,p. 30, 1837 ; Zetterstedt 1838 (partim); Siebke1853, 1863, 1870, 1873, 1880; Wahlberg 1854;Thomson 1870, 1872; Ner6n 1892; Strand 1898a,

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Studies on Scandinavian Bumble Bees9lish. T*- u, fringes of Stn- u bright reddish, thelatter usually of a lighter shade. Otherwise coatblack.Var ia t i o n . Extrernelv colour-stable.Tibia and basitarsus of fore-leg occasionallyreddish fringed, red piles of mid- and hindbasitarsus and tibia more or less replaced byblack hairs. Some individuals have the blackcoat more or less whitish-tipped.MaleMorphological characters. Head about as longas wide. Malar space about as long as distalwidth, longer than A, but shorter than Az+ e.Mandibles bifid with a broad ventral and a smalldorsal tooth. Eyes directed in front of posteriormandibular condyle. Antennae rather short(Fig. 32A) I Au equal to Au or just longeri Aojust transverse to hardly longer than distal widthabout % as long as Au i Au-r, individuallyalmost parallel-sided and about 1/s times longerthan distal width of the segment. Outer surfaceof hind tibia rather convex, alutaceous, and withsparse distribution of hairs of various lengthexcept for an usually well-defined bare area onthe inner distal part of the segment; longesthairs in posterior fringe about twice the greatestwidth of thesegment in unworn specimens;hind basitarsus only slightly narrowed towardsthe base, longest hairs in posterior fringe aboutILA times the greatest width of the segment.St, and genitalia (Figs. 328-C); gonostylusrounded transverse with a small fingerlike processnear the inner base; volsella long, emarginatedat apex; hook of penis valve turned inwards andproduced into a sharp point. Body of mediumsize.Colour pattern Beard dark brownish. Pile offace between and below antennal sockets yellow,pile of vertex with variable admixture of yellow.Collar lernon to greenish-yellow, episternurn withvariable admixture of black and yellow hairs.T n-, bright reddish, Stn- u (or u- u) reddish toferruginous; the remainder of venter, ventralpart of rnid- and hind femora with predominantlyyellowish to pale yellowish-grey hairs. Mid- andhind tibia, n'lid- and hind basitarsus fringedreddish or yellolvish. Ctherwise coat black.Vorfatioru. Yellow coilar varying in width. Theproportion of btrack and yellow hairs in pile ofvertex, episternum, and verlter varying. Scutellum,T, with variEble acl;iiixture cf -vellow hairs.Distribution (Fig. 65)Norway. Confined to southeastern lowlandsand the extreme coast north to 61' N, i.e. notobserved north of Sognefjord. Moreover accidentlyobserved inland further north. It isquestionable, however, whether a queen wascollected near the Polar Circle, being far northof the present known distribution, or the recordis due to erroneous labelling. To my knowledgethe collector, Frits Jensen, did not always makenotes during the collecting trips, but relied onhis memory having returned home. The speciesis locally frequent. Recorded 500 m s.m. andonce 950 m s.m. cf. below.Biotopes: Meadows, gardens, roadsides, pastures.A total of about 940 specimens was exarnined.List of localities. Ostfold.' Hvaler: ?loc. ZMO,Akeroy, Asmaloy, Herfol, Kirkoy, Sondre Sando;Onsoy: Dypeklo, Ellingflrd, Lervik, Ramseklo, Rorvik,Skjreloy, Torp; Sarpsborg: ZN4O ZMB; Halden:NRS, Asak, Berg kirke, Hakelund, Sponvika; Marker:Dybedal, Rodenes; Rflde: Fuglevik, Oven, Tom;Moss: VCA, Jeloy VCA ZMB; Eidsberg: Holm;Trogstad: Monster bro, Rud. Akershus: As: Vollebekk;Frogn: Drobak VCA ZMB; Asker: Bronnoy;Brerum: ? loc. TRM, Egne Hjem, Hovik ZMO, Kolsis,Lysaker Z}l4O, Ostoy; Oslo: NMW TRM VCAZ}J|rIB ZMO; Rrelingen : Fjerdingby ; Aurskog-Holand :Bjorklangen, Gangsnes ; Fet: Bjorkefliten; Skedsmo :Strommen Z}l4O; Nes: Grinkelsrud, Vormsund.Hedmark ( HEs) ; Eidskog: Magnor SMS; Sor-Odal:Mirud gird ZMO; Grue: Finnskog; Hamar: ZMBZlrdo. Oppland (Os).' Gran: Granvollen; Lillehammer;Nord-Aurdal: Valdres ZMO; Oyer: Skfli; Ringebu:Ringebu. On: Dovre: Hjerkinn 950 m ? 1902(Lysholm) KMT. Buskerud (Bd).' Hurum: Hurum,Finadalen, Sagene, Storsand, Vannsbrekka; Drammen: ZMO; Lier: I-okke gflrd; Flesberg: Oydegarden;Kongsberg Ki\{T TRM, Skollenborg. ,Bv.' Sigdal:Nedre Eggedal; Nore og Uvdal: Uvdal stavkirke500 m; At: At ZnnO. Vestfold: Borre: Adal, Nykirke;Rarnnes: Kjrer, Lurnde, Orrevf,l; Andebu: Kodal;Stokke: Lango" SanC, Veierland; Tcnsberg: Prestercdkiler-r; hictteroy : Te it: ; lonrc: Brotsoy, Kjrere,

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Studies on Scandinavian Bumble Bees 93VCA; Sund: Glesvar, Telavflg; Fjell: Litleholmen;Bergen: TRM VCA ZldB; Askoy: Herdla; Fedje;HO i : Voss : Voss. I{ord-Trc)ndelag ( Ir{fy ).' Snisa :Finsfl ? 2 July 1951 (Bakke) ZMO. ?Nordland (I,{si):Rana: Mo ? 28 June 1934 (Jensen).Corrections. Records from Oppland (On):Dovre: Toftemo; Hordaland (HOV) : Bergen;More og Romsdal (MRy): Orskog (Siebke 1880)were revised to B. w, mastrucatus, likewise severalof those from Buskerud (Bv): At (Strancl 1898b).Sweden Wide, abundant distribution in thelowlands, along the eastern coast reaching 63"30' N in Angermanland.A total of about 3000 specimens was examined.List of localities. Skdne: Andrarum ZMB, ArlovZMA, Barkikra NRS, Barsebtickhamn ZMB, Bjiirred,Bjornstorp, Bonderup ZMB, Boksko gen ZMB,Bromolla ZN{A ZML, Bulltofta ZMA, BiickaskogNRS, Bokeberg, Dalby ZMB ZML, Falsterbo NRSZM.B, Grimslov IVU, Haviing NRS, Herrevad KlosterGNM, HjArnarp NRS, Hovs hallar NRS, HiilsingborgNRS VCA, Htissleholm NRS, Hoganiis NRS,Hflslov, Ilstorp, Ivo NRS, Kivik NRS, KristianstadNRS, Kullaberg, Kyrkheddinge, Kdvlinge, Landskrona,Ljungbyhed NRS, Lomma IVU, Lund GNMNR.S ZMAZMB ZML, Loderup NRS, LoddekopingeZMB, Maglarp ZMB, Malmo NRS ZN.{B ZML, MelarhusenNRS, Norra Mellby, Nosaby, NymomossorZMB, Roddinge NRS, Sandhammaren NRS, Silvikra,Simrishamn NRS, Sjobo NRS, Stenshuvud NRS,Svalov IVU, Sodra Sandby ZMA ZN.{{B., SovdeborgNRS,, Teckomatorp NRS, Torekov NRS, TosterupNRS, Trelleborg GNM Ztr IB, Vallikra, Veberod,Vik NRS, Vinslov, Ystad NRS ZML, Oved ZMÂZML, Angelholm NRS, Ahus NRS ZMB ZML. Blekinge:Aspo NRS, Briickne-Hoby Zl|l{A, Horviken,Jtimjoslilft ZMU, Karlshamn NRS ZML, KarlskronaNRS, Kristianopel NRS ZML, Ronneby NRS, RodebyNRS, Sibbaboda NRS, Sandhamn NRS, SolvesborgZMA, Torhamn NRS, Angelholm NRS. Halland:Falkenborg, Fjtirfls, Frosakull NRS, GetingeNRS, Halmstad NRS ZML, Laxvik, Roskiir NRS,Snostorp NRS, Steninge NRS, Tronninge, TylosandNRS. Smdland: Almvik NRS, Aneboda ZMB, BergkvaraNRS, Bla Jungfrun NRS, Brinstorp NRS,Dorarp NRS, Eksjo NRS ZMU, Grinna NRS ZMU,Hullaryd NRS, Horeda NRS, Jiirsniis NRS, KalmarNRS, Ljungarum NRS, Ljungby NRS, Ljungbyholm,Markaryd, Misterhult, Nye NRS, Oskarshamn NRSZML, Pflryd, Piskallavik, Ryssby ZMU, Skiro NRS,Smfllandsstenar NRS, Sommen NRS, Sodra LjungaNRS, Tranfls NRS, Visingo NRS, OsterkorsbergaNRS, Yxnands ZMA, Aseda NRS. Oland: AlgutsrumZN..[B, Borgholm NRS, Byrum NRS, ByxelkrokNRS, Boda NRS ZN[{4B, Ekerum NRS ZMB, GrankullaZN[d.{B, Gflrdby ZN[4B, Halltorp NRS, HornsjoZMB, Hogsby NRS ZMU, Hogsrum NRS, KalkstadZMB, Kastlosa NRS, Kiilla ZN{B, Morbyllnga NRS,Skogsbyn NRS, Torsiunda Zl'.{B, Vickleby Z}/^B..Gotland.' Ardre NRS, Austers NRS, Burgsvik, Burs,Fardume NRS, Frojel, Fflro NRS, Gammelgarn,Kappelshamn NRS, Kyllej NRS, Ljugarn NRS, RomaNRS, Slite, Sudersand NRS, Ulla Hau NRS, VisbyNRS, Vtimlingbo. Ostergdtland: Alvastra NRS, BohyttanZMB, Borensberg GNM NRS, Borsjon NRS,Fjuk NRS, Harg NRS, Hiistholmen ZMA, HogbyZMB, Kimstad NRS, Kisa NRS, Klinga NRS, KrokekNRS, Kvarsebo NRS, Linkoping NRS Z}dBZML, Malmsltitt NRS, Norrkoping NRS Z}l/.A,Norsholm, Ringarum NRS, Skonberga NRS, StavsjoNRS, Stjiirnorp ZN{IB, Ulrika NRS, Vadstena NRS,Vretakloster ZMB, Vdversunda NRS. Vcistergtitland:Alingsis ZMIJ, Brandstorp NRS, Brastorp, FalkopingNRS, Griistorp NRS, Goteborg NRS, Hornborgsjo, Karleby NRS, Kinnahult, Kymbo NRS,Lricko NRS, Nolhaga NRS, Ribrick NRS, SkaraGNM NRS, Skinnarhult in Boris NRS, Skovde NRSZML ZMU, Stenum NRS- Toreboda NRS ZML,VartoftaZN[d{{B, Vinninga NRS, Viittlosa NRS, Vimb,Virgirda ZMIJ. Bohuslcin: Bovallstrand NRS, Brastad,Dingle NRS, Grebbestad NRS, Kareby NRS,Munkedal, Stromstad NRS, Syd-Koster NRS, TanumNRS, ljorn GNM, Ytterby NRS. Dalsland: BengtsforsGNM NRS ZML, Gesiiter Z}l4B, Kopmannebro,Nossemark ZMB, Skillerud NRS, Sk6pafors NRS,Amil NRS. Ncirke: Laxil NRS, Stora Mellosa Z}/rB,Orebro NRS ZMB. Sc)dermanland: Bjornlunda NRS,Botkyrka NRS, Brandalsund NRS, Enhorna NRS,Fittja NRS, Handen NRS, Herrhamra NRS, JulitaNRS, Molnbo NRS, Nygiirdet NRS, Nykoping NRSZML, Nyniis ZMB, Nyniishamn NRS, NeivekvarnNRS Oxelosund NRS, Rekarne NRS, Salem NRS,Sjosa ZN ..{IB, Sparreholm NRS, Stjiirnhov NRS, StoraSundby NR.S, Striingniis ZMB, Svdrdso NRS, SodertiiljeNRS, Torshiilla NRS, Toro NRS, Trosa, TullgarnNRS, Turinge NRS, Tyreso NRS, Uttran NRS, UtoNRS, Valla NRS, Viksberg NRS, ViisterhaningeZII{B, Alberga NRS ZldB. (Ippland; Adelso NRS,Almunge NRS, Angarn NRS, Bjorklinge NRS, BondkyrkjaNRS, Brudniis NRS, Bfllsta NRS, DanderydZNdf.B, Djuro NRS, Drottningholm VCA, EkeroNRS, Eldgarn NRS, Enkoping NRS, Gimo NRS,Garnviken NRS, Grisslehamn NRS, Griiddo NRS,Habo-Tibble NRS, Hfltuna NRS, Ingaro, KnutbyZMB, Kolstrom NRS, Kungsdngen NRS, LennartniisNRS, Ljustero NRS, Ldnna NRS, Moja NRS,Norrviken NRS, Penningby NRS, Rimbo NRS, Roslagen-KullaNRS, Ro NRS, Ridmanso NRS, SigtunaNRS, Skokloster NRS ZIV..{B, Stavsniis NRS, StockholmGNM IVU NRS VCA ZMB ZML ZMU,Tensta ZMB, Triibygget NRS, Teljo NRS, UppsalaBML IVU NRS ZMAZMLZMU, Vallentuna NRS,Valsiitra IVU, Vassunda IVU, Vaxholm NRS, VindoNRS, Vaddd NRS, Viirmdo NRS, Yxlan ZMB, OregrundZ}i.dB, Osseby-garn s: n NRS, Osterskdr, OstraRyd NRS. Vristmanland: Arboga NRS ZML, Ding-7. Bumble Bees.

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Studies on Scandinavian Bumble Bees 95Taxonomical remarksThe interspecific relationship in the unit isindicated in a triangular graph (Fig. 66) based onScandinavian material. The spreading of themarkings indicates clearly the specific positionof the four taxa involved, even though there isa rather pronounced overlapping between B. alpinusand B. Arcticr,ts. The European occurrence ofB. arcticus was overlooked until it was recognizedby Richards (1931). The graph moreover includeseight ,8. arcticus ?? from ArticCanada to illustratethe taxonomic position of the Canadianand the European population, viz. B. A. arcticusand B. ct. diaboliczs Friese ( - B. a. alpiniformisRichards) respectively. The fact that both populationsoccupy the same area on the graph favourstheir conspecific status. The available Canadianmaterial thus confirms the subspecific rank of theEuropean population and consequently refutesits elevation to a species as suggested by Pittioni(1939b).Biological remarksOwing to the very few records of workers andthe absence of informationabout nests, it hasbeen suggested that colonies of Alpinobombusspecies, all of which are confined to arctic andO - B. ATPINUSE - B. ARCflCUS ABCTICUS* -a. ARcncus DtABoltcusO - B. BALTEATUSX - B. HYPERBOREUSFig. 66. Triangular graphing of malar space (A), 'radial length' (B) and interalar width (C) of four species ofthe subgenus Alpinobombus Skorikov.

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Studies on Scandinavian Bumble Bees 97Fig. 67. B. alpinus (Linnaeus). Legend as in Fig. 50.width, about as long as A2a 3. A, as long as Anor just longer i An about as long as distal width;Au-r, individually almost straight and nearlytwice as long as distal width. Outer surface ofhind tibia scarcely or not alutaceous. Hindbasitarsus 3 to 3 /2 times longer than distal width(Fig. 24A). St, and genitalia (Figs. 2aB-C);distal part of St s more or less regularly taperingtowards apex, which is usually rounded, rarelypointed; apex of volsella, seen from beneath(Fig. 24D); penis valve with subapical tooth.Body of moderate to large size.Colour pattern Hairs of T r-, ferruginous.Posterior fringe of hind tibia, hind basitarsusferruginous or tipped so. Otherwise coat black.Variation. A few individuals with a slightadmixture of yellow hairs in pile of vertex and/orwith indication of yellow collar were examined.

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Studies on Scandinavian Bumble Bees 99BiologyNest. Pittioni (1942, p. 178) states that nestsof .8. alpinus are not recorded at all, which mayjustify the mention of the three colonies below.A nest was located in regio alpina in SouthernNorway (Buskerud (Bv): Hol: 1.5 km NWHaugastol 1150 m 23 July 1967). The nestingsite, slightly sloping towards northwest, wascharacterized by growth of Betula nana Linnaeus,Vaccinium myrtillus Linnaeus and Empetrum hermaphroditumHagerup. The colony was foundedin an old rodent nest situated just below thesurface of the groind, the length of the accesstunnel being only about 2 cm. The entrance tothe nest was in between two rocks, one of whichpartly covered the colony. The nest material consistedof fragments of hay with some leaves ofEmpetrum admixed, partly plastered together bywax. The queen and most of the workers offirst brood were captured on returning from thefield loaded with pollen, i.e. the founder stilltook part in the field activity. The remainingworkers, & few of which had just emerged, weretaken as they issued from the nest or on thecomb.The data obtained from the colony are summarizedin Table IV. Two first brood workerswere missing. The cocoons from the third andfourth broods were opened and revealed thatseveral individuals from each of the batcheswere ready to emerge. Thus females and maleswould occur simultaneously in this colony. Theeggs were distributed in three egg cells individuallybuilt on the top of one female and two malecocoons. The egg cells contained 6, 6, and 5 eggsrespectively. If the healthy colony was not destroyed,all stages of young, including the eggs,might fulfil the development, emerging to queensor males. The total number of the progeny wouldthen reach 67 , 18 VE, and 49 ?? and dd respectively.Pollen bread stored beneath and adjacent tolower layers of masses of worker brood and,moreover, stuffed beneath some of the cocoonsabove, indicates that B. alpinus is a pocketmaker.Ffowever, a big lump of pollen, formedas pollen cylinder, placed directly on the groundjust apafi from the comb, indicates that the youngwere fed a mixture of pollen and nectar as well.The behaviour of the species may perhaps be asmentioned by Hobbs (1964) for another specieso.oE the subgenus, -8. balteatus Dahlbom: . . apocket-maker when rearing worker larvae to thelast instar and a pollen-storer when rearing lastinstar lawae, male, and queen larvae'.A palynological survey of the colony, set outin Table V, reflects the unusually delayed vegetationperiod that season . Salix was the predominatingfood plant up to the time the nest wasexcavated on 23 July. Pollen loads from corbiculaof worker number I and worker number 8 indicatethe start of a preference for Vaccinium andLotus corniculatus Linnaeus, that is, a change infood plants. V. myrtillus Linnaeus and V. uli-Table IV. Contents of a colony of B. alpinus (Linnaeus) excavated in Norway: Buskerud (Bv): 1.5 km NWHaugastdl ll50 m s.m. 23 Jttly 1967Queen, founder of the nestOffspring:l st brood : workers 82nd brood: workers 8imagines 16worker cocoonsworker cocoons3rd brood: male pupae4th brood, lst batch: male prepupae/pupae4th brood, 2nd batch: queen prepupae/pupae1086t2+ I emerged queen 65th brood, lst batch: queen or male larvae 8eggs in 3 batches 17Total number of progeny 6714 cocoons used as honey-potsLarge pollen-cylinder on the ground apaft from the comb

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Studies on Scandinavian Bumble Bees101Norwegian and Swedish specimens, comprises,however, 25 individuals from the two northernmostcounties in Norway, i.e. Troms and Finnmark.At least two worker broods raised in eachof the colonies above and the number of workersobserved throughout Scandinavia indicate thatB. alpinus regularly produces worker caste andto the northern limit of its distribution.BOMBUS (ALPINOBOMBUS) ARCTICUSKIRBYBombus arcticas Kirby, 1821, p. ccxvi nec Dahlbom1832 nec Zetterstedt 1838. Type ? BML!,type area Arctic Canada: Boothia.(Bombus arcticzs Kirby; Meidell 1934a, p. 124;Bombus alpinus: Dahlbom 1832, p. 32 (partim);Zetterstedt 1838 (partim); Bombus polarus Curtis,1835, p. lxiii No. 7, type ? BML!, type areaArcticCanada: Boothia. Bombus alpinus var.Thomson 1872. Bombus balteatus var. kirbiellus:Aurivillius 1890, p. 28 nec Curtis 1835. Bombuskirbyellus: Friese 1935 nec Curtis 1835. Bombusalpinus var. diaboliczs Friese, 1911 : 571 No. 21,holotype ? IZB!, type area USSR: Kola pen.Bombus alpinu^s var. pretioszs Friese, 1911, p. 57INo. 22 holotype d IZB! labelled Kirkenes (excoll. Sparre Schneider). The author designatedNorway: Finnmark (Fn): Nordkapp as typelocality cf. below. Bombus arcticus var. alpini'formis Richards, 1931, p. 13, holotype 6 ZMO!,type area Norway: Oppland (On): Dovre;Loken1960; Bergwall 1970. Alpinobombus tristis (SparreSchneider) nomen nudum cf. below; Skorikov1937 . Bombus alpiniformis Richards; Pittioni1939b; Elfving 1960, 1968; Bombus polaris alpiniformisRichards; Kruseman 1959; Ander 1965.Bombus arcticus diabolicus Friese: Lsken 1966a.)only examined one specimen (Friese & Wagnerl9l2). Sparre Schneider communicated livelywith Friese and supplied him with material. Tomy knowledge Sparre Schneider and his colleaguesnever collected in Nordkapp but spentseveral seasons in Sor-Varanger. Though thereare no records from Nordkapp situated at thenorthernmost coast approx. 71" N and roughlyabout 210 km northwest of Kirkenes, the speciesffi&y, however, teach that far north.Alpinobombus tristis is a nomen nudum asSparre Schneider never published the name.First Scandinavian recordsNorway. Finnmark (Fv): Nordkapp (Friese1911) published as B. alpinus var. pretiosus cf.above.Sweden. Lapland (Kruseman 1959), mentionedas Arctic Sweden.Taxonomical remarksThe Scandinavian population is recognized asa subspecies .8. arcticus diabolicus, presenting acolouring of the coat entirely different from thatof the nominate forrn. The latter has pile oncollar, scutellum and Tr-, ochrous-yellow todull yellow, hairs on Tn- u either black or ferruginous,coat otherwise black. The colouringof ,8. arcticus diabolicus is described below.B. alpinus var. pretioszs is revised to an infrasubspecificform of B. arcticus diabolicus. Richards(1931) mentions five workers (Norway: Oppland(On): Dovre TRM) belonging to a black-tailedform, B. arcticus var. friesei Skorikov, 1908,described from Northeast Siberia. The individualswere not traced and other specimens of thisform were not recorded in Scandinavia.Nomenclatural remarksThe type area of ,8. alpinus var. pretioszs beingNordkapp while Kirkenes, i.e. Finnmark (Fo):Scir-Varanger: Kirkenes, the locality written onthe label attached to the holotype, is due toconfusifig topographical information, as FrieseDistributionFennoscandia. Cf. the subspecies.World distribution. Circumpolar. Europe (Fennoscandia;Kola pen. ; Kolguyev) NovayaZemlya Northern Siberia KamchatkaArctic America Greenland (Skorikov 1937).

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Studies on Scandinavian Bumble Bees 103Fig. 68. ^8. arcticus diabolicrzs Friese. Legend as in Fig. 50.(SFi); Balestrand: Dueskard SW Balestrand 600 m;Luster: Turtagro 900 m TRM . Stir-Trtindelag ( STi ) :Oppdal: Givili 975 m, Jerosbekken 920 m, Knutsho1300 m BML ZIN{B, Kongsvoll 900 m BML ZM.B.,Stolflhollen 1400 m. Nordland (Nsy).' Meloy: SEGlomfjordbreen 300 m. Nsi.' Saltdal: S Lonsdal st.330 m; Fauske: Blimannsisen N Sulitjelma 950 mBML, Duoldakapjavrre NE Sulitjelma 810 m BML.Troms (TRy).' Tromso TRM ZIN{A, Tromsdal TRMZlI./Â. TRi: Gratangen: Myrland; Milselv: taralTRM, Kirkesdal, Mauken ZMO; Balsfjord: OrtaTRM ; Storfjord : Helligskogen; Nordreisa: JavroaiveSW Sappen 900 m; Kvanangen: Navetdalen. Finnmark(Fv).' Hammerfest TRM ZMO; Kvalsund:Skaidi, unna Hatteras. Fi.' Alta: Jotkajavrre 400 mTRM; Alta-Kautokeino DCL. Fn: Porsanger: FestningsstuaTRM, Kistrand TCL ZN{O, Lakselv ZIN{O1'Nesseby: Mortensnes ZMC. ^Fb'; Sor-Varanger: Grense-JakobselvTRM, Kirkenes TRM IZB, Neiden TRM.Sweden Scattered occurrence along the mountainchain from Hiirjedalen and north throughoutLapland.

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Studies on Scandinavian Bumble Bees 105BOMBUS (ALPINOBOMBUS) BALTEATUSDAHLBOMBombus balteatzs Dahlbom, 1832, p. 36 No. 8.The description is based upon a single queenwhich has not been traced. Lectotype V designatedby Milliron (1960) not valid (Ander 1967).Type area Finland: Lapland (Le): Enontekis.(Bombus balteatus Dahlbom; Siebke 1863,1870; Richards I 9311' Helldn 1933; Meidell 1934a;Barendrecht l94l; Pittioni 1942; Brinck & Wingstrand19491' Laken 1949, 1950, 1966c; tsrinck1951; Kruseman 1959; Elving 1960, 1968; Ander1965. Bombus nivalis Dahlboffi, 1832,p.40 No. 16,lectotype ? ZCL! designated by Milliron (1960),type area Sweden :Lapland(T. Lpm.): LakeTornetrdskarea;Dahlbom 1837 ;Zetterstedt 1838 ; Boheman1844, p. 103, 1857, p. 22; Wahlberg 1854,1855 ; Siebke 1863, 1880; Thomson 1870, 1872;Sparre Schneider 1889, 1895b, 1898 ; Strand 1898a,1898b ; Lie-Pettersen 1901 , 1907 ; Bengtsson 1908 ;Ander 1967. Bombus tricolor Dahlborn, 1832,p. 4l No. 17, lectotype d ZCL! designated byAnder (1967), type loc. Sweden: Lapland (T.Lpm): Karesuando ; Zetterstedt 1838 . Bombuskirbiellus Curtis, 1835, p. lxii No. 6 : Bombuskirbyellus Curtis, type not traced, type areaArctic Canada: Boothia; Aurivillius 1903;Bengtsson 1904, l93l; Sparre Schneider 1909;Friese & Wagner l9l2; Ringdahl 1915; Soot-Ryen 1925; Forsslund 1929; Gaunitz 1929. Abibliographic reference to the species is presentedby Pittioni 1942.)First Scandinavian recordsNorway. Oppland (On): Dovre: Hjerkinn andSor-Trcindelag (STi): Oppdal: Drivstua andKongsvoll (Siebke 1863) published as ,8. balteatusand also as ,8. nivalis.Sweden Lapland (T. Lpm.): Tornetrdsk andKaresuando (Dahlbom 1832) published as B.balteatus, B. nivalis and B. tricolor.Taxonomical remarksThe Fennoscandian population belongs to thenominate subspecies.The unstable colouring of the coat has givenrise to a vast number of designations (Pittioni1942). Scandinavian variants mentioned as B.balteatlts vat. nivalis Dahlbom, B. kirbyellus var,tristis Friese, \902 nec Seidl 1837 (: B. kirbyellusvar. similis Friese, 1911), B. kirbyellus var. pyropygusFriese, 1902, B. nivalis var. lysholmi Friese,1905 and B. kirbyellus var. grnelini Skorikov,l9l4a (Friese & Wagner l9I2; Skorikov l9l4a;Soot-Ryen 1925; Richards l93l; Meidell 1934a;Laken 1949, 1950; Brinck 1951) are infrasubspecificforms only. Aurivillius (1903) moreovermentions a black-tailed variety resembling B.hyperboreus. It might correspond to the frieseiform revised by Richards (1931) to B. arcticus,to which, however, no Fennoscandian specimenswere traced, cf. p. 101.Queen, workerMorphological characters. Head slightly longerthan wide. Malar space markedly longer thandistal width, yet less than lr/2 times this width,about as long as A3a a. Clypeus centrally almostimpunctate or with very fine puncturing, otherwisewith rather sparse uneven puncturing exceptfor the small impunctate anterior area betweenmoderately defined, coarsely punctured laterulimpressions. Ocelli just in front of supra-orbitalline, occasionally touching this line. A' at leastlr/z times longer than distal width and about% the length of An* r or nearly so. An about aslong as distal width, slightly shorter than Ab.Surface of vertex near and above supra-orbitalline smooth with sparse, coarse puncturing admixedwith very fine ones. Outer surface ofhind tibia slightly to moderately alutaceous;distal inner dorsal process of hind tibia usuallymoderately produced. Hindbasitarsus longerthan twice its greatest width but shorter than2r/z times this width. Tn-u slightly to stronglychagrinated with rather fine dense puncturing.Body rather robust, coast shaggy.Queen measurements: N : 201' NorthernNorway; malar space: 0.96 mm (+O.O++0.01)range: 0.88-1.00 mm; 'radial length': 5.00 mm(+0.14+0.03) range: 4.75-5.25 mm; interalarwidth: 5.80 mm (+0.17+0.04) range: 5.38-6.10 mm. Body of large size.

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Studies on Scandinavian Bumble Beest07-{r,: /^- l*t Jl( -/=&1 ..-L.1.'n=aFig. 69. B. balteatus Dahlbom. Proportion of melanism. The figures refer to the total number of examinedrecords within a county or district thereof. Black : pfoportion of melanics. Small circle : single record.

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Studies on Scandinavian Bumble Bees 109reduced on going north. There is, however, aslight increase in melanism in Finland exemplifiedby the rather large collection from Finland (Le:mainly Kilpisjtirvi area) comprising 7 per centmelanics compared with that from Sweden (T.Lpm.: mainly Tornetriisk area) of 2 per centmelanics only (Table VII). The high frequency ofmelanism in Finland (Lk: mainly Pallastunturi),viz. 2l per cent, is even more striking, but basedon a small collection only.It is noteworthy that the most reliable frequencies,i.e. those referring to Oppland (Os)and Hordaland (HOi), are about the same, viz.17 per cent and 16 per cent respectively (TableVI). The actual areas are situated in the ratherlower mountainous area linking southeasternlowlands to the mountain chain (Os: mainlyOyer: Aksjo area) and at the western slope ofthe elevation of the mountain chain (HOi:mainly Ullensvang: Fossli Sysendalen i.e.the northwestern slope of Hardangervidda), i.e.areas presenting different macroclimate emphasized,for instance, by the great difference inprecipitation (Fig. 1). As far as accumulation ofmelanic alleles is influenced by humidity, thefrequency of melanism which is about the samealong either side of the mountain chain in SouthernNorway, mey, however, be more related tolocal ground moisture and humidity near theground than the actual amount of precipitation.Pittioni (1943, pp. 4-6) states that sex andcaste in B. balteatus display a difference in thevariability of colour pattern, charactefized bythe tendency to melanism being more pronouncedin queens than in males and not recorded inworkers. Melanic individuals examined by Elfving(1960) were males only. Flowever, in thematerial at my disposal, sex and caste exhibitthe same wide range in the colouring, includingmelanism. The statement is exemplified by therecords from Oppland (Os) and Hordaland (HOi)where the frequency of melanisffi, set out inin Table VIII, clearly asserts that melanic queens,workers and males are equally produced. Thusthe great vaiabilityin the colour pattern locallyaccentuated by pronounced tendency to melanismis not sex-linked.Distribution (Fig. 70)Norway. In Southern Norwaywidely distributedalong the mountain chain from 59"N inAust-Agder - Rogaland to 62" 40'N in Scir-Trondelagand further north occurring from about64" 30'N to Arctic proper. The species is confinedto alpinelarctic and subalpine/subarctic zones andlocally observed in adjacent conifer forest aswell. It is also singly recorded at sea level in theinner part of the western fiords and locallyreaches the coast in the northernmost part ofthe country, yet so far only rarely observed onislands off shore. B. balteatzs has a wider distributionthan the remaining species of this subgenus.The abundance is moreover greater andthe frequency higher.Biotopes: Salix biotopes, Vaccinium heaths,mountain meadows, mountain pastures.Table VIII. B. balteatus Dahlbom. Frequency of melanism in sex and casteNo. per cent No. per cent No. per centNo.Totalper centOppland (Os)Typical form:Melanic form:Hordaland (HOi)Typical form:Melanic form:11 92 46 70 11 58 68 70I 8 2 0 3 0 8 4 2 2 9 3 0l2 100 100 t9 100 10010 s3 42 72 19 53 71 639 4 7 1 6 2 8 1 7 4 7 4 2 3 7t9 100 58 100 100 113 1008. Bumble Bees.

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Studies on Scandinavian Bumble Bees 111Tinn: Borsjoen 1100 m. Aust-Agder (AAi); Valle:Kvestad, Rygnestad; Bykle: Breive 700 m. Rogaland(Ry); Hjelmeland: Fister sea level Q 27 Aug. 1933(Mei). Ri.' Forsand: Meling sea level H 16 July 1931(Mei); Suldal: Britveit, Jonstdl 700 m, Kyrkjesteindalen900 m, Mostol 600 m, Mosskar 700 m, Steinkilen,Ullsneskvelven 750 m, AimjoOlonuten 700 m.Sauda: Ovre Sandvann 1100 m. Hordaland (HOi):Odda: Mittleger 1150 m, Valldal 900 m; Ullensvang:Berdola 725 m, Bjoreidalshytta 1100 m, Fossli 750 m,Grythorgi 900 m, Grytskardseter 850 m, Hedlo1000 m, Hjolmo, Lofthus sea level I 23 June 1949(Ls), Rjoto 1000 m, Storliseter 900 m, Sysendalen750 m TRM ZMB, Viveli 875 m; Ulvik: Finse 1200 mVCA Z}ldB. Sogn og Fjordane (SFy); Gaular: Slotten;Jolster: Hamar. SFi.' Vik: Hestavollen 1000 m, Krflktjern850 m; Aurland : Uppsete 850 m, Ovstebo800 m ; Lnrdal: Breistol 650 m, Eggestol 700 m, Maristova800 m; Leikanger: Hermansverk sea level ? 30May t948 (Ls); Balestrand: Dueskard W Balholm600 m, Horpedalen; Luster: Fflbergstolen 600 m,Turtagro 900 m TRM ZMB; Stryn: Videdalen 400 m,Videseter 600 m. Mdre og Romsdal ( MRi) .. Stranda:Kvanndalsetra 700 m. Srir-Trrindelag (STi); Oppdal:Dalsbekk 600 m, Gravbekken KMT, Gflvili 975 m,Jerosbekken 920 nBML ZMB, Knutshd 1200-1400 mBML ZMB, Kongsvoll 900 m TRM VCA ZM3 ZMLZMO ZNIU; Roros: Evavoll 700 m, Myrmoen 700 m.Nord-Trt)ndelag ( NTi ).' Namskogan : Atteiavre ; Royrvik:Bjorkhaug, Namsvann KMT. Nordland (Nsi):Rana: Bjellines ZMO, Krokstrand, Randalsvollen,Umskarstjern SE Mo; Saltdal: ? loc. ZMO, Lonsdal;Fauske: Duoldakapjavre NE Sulitjelma 810 m BML,Sulitjelma 600 m BML . Nnci: Tysfjord: Tysfjord sealevel ZMO; Ankenes: Seterfjell; Narvik. Ir{nv.' Sortland:? loc. TRM. Troms (TRy); Kvrefjord: Borkenes;Tromso: TCL TRM VCA ZMO, TromsoysundTRM VCA ZMO; Karlsoy: Vanno TRM. TRi: Gratangen:Gratangen ZMA; Bardu: Altevann, StromsmoVC A ZMO; Milselv: Bjerkeng, Kirkesdal, KlettenZMO, Mauken TRM ZMO, Nordmo TRM;Balsfjord: ? loc. TRM; Storfjord: Signaldalen; Nordreisa:Gapprusfjell 700 m, Sappen. Finnmark (Fv):T alvik: Kvalfj ard ZMO; Kvalsund: Kvalsund TRM,Skaidi, unna Hatteras. Fi.' Alta: Bossekop TRM ZMLZMO, Jotkajavrre TRM, Kvalfjord ZMO. Fn: Porsanger:Hamnbukt, Lakselv TRM ZMB, StabbursnesZMO; Tana: ENE Leirpollen; Vadso: Makkanes WVadso, Vadso BML NMW. ^&D'; Sor-Varanger: ? loc.TRM ZMO, Grense-Jacobselv TRM, Kirkenes NRSTRM VCA ZM.B, Neiden TRM, Ropelv TRM.Records from Oslo are questioned. The male,labelled 'Chria 1844' (- Oslo), is a specimenDahlblom apparently obtained while attendinga meeting of naturalists held in Oslo that year.The workers are both labelled (1) Kristiania; (2)Siebke. The modern spelling of the city nameproves a labelling of the specimens after siebkedied in 1875 and the record is therefore notreliable.corrections. Records from the southern coast,viz. Aust-Agder (AAV): Tvedestrand: Skipvik,Stensciy (strand 1898a) were revised to B. lucorumwhich agrees with previous revision (SparreSchneider 1909, p. 146). The tendency to melanismin Arctic Norway discussed by Sparreschneider (1909) is based on misidentificationof three dark males from Troms (TRV): Tromsci:Floyfiell TRM. Two of the specimens wererevised to B. arcticus diabolicus and the thirdone is not traced but might also belong to thistaxon. Another announcement of melanic arcticindividuals (Richards 1931) is due to misinterpretationof localities being situated in SouthernNorway.Non-melanic form of B. balteatus has beenconfused with B. jonellus and melanic ones withB. mastrucatus and melanic B. hortorum. Unrevisedrecords are therefore not included herein.Sweden Occurring from 61" 30' N in Dalarneand northwards throughout Lapland with adisjunction corresponding to that separating thesouthern and northern part of the mountainchain. Confined to montane areas and locallyalso occurring in adjacent forest zone.A total of 525 specimens was examined, viz.23s ?? 1e4 W and 96 66.List of localities. Dalarne.' fdre, Harrsjon NRS,Mora H 12 July 1947 (Erlandsson) NRS, SlagufjiiUNRS, Stiidjen, Tofsingdal National Park NRS. Hcirjedalen:Hamrafjail NRS ZML, Ljungdalen, Skorvdalsfja[NRS, Tiinndalen NRS ZI\4L. Jcimtland: Enafors,Handol NRS ZML, Leipikv. NRS, Skalstugan NRS,Stalltjiirnstugan DCL, Storlien, Svenstavik NRS,Undersiker, Vallistafja[, Are NRS ZML ZMIJ. Lapland(Ly. Lpm.).' Aitelnafors NRS, Bjorkfors, Joestrom,Sorsele mountains NRS ZML, Tjultriisk BMLNRS ZML ZMU, Ttirna NRS ZML, VindeldalenNRS, Viinnds NRS, Vtistansj6. P. Lpm... PeskehaureZMU. Lu. Lpm.: Gtillivare, Kvikkjokk NRS ZML,Malmberget ZML, Meatajaure NRS, Nuonjes VCAZML, Sallojaure NRS, Sarek NRS, Situojavre, Sjofallet,Tarradalen NRS, Virihaure (Ka ZMU - RapaZMU -StaloluoktaZMLZildu - unna Titir). T.Lpm.:Abisko BML IVU NRS ZMAZMLZMIJ, BjorklidenNRS ZMA ZML, Jebrenjokk, Kaisepakte NRS, Karesuando,Kiruna, Kopparisen BML, NakerijiirviNRS, Selkavaare BML, Sinnukasjiirvi ZMU, Sjangeli900 m BML, Sjanglitjikko 1000 m BML, Tjuonatjokko

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Studies on Scandinavian Bumble Bees 113heathery hillside exposed to the southeast. Thecolony was established in an abandoned rodentnest about 15 cm below the surface of the ground.A roof, consisting of fibrous fragments of plantsplastered together with wax, covered the comb,which had a disorderly appearance. The dataof the nest content are shown in Table IX. Oneof the queens and all the workers display apredominantly melanic appearance, the queenwith hindmost tergites whitish-haired while thoseof the workers being pale ferruginous-haired.The second queen displays a typical, thoughstrongly faded colour pattern. She was founddead in the nest, away from the comb, andmight have been an intruder. The dark queenand the workers of the first brood were capturedon returning to the nest. The former had nopollen and the latter had their corbicles onlypartly filled with pollen. The remaining fiveworkers, two of which had just emerged, werefound on the comb.The honey-pots, being larger in size than thecocoons but of similar shape, were all built atthe same level and adjacent to the first brood.Additionally five of the vacated cocoons wereused as honey-pots. A rather large pollen-cylinder,also termed pollen-pot, was built adjacentto the third brood, though detached from thecluster of cells. It contained only a little pollen.The dipterous cocoons listed in Table IX werein separate cocoons of the second bumble beebrood.A Swedish nest recorded by Boheman (Wahlberg1854, p. 202) contained melanic and nonmelanicindividuals, but is otherwise not commentedupon.In addition to the nests above, Hasselrot (1960Table pp. 158-162) succeeded in getting a queento establish a colony in an observation box.The colony was initiated 19 June and kept underobservation until I October when the developmentwas almost completed. The total numberof offspring was 8 VH 3 dd 3 ?? and one unhatchedworker pupa. Only one worker brood was raised.The author does not comment upon the longperiod of development.Flight seeson. From last half of May to end ofAugust. Queen: 16 May-27 Aug.; worker: 15June-28 Aug.; male: 4 July-z3 Aug.Biological remarksHobbs (1964) succeeded in getting hibernatingqueens to establish nests in hives set out in theTable IX. Content of a colony of B, balteatus Dahlbom, excavated in Norway: Rogaland (Ri): Sauda: OvreSandvatn, 1050 m s.m., 7 July 1936 by O. MeidellQueen, founder of the nestQueen, intruderOffspring:lst brood: workers2nd brood, lst batch: workers2nd brood, 2nd batch: workersimagines4 worker cocoons 53 4 worker cocoons, 2 worker pupae 62 3 worker cocoons, 3 worker pupae 69 3rd brood, lst batch: 2halfgrown separated worker larvaeon top of cocoon lst brood 23rd brood, 2nd batch: t halfgrown worker larvaeon top of cocoon 2nd brood, lst batch I4th brood, lst batch: 2 small larvae on top of cocoon2nd brood, lst batch 24th bro od, 2nd batch: 2 small larvae on top of cocoon2nd brood, lst batch 25th brood, lst batch: 7 eggs in cell on top of cocoon2nd brood, 2nd batch 7Total number of progeny 315 honey-potsI pollen-cylinderParasites: 2 dipter cocoons ?Brachycome devia (Fall6n)

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Studies on Scandinavian Bumble Bees 115longer than Aa+ a. Clypeus with rather even,dense distribution of coarse and fine puncturesexcept a small, almost impunctate atea betweenwide, well-defined impressions. Ocelli usually dorsallytouching supra-orbital line. A, at least lyztimes longer than distal width, and about %the length of Aa+ s. ,A.4 about as long as distalwidth, slightly shorter than Au. Outer surfaceof hind tibia alutaceous; distal inner dorsal processof hind tibia moderately produced (Fig. 9A).Length of hind basitarsus about zyz times thegreatest width of the segment (Fig. 9B). T** usmooth or inconspicuously alutaceous, withrather sparse puncturing. Body robust, coat longanC somewhat shaggy.Queen measurements: N : 201' SouthernNorway; malar space: l.l4 mm (+O.O:+0.01)range: 1.05-I.20 mm ; 'radial length' : 5.53 mm(+0.16+0.04) range: 5.20-5.85 mm; interalarwidth: 6.20 mm (+0.17+0.04) range: 5.75-6.53 mm. Body of larger size than any otherScandinavian Bombus species.Colour pattern Collar, scutellum, Tr-, deeporange to ochrous-yellow. Otherwise coat black.Parallel-sided interalar band not wider than thebroad collar.Variation. Extremely colour-stable. Only minutevariations in the extension of collar, thewidth of interalar band, etc.MaleMorphological characters. Head markedlylonger than wide. Malar space exceeding l%times the distal width, about as long as An* u.A, hardly longer than An i An as long as distalwidth or just longer I Au-', individually straight,nearly twice as long as distal width. Outer surfaceof hind tibia scarcely alutaceous. Hindbasitarsus about 3yz times longer than distalwidth. St, and genitalia (Figs. 27A-B); insideof volsella, seen from beneath, strongly projectinginwards; subapical tooth of penis valve absentor much reduced. Body of large size.Colour pattern Pile of vertex predominantlyorange to ochrous-yellow; otherwise colouringas in the female.Variation. As in the female.Distribution (Fig. 71)Norway. Scattered occurrence in SouthernNorway restricted to pafi of the mountainousmassive ranging from Jotunheimen to the northernfringe of Dovrefiell and in Northern Norwayto oislets' north of Polar Circle. Scarce.In Southern Norway once recorded 400 m s.m.,otherwise occurring 900-1300 m. s.m.Apparently conflned to limestone biotopes.A total of 72 specimens, i.e. 44 ?? I H 27 d6was examined.List of localities. Oppland ( On) : Yhgil: Gjendesheim10C0 mC 15 Aug. 1959 (I. Meidell), Jondalen900 m 2 66 12 July 1953 (Hackman,La), Vigimo400 m 3 12 July 1953 (Kontuniemi) Zli.d{}{, Ovre Sjodalsvann950 m ? 9 Aug. 19571fo); Dovre: Dovrefjell1000 m d 18 July 1832 (ex coll Smith) BML ? l9O2(Lysholm) KMT 2 ?? 466 July 1832 (Boheman) NRS? 1902 (Lysholm) TRM J @smark) VCA ? 2l June1926 (ex coll Mei) ZMB 336 (Mus Drews) ZMC 4 ??2 66 July 1832 (Rosenskiold) DCL TCL 4 ?? 5 dC HJuly 1832 (Esmark) Zll4.O 2?? (Moe) ZII{O ? 2 33(Siebke) ZMO ? 1877 (Schoyen) ZMC 2 ?? July 1832(Boheman& Rosenskiold) ZMU, Fokstua 930 m d6 Jutry 1933 (Nordstrom) ZML, Vfllflsjo 950 m d14 July 1953 (Bakke) ZMO. Srir-Trrindelag (STi):Oppdal: Drivdalen ? 29 Aug. 1948 (Knaben), Kaldvelldalen120C m ? 26 July 1968 (Alendal), Knutsho1200-t300 m 2 ?? 1954 (Holaker) ? 20 June 1967 (Ls)Zl{B ? 16 July 1909 (Wretlind) ZMU, Kongsvoll900 m ? 1 July 1910 (Sch) TRM ? 8 July 1934 (Nordhagen)? 17 June 19671t-a)67 Jluly 1933 (Nordstrom)ZML?20 July 19C6 (Wretlind) ZMU. Nordland ( Nsi) :Saltdal: ?loc. 2 ?? (Schoyen) Z]N{{O ? (Sch) VCA.Troms ( TRi ).' Bardu : Altevann 500 m 4 ?? I 8-2I July1893 (Sch) TRM; Nordreisa: Gapprusfjell 700 m 3 ??22 July 1955 (La), Javroaivve 900 md 12 Aug. 1968(Ryvarden), Kvrenangen: Jokelfjord ? ?sea level(Zetterstedt) ZML. Finnmark (Fn).' Kistrand: Festningsstua50C m 3 ?? July 1924 (A. Strand) TRM.Unrevised records : Troms ( f Ry ).' Tromso : FloyfjellQ and Finnmark (Fd); Sor-Varanger: JacobselvH (Sparre Schneider 1909, p. 89). Oppland (On).' Jotunheirnenand Rondane, sex and number of specimensnot mentioned (Meidell I931a).Sweden Except for an isolated occurrence inJiimtland the species has only been recorded inrestricted areas in the mountainous Lapland(Ly. Lpm - T. Lpm.).A total of 57 specimons, viz. 43 ?? 3 VH 11 Cdwas examined.

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Studies on Scandinavian Bumble Beesll71923 (Winblad) ? (Malaise), Tornetriisk - Vassijaure2?9 8, 12 Aus. 4 66 13-20 Aus. H 14 Aus. 1908 (Roman)NRS, Vassijaure ? 6 Aug. 1927 (Erl.) NRS.Unrevised records: Lapland (Lu. Lpm.).' Virihaurearea: Hildomvare ?, Kappaluobal ?, Vuolle PoulejaureI (Brinck 1951).Finland. Recorded in Lapland (Le): Kilpisjflrviarea only. A total of about 60 ?? and ddso far observed (Elfving 1968).World dis trtbution. Circumpolar. Fennoscandia- Kola pen. - Arctic Russia - NovayaZemlya -Arctic Siberia Alaska Arctic CanadaGreenland (Richards 1 931; Skorikov 1937).Correction. B. hyperboreus workers fromGreenland (Aurivillius 1S90) referred to bySparre Schneider 1909, P. 90) concern four specimensfrom Nordenskiolds Exp. The only individualso far traced NRS has been revised to B.e. arcticus and the description of Aurivilliusindicates that the remaining individuals also belongto this species.BiologyNest. There are no reliable records of the -8.hyperboreus colony.Flisht season. Middle of June to the end ofAugust. Queen: 17 June-29 Aug. ; worker: ?;male: 7 July-ls Aug.IBiological remarksThe suggestion that nests are established deepin the ground (Friese & Wagner 19t2, p. 162ff;Pittioni 1942, p. 185) is apparently based uponobservations from Novaya Zemlya (Friese 1904,p.413 referring to Jacobsen in litt.). As, however,only entrances were observed, there is no wayAug. 1930 (Knaben) ZMO), identified as B.hyperboreus by Friese (1935), is revised by meto a colony of B. a. arcticlt,s usurped by B. hyperboreus.According to Knaben (in litt. 1968) thecolony contained eight imagines, all of whichwere captured. Five individuals were traced, viz.B. hyperboreus Q and B. a. arcticus 4 HH ZMO.Two colonies of B. a. arcticus located in NorthernCanada (Northern Ellesmere Island 10-13 Aug.1961) were likewise usurped by B. hyperboreus(Milliron & Oliver 1966) and additionally acolony of B. jonellus (Sweden: Lapland (Lu.Lpm.): Staloluokta in Virihaure 23 July 1964)was invaded by this species (Bergwall 1970).The phenomenon of usurpation presentedabove supports the suggestion of Milliron &Oliver (1966) that '. the short season is notthe only reason for the absence, or paucity, ofworkers in hyperboreus'. That is, a total of fourrecords of usurpation so far gives reason toquestion whether the theory of adaptation tosolitary living in species of the subgenus Alpinobombus(cf. p. 96) applies to B. hyperboreuseither. The workerless condition may insteadbe due to interspecific usurpation. Usurpation,being interspecific or intraspecific, does, however,occasionally occur in Bombus spp., generallyproducing a worker-caste (Free & Butler 19591'Hobbs 1967, p. 1288). It is therefore uncertainwhether the records above are due to casualevents or to B. hyperboreus being a facultativebut not obligatory inquiline, the latter being therule in the genus Psithyr,as. The fact that a totalof four Scandinavian workers were examined,that one worker was mentioned by SparreSchneider (1909), and that supposed specimensof this caste have also been recorded outsideFennoscandia (Kola pen. Svjiitio, the numbernot given (Enwald 18810 P. 255), Gavrilovo VZIi{IJ (Elfving 1960); ?Novaya Zemlya cf.above; with no data 2 VH (Skorikov 1922b,p. 79); Russian Lapland number not given(Friese 1923)) proves that workers are at leastof controlling whether these were virtually associatedwith colonies of ,8. hyperboreus. A totalof 30 bumble bees from the expedition of Jacobson(1899) were revised by Friese (1908) and occasionally produced.comprised no workers of this species. In a survey Within its entire area of distribution B. hyper'of the bumble-bee fauna of Novaya Zemlya boreus occurs in irregularly spaced 'islets', gene-(Friese 1923),theworkercasteof!. hyperboreus rally presenting a dispersed growth of foodis not mentioned.plants. Flying activity is consequently scatteredA nest located on Greenland (Vegasund 17 except perhaps where the early blooming Sallx

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Studies on Scandinavian Bumble Beest19o - 3. coNsoERtNUstr - B. GERSTAECKERIX - B. HORTORUM. - B. RUDERATUsFie.72. Triangular graphing of malar space (A), 'radial length' (B) and interalar width (C) of four species ofthe subgenus Megabombus Dalla Torre.BOMBUS (dahl l9l5; Lundblad 1924; Gaunitz 1929, 1938;MEGABOMBUS ) CONSOBRII\,IUSMeidell 1934a; Brinck & Wingstrand 1949;DAHLBOMLaken 1949,1950, 1960, l96lb; Kruseman 1959;Bombus consobrinus Dahlboffi, I 832, p. 49 No. 30, Elfving 1960, 1968 ; Ander 1965. Bombus hortorumneotype ? ZML! selected by Lsken (1966b), var. /. Morawitz, 1881 , p. 240. Bombus hortorumtype area Sweden: Lapland (Lu. Lpm.): Virihaure.Friese & Wagner 1912. Bombus consobrinus sahl-var. consobrinus Dahlbom; Lie-Pettersen 1901 ;(Bombus consobrinus Dahlbom; Dahlbom 1837 ; bergi Skorikov, I9l4b; Brinck 1951.)Wahlberg 1854, 1855 ; Boheman 1857 , p. 22;Siebke 1863, 1870, 1880; Thomson 1870, 1872;First Scandinavian recordsAurivillius 1887, 1903; Sparre Schneider 1895b,1898, 1909, 1918 ; Strand I 898a, I 898b ; Lie- Norway. Oppland (On): Dovre: Fokstua andPettersen 1907; Friese 1902; Bengtsson 1904, Sor-Trondelag (STi) : Oppdal: Drivdalen (Siebke1908; Vogt 1909, 1911; Skorikov l9l4b; R.ing-I 863).

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Studies on Scandinavian Bumble Beest21with variable mixture of black and greyish-whitehairs. Fringes of sternites predominantly greyishwhite.Otherwise black.Variation. Black hairs of head more or lessreplaced by yellowish-brown ones . Lateral blackpatches of T, more or less pronounced, occasionallythe hairs of T2 entirely black or entirelyyellowish-brown. Hairs of episternum (except foryellowish-brown pile on the dorsal part), andTn-u, are individually ranging from entirely greyish-whiteor pale greyish-yellow to entirely black.Corbicular fringes with an admixture of yellowish-brownhairs or tiPPed so.MaleMorphological characters. Malar space usuallyexceeding twice the distal width and about aslong as Ar* 4+ b. Longest hairs in pile of faceabout as long as antennal scape or longer. Hindtibia with bristles exceeding the distal decumbenthairs of margin; longest hairs in posterior fringesabout twice the greatest width of the segmentor nearly. Hind basitarsus slightly or not widerdistally than at base. St, and genitalia (Figs.a0A-B); lateral depressive of gonocoxite moderate(Fig. 40C); gonostylus larger than in B.hortorum, its posterior margin, seen from behind,being almost straight; boot-like apex of volsella(Fig. 40D) varying in shape but usually more or\ less different from that of B. hortorum (Fig. 39F).Coat shaggy. Body in general of large size.Microsculpture of Tr-u rather smooth with fine Colour pattern. Pile of vertex with variablepuncturing. Body slender, coat shaggy, long- mixtureofblackandyelloworyellowish-brown.haired.Episternum and T' predominantly yellow, T7Queen measurements. N: 20; SE Norway; black-haired. Otherwise colouring as in the femalarspace: 1.44 mm (+0.03+0.01) range: male.1.40-1.50 mm;'radial length':4.82 mm (+O.OS Variation. Individuals with hairs of Tu-' en-+0.02) range: 4.65-5.00 mm; interalar width: tirely or predominantly black. Otherwise varia-5.27 mm (+0.06+0.01) range: 5.15-5.35 mm. tions as in the female.Body of large size.Workers vary greatly in size, the largest onesbeing difficult to distinguish from the queens.Colour pattern Pile of vertex with variablemixture of black and yellowish-brown. Dorsumof thorax, at most dorsal third of episternum,Distribution (Fig. 73)Norway. Widely distributed inland between59' and 68'30' N. Only north of Trondheimsfiord,Tr-, (except laterally) with yellowish-browni.e. north of 63" 30' N, reaching the extremehairs; remainder of thorax, hind femur, Tn-u coast. In Southern Norway confined to the easternsubalpine elevation along the mountain chainand adjacent forest zofle, and occasionallyreaching the lower alpine belt. Occurring in afew canyons west of the watershed divide, locallyreaching sea level in the inner pafi of severalfjords which are the western limit of the distribution.Moreover, extending throughout thesoutheastern lowlands where the population is,however, decreasing, probably because of intensivecultivation and industrialization in the lastdecades. ,8. consobrinus and the main food plantAconitum septentrionale are locally eradicatedand the southern limit of their distribution isconsequently on the retreat. Locally frequent.In Southern Norway recorded to 1300 m s.m.Biotopes: Aconitum biotopes and adjacentfields.A total of about 770 specimens was examined.List of localities. Ostfold; Eidsberg: Holm; Trogstad:Monster bro, Asen. Akershu^s.' Asker: Lushattdalen;Oslo: Slemdal TRM . Hedmark ( HEs) : Hamat:Helgoya ZMO. HEn: Trysil: Enga, Vestby; Amot:Glesubekken; Tynset: Haugsvangen gflrd, Tyldal;Tolga-Os : Hodalen 700 m; Engerdal: Risbakken750 m. Oppland (Os).' Gran: S Einavold, Gjervika,Lander; Ostre Toten: Nrerum, Stubdalen; SondreLand: Fluberg; Nordre Land: Kinn; Etnedal: Fladoydegard;Sor-Aurdal: Breidablikk, Ellingseter 820 m,Reinli, Skard 700 m; Nord-Aurdal: Knutshaugen,Sabuoygard N Fagernes, AUjor; Oyer: Aksjo 210 -'Skei StjO m, Oyer; Ringebu: ? loc. ZMO. On: OystreSlidre: Skammestein 700 m; Vang: Eidsbugarden1150 m, Nystova 1000 mZM3 ZN.4O, NE Otrovann1000 m, Tyin 1100 m, Oye USU; Fron: ?loc. ZN,{O:,Sel: Heidal kirke, Klaksh&ug, Leirflata, Mysuseter

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Studies on Scandinavian Bumble Beesr23Vilen. Aust-Agder ( AAil ; Bykle: Breive 700 m. Hordaland(HOi); Odda: Austmannli 800 m, Seljestad;Ullensvang: Hardangervidda VCA, Hjolmo, Fossli750 m, Isdalen 840 m, Mebddalen, Sysendalen 750 m;Ulvik: N Osa, Rubbeliseter. Sogn og fiordane (SFi):Vik: Vik; Aurland: Berekvam, Kvammadal 950 m,Uppsete 850 m VCA ZN4B, Vatnahalsen 800 m,Ovstebo 800 m; Lrerdal: Grote, Horgje, LnrdalTRM; Leikanger: Hermansverk; Balestrand: Fjrrland,Horpedalen, Suphellesetra; Luster: Dalsdalen,Fibergstolen, Turtagro NRS TRM VCA ZNII{.IB ZMO;Stryn: Videdalen, Videseter 600 m. Mrire og Romsdal( MRi ).' Stranda : Kvanndalsetrene 700 m, Vollset;Norddal: Indreeidet, Krike bru; Sunndal: Groa; Surnadal:Stangvik; Rindal: Bolme. Sdr-Trtindelag(STy).' Rissa: Stadsbygd; Afjord: Molsletten. STi:Oppdal: Dalsbekk, Driva st., Drivstua 850 m ZIN'4OZMU , Knutsho 1200-1300 m, Kongsvoll 900 m VCATRM ZM3 ZMO ZMU, Kvammen 1200 m, Oppdalst., Skansen, Virstigen 900 m; Rennebu: Sorflfl; Alen:Berglund; Roros : Evavollen 700 m, Myrmoen 700 m;Tydal: Mo bro, Sakrismoen, Vekterstua; Selbu:Heggset bro; Trondheim: VCA Zll4'B. Nord-Triindelag(NTy).' Neroy: Saltbotn, Skogenga, Teplingan.NTi: Merflker: Merflker; Stjordal: Jullum; Levanger:Bergsve; Verdal: Sandvika, Stiklestad, VerdalsoraDCL; Steinkjer: Sem; Overhalla: Gryten; Grong:Trangen; Snisa: Hegge , Ler6" bro, Asmulen; Lierne:Kvelia, Seterhaug, Sorli KMT; Hoylandet: Hoylandet,Skilleberget; Namskogan : Brekkvasselv KMT,Finnvollan; Royrvik: Namsvatn KMT. I,{ordland(Nsy).' Somna: Somnes TRM, Vik; Nesna: Nesna,Oldersletten; Meloy: Dalen; Gildeskfll: Finnes, Gildeskfll,Jelstad, Storvika; Bodo : Falkflaugdalen, Frostffio,Planteskolen, Lopsviken. Nsi; Hattfjelldalen:Bjorkisen brygge; Grane: Bifjellmo, Grane; Vefsn:Store Bjornflvann ; Hemnes : Bleikvassli, Finneidfjord,Korgen ZMA; Nord-Rana: Dunderland st., KrokstrandNRS TRM, Steinkjonnli TRM, Strombotn,Umskarstjern; Saltdal: ? loc. TRM ZM.O, Drageid,Junkerdalen, Junkerdalsura TRM, Saltsjo near SetsflVCF, Solvigfjell 800 m, Storjordet TRM VCA ZMO,Vensmoen; Fauske: Fauske. Nnr): Sorfold: Bonni,Bonnisjoen, Rosvik; Hamaroy : Sandnes; Ballangen:Bakkerud, Hesjeli, Myrbakk. Troms (Z.Ri); Skfln-Iand: Lavangseid.Unrevised records: Rogaland (Ri).' Suldal: Kvandal;Sauda: Slettedalen (Meidell 1934a).Corrections. Records from Aust-Agder (AAy):Tvedestrand: Lyngcir, Stensciy and Buskerud(Bv): At (Strand 1898a) are all revised to B.pascuorum (Scopoli). Five specimens in Vogt'scollection, captured by Sparre Schneider 1896,were wronglyy'abelled Tromsci and may havecontributed to the misunderstanding that B.consobrinus is an Arctic species (Friese & Wagnerl9l2; Pittioni 1938, p. 49; etc.). The labels,apparently written by Friese, are a misinterpretationof the name Turtagrci, being written onthe original labels attached to adjacent specimens.Tromso is situated at 69" 40' N, i.e. about 150 kmbeyond the northern limit of the distributionof the species. B. consobrinus is well known fromTurtagrci, being situated in Southern Norway,viz. Sogn og Fjordane (SFi) : Luster: Turtagrd,and was, moreover, visited by the collector in1896 (Sparre Schneider 1898).Sweden Occurring from Jiimtland north toVirihaure in Lapland (Lu. Lpm.) and togetherwith Aconitum extending east to the Gulf ofBothnia within restricted areas in MedelpadVtisterbotten. A specimen from Lapland (T.Lpm.): Abisko, is the only record outside thedistribution of Aconitum.A total of about 250 specimens was examined.List of localities. Medelpad: Jackvissle NRS, Sundsvall.Hcirjedalen: Hamrafjiill NRS, Tiinndalen. Jcimt-Iand: Berge in Alsen TCL, Berge, Bugirden DCL,Bydalen, Doda Fallet NRS ZML, Faxiilv TCL DCL,Gtiddede NRS, Hallen, Hfllland NRS ZML, Jormlien,Morsil TCL, Nyland DCL, Oviken ZN'4B ZML,Sandven DCL, Storlien Z1N4B ZML, Storsjon, Sdter,Undersiker NRS ZML, Are TCL, Areskutan DCL.Angermanland : Bingntis SW Hiirnosand, Hiirnosand,Ramsele GNM. Vristerbotten: Vindeln NRS. Lapland(As. Lpm.); Bflngnds, Dorotea NRS, VilhelminaNRS. Ly. Lpm..'Ammarnds NRS ZMLZMU, Bjorkfors,Fflrkammaren in Sorsele NRS, Gargntis, GiltjaurNRS, Htiggis NRS, Jiirnforsen GNM, Laxfjail NRSZML, Rodingsbiick NRS, Slussfors, Sorsele NRS,Tjultriisk GNM NRS ZMU, Tiirna NRS ZML, VindelforsenZMU, Vuvosbticken, Viinniis NRS, ViistansjoZM A ZML, Ornds NRS. Lu. Lpm.: KvikkjokkZMC ZML, Lammeholm, Njunjes ZMA, Sarek NationalPark, Tarradalen NRS, Tjeurak NRS, VirihaureNational Park ZML ZMU. T. Lpm.; Abisko $20 July-l0 Aug. 1950 (Dalenius).Unrevised records (Ander in litt.): Medelpad: SErikslund, Friinsta, Liden, Tynderci, Stode, Asto.Jrimtland: Follinge, Hammerdal, NW Stromsund.Lapland (At. Lpm.).' Granliden 40 km SSW Vilhelmina.P. Lpm..' Jtickvik.Finland. Isolated occurrence NW Ladoga. Inpresent Finland observed only in Aconitum biotopesin Northern Karelen (Kb).List of localities. Kb: Kitee ZMB,Tohmajtirvi ZN'.4.AZ}MB; Kl: SortavalaZMAZ}i'4H. These localities arealso listed by Elfving (1968).

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Studies on Scandinavian Bumble Bees 125southern Norwegian populations is revealed inTable X. It is significant in two of the measuredfeatures, viz. 'radial length' and interalar width.These apparently clinal variations do not supportsubspeciflc designations.A possible difference between the northern andsouthern Scandinavian population characterizedby the ability to produce melanic individuals,is discussed on p. 135.B. hortorum var. fidens (Harris) ( - Apis fidensHarris 1776), B. hortorum var. harrisellus (Kirby)(- Apis harrisella Kirby 1802), B. hortorum var.nigricans Schmiedeknecht 1878 ( -,-= B. hortorumvar. c Thomson 1872), B. hortorurn ab. quasffidensVogt 1909 and B. hortorum f . balticzs Scholz 1924,one or several of which mentioned as tace orvariety occurring in Scandinavia (Sparre Schneider1898, 1918 ; Lie-Pettersen 1901, 1905, 1907 ;Friese & Wagner 1909 ; Vogt 1909 ; Meidell1934a; Lsken 1949 , 1960 ; Erlandsson 1960)concern infrasr.lbspecific forms only. Two of thedesignations need to be discussed: (1) the descriptionof Apis fidens, supplemented with acolour-drawing (F{arris 1776, p. 130, Table 38Fig. 3) presents a red-tailed bumble bee withyellor,v collar, scutellum and interrupted bandon Tr. The identification is uncertain. The nameis wrongly applied to the completely black formof B. hortorum. Dalla Torre (1896) re-establishedits specific rank, but considered the taxon asa dark form according to his list of references.Since then a number of authors, cf. above, havetreated Apis fidens as a melanic form of B. hortorum,but the colouring and the shape of thebody in the original drawing refer better to aform of B. pratorum than to the slender B. hortorum.(2) Apis harrisella is a melanic form ofB. ruderatus (Fabricius) and the name is thereforeerroneously applied to B. hortorum./Queen, workerMorphological characters. Malar space markedlyless than twice the distal width, occasionallyhardly exceeding lyz times this width, about aslong as A21 3a a. Disc of clypeus almost impunctateexcept for rather sparse, coarse puncturingof feeble anterior lateral impressions. Supra-orbital line usually touching laterul ocelli dorsally(Fig. 17 A) (in workers often just transectingthe ocelli). An transverse. Anterior median hairlessfield of scutellum small, usually dull. Midbasitarsus(Fig. 17B). Outer surface of hind tibiaslightly or not alutaceous. Body slender, coatrather shaggy.Queen measurements. N - 20; SE Norway;malar space: 1.24 mm (+0.04+0.01) range:1.20-1 .35 mm ; 'radial length' : 4.59 mm ( +0.1 1+0.02) range: 4.40-4.80 mm; interalar width:5.1 1 mm ( +0.18 +0.04) range : 4.90-5.70 mm.Body of large size.The workers vary greatly in size, the largestones being difficult to distinguish from thequeens.Colour pattern Collar extending to adjacentmargin of episternurn, crescent-shaped posteriorpaft of scutellurn, Tr, basal margin or anteriorlunate part of T, lemon yellow-haired. Extremeposterior margin of Tr, Tn-u and variable partof fringes of St^-u (or '-u) whitish-haired. Coatotherwise black. Interalar band posteriorly curvedor v-shaped, i.e. including black hairs of scutellum.Variation. Yellow collar varies in width,amount of black hairs on anterior pafi of scutellumand that of whitish hairs on posterioredge of T, display some variations. The strikingvariation is, however, the tendency to melanismwhich is discussed below.MaleMorphological characters. Malar space abouttwice the distal width or nearly, shorter thanAr* n* u but longer than Ar* r* n. Longest hairsin pile of face about as long as antennal scape.Ar-, (Fig. 39A). Hind tibia with a few bristlesexceeding the distal decumbent hairs of margin;fringes of hind tibia markedly longer than thegreatest width of the segment (Fig. 39B). Hindbasitarsus not much wider distally than at thebase. St, and genitalia (Figs. 39C-D); lateraldepressive of gonocoxite well-defined (Fig. 39E);distal margin of gonostylus, seen from behind,curved; boot-like, apex of volsella (Fig. 39F)varying slightly in shape. Coat shaggy. Body ofmoderate to large size.9. Bumble Bees.

\os(,o\ooN)N)@O{OON)(,5N)@(,5" FE?eSeWFloF frfttr HE F.? ff Lt tt tr'' ? E s e F h c:-F -+-toB, It r l i l d = N ) l r f doF.. rE. Q\ lP t9 Lh H N) O\ N \O { Oo O rr Fr N) (, (,C) A O\ H (r l\) \O O\g.?Hoo(/l o\ 5 (, o\ o u c\ f-.-. o\ Lh (, o\ o\ o o \o o \o o\ o\ w oo (, - o trr \o ur rooFDo\o Xr \r+s '-Gs5=, 5 | , NlH-. rr riN) d eI rr N O t9 oo'r \O A rr ao\ 5 (, c) r- tJ or I trr I N) { 5 o q (.,t (, o\ N) o\ t9 -zo o o o o o C) rr (,rr .F, (, lrc) o N) o\ Lh o\ c) o\ oo nF H H \ori -{ (, | | I | | | | | | | | | | ,---$ | | | ,.- | | I I I I \o | | I IFA tJ tr(ro\oNo\\ortf$'NNO\ = =lr Fr F 1.1e \O Lrr C\ C) F{ 6 rr O O O O O O O O S O O C) e C) m O rr rr { (, b) O Oo \Oli(r\O,It l l l t l l l l t t I l rro\OLrr I | | | | | | | | l s r l I | |rr.tr ^-S urFdN)o\Gro;H*8 sFrsLrr XSs{G-sG--i oo Lh o\ 5 t-*ro\ q oo o\ |r Fr o\ (, rr c\ s o rr.tr -*I ri 6 N +' \o Q rr { (,+. \oO H9 3F..= o ! J' traH \ JHfr. iJoblax'*85'8 nF FstiJ l-(i i . , HolJ* 3,$oiJPcroFtFna.P.FDq)-lo!0O r-1!1 FD5'go o6'X5 !- o qto | N { t c) o | F f - N) fr (, (, 5 N ) \-l o - f f d F rr.n a- --J oo (, A \.I o\ \o oo ur s \o N)a F d | \/ v \/ .- O O N) +. O O rr ir -{ , N) r- r- I U"r I O { OO 5 O\ (Jt OO (, (/tv v v v v^H(r{O\ r Lr), N)-. frFrI | 5 \ o o o or . , . r I O O O O O O o \ O O O r r O o o - o d U i o o \ u r r u r S t . . r O o \ o o N) OO O(r(ro\{ (, N)N)F HN)r-N) O\ \O \O -i t'J \O O (, O O -.f O O O tl.) O qrr O { { O\ O\ 5 6 O OA F rv Fts.= o i l. H Aa\ \,a-f-{ \-,a EE!a '*' Ai - 9 9I ' i Oo rct?8 B= o c >. - Ag) \Ji J F 1a 6'6=:rq'' , x oo )dct)l+l ' voF-tooF!oo t')T1qc\o\oY ():-ilo X80o O.ua4o7'v9Zr

Studies on Scandinavian Bumble Bees 127Colour patternPile of vertex with variablemixture of black and yellow. Yellow collarextending more or less down episternum. Basalpart of Tu, Str-u whitish-haired, remainder ofTe, T? black-haired. Otherwise colouring as inthe female.Variation. In general as in the female.Dtscussion on melanismMelanic forms occur locally, with the typicalform throughout Scandinavia except for thenorthern pafi of the peninsula. All degrees ofmelanism have been observed, from individualswith variable amount of yellow hairs replacedby black ones. to completely black specimens.The frequency and the degree of melanismthroughout the peninsula are illustrated on the piechart (Fig. 74) based on the figures set out forthe examined collections in Table XI, XII. Specimenswith trace of yellow hairs are grouped asincomplete melanics. Entirely black or brownishblackindividuals as well as those with no traceof yellow hairs visible by naked eye, but withhindmost tergites presenting a variable amountof whitish hairs, are grouped as complete melanics.The pie chart demonstrates a more pronouncedmelanism in the population inhabitingthe swedish islands oland and Gotland than inthe neighbouring mainland. There is, moreover,a distinct, apparently clinal, decrease in melanismTable XII. B. hortorum (Linnaeus). Proportion and degree of melanism throughout Sweden, based on collections.Area : county or district thereof (cf. Fig. 99)Areatyp.formNumber of individualsincompl.mel. formcompl.mel. formTotalnumbertyp.formPer centincompl.mel. formcompl.mel. formsk.Bl.Hall.Sm.Ol.Gtl.Oe.Ve.Boh.Dlsl.Nrk.Sdm.Upl.Vstm.Vrm.Dlr.Gdstr.Hls.Med.Hrj.Jmt.Attg.vb.Nb.As. Lpm.Ly. Lpm.P. Lpm.Lu. Lpm.T. Lpm.392t62l253861038694l150226273333r3778r23 /81046112126820t371320955t4900t24018I2092450l5500001002000005II11t91320I50I30840000100000000406l7222761454l4l7l42164228278341137101132810461121468201391320955t42869 125 75 306997949692592493979892999898100779310010010010099100100991001001001001002004284423061II01540000I00I00000I6541332502300I0830000100000000

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t--lt(.- a \gi\J/I-aStudies on Scandinavian Bumble Bees129from the southwestern coast of Norway towardsnortheast, i.e. from an atlantic to a more continentalclimate. Yet the frequency of melanicscannot clearly be correlated with climaticalgradients; cornpare, for instance, the precipitation(Fig. 1). The proportion of melanism in therather continental Southeastern Norway is noteworthy,whereas dark coloured specimens havenot been observed along the western coast northof about 61' N.The evaluation of melanism is based on astudy of 2834 Norwegian and 3069 Swedishcollected specimens. A total of 1682 observed,but not collected Norwegian individuals quotedby O. Meidell (minor part) and me as belongingto the typical, incomplete, or complete melanicform, has also been considered. As the availablematerial was not recorded at randoffi, the strikinglydark individuals may have been collectedmore often than those of the comrnonly knowntypical form. The proportion of melanism basedon the total of collected and observed, but notcoliected material f;gured in separate columnsof Table XI, XIII-XIV, favours this suggestion,i.e. the frequency distribution of the melanicforms presented in collections is in general higherthan in the actual population.The only at random study was carried outbetween 31 July-4 Aug. 1963 and 5-7 Aug. 1966in various fields in Jaeren inrRo9aland (Ry),an area where the proportion of melanismreaches a maximum (Fig. 74). Every observedspecirnen of B. hortorum was then couuted andquoted as belonging to the typical form, incompleteor complete melanic form respectively. Inorder to save the beneficial insects, only a partof the records was collected and kept. The resultis demonstrated in Table XIII,which also includesthe corresponding data for the remainingmaterial (not recorded at random) from theouter district of Rogaland. The frequency registersof the total of collected, and observed butnot collected individuals should reflect fairlywell the frequency and degree of melanism inthe local population. The difference in the proportionof incomplete melanics, i.e. 5% and 7%in 1963 and 1956 respectively, in relation tootherwise 13"A, may partly be due to subjectivejudgement in grouping, particularly of recordedaC)U^-:'FFrn.9aq)a().2()()8dt_r,_i>'=rl .=f-lIv d' o d(-r- ) O:v +)9 0$i t-'to Fz6C).Et ()E3A s ,.i \JOtrar - 6r, A- vvO ?q.)Xt- \.Jtrn''- vgr9q ) L )-'o c)sEec)4 r v t ld a )FA f _v . -'-: , HL i i/1 cgXXli tlA, rla a ,F{ \JF ' f \C)\J\vtrbo'r q)-Jsr..-z$FF \.,G\ F T\ 4 +H \ Vbs: A_\.,: c sr\.. dAA \\J\ s. eF l vx&F?- )(.) c\3Fa LV..- +_)- aO,()Fa()Izo :(.)

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Studies on Scandinavian Bumble Beesl3lmoreovet, a non-sex-associated variation, yet thefrequency of melanic males is higher than thatof the females. The proportion of melanisffi,based on the representative material from Oppland(Os) and Rogaland (RV) (cf. above), andthe great number of records from the fairlywell investigated Hordaland (HOy), is set outin Table XV and demonstrates that melanismis produced in sex and caste. The queen/worker/male ratio of complete melanics beine slqn inOppland (Os), 70169193 in Rogaland (Ry), and32129151 in Hordaland (HOV) is noteworthy andemphasizes the statement above. The frequencyof melanics in per cent is everywhere about thesome in queens and workers, and markedlyhigher in males. A difference in sex is alsoaccentuated by the degree of melanism beingmore pronounced in males. By definition, individualsgrouped as complete melanics comprisethose with a black coat with various amounts ofwhitish hairs on hindmost segments admixed,and entirely black specimens. The proportion ofcompletely black individuals is, however, largerin males than in females; entirely black queensand workers have, in fact, rarely been recorded.The observations agree with those of Lie-Pettersen(1901) and Sparre Schneider (1918), whoboth observed a non-sex-limited local tendencyto melanism characterized, however, by a highfrequence of strikingly black males.Distribution (Fig. 75)Norway. Widely distributed from the extremesouthern coast north to about 70" N. In SouthernNorway occurring from sea level to subalpinevalleys and occasionally penetrating the lowerbelt of alpine zone. In Northern Norway confinedto luxuriant subarctic biotopes. The frequencyis locally high. In Southern Norway recorded1300 m s.m.Biotopes: Gardens, orchards, leguminosaefields and other biotopes providing proper foodplantsas Aconitum, Galeopsis, Vicia, etc. Heatherand barrens are avoided.A total of about 2900 specimens was examined.List of localities. Astfold.' Hvaler: Herfol, Kirkoy,Sondre Sando; Krflkeroy: Odegflrd; Onsoy: Ellingird,Ramseklo, Skjreloy; Sarpsborg: ZMO; Halden:Sponvika; Marker: Dybedal, Rodenes; Rakkestad:Rorvik; Ride: Oven, Tom; Moss: VCA, Jeloy VCAZMB; Eidsberg: Holm; Trogstad : Monster bro,Bistad. Akershus: As: Vollebekk; Frogn: Drobak;Table XV. B. hortorum (Linnaeus). Frequency of melanism in sex and casteNo.dTotalper cent No. per cent No. per cent No. per centOppland (Os)Typical formIncompl. mel. f.Compl. mel. f.305281t4537 1006943769l5410011I6186l63310011010111318488100Rogaland (Ry)Typical formIncompl. mel. f.Compl. mel. f.Hordaland (HOV)Typical formIncompl. mel. f.Compl. mel. f.34 22 6l 23 5 6 100 2013 8 22 8 1 I 36 7110 70 184 69 74 93 386 73157 100 267 100 80 100 s04 10060133456l2329l224757t42935114837I251186461,29511336r07 100 160 100 100 361 100

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Studies on Scandinavian Bumble Bees 133ZMO; Lom: Galdesand, Solell 1200 m; Lesja: Lesja,KMT; Dovre: Dovrefjell ZMCZML. Buskerud (Bri) :Hurum: Filtvedt TRM, Pinadalen, Storsand; Royken:Aros; I)rammen : ZMB ZMO; AOat: Hen; Krodsherad:Bjorkerud, Brfrtfln, Fagerheim, Glesne, Veikflker;Modum: ? loc. ZMO, Snarum; Flesberg: Lampeland,Oydegflrden; Kongsberg: TRM ZMO, Komnes,Lintvedt, Skollenborg. Bv: Sigdal: Kopseng,Nedre Eggedal, Nordbygda, Prestfoss, Sandsbrflten,Sigdal; Nore-Uvdal: Hegnum seter, Norstebo 700 m,Tunnhovd, Uvdal stavkirke; Flfl: Kve gArd; Nes:Bergheim bro, Bortnes. Eidal; Gol: Gol, Randalseter940 m; Flemsedal: Lykkja 900 m; Al: ? loc. ZMO;Hol: Geilo 800 rn VCA ZI[{B, Haugastol 1000 mVCA ZM.B, Nygird 1000 m, Seirn, Ustaoset 1000 m,Vikastol 1000 m. Vestfold: Sande: SW Sande; Holmestrand: Angerskleiv; Borre: Adal, Nykirke; Vflle:Ryk; Ramnes: Lunde; Lardal: Styrvoll; Andebu:{ndebu; Stokke: Lango, Sand, Veierland; Notteroy:Teie; Tjome: Kjrere, Tjome; Sandefjord: Austeroy;Hedrum: Ringdal; Brunlanes : Dolven, Helgeroa;Larvik. Telemark (fEy ); Skien : Kikut; Porsgrunn :Mule; Bamble: AUv; Ncme: Damtjern; Drangedal:Drangedal, To-myra. TEi: Nctcdden : Gransherad,Tinncset; Seljord: Svartdal; Kviteseid : Brunkeberg,Morgedal; Nissedal: Kyrkjebygda, Lauvviki; Tokke:Amdals verk; Vinje: Bossbo en 970 m, Krossen 700 m;Tinn: Miland, Tinn, Vfllen. Aust-Agder (AAy); Gjerstad:Fiane; Tvedestrand: Eidbu, Nes verk TRM,Osterfl; Moland: Dal, Holmsund, Kilsund; Arendal:ZMO; Hisoy: His; Grirnstad; Landvik: Hombor;Lillesand: Kvflse, Langholmsund, Natvik, Rosnes,Troe. AAi: Bygland: Londal; Valle: ? loc. TRM,Kvestad, Rygnestad; Bykle: Breive 700 m . Vest-Agder(VAy); Kristiansand: ZMC, IV[ovik; Sogne: Aros;Lindesnes: Lindesnes, Rarnsland; Lyngdal: TRM;Kvinesdal: Gjemlestad; Farsund: Fjellesfl, Lista fyr,Lodshavn, Vanse; Flekkefjord: Hidra. Rogaland(Ry); Sokndal: Bu, Hauge i Dalane, Nesvflg, Rekefjord,Vatland, Amot; Eigersund: Birkeland, Eigeroyeidet,Fotlandsvann, Klungland st., Leidland, NedreHetland, Skerpe, Tengs; Bjerkreim: Malmeim; H6:Anisdal, Bjelland, Brusand, Nrerland, Ogna ZMr*ZMO, Opstad, Salte bro, Sirevflg, Stolen, Varhaug,Vigrestad; Klepp: Borsheim, Bore bro, Innre Reve,Gjeishaug, Horpestad, Jreren TRM ZMB, Kflsen,Orre, Orrevann, Reve, Skarsheim, Vik, Oksnevad;Time: Kartevold, Mcssige, Time, Tjflland; Gjesdal:Nese; Sandnes: Figgjo, Ganddalen, Hana, Hornmersflk,Ims, Kjellingland, Myrland; Sola: Reke, Slettehei,Solastrand, Tananger, Olbergstranda; Randaberg:Randaberg, Sande; Stavanger: Hafrsfjord, Hinna,Lindoy, Madla ZN4B ZMO, Sunde; Strand: Tau,Vatne; Finnoy: R.eilstad, Sjernaroy SMS; Rennesoy:Dale; Kvitsoy; Karmoy: Eide, R.isdal, Sandve, Sreveland,Vigsnes, Vikre, Akra; Vindafjord: Frovik; Tysv&r:Leiranger, Nedstrand. Ri; Forsand: Forsand,Meling; Hjelmeland: Fister, Flaugsli, Igland, Kvamme,Steinslandsvann, Vadda, Ardal; Suldal: Leirdalen700 m. Nesflaten. AimioOlonuten 700 m: Sauda:Sauda. Hordaland ( HOy ).' Bomlo : Mosterhavn;Sveio: Forde; Olen: Dommersnes; Fitjar: Koloyholmen,Rubbestadnesset, Sorfonno; Austevoll:Karlsoy, Lunde, Ytteroy; Os: Berge, Hauglandsdal,Kvalsund, Lysekloster, Moldegflrd, Nordstrono, Rod ;Fusa: Ovre Hilandsdal; Samnanger: F{oyseter, Adland;Fana: Biol. st., Bjines, Blomsterdalen, DolvikVCA, Fana kirke, Fjosanger VCA, Flesland, Grimstadnesset,Hjellestad, Krokeide VCA, Milde, MindeVCA, Nesttun VCA, Skipanes VCA ZN4B, Skjold,Smor6sen, Stend, Titlestad; Sund: Bokken, Telavhg,Tyssoy; Fjell: Eide, Fjell, Fossavann, Landro ; Laksevflg:Storingaviken; Bergen: TRM VCA ZNIr-- ZMO;Askoy: ? loc. VCA, Herdla VCA ZMB; Asane: Haukedal;Osteroy: Gjerstad, Havratun, Haus, Kleppe,Nj6stad, Rakanesset, Skolmen; Meland: Brakstad,fo, Jacobsoy, Kjeken, Landsvik; Radoy: Kvalheim,'skjaergirden' VCA TRM ZMB; Lindfls: Fosse, Lauvis,Vollom; Fedje. HOi: Etne: Oslandsvflg, SkinevikTRM ZMB; Kvinnherad: Berget, Guddalsdalen,Lio, Ljosmyr, Prestvannet, Rosendal, Varaldsoy;Odda: Dalen gard, Roldal, Valldal 900 m; Ullensvang:Djonno. Espe, Fossli 750 m, Fresvik, Gnychorgen900 m, Hardangervidda VCA, Fijolmo, Kinsarvik,Lofihus, Mebodalen, Sysendalen 750 m, Tveito, UtneVCA, Viveli 875 m, Ovre Eidfjord; Kvarn : Omastrand,Strandebarm Z}}y'r3, Vartand, Asheirn; Voss:Gossland, Kinne, Rogn, Vcss VCA ZMB; Granvin:Eide VCA, Granvin, Holven; Ulvik: Berge, Ulvik.Sogn og Fjordane ( SFy ).' Gulen : Austgulen, InnreBrekke; Hyllestad: Hatleim, Hyllestad, Skivenes, Alefjell;Hoyanger: Lavik Z ll4B ZMO; Gaular: Bygstad,Eldalsosen; Fjaler: Dale, Strandene$; Askvoll:Aralden, Askvoll, Grytoyri; Flora: Florelandet, Floro,Kinn, Nekkoyna, Rindemyr, Verpevik; Naustdal:Horstad, Naustdal; Forde : Flflten, Hallbreim; Jolster:Alhus; Gloppen: Hjortset, F{ope, Lotsberg, Lote,Sandane; Eid: Kjolsdalen, Naustdal, NordfjorCeid,Sletta, Stirheim; Vigsoy: Kjolstad, Refvik; Selje:Frvik, Lekanger. SFi.' Vik: Vik; Aurtrand: Berek!'am,Flflm, Gudvangefl, Kvamsh agen, Nedre Aurlandsdal,Sinjarheirn 650 m, Uppsete 850 m VCA ZN.{B., Vassbygda; Laerdal : Laerdal ZMO, Ystabo ; Scgndal :Slinde; Leikanger: Fosshagen, F{amrestol, Hermansverk,Leikangerseter; Balestrand : trlesje, Fjrerland,Horpedalen, Suphellen, Suphellesetra; Luster : FortunTRM, F6berg, Gaupne, Jcstedal, Skjolden, TurtagroTRM ZMB; Stryn: Flo, F{jelledalen, Hornindal, Clden,Stryn, Videdalen, Videseter 7C0 m, Ovre Oldenvann.Mrire og Romsdal (MRy); Volda: Bjorkedalsvann,Straumshavn; Hareid: Hjorungavig; Ulstein:Flo, Ulsteinvik, Vonheim; Heroy: Leikanger, Runde;Alesund : VCA; Vestnes: Gjerrnundnes; Molde;Aukra: Aukra kirke, Rod; Fraena: Bud. MRi: Orsta:Viddal, Oy.; Stranda: Fiveistad, Geiranger NMW,Vcllset; Norddatr: Indreeidet, Valldal ZN444r-- ZMO;Raun:a: Veblungsnes, At< zlrO, Andalsnes; Nesset:Torhus; Surnadal: Lahjel, Melhus, Stangvik, OvreSurnadal ; Rindal : Bolme. Scir-Trrindelag ( STy ) :Hitra: Hamn; Rissa: ?loc. VCA, Sotvik; Orland :

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Studies on Scandinavian Bumble Bees 135ZML, Valsiitra IVU, Vassunda IVU, Vindo NRS,Vaddo NRS, Yxlan Zll4B, Ossebygarn NRS, OstraRyd NRS. Vristmanland: Arboga ZN{{{B ZML, FanthyttanNRS, Fellingsbro Zi|dB, Guldsmedshyttan,Hiillefors NRS, Kolbtick NRS, Kdrrbo ZMB, LindesbergNRS, Sala NRS, Skultuna Z},.dB, As NR.S.Vcirmland: Arvelsiiter NRS, Arvika, Frykerud NRS,Gylleby in Sunne NRS, Hammaron NRS, FtrorrsjonNRS, Lflngban NRS, Molkom NRS, Nilsby, SiiffleNRS, Olme. Dalarne; Aspeboda NRS, Avesta ZMB,Brunnsvik NRS, Blyberg NRS, Bingsjo NRS, Idre,Leksand NRS, Ludvika NRS ZML, Mora NRS,Riittvik NRS, Sandsjo, Sjurberg NRS, Smedjebacken,Stjiirnsund, Store Tuna, Sundborn NRS, Siirna, SdterNRS, Transtrand NRS, Vikarbyn NRS, VristanvikNRS . Gcistrikland: Hamringerfjiirden NRS, Hille-Finland. Commonly occurring throughout thecountry north up to the Polar Circle and sporadicallyobserved further north, yet not recordedin Lapland (Le) (Elfving 1968).World distribution. Europe (British Isles; entireWest European continent; European USSR,except semi-desert areas north of Sea of Azovand the Caspian Depression) - Algeria - Turkey- Iran - Caucasus - Transcaucasus - Northernand Eastern KazakhstanWestern SiberiaTransbaykal Baykal Jakutia NorthernMongolia (Panfilov 1957 and in litt.; Reinig 1968).Introduced to New Zealand (Gurr 1964).Z oogeographical re mark sThe tendency to melanism confined to onlypart of Scandinavia justifies the question whetherB. hortorum consists of two Scandinavian populations:(1) a northern post-glacial populationwith gene-pool light, immigrating from the eastand (2) a southern population with gene-pooldark, i.e. with melanic traits, invading the peninsulaForsby NRS, Hogbo NRS, Trodje. Hcilsingland: Bollnds,from south. The theory is favoured by theGnarp NRS, Holmsveden NRS, Los NRS, fact that a tendency to melanism along the easternNjutflnger NRS, Orbaden NRS, Soderhamn NRS.migration route has not been observed (PanfilovMedelpad: Liden ZMB, Mattfors, Stavreviken NRS,Sundsvail, Ange . Hcirj edalen : Hamrafjiill, Ljungdalen, in litt.), whereas it is recorded, but the frequencyTdnndalen NRS ZML. Jrimtland: Bydalen, Doda not estimated, on the continent south of ScandinaviaFallet NRS, Eldsisdalen NRS, Froson NRS, Giiddede(Alfken l9l3; Stoecklrt 1933; Schmiede-NRS, Hallen, Jormlien, Jdrpen, Mattmar, Mullfjtill,knecht 1930; etc.). The present information aboutOttsjon NRS, Oviken, Snasahogarna NRS, Storlien,Svenstavik NRS, Undersiker NRS ZML, Ostersundthe identity of the northern and southern Scandinavianpopulations is meagre; anyway, a dif-NRS, Are, Areskutan. Angermanland: Bjiista NRS,Bjorna NRS, Bondsjo NRS ZML, Forsmo NRS, ference as regards. presence and absence ofHikniis NRS, Kramfors NRS, Ramsele GNM, SollefteiNRS ZML, Sidensjo NRS, Siibri, Ornskoldsvikmelanic traits does not qualify for subspecificdivision of the Scandinavian B. hortorum.NRS. Vcisterbotten' Bodarna NRS, Degerfors NRS,Hiillniis, Jorn, Urnefl NRS ZML. Norrbotten: Anttis,Haparunda, Kalix NRS ZML, Karungi NRS ZML,Lulefl NRS ZML,Lingtrdtsk, Munksund NRS, NybyggetBiologyNRS ZML, Pajala NRS, Rflnei, Overkalix NRS,Nest. Pocket-maker. Nests were recorded atOvertornei. Lapland (At. Lpm.) : Bingnds, DoroteaNRS, Fattmomakke NRS ZML, Frederika NRS,various depths from 50 cm to just below theHunneborg, Vilhelmina NRS, .A,sele NRS. Ly. Lpm.: surface of the ground. Produces large coloniesAmmarniis NRS, Bjorkfors, Lycksele NRS, Sorsele, (Dahlbom 1837; Lie-Pettersen 1901 ; HasselrotTdrna, Vindelforsen NRS, Vtistansjo. P. Lpm.: Skattriisk.Lu.Lpm.: Gtillivare NRS, Kvikkjokk, Malm-1e62).Flisht season. From middle of April to endberget, Njunjes, Sarek National Park, Sitasjaure. T.Lpm.; Abisko NRS, Jukkasjiirvi NRS, Kiruna NRS, of September. Queen: I May-9 Sept. ; worker:SvappaYaara NRS.Ander in litt. : Gotland; Fide, Lirbro. Vristerbotten:10 May-27 Sept.; male: 3 July-Z7 Sept. Twomales collected 10 May 1908 (Akershus: Moss:Hokmark, Lovanger.Jeloy (Barca) ZMA) illustrate that unfertilizedeggs are accidently laid in the first batch oftrrood.BOMBUS (MEGABOMBUS) RUDERATUS(FABRICIUS)Apis ruderata Fabricius, 177 5, p. 380 No. 7 ,lectotype ? BCL ! selected by Richards accordingto Zimsen (1964, p. 415 No. 1074). Type areaMadeira.

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Studies on Scandinavian Bumble Bees 137whitish-haired. Otherwise coat black. Interalarband parallel-sided.Variation. Yellow collar varying somewhat inwidth, yellow hairs of T' ranging from a slightadmixture in lateral patches to the entire tergite.White on Ta more or less pronounced, particularlydistal part of T5 with variable admixtureof black hairs. However, variations are minute,i.e. Scandinavian ?? and VV are colour-stable.MaleMorphological chcracters. Malar space nearlytwice the distal width and about the length ofAr* r* u. Longest hairs in pile of face usuallyshorter than antennal scape. Hind tibia withonly very few bristles, none of which exceedin general the decumbent hairs at the distalmargin; longest hairs in fringes of hind tibiabarely or not exceeding the greatest width ofthe segment (Fig. 38A). Hind basitarsus widerdistally than at the base. St, (Fig. 38ts). Apexof volsella (Fig. 38C) varying slightly in shape.Body of moderate to large size.Colour pattern Pile of vertex usually pronouncedyellow, anteriorly and laterally encroajhedby black hairs. Collar, usually extendingdown episternuffi, T', median anterior lunatepart of T, yellow-haired. Entire or the rnain partof Tu, St1-u and lateral patches of Sto-z whitishhaired;remainder of St u-, with black hairs.The colouring otherwise as in the female.Variation. Gena and venter of head with avariable admixture of whitish hairs, otherwisevariations as in the female.Distribution (Fig. 76)Scandinavia. Restricted to southernmost Sweden,which is the northern limit of the distributionof the species.Biotope: Apparently prefers rather arid areas.A total of about 135 specimens was examined.List of localities. Skdne: Alnarp, Bjiirred, Bjornstorp,Blentarp, Bulltofta, Bokeberg, Dalby, Eslov,Falsterbo, Glimslov, Hohog, Hiilsingborg NRS ZML,Hflslov, Klostersagen, Kniistorp Z}r dB., Krageholm,Kungshult, Kyrkheddinge, Kdvlinge TCL ZML,K6selberga,Lund GNM ZldB ZML, Loddekopinge, LoderupNRS ZML, Malmo, Nosaby, Nytorp" NiisumGNM, Ravlunda, Rea, Simrishamn NRS, Sjostorp,Skarhult, Sodra Sandby ZI|I{B ZML, Torekov, TrelleborgGNM ZML, Veberod, Vellinge, Villands Vflnga,Vinninge, Vitemolla, Ystad NRS, Ortofta, OstratorP,Oved NRS ZML, Ahus, Alabodarna. Blekinge; Hdllevik,Kristianopel GNM. Halland: Falkenberg, Flalmstad,6stra l(arup, Aku[a GNM.The total distribution of the subspecies, B. ruderatuseurynotus Vogt. Europe (cf. the worlddistribution of the species) with exception ofthe Iberian peninsula and the Mediteffaneanisles (Pittioni & Schmidt 1942).BiologyFlight seeson. From the middle of I\4ay tothe end of September. Queen: 12 May-26 Sept.;worker : 7-21 Aug.; male : 24 July-22 Sept.SUBGENUS THORACOBOMBUS DALLATORREThoracobombus Dalla Torre, 1880, p. 40, typespeciesBombus sylvarum (Linnaeus) by designationof Sandhouse (1943).( Chromobombus Dalla Torre, 1880, p. 40,type-species Bornbus muscorum (Linnaeus) bydesignation of Sandhouse (1943). AgrobornbusVogt, 1911 , p. 52, type-species Bornbus agrorltm(Fabricius), 1787 nec Schrank, 1781 by designationof Sandhouse (1943). RuderariobombusKrtiger, 1920, p. 350, type-species Bombus rude'rarius (Mtiller) by designation of Yarrow (1971).Adventoribombus Skorikov, 1922a, type-speciesBombus sylvarum (Linnaeus) "by designation ofYarrow (1971), who refutes the designation ofSandhouse (1943), viz. Agrobonfuts ( Adventori'bombus) adventor Skorikov, 1922 ( - Agrobom'bws adventor Skorikov, l9l4a), as being invalid.)Queen, workerHead longer than wide. Clypeus, malar spacehardly to markedly longer than respective distalwidth, yet less than l1/z times this width (occasio-

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Studies on Scandinavian Bumble Bees 139are often difficult to distinguish from the queens,particularly in B. pascuorum (Scopoli).MaleHead longer than wide. Clypeus, malar spaceslightly to markedly longer than respective distalwidth. Mandible bidentate with dorsal toothsmaller than ventral tooth. Eyes directed infront of posterior mandibular condyle. A, markedlyshorter than Au, Au-r, individually abouttwice the distal width or nearly So, at leastAr-r, slightly to strongly swollen underneath(Figs. 444, 45A, 46A, 47 A, 48A, 49A).Outer surface of hind tibia usually convex withsparse to rather dense distribution of more orless decumbent hairs of various length throughout;Iongest hairs in posterior fringes of hindbasitarsus not exceeding the greatest width ofthe segment. SJ, and genitalia as in Figs. 44 B-D,458-D,468-?C , 47F._.C,4gB-E , 4gB_.F,.Scandinavian speciesThe subgenus is represented by six Scandinavianspecies, viz. B. humilis Illiger, ,8. muscorum(Linnaeus), B. pascuorum (Scopoli), .8. ruderarius(Mtiller), B. sylvarum (Linnaeus) and B. veteranus(Fabricius).BOMBUS (THORACOBOMBUS) HUMTLTSILLIGERBombus humifts Illiger, 1806, p. l7l, No. 48 necYarrow 1968, type V IBZ!, type area Germany:Niirnberg.(Bombus muscorum: Dahlbom 1832, p. 46(partim), 1837 (partim); Wahlberg 1854 (partim);Thomson 1870 (partim), 1872 (partim); Aurivillius1903 (partim). Bombus humilis Illiger; Tjeder1954; Frid6n, Eskilsson & Bingefors 1962. Bombussolstitialis Panzer, 1805 or 1806, paft 99Teil 17 , type not traced; Sparre Schneider 1918 ;Erlandsson 1948; Ander 1953a, 1963, 1965.Elfving 1960, 1968; Hasselrot 1962; Frid6n 1967.Bombus helferanus Seidl, 1837, p. 66, type lost.Bombus variabifts Schmiedeknecht in Radoszkowski,1877 , p. 199, type lost; Schmiedeknecht 1878,p. 424, Bengtsson 1907, 1908; Meidell 1934a;Krtiger 1940. Bombus solstitialis var. auranticusDalla Torre, 1882; Sparre Schneider 1918. Bombusvariabilis var. staudingeri Dalla Torre, 1882;Meidell 1934a. Bombus helferanus hafsahli Vogt,1909, p. 36, type ? VCA, type area Austria:Tirol. Bombus variabilis hafsahli Vogt; Kriiger1940, pp. 281, 386. Bombus humilis hafsahli Vogt;Laken 1960. Bombus humilis hafsahlianus Vogt,1947 , p. 4, lectotype P VCA ! hereby selected.Labels : (1) Drobak; (2) 26 h; (3) collecti C.et O. Vogt; (4) lectotype B. humilis hafsahlianusVogt design. A. Lsken 1969. Type loc. Norway:Akershus: Frogn: Drobak; Kruseman 1959.Bombus humilis hafsahioides Vogt, 1947, p. 4;Kruseman 1959.)Nomenclatural remarksAccording to Sherborn (1902-1933) the dateof B. solstitialis Panzer, Fn. Germ. part 99No. 17 is 1805 or 1806 whereas Horn & Schenkling(1928) gave the date of parts 97-108 as1806-1809. The year 1794 for this designation,as stated by Erlandsson (1948), is due to misunderstanding,&S the author confused number17 with part 17 published 1794. Though it isuncertain whether B. solstitialis or B. humilis isthe senior synonym, the latter is kept for thisstrongly variable taxon, because the identityrefers to preserved type. B. hurrufts is, moreover,used by a number of European taxonomists(Knechtel 1955; Yarrow 1959; Kruseman 1958b,1959, 1960; etc.). The species has been confusedwith B. pascuorum and B. muscoruffi, and thegreat nomenclatural confusion of the three taxais illustrated in references listed by Dalla Torre(1896, pp. 504, 535, 560).First Scandinavian recordsNorway. Akershus: Brerum (Sparre Schneider1e18).Sweden Dalsland. Scidermanland, Vdrmland

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Studies on Scandinavian Bumble Beesr4lfollows: Pile of face, episternum, hind femur,and corbicular fringes pale yellow to yellowishwhite.Yellowish-brownhairs on dorsum ofthorax of paler shade. T2 only slightly darkerhairedthan T, i occasionally hairs on Tr-, displaythe same shade of colour. Tr-u withoutblack bristles. The figures from Skflne (TablesXVI, XVII) emphasize the stable colouring. Allthe individuals examined present entirely yellowish-whitehairs on episternum, femur, and corbicularfringe, and only 5 per cent of the individualsdisplay black bristles on one or severalof the tergites Tr-u. The available material fromNorthern Germany reveals, however, a closeaffinity between the 'hafsahloides' form in Skineand the so-called 'staudingeri' group (Kri.iger 1940)in Germ?ny,thus supporting the theory that SouthernSweden is a transitional area. Therefore onlyone Scandinavian subspecies is recognized, yiz.B. humilis hafsahlianus.Queen, workerMorphological characters. Malar space markedlylonger than distal width, about as longas Ar+ n. Clypeus hardly longer than distal width,on the disc with fine uneven puncturing exceptfor a small impunctate area between lateral,coarsely punctured impressions. Labral furrowmore or less widened towards the base, labrallamella with dull fine sculpturing and moderatelysharp edge. Frons with rather sparse coarsepuncturing, almost impunctate near visible paftof frontal line. Distance from lateral ocellus topreoccipital ridge hardly longer than distancefrom lateral ocellus to eye. Tr-, laterally withstriate-rugose microsculpture at least in theposterior half of the individual segments, i.e.hairs arising from coarse coalescing punctures.Distal keel of Stu moderate. ,Dorsal coat ofthorax markedly shorter and less shaggy thanTable XVI. Percentage of female -8. humilis llliger exhibiting certain characters of colour pattern in populationsranging from Southern Sweden to southern coast of Norway. Area - county or district thereof (cf. Fig. 99)Area Sk. Boh. o AK HEs Os VE TEy AAy VAyNumber of specimens 56233343t635 44 25Pile of faceYellow 98Mainly yellow 2Mainly to all black48302234255229 38 r769 62 8362 4 1 8 470 82 96 100EpisternumYellowish-white 100Mainly yellowish-whiteMainly blackBlack5213221336451532 13t9 13 t779 74 83116 9 1 632 20 2451 7t 60 100Hind femurYellowish-white 100Mainly yellowish-whiteMainly blackBlack6lt72242152l2l25l2816131368l783153829784t288 100Corbicular fringesYellow 98Tipped yellow/yellowish-brownMainly to all black 226304397l684316934663797 93100 1003rd to 5th tergiteWithout black bristlesWith black bristles955485218824961003 5100 97 95 100 10010. Bumble Bees.

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Studies on Scandinavian Bumble Bees 143worthy and indicate that the species occasionallyreach regio subalpina.Biotopes: Meadows, leguminosae fields.A total of about 400 specimens was examined.List of localities. Ostfotd (0).. Hvaler: ? loc. ZMO,Herfol, Kirkoy; Sarpsborg : ZMO; Halden : NRSVCA, Hakelund, Sponvika; Marker: Dybedal; Rflde:Tom; Moss: Jeloy VCA; Trogstad: Monster bro.Akershus ( AK ); As : Vollebekk; Frogn : Drobak VCA,Tusse TRM; Brerum: ? loc. TRM, Hovik Z}dO-Oslo: DCL TRM ZMIB ZMO. Hedmark (HEs)':Grue: Finnskog; Hamar: Zl|dB ZMO; Ringsaker:Vea. Oppland ( Os ).' Gran : Lander; Ostre Toten :NW Skreia; Oyer: Skfli, Oyer. On: Dovre: Dovrefjell2?? H (Schiodte) ZMC, H 1910 (Wretlind) ZM\J.Buskerud (Bd).' Hurum: Filtvedt TRM; Drammen;Modum: Modum ZMO, Snarum. Vestfold (VE) :Andebu: Kodal; Stokke: Lango, Stokke, Veierland;Notteroy: Teie; Hedrum: Hedrum; Brunlanes: Dolven,Helgeroa; Larvik: Byskogen. Telemark (TEy):Bamble: Brevikstranda, Trosby, Aby; Nome: Damtjern,Nos, Risdalen, Akredalen; Kragero: Levang.TEi: Notodden: Tindegrend; Sauherad: Liagrend.Aust-Agder ( AAy ); Risor : Nistevig; Tvedestrand :Dypvflg, Laget, Nes Verk TRM ZMO; Arendal:Salterod; Hisoy: Gjervoldsoy, His; Oyestad: Bjorbekk,Helle. Vest-Agder ( VAy); Kristiansand : ZMO:Lindesnes : Ramsland. VAi: Xr.rur i R;J"d.Sweden Widely distributed throughoutthelowlands northeast to nearly 64" 30' N in Vdsterbottenand penetrating reg. subalpina inDalarne.Moreover occurring in Gotland, but not recordedin Oland.A total of about 1250 specimens was examined.List of localities. Skdne: Barkikra NRS, Bjuv,Bjiirred, Bjornstorp ZI$'4B ZML, Blentarp, BrosarpNRS ZML, Bulltofta, Dalby, Degeberga NRS ZML,Drakemollen NRS, Falsterbo, Figelsing, GlimflkraNRS, Gtirdslov, Hannds ZMU, Hollviksnds, KristianstadNRS, Kyrkheddinge, Kiivlinge, Lack aldnga,Ljunghusen, Lund NRS ZML, Loddekopinge, LoderupNRS ZML, Maglehem NRS ZML, Norra Uglarp,Nosaby, Ndsum NRS, Revingehed, SandhammarenNRS, Silvflkra, Sjobo NRS, Skivarp, SodraSandby ZMB ZML, Torna-Hrillestad, TrelleborgZI.[B, Veberod, Vinslov, Ystad, Ahus NRS ZMIJ.Blekinge: Brdckne-Hoby, Hemsjo, Hiillevik NRSZML, Jtimjosliitt, Karlskrona NRS, KristianopelGNM NRS, Morrum, Sandbdck, Sjoarp NRS, TorhamnNRS ZML. Halland.' Fagered, Falkenberg,Fjiirfls, Halmstad NRS ZML, Harplinge NRS, Mellbystrand,Tronninge, Tylosand NRS, Varberg. Smdland:Brinstorp NRS, Eksjo NRS, Hiiradsdter GNM,Hogsby NRS, Horeda NRS, Jonkoping NRS, KalmarNRS, Ljungarum NRS, Ljungby, Misterhult ZMIJ,Myresjo NRS, Nye, Oskarshamn, Pflryd, Ryssby ZMLZMU, Sommen NRS ZML, Traneryd ZML, TranisNRS, Vimmerby, Osterkorsberga NRS ZML ZMU.Gotland; ? loc. JCL, Atlingbo GNM, Boge GNM.Ostergt)tland: Aska NRS, Borensberg ZML ZMIJ,Boxholm, Frostad, Harg NRS, Hiillestad NRS, HogbyNRS, Kimstad NRS, Kisa, Kvarsebo NRS, LinkopingNRS Zli/B ZML, Malmsliitt NRS, MjolbyNRS, Norrkoping NRS, Norsholm, Skonberga NRS,Stjiirnorp ZMB, Sviirtinge NRS, Vadstena NRS,valdemarsvik GNM NRS, Ostra Skrukeby. vcistergritland;Askim, Falkoping NRS, Goteborg NRS,Hjo NRS, Horred NRS, Hiindene NRS, JonseredNRS, Kinnekulle, Liicko NRS, Molndal, Skara GNMNRS, Skinnarhult NRS, Skovde NR.S, TorebodaNRS, vinninga NRS, oiatreo. Bohuskin: BovallstrandNRS, Bro NRS, Dingle NRS, Edshultshall NRS,Fiskebiickskil, Hilta NRS, Munkedal, Naverstad,Rossohamn NRS, Stromstad NRS ZML, Syd-KosterNRS, Tanumshede NRS, Ulebergshamn NRS, ytterbyNRS, Odsm6l. Dalsland: Ed,NRS, Froskog GNM,Gesdter ZMB, Holm, Kopmannebro, Rostock ZMB,R6ggiird ZMB, Sk&llerud NRS, Skipafors NRS, StenebyniisNRS, Tosse, Amfll NRS ZML. Ncirke:Askersund, Stora Mellosa zMB, Orebro. scidermanland:Bjornlunda NRS, Brandalsund NRS, FagersjoNRS, Fittja NRS, Handen NRS, Ludgo NRS, MjolnboNRS, Musko NRS, Nacka, Nykoping NRS ZML,Niishulta NRS, Orno GNM, Sparreholm NRS ZML,Stjtirnhov NRS, Striingniis ZMB, Trosa, TullgarnNRS, Tullinge NRS, Tyreso NRS, Valla NRS, ViksbergNRS, Vreta, Alta NRS, Alberga ZM.B ZML.Uppland; Adelso NRS, Almunge NRS, BjorklingeNRS, Bjorkniis NRS, Gustavsberg NRS, HammarbyNRS, Harpabol, Hibo Tibble NRS, Hiiggeby, Hfltuna,Ingaro, Jumkil NRS, Jiirlisa NRS, LennartsndsNRS, Rimbo NRS, Roslagen ZMIJ, Ro NRS, SkoklosterZMB, Stockholm NRS ZML ZMIJ, TenstaZMB, Tierp NRS, Teljo NRS, Ultuna ZML IVU,Uppsala BML IVU NRS ZML ZMIJ, Vassunda IVUNRS, Viirmdo, Ossebygarn NRS, Osterskdr, OstraRyd NRS. Viistmanland: Arboga Z}dB ZML, FellingsbroZMB, Guldsmedshyttan NRS, KolbiickNRS, Kiirrbo ZMB, Koping ZM3, Lindesberg NRS,Skultuna ZMB, Valsta Z]|dB. Vcirmland; Alster NRSZML, Arvelsiiter NRS, Arvika NRS ZML, DaglosenNRS, Dalby, Forsvik, Frykerud NRS, Gyllebyin Sunne NRS, Hammaro NRS, Lflngban NRS, MolkomNRS, Munkfors NRS, Molnbacka NRS, NiklasdarnmNRS, Odenstad NRS, Ransby, RottnerosNRS, Siiffle NRS, Olme. Dalarne; Blyberg NRS,Brunnsvik NRS, Falun ZMIJ, Finngiirdet, Hedemora,Hiiggesundet NRS, Ludvika NRS, Mora,Smedjebacken, Stora Tuna NRS ZML, Sundborn,Soderbiirke NRS, Torsing NRS, Tallberg Tje . Gristrikland:Bjorke NRS, Gdvle, Hamrflngerfjiirden,Hille-Forsby NRS ZML, Ockelbo NRS, Torsiker,Trodje NRS. Hcilsingland: Delsbo NRS ZML, KilaforsNRS, Soderhamn NRS. Medelpad: Bergeforsen

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Studies on Scandinavian Bumble Beesr45BiologyFlight season. From beginning of May till endof August. Queen : 14 May- ?; worker: 20 May-22 Aug.; male: 22 June-24 Aug.wise colouring as in the female though oftena lighter shade.Variation. Vast minute variations in additionto the general variations included above.ofDistribution (Fig. 79)BOMBUS HUMILIS HAFSAHLIANU,S VOGTQueen, workerQueen measurements. N : 2O; SE Norway;malar space:0.81 mm (*0.04+0.01) range:0.75-0.90 mm ; 'radial length' : 3.57 mm ( +0.08+0.02) range: 3.45-3.75 mm; interalar width:4.63 mm (+0.14+0.03) range: 4.28-4.90 mm.Body of medium to small size.Colour pattern Pile of face with variable mixtureof black and yellow, or entirely black. Pileof vertex yellowish-brown or with slight admixtureof black. Dorsum of thorax, at most dorsalthird of episternum pronounced yellowish-brownto orange-brown. Episternum otherwise blackhairedas also venter, or with variable admixtureof yellowish-white hairs. Fringes of sternitesblack or greyish-white. All legs black, occasionallytrochanters and hind femur with variableadmixture of pale yellow hairs. Corbicularfringes black or dark yellowish-brown, occasionallytipped so. T1 usually with yellowish-brownpile, occasionally with lateral black patches ormore scattered admixture of black hairs. Pileof T2 dark yellowish-brown to brownish-blackexcept for posterior margin being fringed yellowish-brown.Tr-u dull yellow-haired with variableadmixture of black bristles, occasionallyyellow hairs replaced by yellowish-brown ones,particularly on anterior part of each of Tr-n.Te black-haired, occasionally with admixture ofyellowish-brown hairs.MaleColour pattern Pile of face, episternum, venter,trochanters, femora usually predominantly paleyellow. Fringes of sternites whitish. T6 blackhairedwith variable admixture of yellow oryellowish-brown hairs. Pile of T, black. Other-Scandinavia. Confined to the northern borderareas of the distribution of the species in Scandinavia.Smooth clinal variations and rather sparsematerial fromsome of the districts make itdifficult to indicate the southern limit of thedistribution of the subspecies which of coursein any case is artificial.Total distribution of the subspecies, B. humilishafsahlianus Y ogt. Scandinavia.BOMBUS (THORACOBOMB US )MUSCORUM (LINNAEUS)Apis muscorum Linnaeus, 1758, p. 579 No. 32,type area Sweden: Uppland. A female LSL !labelled muscorum disagrees with the usual interpretationof the taxon.(Bombus muscorum (Linnaeus) in sensu Fabriciust77 5, nec Yarrow 1968 ; Dahlbom 1832,p. 46 (partim), 1837 (partim); Wahlberg 1854(partim); Thomson 1870 (partim), 1872 (partim);Siebke 1870, 1880; Roth 1897; Strand 1898a,1898b; Aurivillius 1903 (partim); Wahlgren 1908,l9l5; Ringdahl l92l; Meidell 1934a, 1946;Wexelsen & Skire 1934; Kruseman 1959, 1964;Elfving 1960, 1968 ; Erlandsson 1960 ; Loken1960; Ander 1963, 1965. Bombus cognatus auctt.nec Stephens, 1846 (partim). Bombus smithianusauctt. nec White, 1851; Friese 1902; SparreSchneider 1902; Richards 1935; Yarrow 1959;Elfving 1960, 1968. Bombus alpinus smithianusauctt. nec White; Lie-Pettersen 1901 . Bombuscognatus var. smithianus auctt. nec White; Lie-Pettersen 1905; Sparre Schneider 1906. Bombusmuscorum var. smithianus auctt. nec White; Lie-Pettersen 1907; Sparre Schneider 1909, 1910;Friese & Wagner 1909, l9l2; Hell6n 1933;Meidell 1934a. Bombus muscorum smithianusauctt. nec White; Laken 1960. Agrobombus

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Studies on Scandinavian Bumble Bees 147mediates (Table XVI[) which show clinal variationsin steps.The Swedish population belongs to the nominatesubspecies except the occurrence in Dalarne.Taxonomical remarks on the subspeciesThe dark populations occurring in the BritishIsles, Scandinavia, Finland and European USSRhave been recognized in turn as a distinct species,B. smithianus auctt. nec White (Forsius 1925;Popov 1930 ; Richards I 935; Elfvin g 1960, 1968),as several subspecies of .8. muscorum (Kruseman1964; Yarrow 1967), and as a single subspeciesof .8. muscorum (Reinig 1970, p.71). Popov (1930)claims the specific position of the dark populations,the 'smithianus' group, from the shadeof the colouring of the coat and the shape ofthe male Str. Specimens examined by me agreewith those studied by Richards (1935), i.e. themale St, in the dark populations and that oftypical B. muscorum display so many variationsin shape that this feature has no specific significance.In addition to the phenotypical studyabove, the triangular graph (Fig. 81), based onmeasurements of malar space, 'radial length',and interalar width, illustrates clearly the subspecificposition of the dark Scandinavian populations.B. m. muscorum, B. m. liepetterseni, andthe specimens from Dalarne present a congruentspreading of the markings on the graph, whichemphasizes their conspecific identity.Thus it seems likely that B. muscorum producesdarker forms in northwestern and northernborder areas and in the south, viz. B. m. pereziellusSkorikov in Corsica (Yarrow 1967).Further information is needed to estimate whetherthe subspecies, designated to the 'fringed'populations, qualify for that rank, should besplit, or reduced to infrasubspecific forms. Thesubspecific position of B. m. liepetterseni is wellTable XVIII. Percentage of female B. muscorum (Linnaeus) exhibiting certain characters of colour pattern inpopulations along the coast of Norway. Area : county or district thereof (cf. Fie. ee)Area VAy Ry Hov SFv MRv STv NTy NsyNumber of specimens 7 25 583031t222Pile of faceYellowMainly yellowMainly blackBlack100 92824102l4532473352455050t4862773EpisternumYellowMainly yellowMainly blackBlack100 100312t750310871387100 100 100Hind femurYellowMainly yellowMainly blackBlack100 10033231347t3805s45l78343571882Corbicular fringesYellowMainly yellowMainly blackBlack5743643622l49553790l6 3384 67 1001882

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Studies on Scandinavian Bumble Beesr49Colour pattern Dorsum of thorax orangebrown.Tr-u dull yellow to greenish-yellow, otherwisecf. the subspecies.MaleMorphological characters. Malar space slightlylonger than distal width, about as long as Az+ r.An hardly longer than distal width of the segment.Au-r, moderately swollen beneath (Fig.48A). Posterior fringes of hind tibia longer thangreatest width of the segment. Str and genitaliaas in Figs. 48B-C; distal part of gonocoxiteproduced inwards; gonostylus inwardly producedinto simple jointed, strongly projecting lamella;distal margin of volsella with a broad not muchprojecting process and a broad subapical triangulartooth (Fig. 48D); penis valve hookedat apex, without or with indication of toothbeneath (Fig. 48E). Coat rather even. Body ofmedium size.Colour patterndetails cf. the subspecies.Distribution (Fig. 82)Resembles the female, forNorway. Restricted to coastal areas from thesouthernmost point of the country, viz. Vest-Agder (VAV): Lindesnes, north to nearly 68" N.Single records from southeastern lowlands., viz.Oslo area and Buskerud (Bo): Kongsborg, concernold finds and it is doubtful whether thespecies exists in these areas nowadays.Biotopes: Erica marshes, sandy fields withgrowth of Leguminosae.A total of about 820 specimens was examined.List of localities. Akershus.' Oslo: ? 2 HV 263 (Siebke)ZMO. Buskerud (Bd).' Kongsberg: ? Aug. 1902 (ex.coll. Schneider) TRM. Vest-Agder (VAy); Lindesnes:Lindesnes ; Farsund : Fjellesi, Lista fyr, Lodshavn,Tjorve, Vanse; Flekkefjord: Hidra. Rogaland ( Ry) :Sokndal: Hauge i Dalane, Nesvig, Sogndal, Vatland;Eigersund: Eigeroeidet, Fotlandsvann, Klunglandst., Leidland, Lysevann; H6: Anisdal, Brusand,Fossfjellet, Fuglset, Narland TRM, Ogna, Opstad,Salte bru, Stolen, Varhaug, Vigrestad, Vatnamo;Klepp: ? loc. TRM, Borsheim, Gjeishaug, Horpestad,Klepp, Kisen, Orre, Orrevann, Reve; Time: Time;Sandnes: TRM, Hana; Sola: Solastrand, Tananger,Olbergstranda; Randaberg : Sande; Stavanger: Lindoy;Rennesoy: Dale; Kvitsoy; Karmoy: Ferkingstad,Fotvann, Risdal, Sreveland, Sondenfl, Vigsnes; Utsira:SMS. Ri; Forsand: Meling. Hordaland ( HOy) :Bomlo: Gjertrudsviken, Rolfsnes, Goddo; Fitjar:? loc., Klaksoy, Risoy; Tysnes: Vernoy; Austevoll:Inneroy, Horgo, Karlsoy, Ytteroy; Fana: DolvikVCA, Krokeide VCA, Minde VCA, Skipanes VCA;Sund : Irroy, Steinsland, Telavflg; Rell: Eide, Landro,Movik, Solsvik, Olveset; Laksevflg: Storingavika;Bergen: TRM VCA ZIN,[B_ ZMO; Askoy: Davanger,Hanoytangen, Hedlesund, Herdla VC A ZIN..IB, Horsoya,Lamoy, Odlandsvik VCA; Asane: Salhus VCA;Meland: Bratshaug VCA, fo, Satre VCA Zli.{B; Oygarden:Njupsholmen, Rongoy VCA, Straumoy;Radoy: ? loc. VCA TRM ZM'B, Fedje. Sogn ogFjordane ( Sty ).' Solund : Faroy, Ospa; Hyllestad :Hatleim, Hyllestad, Skivenes; Askvoll: Aralden,Einen, Grytoyri ; Flora: Askrova, Florelandet, Kinn,Nekkoyna, Reksta, Rindemyr; Eid: Haugland ; Vigsoy:Degnepollen, Hagen, Krikenes, Kvalheim, Miloy,Refvik, Roysen; Selje: Dalsbo, Ervik, Lekanger.Mcire og Romsdal (MRy).' Sande: Gurskevig, Kobbevik,Larsnes, Aran; Hareid: HjorungavAg; Ulstein:Flo, Ulsteinvik, Vonheim; Heroy: Djupsvik, Runde;Alesund: VCA; Aukra: Falkhytta, Kolsholmen, Rindaroy,Rod; Frrena: Bud, Gjendem, Gule Bru, Viken;Eide: Vevang; Smola: Andholmen KMT, Edoy KMT.Sc)r-Trt)ndelag ( STy).' Froya: Froan VCA TRM,Halten KMT; Hitra: ? loc. TRM, Hamn; Orland:? loc. KMT, Bakken, Beian, Hovde KMT. STi:Trondheim: KMT TRM VCA. Nord'Trt)ndelag(NTy); Flatanger: Bjoroyvrer KMT, Halmo TRM;Nreroy: Garmannsviken. Nordland (Nsy); Vega:TRM VCA; Heroy: Syd-Heroy TRM; Luroy: ZMO,Lovund TRM; Meloy: Kunna, Storglomvann 500 mZMO; Gildeskfll: Sor-Fugloy; Bodo: Bodo, Karlsoyv&r,Lopsviken, Store Hjartoy.Corrections. Records from Buskerud (Bv): At(Strand 1898a) were revised to B. distinguendusMorawitz and B. pascuorum Scopoli respectively.Those from Finnmark (Fi): Alta: Jotkajavrreand K6fjord (Hell6n 1933) were revised to B.pascuorum smithianus White. Records from Hedmark(HEs): Vang: Hjellum (Wexelsen & Skire1934) were revised to B. humilis Illiger by Meidell(unpublished). Occurrence in Troms (TRy): Hillesoysituated at 69"35' N (Popov 1930, p. 97in referring to Sparre Schneider 1910) is dueto linguistic misunderstanding.As the species has been confused with B.humilis and B. pascuorum, unrevised records arenot included here.Sweden Locally rather freqUent in 6land,Gotland, and in southernmost part of the main-

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Studies on Scandinavian Bumble Bees 151Yd,Lar, Ystad NRS, Ortofta GNM, Ovedkloster ZM.AZML,Ahus NRS ZML. Ble king e : G ammelstorp GNM,Hiillevik, Torhamn. Halland; Falkenberg, HalmstadNRS. Varberg GNM. Smdland: ? loc. GNMNRS TCL. 6land: Bengtstorp, Frirjestaden, HornsjoZlJ[v[B, Hogsby NRS, Koping ZN{B, MockelmoseNRS, Morbylonga NRS ZML, Norra alvaret, ResmoNRS, Stenflsa alvar NRS ZML, Stora alvaret NRS.Gotland.'Austers NRS, Burs, Fflro DCL, Molner, UllaHau DCL, Visby NRS. Osrcrgdtland: Linkoping ? dZMB ZML. Vristergdtland: Falkoping ? NRS. Bohuslcin:Kristineberg, Skafto NRS. Uppland.' ? loc. ? (oldspecimen, no data) NRS, Nassa 70 km E Stockholm ?30 July 1951 (Nordstrom) NRS, Stockholmd (Roheman)NRS, Soderarm 30 km E Norrtiilje2SV 13 Aug.1932 (? leg.). Vristmanland: Malmor E Koping ?28 June 1929 (Ander) ZIN{{B. Dalarne.' Idre 3 ?? 10July1907 ? 14 July 1907 (Bengtsson).Unrevised records are not included here except thefollowing (Ander in litt.): Gotlqnd: Fide, Fideniis,Fleringe, Hamrashage, Holmhallar, Horte, Niirshamn.Bohuslcin: Ockero . Ostergt)tland: Brflviken,Tikern.Corrections. Records from Vdrmland: Dalbyand Ransby in Klariilvdalen (Wahlgren 1908)were revised to B. humilis. As regards recordsfrom Dalarne: Mora and Hiilsingland: ? loc.(Kruseman 1964 the map) the first was revisedto B. humilis while the latter, being questionedon the map (Fig. 82) was not traced.Finland. Observations so far indicate one populationconfined to the southwestern coastal areas,preferably inhabiting the archipelago, and anotherpopulationrestricted to the coast around65' N. The very disperse additional records, evenas far north as Lappland (Le): Kilpisjiirvi (Elfving1968), are noteworthy.Revised and unrevised records (Fig. 82) referto Elfving (1968) except the following: Al: KlfivskdrAKA; N: Lappvik and Lojo AKA; Om:Ruukki ZML; Ok: Yaala ZI|i/.A and Sotkamo(Popov 1930, p. 98); Ob: Aavasaxa NRS.World distribution. Europe (British Isles; Corsica;on the continent from Spain, Mediterraneannorth to Fennoscandia; European USSR, innorth locally penetrating the taiga almost toPolar Circle and in the south avoiding semidesertareas) Turkey Caucasus TranscaucasusKazakhstan Northern Tien ShanSouthern part of Western Siberia Baykalarea Yakutsk Southern PrimorskyNorthern Mongolia Manchuria (Panfilov1957 and in litt.; Kruseman 1964; Yarrow 1967).Z oogeographical remark sThe islet records in Southeastern Norway, inSweden: Dalarne anC in Northern Finland (Fig.82) may refer to relic occurrence indicating awider distribution of ,8. muscorum throughoutFennoscandia in earlier post-glacial periods.According to Popov (1930) the dark populationsfringing the British Isles, Western Scandinavia,and dispersely occurring in Sweden, Finland,Northern Russia, is a European subarctic speciesrelic in the recent fauna. As indicated in thesubspecific discussion above, these populationsrefer to B. muscorum producing darker forms insome range of its extreme border areas.BiologyNest. Pocket-maker. Nest on the surface ofthe ground is recorded in grass fields, in mossand in marshy Erica - Callune heaths. The coloniesare small and often infested by parasites.The total number of imagines and young atthe peak of the season has not exceeded 40 individualsin investigated colonies (Lie-Pettersen 19011905, 19071' Laken l96la).Flight season. From middle of May to lasthalf of September. Queen: 13 May-18 Sept. ;worker: 18 June-l5 Sept. ; male : 28 July-20Sept.BOMBUS MUSCORUM(LINNAEUS)MUSCORUMThe subspecies is distinguished from ,8. m. liepetterseniby the dull rather pale yellow episternumand venter.Queen, workerQueen measurements. N : 20; Norway:Rbgaland; malar space: 0.76 mm (+O.O++0.01)range: 0.70-0.85 mm; 'radial length': 3.95 mm' ( +0.1 I +0.02) range : 3.7 5-4.10 mm; interalarwidth: 5.03 mm ( +0.12 +0.03) range: 4.85-5.23 mm. Body of medium to large size.

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Studies on Scandinavian Bumble Bees 153fringes of propodeum black with slight admixtureof yellowish-brown in the dorsal part. Tt anteriorlyblack-haired, otherwise dull yellow. Tr-udull yellow. Otherwise coat black.Queen, workerQueen measurements. N : 2O; NorwaY:Nordland: Malar space: 0.82 mm ( +O.O: +0.01)range : 0.75-0.85 mm ; 'radial length': 4.09 mm(+O.Oq+0.02) range: 3.80-4.20 mm; interalatwidth: 5.27 mm (+0.14+0.03) range: 4.90-5.53 mm. Body of large size.Colour pattern Cf. the holotYPe.Variation. Pile of face, fringes of propodeumentirely black or with a few yellow hairs admixed.Femur, corbicular fringes with singlyyellow hairs, occasionally the lattet tippedyellowish-brown. Fringes of sternites with slightadmixture of greyish hairs. Pile on T, occasionallyof a somewhat darker shade than thesubsequent tergites. A few individuals with singleblack hairs in the middle of anterior edge of T,were examined. The variations are rather minute.MaleColour pattern Pile of face with variableadmixture of yellow, occasionally predominantlyyellow. Pile of vertex predominantly yellow oryellowish-brown. Dorsum of thorax and at leastdorsal third of episternum yellowish-brown.Extreme anterior part of Tr black as occasionallyalso lateral patches, otherwise hairs onthis tergite yellow. Hairs on Tr-u dull yellow.At least fringes of Str-u predominantly yellow.Otherwise coat black.Variation. Ventral part of episternum rangingfrom entirely black to predominantly pale yellow.Hind femur with slight admixture of yellowhairs. T, occasionally with single pale yellowhairs admixed.Distribution (Fig. 82)Norway. Restricted to the extreme coast from60' N and northwards.The total distribution of the subspecies, B. muscorumpetterseni n. ssp. NorwaY.BOMBUS MUSCORUM? SSP.The subspecific status of 4IQ from an isolatedlocation Sweden: Dalarne: Idre by, altitude450 m, erroneously identified as B. pascuorumsmithianus White ( - B. agrorum arcticus Dahlbom)by Bengtsson (1908, P. 242), and revisedto unnamed subspecies of B. muscorum grexsmithianus by Kruseman (1964), is in my opinionuncertain. The colour pattern of the somewhatfaded specimens is atypical and neither coincideswith that of the nominate form nor with .8. m.liepetterseni. The striking difference is the colouringof the abdominal coat. Pile of Tr-n is yellowish-brownexcept for lateral black patches,and Tb is black-haired. Yet one specimen presentsentirely black-haired Tr while another hasentirely black-haired Tn. Thus, instead of the dullyellow coat on T1-5 being typical of B. muscoruffi,the abdomen has the same yellowish-browncolour as thorax and additionally a pronouncedadmixture of black hairs. The coat is, moreover,longer and much more shaggy than typical forthe species. The appearance is in fact closer toB. pascuorum than to B. muscoruffi, which explainsthe misidentification of Bengtsson. Thespecimens should for the moment be consideredas aberrants.The isolated records in Finnish Lapland (Le):Kilpisjiirvi 2 ?? (Elfving 1968) ate striking. As'islet'for the remaining occurrence of darkspecimens in Finland (Forsius 1925; Elfving1968), it is distinguished from ,8. m. liepetterseniby markedly shorter coat, smaller size and a fewyellow hairs admixed in pile of face and episternum.The scattered records of dark specimensfrom the coast of Uppland in Sweden acrossGulf of Bothnia to Aland Isles and southwesternmainland of Finland, and those from NorthernFinland, are in fact more related to each otherthan to B. m. liepetterseni and the nominatesubspecies. The sparse material is insufficient toindicate their taxonomical status but may favourthe theory of relic post-glacial distribution mentionedon p. 151.

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Studies on Scandinavian Bumble Bees 155Kirby l82l is hereby re-established. The taxonis a subspecies, B. pascuorum smithianus White.First Scandinavian recordsNorway. Nordland (Nno): Ankenes: Bjerkvik(Dahlbom 1832) published as B. arcticus.Sweden Central Sweden and Vdstergcitland,published as B. agrorum and B. curtisellus respectively(Dahlbom 1832).Subspectfic discussionB. poscuorum exhibits a great variability ofcolour pattern within the entire range of itsdistribution, locally and among inhabitants ina colony, yet geographic variations, some ofwhich form clines, are recognized. A vast numberof European forms were designated (Kirby 1802;Vogt 1909, lgll; Kri,iger 1928, l93l; Pittioni1939a; etc.), some of which are segregated into-geographic subspecies while others ate onlyinfrasubspecific forms.Three Norwegian subspecies have been recognizedin the passed, viz. B. pascuorum smithi'Anus White ( - B. agrorum erlandssoni Kruseman),B. p. sparreanus nov. nom. (: B. a. bicolorSparre Schneider) and B. p. barcai Vogt, consideredas arctic, western and southeastern subspeciesrespectively (Loken 1960). They havebeen singly or together previously treated asraces or local varieties (Schoyen 1881 ; SparreSchneider 1895b, 1906, 1909, 1918; Friese 1902;Bengtsson 1904; Friese & Wagner 1909 ; Vogt1909; Kriiger 1928, l93l; Soot-Ryen 1925; Meidell1934a; Laken 1950). The local variation ismoreover reflected by the division in variantso to g of Thomson (1872) followed by Siebke(1880) and Strand (1898a, 1898b).A total of five subspecies have so far beendesignated in Sweden: (1) B. p. smithianus occurringin Lapland (Boheman 1844; Wahlberg 1854;Thomson 1872; Aurivillius 1903; Bengtsson 1904,193 1 ; Ringdahl 1915 ; Lundblad 1924; Barendrechtl94l; Brinck 1951; Kruseman 1958a,1959; Ander 1963). (2) B. p. sparreanus extendingfrom Dalarne to Norrbotten (Brinck 1951; Kruseman1958a, 1959; Ander 1963, 1965). (3) B. p.romani Vogt restricted to Uppland county (Krii-ger 1928, p. 366, 1931, p. 170). @) B. p. barcaiconfined to Southern Sweden with clinal variationsdirected north to Norway: Oslo area(Krtiger 1931, p. 163; Kruseman 1958a) andadditionally to the Swedish counties Dalarneand Hiilsingland (Kruseman 1958a). (5) B. p.gotlandicus by Thomson (1872) mentioned asB. agrorum var. francillonella, restricted to Gotland(Erlandsson 1953).Four Scandinavian subspecies are hereby recognized,viz. B. p. smithianus, B. p. sparreanus,B. p. pallidofacies Vogt nov. status and B. p.gotlandicus, based on the study below.Generally speaking, the Scandinavian populationsdisplay a shift in the colouring of the coatfrom the palest in Southern Sweden to the darkestin Northern Scandinavia. Parallel with the changeof greyish-white to black hairs on episternumand venter, the dorsum becomes a shade ofbrighter yellowish-brown and turns to orangebrown.An increase in hair length on going northhas not been measured, but the difference betweenthe southern- and northernmost populationin length of the coat of thorax dorsallyis estimated at 0.50 mm.Striking features of clinal variability, viz. thecolour in pile of face, episternum and hind femur,have been studied to evaluate the taxonomicposition of the populations. The data, beinggrouped in steps and in progression from thepalest to the darkest, prove that each of thecharacters form clines directed from southernmostSweden to the western coast of Norway(Table XIX) and north to inner part of Gulfof Bothnia as well (Table XX). The result ispresented graphically (Figs. 83-84) and indicatesa correlation of the features. The levellingof the clines at either end may justify subspecificdesignations, yiz. B. pascuorum pallidofaciesVogt, herewith designated to the population inSouthern Sweden cf. below, and B. p. sparreanusextending from western coast of Norway northeastto the Gulf of Bothnia (Fig. 86). The subspecificidentity of the populations occurring inHordaland (Fig. 83) and in Medelpad-Norrbotten(Fig. 84) is confirmed by a study of thesame three features on individuals from countiesranging in between Hordaland and Norrbotten.The result (Table XXI) shows only slight local

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Studies on Scandinavian Bumble Beests7Table XIX. Percentage of female B, pascuorum (Scopoli) exhibiting certain characters of colour pattern inpopulations ranging from Southern Sweden to western coast of Norway. Area : county or district thereof(cf. Fig. 99)Area Sk. Hall. Boh. O AK Bo Bv HOi HOyNumber of specimens901034445231 35 100Pile of faceYellowMainly yellowMainly blackBlack2849l94205030348453t457292135035574310901189 100EpisternumGreyish-whiteMainly greyish-whiteMainly blackBlack78202405010l756243t6304591031461329t4t44361084100595Hind femurGreyish-whitePartly greyish-whiteMainly to all black10010085696425ll522325t45729639l2100 98Table XX. Percentage ofpopulations ranging fromfemale B. pascuorumSouthern to Northern(Scopoli) exhibiting certain characters of colour pattern inSweden. Area : county (cf. Fig. 99)Area sk. Sm. Og. Sdm. Upl. Gstr. Ffls. Med. Aog. Vb. Nb.Number of specimens906543369l11 55 53 34 55 6lPile of faceYellowMainly yellowMainly blackBlack2849t94115531323549t4330501723649l2455522375411853 297 98 100EpisternumGreyish-whiteMainly greyish-whiteMainly blackBlack78202603822358l9t96lt7318453527327264l51678799t 100 100Hind femurGreyish-white 100Partly greyish-whiteMainly to all black9828695831167227I451836l723602494698s 10029811. Bumble Bees.

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Studies on Scandinavian Bumble Bees 159The designations B. pascuorum barcai Vogtand B. p. romani Vogt concern transitional populations,i.e. they are suppressed as subspeciesfor the following reasons:According to Vogt (1909), B. pascuorum var.barcai occurs in 'Mittleres Sweden und Christiania(und weitere Umgebung).' fn addition tothe holotype, labelled Delsbo, the type materialconsists of 7 ?? 5 HV 7 33 from Sweden: Hiilsingland:Delsbo and 5 ?? 11 HH 7 dd from Norway:Akershus: Oslo and Ostfold: Halden; Moss.According to the present study, Ostfold, Akershus,Hiilsinglandare situated in intergradingareas (Tables XIX, XX; Figs. 83, 84, 86), theformer perhaps in the periphery of the distributionof .8. p. pallidofactes, the latter almostreaching the atea of B. p. sparreanus. The typematerial from Htilsingland, including the holotype,is all intermediates with close affinity toB. p. sparreanus. As stated by Ander (1963),the light-coloured individuals in southernmostSweden do not agree at all with the descriptionof ^8. p. barcai. Revisions reveal that the Norwegianform of barcai concerns intermediates betweenB. p.sparreanus and B. p.pallidofacies,while Swedish B. p. barcai concerns either intermediatesor B. p. pallidofacies.Vogt (1911) states that B. p. var. romanioccurs in 'Uppsala. Gouv. Petersburg.' However,the type locality, viz. Uppland: Uppsala, concernsthe intergrading area or the peripheralnorthern occurrence of the subspecies B. p. pallidofacies(Table XX, Fig. 84). The holotype belongsto the darker individuals of the local population.Pile of face black, episternum with a fewgreyish-white hairs admixed with the black ones,hairs of Tr predominantly yellowish-white, thoseof femora partly greyish. Romani is an intermediateform between B. p. sparreenus and B. p.pallidofacies.As demonstrated in Table XXIII,the fourScandinavian subspecies recognized in this studyare fairly colour-stable within their central areaof distribution.Taxonomical remarksIt might be questioned whether the subspecificdesignation B. p . pallidofacies should be suppressedand the population in Southern Sweden treatedeither as a transitional form linking B. p. sparreanuswith the subspecies further south, or perhapsincluded in this subspecies (Reinig 1939,p. 194 the map). In my opinion the subspecificstatus of the population in Southern SwedenSweden should at least be kept until the subspecificstatus of populations in Germany andadjacent areas has been studied further.Queen, workerMorphological characters. Malar space nearlyl1/z times the distal width, Ionger than Ar* nbut shorter than Ar* r* n. Clypeus markedly longerthan distal width (except in some workers);disc of clypeus moderately punctured except foranterior impunctate area in between lateral,coarsly punctured impressions. Labral furrowwidened towards the front (Fig. 2l); labralIamella shining, with knife-sharp margin. Stuwith distinct distal keel. Coat shaggy. Queenmeasurements, cf. the subspecies.Colour pattern The species is in generaldistinguished by the following characters: Dorsalof thorax with yellowish-brown hairs. Pile onT1 ranging from entirely black to entirely yellowish-white,yellow, or yellowish-brown; oneor several of midmost gastral tergites mainlyblack-haired:' at least Tu-u with yellowish-brownhairs. Corbicular fringes black. The coloursfrequently encroach upon another and exhibita gteat number of variations. Furtherdetails are given in the description of the subspecies.MaleMorphological characters. Malar space markedlylonger than distal width, hardly longer thanAr+ *. As hardly l% times the distal width,,{4 usually as long as distal width, Au-r, (orAn-rr) strongly swollen beneath (Fig. 47A).Posterior fringe of hind tibia markedly longerthan the greatest width of the segment. St, andgenitalia (Figs. a7B-C); gonostylus inwardlyproduced into a long narrow tooth, distal

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Studies on Scandinavian Bumble Beesr61Asker, Bronnoy, Rustad; Brerum: ? loc. TRM, B16-tan, Fornebo ZMO, Hovik ZMO, Lysaker ZI|I4BZMO, Oslo: TRM ZN{4B ZMO; Rrelingen: Fjerdingsby;Aurskog-Holand : Bjorklangen, Gangsnes, Loken,Skogen, Ostegird; Fet: Bjorkflflten; Hurdal: Tomte;Feiring: Feirin g. Hedmark ( HEs) ; Eidskog: Bolfoss,Gjelarisen, Magnor SMS; Kongsvinger: Eidsberg;Sor-Odal: Mflrud ZMO; Hamar: ZMB ZMO; Ringsaker:Gaupen, Vea. HEn: Trysil: Enga, Egernesset,Vestby; Amot: Glesubekken; Rendal: Solbakken SEAsheim ZMO; Alvdal: Atvdal; Engerdal: Risbakken750 m. Oppland (Os).' Jevnaker: Randsfjord USUZM.F ; Gran: Granvollen; Kolbu: Dal; Ostre Toten:Hveem, Skreia, Stubdalen; Sondre Land: Fluberg;Nordre Land: Dokka USU, Torpa; Lillehammer:Jorstadmoen, Lillehammer, Stor-Hflve; Etnedal : Bruflat1'Sor-Aurdal: Begndal, Ellingseter 820 m, Skard,Strommen ; Nord-Aurdal : Knutshaugen, SabuoygardN Fagernes; Gausdal: Follebu, Gisoya; Oyer: Aksjosetra1000 m, Skfli, Oyer Zl/rB ZMO, Astdalen 900 m;Ringebu: Ringebu ZMB ZMO, Venaby gd. On: VestreSlidre: Kinnholt 1000 m, Uppslidre; Oystre Slidre:Kvithovd 800-1000 m ZMO; Vang: ?loc. ZMO,Oy. USU ZMB; Sel: Heidal TRM ZMB, LaurgflrdZMO, Mysuseter 900 m, Otta ZMO; YAgtL: Gjendesheim1000 m, Hindseter 900 m, Klones, Leirungen1050 m, Randsverk TRM ZN4{4-B., Vigimo ZlldB ZMO,Ovre Sjodalsvann; Lom: Galdesand, Lom ZMO:Dovre: Dovrefjell 1000 m ZMO, Fokstua 930 m,Hjerkinn 950 m, Toftemo ZMO, Vilisjo 950 m ZMO.Buskerud (Bd).' Hurum: Filtvedt TRM, Holmsbu,Hurumo Pinadalen, Sagene, Storsand, Vannsbrekka;Royken: Aros; Drammen : ZMB ZMO; Lier: Lier,Lokke gird, Spikkestad, Sylling, Toverud; Ringerike:Smedrud; Krodsherad: Britfln, Fagerheim, Glesne;Modum: ?loc. ZMO, Snarum; Flesberg: Hvila,Lampeland, Oydegarden; Kongsberg: Komnes, Lintvedt,Skollenborg. Bv; Sigdal: F16gan, Hagle, Haglebu,Kopseng, Nedre Eggedal, Nordby gda, Sigdal;Nore og Uvdal: Norstebo, Rodberg, Solheimstul1000 m, Tunnhovd, Uvdal; Fli: Grosland; Nes: Eidal,Langeslet; Gol: Gol; Hemsedal: Lykkja; Al: ? [oc.ZMO, Levell 700 m; Hol: Geilo VCA ZMB,Halfardokkistolen860 m, Hovet, Krisle, Nyestolen 1000 m.Vestfold.' Sande: Sande kirke; Holmestrand: Angerskleiv,Bogen, Holmestrand ZMO; Borre: Adal, Nykirke;Vflle: Ryk; Ramnes: Kjrer, Lunde, Orrev6l;Lardal: Styrvoll; Andebu: Andebu kirke, Kodal;Stokke: Lango, Sand, Stokke, Veierland; Tonsberg:Prestrodkilen; Notteroy: Teie; Tjome : Kjrere, Solvang;Sandefjord: Austeroy; Hedrum: Kvelde, Ringdal;Tjolling: Viksfjord; Brunlanes : Berg, Dolven,Helgeroa, Tronsrod; Larvik. Telemark (TEy); Skien:Fjelldalen, Klkut; Porsgrunn: Langangen, Mule, Nystrand;Bamble: Trosby, AUV; Nome: Damtjern,Tyri, Vommstol; Drangedal: Drangedal, Holmen,Naksjo, Nordbo, Nos, Omnes, To-myra, Akredalen;Kragero: Kragero, Levang . TEi: Notodden: Gransherad,Tindegrend, Tinnoset; Sauherad : Liagrend ;Bo: Lifjell, Sande, Vatnar; Seljord: Seljord, Svartdal;Kviteseid: Brunkenbergo Eidstad, Kviteseid, Morgedal,Vridal; Nissedal: Kyrkjebygda, Lauvviki, Lia,Tjonnefoss, Treungen ZMB ZMO; Fyresdal: Hegglandsgrend,Moland, Veum; Tokke: Bandaksli, Grimdalen,Vriliosen, Amdals verk; Vinje: Bosboen 970 m,Edland, Haukeligrend, Krossen, Rauland kirke, Serengrend,Va, Vigsli, Amot; Tinn: Austbygda, Borsjoen1100 m, Mosdammen, Tinn . Aust-Agder ( AAy) :Gjerstad: Fiane; Risor: Brflten; Vegirdshei: Ljostad;Tvedestrand: Askeroy, Boroy, Dypv6g, Eidbu, Fiane,Hanto, Laget, Lyngor ZMO, Nes verk TRM, StensoZMO, Strengereid, Tvedestrand, Oster6; Arendal:Arendal ZMB ZMA, Salterod; Moland: Dal, Hannholmen,Holmsund, Kilsund, Staubo, Tverrdalsoy,Voje; Tromoy: ZMO; Hisoy: Flodevigen, Gjervoldsoy,Hisoy ZM.B ZMO; Fjrere: Fjere kirke ZMO;Grimstad; Oyestad: Bjorbekk, Helle, Rykene; Froland:Hynnekleiv, Svenes; Landvik: Havnevig, Horte,Molland, Tonnesol; Lillesand: Blikksund, Hrestad,Hovig, Kjostvedt, Kvise, Langholrnsund, Natvik,Troe, Ulvoy, Amh; Birkenes : Sagen, Svaland; Iveland:Frikstad, Ivedal. AAi; Evje og Hornes: Hornes,Lauvland, Syrtveit; Gjovdal: Gangsei, Gjovdal,Smedland, Amh, Arh; Bygland: Aust ad ZMO, Londal,Ose, Setesdal TRM; Valle: Hylestad, Kvestad,Rygnestad, Valle; Bykle: Breive 700 m, Breivann900 m, Bykle. Vest-Agder ( VAy); Kristiansand:Buane, Erkleiv, Holskogen, Kristiansand, Movik,Stromme; Marnadal: Ask, Finsland, Gretteland,Royris, Royseland, Solis; Sogne: I-angeneset, Sogne,Aros; Mandal: Kvisla TRM, Mandal TRM ZMB,Ramso TRM, Sinum, Tregde; Lindesnes: Ramsland,Reine; Lyngdal: Rom; Kvinesdal: Feda, Gjemlestad,Kvinesdal, Solfls, Oye; Farsund: j-ellesfl, Hanangervann,Lodshavn, Ore, Vanse; Flekkefjord: Dragoy,Ersdal, Fedflk, Langevann, Loga, Nordheim, Nuland,Sandvang, Solvang. VAi: Hegebostad: Skeie; Aseral:Espelid, Rosseland, Aseral; Sirdal: Rekevik, Sinnes,Adneram 800 m. Rogaland (Ry).' Lund: Moi; Sokndal:Bu, Hauge i Dalane, Nesvig, Rekefjord, Sogndal,Vatland, Amot; Eigersund: Egersund, Eigeroyeidet,Fotlandsvann, Sjines, Skjerpe, Tengs; Bjerkreim:Asseimvann, Hovland, Ivesdal, Malmeim; H6:Anisdal, Brusand, Ogna, Salte bro, Sirevflg; Klepp:Borsheim, Klepp, Orre, Orrevann, Orstad, Revtangen,Vik, Oksnevad; Time: Mossige;Gjesdal: Madland SMSNese; Sandnes: Austrhtt, Bergsagel, Dale, N Ganddal;Sola: Gimre, Reke, Slettehei, Tananger; Randaberg:Kverneviken; Stavanger: TRM SMS ZMB,Lindoy; Strand: Vatne; Finnoy: Reilstad, Sjernaroy;Rennesoy: Dale; Karmoy: Eide, Indre Eide, Risdal,Sandve, Seveland, Vikre; Vindafjord: Imsland; Tysv&r: Nedstrand. ( Ri ).' Forsand : Forsand, Fylgjesdalen,Lysebotn, Meling, Songedalen; Hjelrneland:Fister, Igland, Kvamme, Ardal; Suldal : Berge, Hivarstol,Jelsa, Jonstol, Krokvassheia, Kvanndal, Mostol,Nesflaten, Roaldkvam, Svortebekk, Ullsneskvelvene,Aimjodlonuten; Sauda: Hellandsbygda, Saudasjoen,Slettedalen. Hordaland (HOy).' Bcmlo: Espever,Goddo, Langev&gen, Rubbestadnesset, Royksund;

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stad; Meland: Brakstad, Lansvik, Meland; Oygarden:Straumoy; Radoy: Kvalheim; Lindfls: Alversund bro,Herland, Lauvis, Romarheim, Srevrisvflg, Vollom;Fedje. HOi: Etne : Fjrera, Oslandsvig, Skflnevik;Kvinnherad : Fossberg, Gamlestolen, Gjermundshamn,Gronningseter, Guddalsdalen, Ljosmyr, Prestvann,Rosendal, Seimsfoss, Skeie, Varaldsoy; Odda: Dalengird, Hordasvingen 1000 m, Roldal, Seljestad; Ullensvang:Aga, Borve, Djonno, Espe, Fossli, Fresvik,Grythorgen 900 m, Hjolmo, Kinsarvik, Lofthus, Mabodalen,Nesheim, Sekse, Storliseter 900 m, Tveito,Utne VCA ZMB; Kvam: Djonne, Kvamskogen,Mundheim, Norheimsund, Omastrand, Vangdal, Oystese,Atvit USU Z}dB, Asheim; Voss: Armot, Bulken,Bomoen, Hangurfjellet 800 m, Kinne, Liland,Mjolfjell, Vossevangen; Granvin: Granvin, Skjervet,Vindal; Ulvik: Ulvik. Sogn og Fjordane (SFy).' Gulen:Austgulen, Takle; Hyllestad: Dale, Hatleim,Hyllestad, Risnes, Skivenes, Skor, W Alefjell; Hoyanger:Gare bro, Lavik ZM3 ZN{O, Stoldalen; Gaular:Bygstad, Slotten; Fjaler: Dale, Hellestrand:Strandenes; Askvoll: Aralden, Askvoll, Einen; Flora:Florelandet, Nekkoyna, Skorpeidet, Verpevik ; Naustdal:Frammarsvik, Furehaugstolen, Heia, Horstad,Trollstua, Ovre Hella ; Forde : Fliten, Hallbreim;Jolster: Hamar, Ahus; Gloppen: Hjortset, Hope,Hyen, Lote, Lotsbergfjell 700 m, Sandane; Eid: Haugland,Naustdal, Nordfjordeid, Stflrheim; Davik: Kjolstad,Kjolsdalen. SFi: Vik: Seljedal, Vik; Aurland:Flim, Gudvangotr, Vassbygda; Lnrdal: Grote, LrrdalZMO, Ystadbo; Sogndal: Slinde; Leikanger:Hermansverk, Leikanger, Suphamar; Balestrand: BalestrandNMW, Fjrerland, Flesje, Horpedalen, Mel,Suphellen; Luster: Dalsdalen, Fortun TRM, Skjolden;Jostedal: Fflberg, Fibergstolene 600 m, Hafslo,Okkleviki; Stryn: Flo, Gillesdal, Heggjabygda, Hornindal,Hornindalsetra, Olden, Stryn, Stoverstein,Vedvik, Videdalen, Ovre Oldenvann. Miire og Romsdal( MRy ).' Sande : Gurskevig, Aram; Vanylven :Thue; Volda: Bjorkedalsvann, Folkestad, Straumshamn;Hareid : Mork; Heroy : Djupsvik, Runde;Alesund:-BMl VCA, Ase; Orskog: Skodje ZldbO;Vestnes: Gjermundnes, Vikebukt; Molde: TRMZMB ZMO; Aukra: Julnesset, Nord-Aukra; Frana:Gjendem, Hollinholm; Averoy: Vevang. MRi; Orsta:Lyngstolvann, {.Irke, Orstavik, Oy"; Stranda: Fivelstad,Fredheim, Geiranger NMW ZN{B, Norangdalen,Sunnylven ZMO, Vollset ; Norddal : Indre-eidet, Valldal,Ytterdal;Rauma: Andalsnes NMW ZMB; Nesset:Torhus; Sunndal: Brandstad, Jordalsgrend, SunndalBML; Surnadal: Kvanne ZiN{dO, Melhus, Stangvik,Ovre Surnadal; Rindal: Bolme. Sor-Trdndelag ( STy) :Hitra: Hamn, Lya KMT; Agdenes: W Hambora;trtissa: Gafsetisen, Sotvik; Orland: Beian; Bjugn:Kotengsvann, Ryvann, Sreter KMT; Afjord: Molsletten,Morreaunet. STi: Oppdal: Dalsbekk, Driva st.,Givili 975 m, Hogsnydda 1150 m, Knumho 1200 mBML, Kongsvoll 900 m BML TRM ZMBZMLZMO,Oppdal st., Skansen; Midrre Gauldal: Aune; Alen:Reitan; Roros: Adamsvoll 820 m, Myrmoen 700 m,Studies on Scandinavian Bumble Bees 163Naustervoll 700 m, Storelvvoll 800 m; Tydal: Mobro,Sakrismoen ; Selbu: Medbus; Trondheim: KMTTRM VCA ZN.4B ZML ZMO; Skaun: Hestjo, SornypanKMT. Nord-Trtindelag ( Nfy).' Namsos: TRM,Strandmoen; Nreroy: Breiviken, Dalene, Garmannsviken,Kvalviken, Saltbotn; Vikna: Rorvik. NTi:Meriker : Meriker, Tovmodal; Stjordal : Draveng,Hegra st., Lflnke, Tilleraunet, Vikan KMT; Frosta:Holmberget; Leksvik: Storvatn; Levanger: Avdal;Verdal: Billingflaten KMT ZMB, GodbekktjernKMT, Sandvika KMT ZIN{IB, Stiklestad, Sulstua,Vuku; Sandvollan: Inderoy ZMO, Kirkenesvig,Skjelvigen; Verran: Sela; Steinkjer: ZMA; Kvam:Sem, Snisavann ZMA; Namdalseid: Hovlia KMT;Overhalla: Sellotoren; Grong: Ekkerseter, Fjerdingetr,Grong SMS ZMB; Snflsa: ?loc. TRM, Bronstad,Hegge, Snisa, Strindmoen, Telnes; Lierne: Kvelia; Hoylandet : Hoylandet, Skilleberget ; Namskogan :Brekkvasselv KMT ZMA, Finnvollan, NamskoganZII'IAZNjN{{B. Nordland (Nsy).'Bronnoy: Flatmo TRM,Nevernes ZMA, Sandvflg TRM, Somna TRM; Vega:TRM ZN{..{B; Vevelstad: TRM; Alstahaug: TRM;Heroy: Syd-Heroy TRM VCA; Nesna: Hamaroy,Oldersletten; Luroy : ZMO; Meloy: Dalen, Kunna,Spilderen, Storglomvann ZMO; Gildeskfll: Finnes,Gildeskil, Gilset, Inndyr, Jelstad, Skauvoll, Storvika,Sor-Fugloy; Bodo : Bodo BML, Falkflaugdalen,Frostmo, Lopsviken, Planteskolen, Skau, Urskarsdalen.l/si; Grane: Grane, Majavatn, Rotstokkmobekken,Skomstad; Vefsn: Fustvann, Luktvann, Ravassbakken,Ravni ZMO, Store Bjornivann; Hemnes:Finneidfjord, Korgen ZMA, Krokselvmoen,Strombotn; Rana: Dunderland st., Mo ZML. Rosvoll,Rovassdal TRM, Stokkvik TRM, Storforshei,Umskarstjern, Aenget; Beiarn: Grfltidalen; Saltdal:?loc. TRM ZMO, Drageid, Lonsdal, Solvigtind800 m, Storjordet TRM ZJN{B., Vensmoen; Fauske:Fauske, Kjeldvatn BML, Sulitjelma 600 m BML,Vargflsen. Nni): Sorfold: Bonnflsjoen ZIN4A ZMB,Rosvik TRM, Tennvann; Hamaroy: Brennvik, Hamaroy,Flamnes, Hansbakk, Innhavet, Kildetoppen850 m BML, Krikmo SMS ZMB, Skutevik, Tranoy;Tysfjord: ? loc. ZMO; Ballangen: Ballangen kirke,Dyrhaug, Heggemoen, Myrbakk; Ankenes: BjerkvikZML, Seterfjell; Narvik: ZMA ZMB; Evenes: Jansbakk.Nnv: Vestvigoy: Knutstad, Skulbru, Stamsund,Valberg; Vflgan: Store Molla, Svolvrer BML ZINI.AZMO, Orsvig; Sortland: ?/oc.; Andoy: Dverberg,Risoyhamn SMS ZM'B. Troms (Try); Kvrefjord:Borkenes, Vik; Bjarkoy: TRM; Lenvik: Finnsnes;Trornso: Breivik SMS, Tromsdal NRS TRM, TromsoKMT NRS PCL TRM VCA ZN{B; Karlsoy: Karlsoy,Vanno TRM; Skjervoy: Ravelseidet-Langslet.Tri: Skflnland: Lavangseid; Gratangen: GratangenZMA; Bardu: Altevann, Bardu, Dypdal, Seternes;Milselv: ? loc. TRM, Aiapmo, Andselv, BjerkengTRM, Kirkesdal, Kjellmo, Moen-Olsborg DCLZMB, R.undhaug TRM ZNd4B ZM.O; Balsfjord: ? loc.NRS TRM, Laksvatn SMS, Sjivikor TRM, TakvatnTRM, Tamok TRM; Nordreisa: Bakkeby, Sappen;

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Studies on Scandinavian Bumble Bees 165ZMIJ, Viirmdo NRS ZML, Yxlan ZMB, Oregrund Overkalix NRS, Overtornefl NRS ZML, Ovre SvartlflZMB, Osseby-garn NRS, Osterskdr, Ostewiia ZMB, NRS. Lapland (At. Lpm.); Bflngnds, Dorotea NRS,Osterflker NRS ZMIJ, Overgran NRS . Vcistmanland: Frederika NRS, Hiillefors in Bflngnds, Kittelfjiill,Arboga ZIV4B ZML, Dingtuna NRS, Fanthyttan Malgomaj NR.S, Njakafjiill, Ormsjo NRS, RisbiickNRS, Fellingsbr o ZMB, Guldsmedshyttan NRS ZML, NRS, Saxnds, Vilhelmina NRS ZML, Asele NRS.Hjulsjo NRS, Hlllefors NRS, Kolbiick NRS, Kolsva Ly. Lpm.; Bjorkfors VCA ZML, Brattikkfjall, LyckseleNRS ZML, Myrliden NRS, Rusksele ZMU,Z}Jli4B, Ktirrbo ZM.B, Koping ZMB ZML, LindesbergNRS, Malmon ZMB, Nora stad, Nordbergsklacken Sickelsberg NRS, Sorsele NRS ZML, Stensele, TjultriiskBML, Tdrna, Umfors, Vdnnds GNM, ViistansjoNRS, Runskiir ZMB, Sala NRS, Saxhyttan NRS,Skultuna ZN{B. Vcirmland: Arvika, Dalby, Ed, FilipstadNRS, Forsvik, Frykerud NRS, Grums NRS, Ealkovare NRS, Brattfors, Skattrdsk. Lu. Lpm.: Jokk-NRS. P. Lpm.; Abborrtrdsk, Arvidsjaur IVU ZML,Gustavsfors NRS, Gylleby NRS, Gflsborn NRS, mokk, Kutjaure NRS, Kvikkjokk, Malmberget, MuddusNRS, Virihaure area (Killanjaure - StaloluoktaKarlstad NRS, KiI ZMU, Lesjofors, L6ngban NRS,Molkom NRS, Nilsby, Odenstad, Rdmmen, Torsby ZML ZMU). T. Lpm.; Abisko NRS Zll4.A ZML,NRS. Dalarne.' Avesta ZMB, Bingsjo NRS, Blyberg Bjorkliden NRS ZM.A ZML, Jebrenjokk NRS, KirunaNRS, Masungbyn VCA, Mell. Merasjtirvi, Njuon-NRS, Brunnsvik NRS, Elvdalen, Falun NRS, FlodaNRS, Fulufjtill NRS, Hjortniis Tje, Hiillsjon NRS, jevare NRS, Ortojokka NRS, Vassijaure NRS, VietovareNRS, Vittangi ZML, Ovre Sopporo NRS.Idre, Leksand NRS Tje, Lima NRS, Ludvika NRSzML, Mora NRS, Nittsjo NRS, Rtittvik NRS Tje,Sandsjo, Siljansborg NRS, Sjurberg NRS Tje, Smedjebacken,Sollero Tje, Stjtirnsund, Stollen NRS, StoraCorrection. Records from Dalarne: Idre(Bengtsson 1908, p. 2aD treated as .8. p. arcticusTandfln NRS, Stora Tuna, Storsiiter NRS, Stiidjan,Sundborn NRS ZML, Sviirdsjo NRS, Stifsen NRS, were revised to B. muscorum.Siilen ZMA, Sdrna NRS ZI|y4A ZML, Siiter NRS, Finland. Commonly occurring throughout theSoderbiirke NRS, Sflngfln Tje, Transtrand NRS, TallbergTje, Vikarbyn NRS . Gcistriklarcd: Bjorke, Hillecountry (Elfving 1968).World distribution. Europe (British Isles; Sicilia;Corsica; entire continent except in EuropeanZ}/I., Hille-Forsby NRS, G6v1e, Hamringerfjdrden,Hedesunda NR.S, Ockelbo ZNAB, Storvik NRS, TrodjeNRS. Hcilsingland: Bergvik, Bollnds NRS ZML, USSR not occurring south of districts of Moldavia,Kharkov, Stalingrad, Tsjkalov) - Turkey -Delsbo VCA ZML, Fredriksfors NRS, Jiirvso NRS,25 km N Hudiksvall ZMB, Kilafors, Ljusdal NRS,Iran - Northern Kazakhstan - Siberia (SouthernOrbaden NRS, Soderhamn NRS ZM.B. Medelpad:part of Western Siberia, Southern Yakutsk)Bergeforsen NRS, Havero NRS, Kolsillre NRS, LeringeNRS, Liden NRS ZMB, Sundsvall NRS ZMB, Baykal China (Kansu) (Reinig 1939, p. 194;Tuna NRS, Viforsen NRS, Overturinge NRS, Ange. Panfilov 1957; Yarrow in litt.).Hrirjedalen: Anefjail NRS, Funiisdalen ZMB, Ljungdalen,StorvarCen, Sveg NRS ZML, Tiinndalen NRSZML. Jcimtland: Bispfors ZMB, Bispgirden NRS, BiologyBrdcke, Doda Fallet NRS, Hallen, Hammerdal,Handsjo NRS, Jormlien, Jdrpen, Leipikvatten NRS, It{est. Pocket-maker. Nests on the surface ofMattmar, Ottsjon NRS, Oviken, Ragunda, SnasahogarneNRS, Stadsforsen NRS, Storlien, Stromsund moss, accidently in decayed stumps of trees, etc.the ground, recorded in grass of hayfields, inNRS, Undersiker NRS ZML, Ostersund ZMA ZML,Produce colonies of various size, the number ofAre NRS ZMA ZML, Areskutan ZML. Angermenland:Aspele NRS, Aspefl NRS, Bindsjo, Borgsjo, workers may surpass 100 (Dahlbom 1837; Lie-Graninge NRS, Gdrsta, Hoting NRS, Hiirnosand Pettersen 1907; Meidell 1934b; Hasselrot 1960,NRS ZML, Hornefors NRS, Hiknds NRS, Lugnvik re62).NRS, Niiske NRS, Orsjon, Ramsele GNM, SoliefteflFli7ht season. From the middle of April toNRS ZML, Stennis, Siibrfl, Ull6nger NRS, OrnskoldsvikNRS, Overldnnds NRS. Vcisterbotten: BodarnaNRS, Botsmark NRS, Degerfors NRS, Hiillntis worker : 2 llr4ay-l I Oct. ; male: 18 June-l 1 Oct.first half of October. Queen : 16 April-l4 Oct.;NRS, Jorn, Kroksjo NRS, Lillekigetrdsk NRS, PiparboleNRS, Stromsor NRS, TtifteA ZMU, Umefl,Vlisterhiske NRS, Abyn Z}/'F.. Norrbotten: Anttis,Boden NRS, Edefors NRS, Gammelstadviken, Giiddvik,Haparanda, Kalix NRS ZML, Karungi NRSBOMBUS PASC UORT.IM SMITHIAI'IU,SZML, Kukkola NRS, Lapptrask, Lillpite, Lulea NRSWHITEZML, Nybygget NRS ,Pajala NRS VCA ZML ZMU, ( - Bornbus agrorum erlandssoni Kruseman)Pello NRS, Periijiivaara, Pitefl NRS ZML, StorsundNRS ZMI-, Sodra Sunderbyn, Tore ZMB, Vita NRS, The subspecies is distinguished from B. p. sparre-

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Studies on Scandtnavian Bumble Beesr67on the disc. Pile on TB yellowish-brown withdistinct lateral black patches. Pile on To-u yellowish-brown,gastral sternites with mainly greyishfringes. Otherwise coat black.Queen, workerQueen measurements: N - 15; Norway:Hordaland (HOV); malar space: 0.89 mm (+O.Of+0.01) range: 0.85-0.95 mm ; 'radial length' :4.07 mm (+0.08+0.02) range: 3.95-4.20 mm;interalar width : 4.84 mm ( +0.15 +0.04) range:4.65-5.18 mm. Body of moderate to large size.Colour pattern Cf. description of holotype.Variation. Pile of black. Pile of vertex withvariable mixture of yellowish-brown and black.Occasionally dorsum of thorax with single blackhairs on the disc. Hairs on T, ranging from entirelyblack to predominantly greyish-white oryellowish-white. Anterior lunate part of T, varyingin size, varying in colour from yellow toyellowish-brown, occasionally T2 entirely blackor entirely yellowish-brown. Hairs on T r-nforming a black band of various width, otk(rwiseyellowish-brown; occasionally black hairsreduced to small lateral patches or expanded toentire tergites. Hind femur with a slight admixtureof greyish hairs, and the fringes of sternitesvarying from black to greyish-white. The shadeof yellowish-brown varying but usually darker,brighter than B. p. pallidofacies. Three HY (Rogaland(RV): Klepp: Orre 3-4 Aug. 1963 (Ls)) arecompletely brownish-black with exception of T'and anterior lunate part of T, having pale yellowhairs.MaleBody usually of medium size.Colour pattern T? with yellowish-brown hairs,fringes of sternites predominantly greyish or paleyellowish. Colouring otherwise as in the female.Variation. Short plumose hairs in pile of facewith variable adrnixture of pale yellow hairs.Pile of vertex black or almost so. Episternum,venter, proximal segment of legs more or lesspronounced greyish-haired. Hairs of T' entirely(greyish or yellowish-white. Rarely Tr- , all blackhaired.Otherwise variations as in the female.Meidell (1934a) describes a melanic male(Rogaland (RV): Rennesciy 24 Aug. 1931) beingalmost black except for reddish-yellow hairs onT*-u. He indicates a local occurrence of B.pascuorum var. mniorum (Fabricius), a commonform in Denmark. Together with the darkworkers mentioned above, the record indicates,however, only a sporadic melanism at the extremesouth-western coast.Distribution (FiS. 86)Scandinavia. From the west coast of Norwayacross the mountains eastwards throughout theconifer forest to Gulf of Bothnia.Finland. According to the description of thecolouring of the coat (Elfving 1960), and thematerial examined by me, this subspecies prevailsthroughout the country except Lapland andapparently the extreme southern coast.Total distribution of the subspecies, B. p. sparreanusnov. nom. Fennoscandia. ? Further eastinto the taiga in USSR.,BOMBUS PASCUORUM PALLIDOFACIESVOGT NOV. STAT.( - Bombus agrorum barcai Yogt (partim))The pallidofacies form of Vogt (1911), consideredby Krtiger (1928, p. 366) and Reinig (1939, p. 194the map) as a transitional form between B. p.romani Vogt and the subspecies on the continentB. p. ? ssp - B. agrorum typicus Vogt nec Fabricius,is herewith given subspecific rank.The subspecies is distinguished from B. p.sparreanus by predominantly light yellow pileof face, entirely or predominantly whitish-hairedepisternum, venter, hind femora, and a shortercoat.Type materialHolotype: typeof .8. p.f. pallidofaciesVogt -? Uppland 17 July 1909 (Roman) VCA!. Paratypesherewith selected: Sweden: Skf,ne: 3 aa

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Studies on Scandinavian Bumble Bees 169BOMBUS PASCUORUMERLANDSSONGOTLANDICUSBOMBUS (THORACOBOMBUS)RUDERARIUS (MULLER)Distinguished from B. p. sparreanus and B. p.pallidofacies by pile on Tr-u being continuousorange-brown except lateral patches of Tr-r.Separated from B. p. smithianus by venter beingpredominantly greyish-haired. Shorter, less shaggycoat than the remaining Scandinavian subspecies.Queen, workerQueen measurements : N :- 15 ; Gotland;malar space: 0.83 mm ( +O.O: +0.01) range:0.80-0.90 mm; 'radial length': 3.82 mm (+0.00+0.02) range: 3.70-3.95 mm; interalar width:4.41 mm (+0.10+0.02) range: 4.25-4.60 mm.Body of small to medium size.Colour pattern Pile of face yellow, occasionallywith slight admixture of black hairs. Pileof vertex orange-brown. Dorsum of thorax, atmost dorsal third of episternuffi, Tr-u withorange-brown hairs except for pale yellow lateralpatches of T1 and black lateral patches of Tr.Ventral part of episternum greyish-white-haired,usually with slight admixture of black hairs.Venter, trochanters, hind femur with greyishwhitehairs. Pile on fore- and mid-femora entirelyblack or the proximal parts greyish-white. Otherwisecoat black.Variation. In addition to the small variationsabove, pile of vertex may anteriorly and laterallybe encroached on by black hairs. All femoraoccasionally greyish-white; corbicular fringestipped orange-brown or with admixture oforange-brown hairs. T1 rarely with admixtureof black hairs. Otherwise colour-stable.MaleBody of small size.Colour pattern Pile of face with great admixtureof black, otherwise colouring as in thefemale.Variation. As in the female.Distribution (Fig. 86)Sweden Gotland.Apts ruderaria Mtiller, 1776, p. 165 No. 1922,no type exists, type atea not mentioned. Denmark:Sjelland hereby designated.( Bombus ruderarius (Mtiller); Sparre Schneider1918 ; Gaunitz l93l ; Wexelsen & Skire 1934;Tjeder 1954; Laken 1958b; Kruseman 1959;Elfving 1960, 1968; Erlandsson 1960; Frid6n, Eskilsson& Bingefors 1962; Hasselrot 1962; Ander1963, 1965; Frid6n 1967. Apis derhamella Kirby,1802, p. 363 No. 105, holotype d KCL! labelledby Yarrow (1968), type loc. England: E. Suffolk:Barham. Bombus derhamellus (Kirby);Dahlbom 1832, p. 44 (partim); Boheman 1844;Aurivillius 1903 ; Wahlgren 191 5 ; Meidell 1 934a;Laken 1949. Apis raiella Kirby, 1802, p. 367No. 107 , holotype Q KCL ! labelled by Yarrow(1968), type loc. England: E. Suffolk: Barham.Bombus rajellus (Kirby); Dahlbom 1832, p. 33;Siebke 1863, 1880; Thomson 1870, 1872; Strand1898b; Lie-Pettersen 1901 , 1907.)First Scandinavian recordsNorway. Sor-Trcindelag (STi): Oppdal: Drivdalen(Siebke 1863). The record is doubted(Sparre Schneider 1918) and no voucher specimenis traced. First reliable records were Akershus:Oslo and Hedmark (HEs): Sor-Odal (SiebkeI 880).Sweden According to Dahlbom (1832, pp. 33,44) B. rajellus is rare in Sk6ne, Sm6land, Vtistergotlandand B. derhamellus is not frequent inSmiland, Ostergcitland, V[stergotland, Upplandand only once recorded in'Lapponia'. fn the presentDahlbom collection no specimens are abranged as .8. derhamellus, and those affanged as.B.rajellus demonstrate confusion with melanic B.sylvarum. First reliable record is from Norrbotten:Heden (Boheman 1844).Taxonomical remarksThe Scandinavian population belongs to thenominate form. B. ruderarius var. integer Alfken,l9l3 (Sparre Schneider 1918) is an infrasubspecificform.

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Studies on Scandinavian Bumble Bees17lFig. 87. B. ruderarizs (Mtiller). Legends as in Fig. 50, 54.Corrections. A worker (Oppland (On): Dovre(Schciyen) ZMO) previously revised to B. ruderariusby Sparre Schneider (1918) is a melanicB. sylvarum. B. ruderarizs has in fact been confusedwith B. w. mastrucatus and melanic B.balteatus in addition to B. sylvarum. Unrevisedrecords were therefore not considered.Sweden Widely distributed throughout thelowlands from the southernmost coast northeastto nearly 66" N in Norrbotten. Apparentlyavoiding the wooded hills in Southern Sweden.Occurring in Ohnd but not observed in Gotland.A total of about 600 specimens were examined.List of localities. Skdne: Baskemolla NRS, Benestad,Blentarp, Bokskogen ZM'B, Bonderup ZIN'.IB,Btickaskog NRS, Degebergd, Figelsflng, HiilsingborgVCA ZML, Ivo NRS, Kyrkheddinge ZM.B ZML,Kiimpinge GNM, Kiivlinge, Linhalnn, Lund ZIN{A7ZlJ[,/iB ZML, Lyckfls NRS, LoderuP NRS, MalmoZMIB, Markiehage GNM, Nytorp, Reslov, Sege,

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Studies on Scandinaviqn Bumble Bees 173Hasselrot 1962; Ander 1963, 1965; Frid6n1967. Bombus derhamellus: Dahlbom 1832, p.44 (partim). Bombus sylvarum var. D Thomson,1872; Siebke 1880; Strand 1898b. Bombussylvarum var. nigrescens P€rez, 1879, p. l2l,lectotype ? MNP !, type area the Pyrenees;Gaunitz l93l; Loken 1958b, 1960; Erlandsson1960 ; Kruseman 1960 ; Ander 1963 . Bombusnigrescens P6rez; Kruseman 1958b ; Faester 1959.)First Scandinavian recordsNorway. Akershus: Oslo (Siebke 1880).Sweden Uppland (Linnaeus 176l).have some pale hairs on episternum and/or thefirst tergite, and one worker displays a pronouncedintermediate colouring. The designationnigrescens P6rez is, moreover, erroneously usedfor the North-European population but needs noreplacement as far as the melanic form does notqualify for nomenclatural rank.B. sylvArum produces dimorphic infrasubspecificforms with sympatric occurrence not only inthe Pyrenees, Scandinavia and Denmark butalso apparcntly in other fringes of its distribution,e.g. in the Caucasus (Schmiedeknecht 1883, p.87 (337)). A few melanic specimens are alsorecorded in England (Mortimer lg22; Yarrow1 959).Taxonomical remarksThe Scandinavian population belongs to thenominate subspecies.oThough Thomson (1872) describes a melanicform as var. b, the dark Swedish individualswere nevertheless later misidentified as .8. ruderariusuntil they were recognized by Gaunitz(1931) and treated as B. ^r. var. nigrescens.Though Siebke (1880) and Strand (1898b) listedvar. b Thomson as occurring in Norway, themelanic specimens were all found under B. ruderariusin the collections when recognizedby Loken(1 958b).P6rez (1879) designated, however, B. ^r. var.nigrescens as the melanic form of the Pyrenees,where it likewise occurs together with nonmelanicforms, and therefore Kruseman (1958b)doubtfullyrecognized it as an endemic species,B. nigrescens. CorrespoRding dimorph forms alsooccur in Denmark where Faester (1959), statingthat intermediates were never observed, likewisegave the melanic form specific rank, B.nigrescens. Ffowever, inhabitants from two B.sylvarum colonies (Sweden: Sk6ne: Norra Nobbelovin Lund 1932 (Kemner) ZML) present bothtypical and melanic forms, which proves theconspecific status. The preserved material consistsof the old Q, 5 VH 4 36 of the typical form and4 melanic HH from one colony while 4 typical Vqand 4 melanic HH from the second colony. Inboth colonies several of the melanic workersQueen, workerMorphological characters. Malar space slightlylonger than distal width, about as long as Asa a.Clypeus hardly longer than distal width; discof clypeus convex with rather flne, dense puncturingexcept for anterior, usually impunctatearea between well-defined lateral impressions,which are rather coarsely punctured. Labral furrowmoderate to well-defined, almost parallelsided,queer (Fig. 224); labral lamella smooth,shining with the edge knife-sharp. Mandibleshort (Fig. 228). Surface of Tn-u smooth,shining with moderate to rather sparse puncturing.Distal keel of St u usually reduced to apoint. Hind basitarsus almost parallel-sided. Coatrather short, rather even.Queen measurements. N - 201' SE Norway;malar space: 0.77 mm (t0.03 +0.01) range:0.70-0.80 mm;'radial length': 3.36 mm (+0.09+0.02) range: 3.15-3.50 mm; interalar width:4.53 mm (+0.13*0.03) range: 4.23-4.78 mm.Body of small to moderate size.Colour pattern Pile of face yellowish-grey,more or less encroached by variable mixture ofblack. Pile of vertex with mixture of yellowishgreyand black, occipital ridge otherwise blackhaired;hairs on head otherwise ranging frommainly grey to predominantly black. Thoraxclothed yellowish-grey with exception of broadinteralar band encroached with yellowish-greyhairs in front, laterally and behind, and additio-12. Bumble Bees.

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Studies on Scandinavian Bumble Bees177Table XXVI. B. sylvarum (Linnaeus). Frequency of melanism in sex and casteNo. per cent No.dTotalper cent No. per cent No. per centSkflneTypical formMelanic formUpplandTypical formMelanic forml5l84555334642s833 100 79 100 25 100 137 10046l2792l230837327t8t774333268693l56813s012858 100 313 100 t07 100 478 1004ls97327examined material fromSweden: Skine andUppland set out in Table XXVI, clearly demonstratesthat rnelanism is almost equally producedin both sexes and castes.Distribution (Fig. 89).Norway. Restricted to the southeastern lowlandsand the extreme southern coast west toVest-Agder (VAV): Farsund. A specimen labelled'Dovre, Schoyet', misidentified to B. ruderariusby Sparre Schneider (1918), is noteworthy,being the only record from a subalpine or alpinelocality. Pronounced local distribution.Biotopes : Pastures, Ieguminosae biotopes, roadsides,etc.A total of 341 specimens was examined.List of localities: Ostfotd (0)tHvaler: Akeroy,Asmaloy, Herfol, Kirkoy, Sondre Sando; Onsoy:Dypeklo, Ellingflrd, Lervik, Ramseklo, Rorvik, Skjeloy;Halden: Asak, Hakelund, Halden, Sponvika;Marker: Rodenes; Rakkestad: Rorvik; Rflde: Oven,Tom; Moss: Jeloy VCA ZMB; Trogstad: Monsterbro, Rud, Vassbotn. Akershus (AK); As: Vollebekk;Frogn: Drobak; Brerum: ? loc., Hovik; Oslo: TRMZlj|dB ZMO; Aurskog-Holand: Bjorklangen; Skedsmo:Strommen ZMO. Hedmark ( HEs).' Hamar.'ZldB ZMO; Ringsaker: Gaupen. Oppland (On) :Dovre V (Schoyen) ZMO. Buskerud (Bd).' Hurum;Royken: Aros; Drammen; Kongsberg: Skollenborg.Vestfold (VE); Vile: Ryk; Ramnes: Kjrer, Lunde,Orrevil ; Lardal: Styrvoll; Andebu: Andebu, Kodal;Stokke: Lango, Stokke, Veierland; Tonsberg: Prestrodkilen;Notteroy: Teie; Tjome: Eidene, Mo, Mostranda,Vasskalven Joh; Sandefjord: Austeroy;Hedrum: Hedrum; Tjolling: Viksfjord; Brunlanes :Berg, Dolven, Helgeroa; Larvik. Telemark (TEy):Porsgrunn : Mule; Bamble : Trosby; Kragero. TEi:Nissedal: Kyrkjebygda. Aust-Agder (AAy) : Gjerstad:Fiane; Moland : Dal; Hisoy: Gjerdvoldsoy, His;Landvik : Havnevhg,Homborsund ; Lillesand : Natvik.Vest-Agder ( VAy ); Kristiansand : Stromme ; Farsund :Tjorve.Sweden Widely distributed on the mainlandfrom the extreme southern coast northeast tonearly 62" N in Hiilsingland. Moreover, abundantin Oland whereas the oOcurrence in Gotlandis questionable. (Two HH of the typical form,labelled Gl, from Liljeborg' collecti on ZMU maybe from this island and then the only specimenstraced so far.) The distribution agrees roughlywith that presented by Erlandsson (1951). Asingle worker recorded as far north as in Jtimtland,cf. the list below, is noteworthy.A total of about 1300 specimens was examined.List of localities. Skdne: Arild, Arkilstorp NRS,Barkflkra NRS, Baskemolla NRS, Benestad, Bjlirred,Bjornstorp, Blentarp, Bokskogen ZMB, BrantevikNRS, Brunnby, Biickaskog NRS, Bokeberg ZN{{.f.BZML, Bistad, Degeberga ZML ZMU, Eriksdal,Falsterbo NRS ZML, Glimflkra NRS, Gorslov, HaviingNRS, Herrevadkloster GNM, Hohog, HiilsingborgNRS VCA ZML, Hoganiis NRS, Hislov, Ignaberga,Kivik NRS, Klagstorp GNM, Klippan NRS,Krapperup, Kristianstad NRS, Kullaberg, Kyrkheddinge,Kiimpinge GNM, Kdvlinge, Kflgerod, Limhamn,Listerum NRS, Ljungholm, Lund GNM ZMAZ}o{B ZML,Loddekopinge ZIV4B ZML,Loderup NRS,Maglarp ZMB, Maglehem ZMU, Malmo NRS ZML,Miilarhusen, Molle NRS ZML, Nosaby ZML, Nyby,Nyhamnsldge, Nymo mosser ZMB, Niisum NRS,

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Studies on Scandinavian Bumble Bees 179Ostergdtland: Borensberg GNM ZMIJ, BorghamnNRS, Borsjon NRS, Gryt GNM, Heda Z} .4B, HiillestadNRS, Kvarsebo NRS, Linkoping NRS ZM.r-ZML, Malmslatt NRS, Mjolby NRS, Motala NRSZMB, Norrkoping NRS VCA, Sankt Anna GNM,Slaka, Stjiirnarp ZMB, Soderkoping, Ttillmo ZldB.,Tollstorp, Uggelo GNM, Valdemarsvik GNM, VinnerstadZIN4B., Vretakloster ZN[[{{B, VSversunda, VingaZI|'[B. Vcistergcitland: Alingsis NRS, Askim, FalkopingNRS, Grdstorp NRS, Goteborg GNM, HerrljungaNRS, Horred NRS, Karleby NRS, Kinnekulle,Lidkoping NRS, Ribiick NRS, Skara GNM NRS,Skovde NRS, Toreboda NRS, Vinninga NRS, VattlosaNRS, Oietred. Bohuskin: Bovallstrand NRS,Dingle NRS, Edshultshall NRS, Havstenssund NRS,Hunnebostrand NRS, Hiillevadsholm NRS, LysekilNRS, Munkedal, Nosund NRS, Stromstad NRSZML, Syd-Koster NRS, Tanum NRS, TanumshedeNRS, Tjuvkil NRS, Uddevalla, Ulebergshamn NRS,Ytterby NRS. Dalsland: Bengtfors NRS, Kopmannebro,Rostock NRS, Skflllerud NRS, Amfll NRS.Niirke: Liirbro NRS ZML, Stora Mellosa ZMB,Orebro NRS ZMB, Atorp NRS. Si)dermanland:Brandalsund NRS, Enhorna NRS, Fittja NRS, HandenNRS, Hiiringe NRS, Liiggestad NRS, MuskoNRS, Molnbo NRS, Nykoping NRS ZML, NyniisZMB, Ndvekvarn NRS, Oxelosund NRS, Sjosa ZM.B,Stjtirnholm NRS, Strdngnds ZN'{B, Svlirdso NRS,Sviirtagird NRS, Sodertiilje NRS, Trosa, TuringeNRS, Uttran NRS, Viksberg NRS, Visvass ZMB,Visterhaninge ZMB, Alta NRS, Alberga ZMB. Uppland:Adelso NRS, Bogesund NRS, Brudnis NRS,Dannemora, Dyvik NRS, Ekolsund NRS, ElmstadZMB, Enkoping ZMU, Estuna ZMU, Gimo NRS,Hiiggeby NRS, Hibo-Tibble NRS, Hfltuna NRS,Ingaro, Jumkil NRS, Knutby ZMB, KungsdngenNRS, Lennartsniis NRS, Lidingo NRS, Lovon NRS,Munso NRS, Morby NRS, Norrtiilje, NorrvikenNRS, Rimbo NRS, Roslags-Niisby NRS, Ro NRS,Ridmanso NRS, Skokloster ZIN{B, Stockholm IVUNRS VCA ZMAZML ZMU, Svartsjo, Tensta ZIN{4B.,Ulltuna IVU, Uppsala BML IVU NRS VCA ZML,Vallentuna NRS, Vassunda NRS, Yxlan ZMB, OssebygarnNRS, Osterskdr, Ostra Ryd NRS . Vcistmanland:Arboga ZMB ZML, Fanthyttan NRS, FellingsbroZMB, Guldsmedshyttan NRS, Hiillefors NRS,Kolback NRS, Kvicksund NRS, Kiirrbo ZMB, LindesbergNRS, Malmor ZMB, Munktorp NRS' Sala,Saxhyttan NRS, Skultuna ZMB, As NRS. Vcirmland:Alster NRS, Charlottenberg NRS, Daglosen NRS,Gylleby in Sunne NRS, Hammaro NRS, Kil NRS,Lingban NRS, Persberg NRS, Siiffle NRS. Dalarne:Avesta ZMB, Borldnge ZMA, Karlsbyn, LudvikaNRS, Riittvik NRS, Sjurberg, Sundborn, VikarbynNRS. Gcistrikland: Bjorke, Hedesunda NRS, Hille-Forsby NRS ZM3 ZML, Ockelbo NRS, Trodje NRS.Hcilsingland: Njutflnger NRS, Soderhamn NRS. Jrimt-Iand: Are ? 13 July 1957 (Hedstrom) ZMIJ.Finland. Confined to southeastern parl of thecountry. Rather rare. Records (Fig. 89) weretaken from Elfving (1968), also those southeastof present Finland.World distribution. Europe (British Isles; onthe continent ranging from NW Spain, the Pyrenees,Italy, the Balkans north to SouthernFennoscandia; in Central European USSR northto districts of Leningrad, Kostroma and Bashkir)- Turkey - Iran - Caucasus - Transcaucasus- Southern Ural - Altai - ? Siberia east toTransbaykal. (Reinig 1937, 1968; Pittioni 1938;Panfilov 1957; Yarrow 1959 and in litt. ; Tkalcu1960). An occurrence in Siberia (Reinig 1937,p. 78) is questioned, not being stated by Panfilov(1957, in litt. 1968).BiologyNest. Nests on the ground, producing coloniesof moderate size (Hasselrot 1962). Queen andworker are in the field distinguished from otherBombur spp. by their nervous, restless flight anda distinctly higher tune in their buzzing.Flight season. From middle of Aprilto endof September. Queen : 17 April- ?; worker:23 May-13 Sept. ; male : 20 July-26 Sept.BOMBUS (THORACOBOMBUS)VETERANUS (FABRICIUS)Apis veterana Fabricius, 1793, p. 324 No. 45;Zimsen 1954, p. 417 No. ll04,lectotype V KCC!selected by Laken (1966a), type loc. Germany:Kiel.(Bombus equestris auctt. nec Fabricius 1793;Thomson 1870; Kruseman 1959; Elfving 1960,1968; Erlandsson 1960; Hasselrot 1962; Ander1963, 1965. Bombus autumnalis: Dahlbom, 1832,p. 45 nec Fabricius 1793. Bombus arenicolaThomson, 1872, p. 3l No. 9, lectotype ? TCL!selected by Ander (1967), type area Sweden:Skflne; Schenck I 875 ; Aurivillius 1903 ; Muchardt1904.)

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on Scandinavian Bumble Bees181Fig. 90. B. veteranus (Fabricius). Legends as in Figs. 50, 54.land: Markaryd. Vtistergiitland: Utbyniis in GoteborgV 24 Aue. 1938 (Lewin).Unrevised records are not included here except thefollowing (Ander in litt.); Skdne.' Blentarp, Bjornstorp,Bokeberg, Hdllestad, Kiivlinge, Nosaby, Sege,Vinninge, Ostra Sonnarsltiv, Ahus.Corrections. Occurrence in Ostergcitland (Auri-Studiesvillius 1903) and a distribution of the speciesnorth to the southern border area of regio coniferina(Kruseman 1959, Hasselrot 1962) are dueto misidentifications. Revision supports Ander(1965), who states that the species has beenconfused with ,B. sylvarum (Linnaeus) and alsoB. distinguendus Morawitz.Finland. Sparsely distributed in Southern andCentral Finland, recorded in all counties southof 64" N (Elfving 1960, 1968). The followingrecords, plotted on the map (Fig. 90), wererevised: Ab: Lojo ZML. Kl: Sordavala ZML.Oa: Vasa ZIN.{H. Sb: Kupio ZN,.{IJ. Kb: Ham-

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Studies on Scandinavian Bumble Bees 183nec Linnaeus, 1761. Bombus pratorum (Fabricius);Zetterstedt 1838. Apis fragrans Kirby, 1802,p. 329 No. 83 nec Pallas, 1771. Bombus fragrans(Kirby); Dahlbom 1832, p. 46; Siebke 1853,1870; Wahlberg 1854; Thomson 1870. Bombusdistinguendus Morawitz; Thomson 1872; Siebke1880; Ner6n 1892; Strand 1898a, 1898b, 1910;Lie-Pettersen 1901 , 1907; Aurivillius 1903 ;Bengtsson 1903; Ringdahl 1915; Sparre Schneider1918; Meidell 1934a; Wexelsen & Skire 1934;Perkins 1942; Loken 1949; Kruseman 1959;Elfving 1960, 1968 ; Erlandsson 1960; Frid6n,Eskilsson & Bingefors 1962; Hasselrot 1962;Ander 1953b, 1963, 1965; Frid6n 1967.)First Scandinavian recordsNorway. Oppland (On): Fron: Storfjellet(Siebke 1853). No specimen is traced, but theidentification of the easy recognisable speciesmight have been correct. Specimens from Oslo;Hedmark (HEs): Sor-Odal; Oppland (On): Seland Dovre; Telemark (TEV): Skien (Siebke 1880)arg kept ZMO.Sweden Skflne, Smiland, Ostergotland, Vdstergcitland(Dahlbom 1832).Queen, workerMorphological characters. Malar space hardlylonger than distal width (Fig. l9),just exceedinglength of Ar+ r. Clypeus rather flattened towardsthe base. Outer surface of hind tibia moderatelyto strongly alutaceous. T6 laterally and distallyrather granulate, densely haired, while centrallyand anteriorly smooth to feebly alutaceous withfine sparse puncturing. Coat even, rather short.to olive-yellow. Collar, scutellum (except blackhairs at anterior margin), T,.-u olive-yellowhaired of which Tr-u usually is of lighter shade.Hairs on episternum predominantly pale yellow,fringes of sternites pale yellow to whitish. Otherwisecoat black except interalar band laterallywith an admixture of pale yellow hairs. Collarbroad, about as wide as interalar band whichis posteriorly slightly curved, i.e. including blackhairs of scutellum. The transition collar to interalarband indistinct, i.e. black and olive-yellowhairs more or less encroaching.Variation. Ventral and posterior hairs ofepisternum ranging from entirely black to dullpale yellow or whitish. Trochanters, femoraparticularlyventrally with variable admixtureof whitish hairs. Individuals with a few blaekbristles on Tn-u were examined. In general ratherminute variations.Body slender, wings evenly infuscate.Queen measurements. N : 20; SE Norway;malar space: 0.96 mm (+0.04+0.01) range:Distribution (Fig' 9l)0.90-1.00 mm; 'radial length': 4.40 mm (+0.09 Norway. Wide, local distribution in south-+0.02) range: 4.30-4.60 mm; interalar width: eastern lowlands and sporadically recorded in5.18 mm (+0.12+0.03) range: 5.00-5.43 mm. adjacent subalpine vallies. Also observed in theBody of large size.lowlands north of the mountainous massive,Colour pattern. Plumose short hairs in pile viz. in the inner districts of S6r-Trtindelag andof face predominantly pale yellow, pile of vertex Nord-Trcindelag north to nearly 64'20' N. It iswith a variable mixture of black and pale yellow uncertain whether an isolated population onMaleMorphological characters.Antennae (Fig . 42 A).Stu distally gently rounded, with distinct lateralswellings. St' and genitalia (Figs. 428-C);preapical process of penis valve usually bidentatewith one tooth strongly projecting forwards; iftrident ate, still only one strongly projectingtooth. Coat rather even, moderately long. Bodyslender and of medium size.Colour pattern Venter of head, trochanter,at least ventral part of femora predominantlywhitish-haired. Tz entirely or predominantlyblack-haired. Otherwise colouring as in the female.Variation. More or less diffuse interalar band.Tn-u withvariable admixture of black hairs.

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sen), Time: ? loc. H TRM; Kvitsoy: I June 1930 (Mei).Siir-Trcindelag (S7|:i); Oppdal: Driva st.; Trondheim:VCA ZMB. Nord-Trtindelag (N7|i); Stjordal: Lflnke.Unrevised records: Oppland (Os).' Gausdal 760 m? and l{ord-Trtindelag (NZ/).' Snisa 2 ?? (SparreSchneider 1918); Aust-Agder ( AAy).' Tvedestrand(Meidell 1934a).Sweden Widely distributed from the southernmostcoast throughout the lowlands northeastto the Finnish border. Singly recorded in subalpinezone from northern part of Dalarne toLapland.A total of about 1200 specimens was examined.Studies on Scandinavian Bumble Bees 185Fanthyttan NRS, Fellingsbro ZNIf.B, Guldsmedshyttan,Kolbiick NRS, Kolsva ZMB, Kdrrbo ZIN[.4B,Koping ZM.B, Munktorp NRS, Sala NRS, SkultunaZMB, As. Viirmland: Arvelsiter NRS, Arvika, BrunskogNRS, Ed, Forsvik, Gylleby NRS, LflngbanNRS, Nilsby, Siiffle NRS ZML, Torsby NRS, Ohe.Dalarne.' Avesta ZMB, Falun, Floda NRS, FurudalNRS, Harnds NRS, Idre, Ludvika NRS ZML, Smedjebacken,Store Tuna, Soderbiirke NRS, Vangsbro.Gcistrikland: Hamringerfjiirden NRS, Hille NRS,Ockelbo NRS, Trodje NRS. Hrilsingland: BollniisNRS, Delsbo, Soderhamn NRS. Medelpad: SundsvallNRS ZML, Timrfl NRS, Ange. Hririedalen: FunisdalenZMB. Jcinttland: Doda-Fallet NRS, FrosonNRS, Hallen, Lockne, Oviken, Snasahogarne NRS,Stromsund NRS, Ostersund NRS. Angermanland:Bjiista NRS, Bjorna NRS, Forsmo NRS, HiirnosandNRS, Hflkniis NRS, Solleftefl NRS, OrnskoldsvikList of localities. Skdne: Alnarp, Balsby, Barselb?ickhamnZMB, Bjornstorp, Brunnby, Bilrslov ZMB,Bokeberg Z}dB ZML, Dalby, Falsterbo NRS, Fialkinge,Fulltofta, Grevie NRS, Gorslov, Hardeberga,Herrevadskloster GNM, Hyby, Hiilsingborg, HistadZ}l4.B, Jdrrestad NRS, Krageholm, KrapperuP, Kronovall,Kungshult, Kyrkheddinge, LoddekopingeZMB, Loderup NRS, Norra Nobbelov, Nyhamnsldge,Reslov, Romele klint, Sandhammaren NRS, Lycksele NRS. Lu. Lpm.; Kvikkjokk V 1843 (Anders-d (Wahlb.) NRS. As. Lpm.; Asele NRS. Ly. Lpm.:Sege, Silvikra, Simrishamn NRS, Skarhult, Skiilderviken,Skire ZMB, Svalov IVU, Sodra Sandby ZM.B Unrevised recor ds: Ostergdtland; Skiinninge (Ner6nson).ZML, Torup, Trelleborg GNM, Vallikra, Veberod, 1892). Jrimtland: Edsflsen (Ringdahl 1915). Gotland:Vegeholm, Viderup, Vik NRS, Vitemolla NRS, Niir; Medelpad: Tyndero ; Hcirjedalen: Fjdtllniis:' Vcisterbotten:Lovitnger (Ander ih litt.).Vomb ZWB,Viideron, Viilling€, Viisby, Ortofta, AhusNRS ZMLZMU. Halland; Falkenberg, Getinge NRS,Halmstad NRS, Tronninge. Smdland: Alingsfls, BarkarydNRS, Eksjo ZMU, Gasslanda ZMU, JiirsniisFinland. Occurring throughout the countryNRS, Jonkoping NRS, Kalmar NRS, Ljungarumnorth to about 67' N, yet nowhere frequent.NRS, Ljungby ZMU, Markaryd ZMU, Nye NRS, (Elfvine 1960, 1968).Piskallavik, Osterkorsberga GNM NRS ZMU. World distribution. Europe (British Isles; onOland: Bengtstorp, Langlot, Morbylinga. Gotland:the continent ranging from Central Europe,Fide, Firo, Hogklint, Klintehamn, Roma NRS,Transylvanian Alps and north to Fennoscandia;Stinga. Gotska Sanddn. Ostergdtland: BorensbergGNM ZMIJ, Borghamn NRS, Hogby ZMB, Kisa, in European USSR extending from White Russia,Kungsiingen NRS, Kvarsebo NRS, Linkoping, NorrkopingNRS, Ringarum NRS, Vdversunda NRS. lov and north throughout the taiga, almostdistricts of Kursk, Voronezh, Kuybyshev, Tsjka-Viis t er g t) t land : Finnekumla NRS, Kinna, Kinnekulle,reaching the tundra) - Western and SouthernSodra Kirketorp, Ojehed. Bohuslciz.' Brastad, DingleNRS, Munkedal, Naverstad NRS, Syd-Koster NRS, Siberia - Northern Kazakhstan - Tien ShanYtterby NRS. Dalsland: Bengtsfors, Elleno NRS, Altai - Primorskiy - Sakhalin (Knechtel 1955;Kopmannebro, Rostock ZMB, Stenebyniis NRS, Panfilov 1957; Yarrow 1959).Vensborg, Amfll NRS. Ncirke: Askersund, Skrimstad,Skolberga NRS, Orebro. Siidermanland: BjornlundaNRS, Musko NRS, Molnbo NRS, Nykoping, Trosa,Tyreso NRS, Vreta, Viisterhaninge Z}dB,, Alberga. Taxonomic and faunistic remarksUppland.' Ekolsund NRS, Eldgarn NRS ZML, EnkopingZMI), Gimo NRS, Hammarby NRS, Hibo- Meidell (1934a) asserted that the isolated populationrestricted to the outmost sandy coast ofTibble NRS, Hfltuna NRS, Ingaro, Ljustero NRS,Munso NRS, Norrtiilje NRS, Ro NRS, Skokloster Rogaland, viz. Jreren and a few islands northZMB, Stockholm NRS ZML ZMU, Tensta ZMB,of Stavang€r, might be a maritime form. HeUltuna IVU ZMB, Uppsala IVU, NRS ZN.{.IB ZMLZMIJ Oregrund ZN.4B, Osterskiir, Osterflker ZMU. found the individuals larger, more robust thanVdstmanland: Arboga Z} '4B ZML, Dingtuna NRS, those of the eastern Norwegian population. HisNRS. Vcisterbotten: Bergsbyn NRS, Bodarna NRS,Degerfors NRS, Hiillnls NRS, Kroksjo NRS, PiparboleNRS, Vindeln NRS, Viinniis NRS, ViisterhiskeNRS. Norrbotten: Boden NRS ZML,, Kalix NRS,Karungi NRS, Lulei NRS ZML, Nybygget NRS,Pajala NRS, Pitefl NRS, Vita NRS. Laplaid.' ? loc.

-pe>lJ€ru eceds Jelel I'statzn"toq? p)!3o7ot1d"toryD4tO(^'UaanQ'urJoJ eleuil,uou eql pelues. -eldeJ Jo>lJoA\ pue e1etueJ >lJep eql lerll lceJ er,IlJo ere^\e lou ere/y\ feql 'u4g pe^\olloJ @vtaDfieple141 pue 'glg 'lqceulepeprqcsI slparoq'J?A Snauohtalqns 'gol peleleJ eJeJ cruelelue sr uorl€lndod uerEezluoN etll Wqt pelecrpur(gt01) repreuqcs e;redg ueq1\ 'f,qrl) Jo ruroJ.r,q-lailla4ol eql qll/y\ eletu egl pu? uorldrrcsep u€eeu-ulf ew qlyyr seerE€ el€tueJ eql'urJoJ eleuruoueql ol sEuoleq uorluFdod u€rneurpu€cs eqls 4,t0u/ a t r)cltu ouo xoJ'(8SrI sneuuulT) puelddn 'uapaws'(816I) reprouqcs erredg.{q po}ecrpu osle se suor}ecgl}ueprsFu o} onpeq lsntu peuorlueul,(lsnoeueJoduloluoc serlrleoolre{}O '(088I o{qels) olso : snqsre>lv 'r(outo1i1spn)at u0l\Durpuols ls,trfl('SE 'd 'ZE8t uroqlq€Cl : stsuaaotos snqwog' L06I'S06 I' 106 [ uesre]led-elf : t tgI >loueqrs: fSg 1 Ereqlqe/v\ i et 'd 'ZEyI uroqlqeq i (,(qrry)srylailla.,rtol snqwog 'ureqreg : {loJns'E : puelEuEeere ed.(t '(ggOt) ^\orre1 ,{q pelleqel i.IJ)P ed.(lolorl '?g 'oN 0€t 'd 'Z0gI ',{qrl) oilaruau-q sgv 'puelddn: uepa^\S eere eddt'VE 'oN'd 'gS 'sneeuur-I6LgLI tantona)o stdv:tgel ugplrg :S96I '896I ropuv iZgel lorlosseH:Zgel sroJeEulg T uossp>lsg 'u?plrC :Ogel uoss-puelrg'896 I'0961 Suptgli1: 6S6I u€tuesnr) :VSetrepetg iVrcI orB{S T ueslexel6 le VE6I IIepleW: g t0 [ reprouqcs erredg | 5rc1 uerE1q€^/$, :VOAylpr€I{cn61 I E06I snrllr^lrnv : qgOg 'eg6g I I puerls: rcgI qlog :ZASI ugroN ! Ogg I e{qels :Ztgt'018I uostuoql !lSg1 EreqFIeA\ lZt'd'T,E9Iutrroqlq€C !(sneeuulT) snauo"t,tatqns snqwog)'uoxel eql Jo uorlelerd-Jelur lensn eql q]ll\ seerEe oauoilatQns pelleqeli-IST eletuoJ 'puelddnV : uepe^\S eere ed,{t'Sg 'oN '8S21'sneeuulT 6LS'd oaao.t.tatQns sldV(snEvNNrD snsNvvtrsJsns(snswososNvvvsJsns ) snswos'ldes Z-KIry SI eletu !'1deg tl-eunf g :re>lro^\!'Eny 6-,{entr E : ueen}'requretdes Jo elppFuol ,{eIN Jo EutuulEeq tuoJC 'tlosoas fi.!8r19'u^\op ue>loJq oJeA\suoococ lseplo eql s€ Lg pessedrns el€q lsntupecnpord slu€trqequr Jo reqrunu [e]o] eq] snql'peqcleqle,{ }ou p"r{ suoococ eleu/re>lJo^\ Lpue sueenb tI qcFlr!\ Jo suoococ elewfwlroi!\St pue suoococ ueenb ZZ peureluoc '(806IuosslEuefl) 0061 lsnEny g uo pezf,leue ',{uo1ocpezrlrsemd 'punorEV eqt ur slseN'$aNrtSolorg'uI 000I ]noq€ epnlrlle'nee1e1d snour€lunou eql ssed ^(eul snpuan&ul|slp'gleql ,(lnqlssod et{l epnlcxo 'relezlloq o}ouop 'enoqe tsfl et{t'lepddgTc :(lfS) Eelepugrl-rgs pue orloq :(uO) puelddo q'e'l 'llefgerzroqJo eEuug ureqlrou pue uJoqlnos eql te spugelEurg 'ur€r{o ure}unour eq} go ped uJeqgoo .q}pue ureqlnos eql uee^\leq de? eql ur seEes-sed pelceJrp-lsem-lsee qEnorql uepea\S ruo4uorseAul ue el€Jrput ^{eur 'q}rou JerI}JnJ usgusur pu" prolgsulreqpuorl Jo epls ureq]nos eqtuo 'e'r 'r{ezlrro51 ur pJoceJ }souruJor{}Jou eql'lno pedrmflqrssod ro pecnper Eureq uorlelndod [ecol erll roJdtryqlsuodser eql eJeqs deul 'ecuelsur ro3i 'uererfJo uorlelrl[nc elrsue]ul 'sopecep ]se[ eql EulJnpedecspuel ur seEueqc eql qlll\ peseeJcur ,(lsnorzr-qo eêq suorlrpuoc elqeJnoêJun eseql'eeqolqulnq eql JoJ suolllpuoc 8ur,rq lsroly\ oql luoserd'sercecls oql Jo uorlnqlrlslp orll Jo lpu11 urorlpou'suorle8rlselorrreJlxe eql le Eureq 'seeJe eql-ur lueceJ Euunp pepJoceJ eJea\ sueurrceds oN'JoEuelels Jo glrou spuelsr eql ol protlolso ruo{lseoc eql Euole ,{llecrperods sJncJo snpuan&ull-s!p 'Br JeqleqÂ ureueJun olor{A\ et{t uo sr }I'eôqe Jc'denuoN ES ur uollelndod eql roJ ercp Eurpuods-eJJoc eqt Jo eEuer eql utqlrl\'lenezlroq 'eteIIe'urur SE'S :qlpl^\ rele;elul pue urru SE V :.ql8ue1IerpeL 'urur S6'0 : eceds Jeletu '4L 'ueenb eql JoslueutueJnseour eql'sree.( lueceJ ur eote leql urpelJesqo ueeq ]ou seg serceds eql prre 'suorlcefiocetlt ur peJeJl eJe.{\ sJe1rolr eeJq} pue ueenb e,{1uo se 'leneznoq 'pelenlele eq }ouuec ,{"loeqtruDIoT'V 98I

Studies on Scandinavian Bumble Bees 187ly longer than distal width, about as long as Ara r.Clypeus bending towards the base. Outer surfaceof hind tibia barely or not alutaceous. To rathercoarsely granulated. Coat even, very short. Bodyrobust, wings evenly infuscate.Queen measurements. N - 15; Sweden:Skine ; malar space: 1 .05 mm ( +0.05 +0.01)range: 1.00-1.15 mm; 'radial length': 4.38 mm(+0.11+0.02) range: 4.25-4.55 mm; interalarwidth : 5.47 mm ( +0.18 +0.05) range : 5.23-5.73 mm. Body of large size.Colour pattern Tn-u with variable mixture ofblack and greyish-white or pale dull fuscoushairs. Otherwise coat black or brownish-black.Variation. Individuals with indication of yellowcollar, often reduced to lateral yellow patches,were examined, likewise those with some yellowhairs on scutellum. Workers with yellow hairson T1 were also traced.MaleMorphologicalcharacters. St u distally gentlyrounded and even, i.e. without lateral swellings.St, and genitalia (Figs. 43A-B); flattened preapicalprocess of penis valve tri-or quadridentatewith lateral teeth strongly projecting (Fig. 43C),occasionally bidentate with only one toothstrongly projecting. Coat rather even, moderatelylong. Body of medium size.Colour pattern Pile of vertex ranging from allblack to predominantly yellow. Collar, scutellum(except for a small black-haired anterior medianarea), Tr, lunate anterior pafi of T, with paleyellow hairs. At least fringes of Tu-u yellowishwhiteor whitish. Episternum, mid- and hindfemora with variable but usually predominantadmixture of whitish hairs. Sternites usuallywhitish-haired. Otherwise coat black. Interalarband, broad, occupying more than half ofmesonotum, posteriorly more or less curved, i.e.including black hairs of scutellum.Variation. Gastral tergites individually witha vafiable mixture of dark and light hairs, oftenfringed whitish. Occasionally Tr-, predominantlyblack-haired andlor To-u entirely yellowishwhite.Slightly melanic individuals were alsoexamined.Distribution (Fig. 92)Norway. Rare. With exception of two queensfrom the western coast I have only examinedspecimens from scattered localities in the southeasternlowlands, in fact restricted to the areasaround Oslofiord. The list below reveals norecent records and the present distribution isuncertain.It is unlikely that the abundance of B. subterraneushas ever been great in the extreme westernarchipelago northwest of Bergen as indicated byLie-Pettersen (1905, p. 10).Theauthor musthave confused the species with melanic B. hortorumeven though he pointed out that bothoccurred together in the actual area. No B.subterraneus individuals were traced in his collection.A total of 23 specimens was examined.List of localities. Ostfold.' Moss: Jeloy ? 24 May1908 (Barca) VCA. Akershus; As: Vollebekk ? July1949 (?); Oslo: ? 27 July 1844 (Dahlbom) ZML 3 99(Siebke)? H (Esmark) ZMO, Slemdal ? 30 May 1910(Sch.) TRM, Toyen 4 HH (Siebke) ZMO, V. Aker 2W(Siebke) ZM.O. Buskerud (Bt,).' Hurum: Filtvedt 3 VVd 8 Aug. 1910 (Sch.) TRM. Vestfold; Notteroy: Teieq 25 Aug. 1934 V 2 Aug. 1935 (Bache). Hordaland(HOy).' Fana: Skipanes ? 28 July 1907 (Barca) VCA.HOi: Etne: Etne ? 2l June 1923 (Mei.).Unrevised records: Vest-Agder ( VAy); Flekkefjord:Loga and Hordaland (HOi): Ullensvang: Ullensvang(Meidelll934a) are questioned (Fig. 92).It is uncertainwhether the author, who obviously recognized thespecies, examined the actual individuals himself.Correction. Records from Hedmark (HEs):Odalen and Sor-Trcindelag (STi): Oppdal: Drivstua(Siebke 1880) were revised to B. ltortorumand those from Buskerud (Bv) : Al (Strand 1898a)likewise to B. hortorum and B. sporadicus respectively.Record from Hedmark (HEs) : Vang:Hjellum (Wexelsen & Skire 1934) was revisedto B. hortorum (Meidell unpublished).Sweden Widely distributed throughout thelowlands northeast to 60o 30' N in Uppland, alsooccurring in 6land and Gotland. The presentstudy agrees with Kruseman (1959) that thenorthern border area coincides with that of regioquercina (cf . Figs. 2 and 92).Therefore a recordof this species in Dalarne, in the forest belt ofHtirjehhgna mountains, cf. the list below, is

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Studies on Scandinavian Bumble Bees 189Siivsjo NRS, Viirnanas. Oland: Bengtstorp, BorgholmNRS, Boda NRS, Hogsrum NRS, Kleveby, MorbylflngaNRS ZML ZMU, Norra alvaret, SegerstadNRS, Skogsbyn NRS, Vickleby NRS. Gotland; AustersNRS, Firo NRS ZML, Garda, Hogklint, Klinte,Kyllei NRS, Lilla Karlso, Ljugarn Zi|l{A, RomaklosterNRS, Slite NRS, Sudersand, Visby NRS,Viinge. dstergritland: Alvastra NRS, BorensbergGNM, Dagsberg NRS, Hdllestad NRS, Hogby ZMB,Linkoping NRS ZMB ZML, Ljung NRS, MalmsliittNRS, Norrkoping NRS ZM.A, Skonberga NRS,Slaka, Tollstorp, Valdemarksvik NRS. Vristergdtland :Finnekumla NRS, Rflbiick NRS, Skara GNM. Bohus-Irin: Skafto NRS, Syd-Koster NRS. Dalsland; Bengtsfors,Bickefors, Kopmannebro, Mellerud. Ncirke:Stora Mellosa ZN.[B. Stidermanland: Molnbo NRS,Nykoping, Ndvekvarn NRS, Rekarne NRS, SjosaZMB, Strdngniis NRS ZMB, Trosa, Uto NRS, VallaNRS, Viisterhaninge Z}dB, Alberga ZMB ZML.Uppland; Adelso NRS, Angarn NRS, Danderyd NRSZMB, Elmsta ZMB, Enkoping ZMU, Gimo NRS,Hammarby NRS, Knutby ZMB, Lidingo NRS, Lovo-Drottningholm NRS, Rydbo, Skokloster ZMB, StockholmNRS ZMA ZML ZMU, Tensta ZMB, UltunaIVU, Uppsala IVU NRS ZM.A ZN{{B ZML ZMU,Viirmdo, Yxlan Z}i..{B, Osterskiir, Osteriker ZMU.Vcistmanland: Arboga ZN{{B ZML, Fellingsbro ZMB,Kiirrbo ZMB, Munktorp NRS, Sala, Skultuna Zll.fB.Ytirmland: ? loc. NRS, Forsvik, Torsby NRS. Da-Iarne: Htirjein, Hiirjehigna ? 27 July 1927 (Forsslund)NRS.BiologyNest. Nests well below the surface of the ground(Thomson 1872), producing rather large colo-.nies (Hasselrot 1962). Yet a colony (Sweden:Sk6ne: Norra Nobbelov in Lund 8 ?? 24 HH3 dd 6 Aug. l93l (Ander & Kemner) Z}t/.L)was established in the moss (Ander in litt.).The total number of individuals raised is unknown.Flight season. From beginning of May untilthe end of September. Sweden: Skflne - Oland;Queen: 16 May-5 Sept. ; worker: 20 May-26 Sept.; male: 13 July-l4 Sept.SUBGENUS RHODOBOMBUS DALLATORRERhodobombus Dalla Torre, 1880, p. 40, typespeciesBremus pomorum Panzer - Bombuspomorum (Panzer) by designation of Sandhouse(re43).(Pomobombus Krtiger, 1917, p. 65, type-speciesBombus pomorum (Panzer) :_ Bremus pomorumPanzer by designation of Sandhouse 1943.)Specimens from Htilsingland (Hasselrot 1962)were not traced. Owing to confusion with melanicB. hortorum, unrevised records were not (PANZER)BOMBUS (RHODOBOMBUS) POMORUMincluded here.Finland. Restricted to the extreme southern Bremus pomorum Panzer, 1802-1805, part 86coast. Rare. Records (Fig. 92) were taken from Teil 18, type not traced.Elfving (1968).(Bombus lefebvrei Lepeletier, 1836, p. 461,World distribution. Europe (Southern England ;on the continent ranging from France, Italy,Albania, Yugoslavia, Bulgaria north to SouthernFennoscandia; in European USSR extendingfrom districts of Kamenets - Podolskiy, Poltava,Kharkov, Voronezh, Saratov, Kirovgrad andnorth to districts of Leningrad, Yaroslavl, Perm;Crimea) - Turkey - Caucasus - TranscaucasusSouthern Ural Northern and Eastern Kazakhstan- Tien Shan - Altai - Baykal - NorthernMongolia (Sagemehl 1882; Panfilov 1957;Yarrow 1959; Tkalcu 1960, 1969; Reinig 1967,1968). Introduced to New Zealand (Gurr 1964).13. Bumble Bees.holotype ? MNP, type loc. France: Gray;Tkalcu 1969.)First Scandinavian recordIt is with doubt that B. pomorum is herebypublished for the first time and from Sweden:Ostergotland: Atvidaberg about 58" 10' N and16" E. The record is based on a single smallworker kept at Zoologiska Institutionen Lund,and has the following labels: (1) O Atvid. vidsjostranden l2-7 -ll; (2) det. Bgtsn; (3)

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Studies on Scandinavian Bumble Beest9Ihere comprises .more than half the number ofEuropean species. It exceeds that of the neighbouringcountries of which Finland is representedby 25 species (present revision), comprising 24Scandinavian ones in addition to the rarelyrecorded B. semenoviellus Skorikov, and Denmarkis occupied by 19 species (Hammer &Holm 1970), all of which occur in Scandinavia.Although some areas are still insufficientlyinvestigated and further observations could improvethe knowledge of the distribution of someof the species, the present study reflects to agreat extent the composition of the Scandinavianbumble bee fauna. The distribution, illustratedon the dot maps, refers, however, to more thanhundred years of collecting, which might giverise to the question whether it corresponds tothe recent occurrence of the individual species.The large collections from the very last decadeshave, however, in general implied an extensionof area, which indicates that the dot maps couldwell depict the actual distribution. In two casesonly, viz. in B. distinguendus and B. subterreneus,peripheral records of older date, not being confirmedduring recent observations, indicate possiblefaunistic changes of negative nature. Thosecases do not, however, influence a broad zoogeographicalgrouping of the species outlined below.Species with restricted southern distributionThis group includes four species , viz. B. cullumanus,B. ruderatus, B. veteranus and B. pomorum,of which the three former are confined to southernmostSweden (Fig. 93). The population ofB. cullumanus (Figs. 58, 93 A) is restricted tosouthernmost Skine and 6land. The species hasa disjunct distribution in Europe and West Asia.The Scandinavian population belongs to thenominate subspecies which has a relic distributionin Europe, being confined to SoutheasternEngland and the extreme coast from theNetherlands to East Germany. The main occurrenceof the species is represented by B. c.serrisquama extending from Central Russia, eastto western part of Southern Siberia and southeastto eremials in Turkestan and Afghanistan.B. cullumanus is otherwise recorded in NorthernSpain, France, Poland, Hungary and NorthernCaucasus. B. ruderatzs (Figs.76,93 B), occurringmainly along the coast from Halland in westto Blekinge in east, is a Mediterranean faunaelement with a Euromediterranean distribution.It is widely distributed throughout the Europeancontinent except in the southeastern part, andmoreover reaches the British fsles, Madeira, andNorthwest Africa. B. cullumanus and B. ruderatusare restricted in Fennoscandia to southernmostSweden, while .8. veteranzs (Figs. 90, 93 C) alsoscantily occurs throughout Southern and CentralFinland. The latter has a southern Eurosiberiandistribution, ranging from the British Channelthroughout Central Europe east to Predbaykaljeand Northern Mongolia. Owing to a singlespecimen of B. pomorum, which according tothe label was caught in Ostergotland: Atvidaberg,this species is added here. It is a Mediterraneanspecies with recent westpalearctic distributionranging throughout Central Europe eastto the Southern Urals and southeast to theBalkans. It is also known from Turkey.Northerly advanced southern species\The term 'northerly advanced southern species'as used here, applies to species with recent distribrrtionin Southern Scandinavia, i.e. broadlyspeaking the area represented by regio faginaand regio quercina (Fig. 2). If also advancedfurther north, the species does not occur inregio alpinalarcticA.As many as ten species are included here.The consecutive northern limits of nine of themare noteworthy (Fig. 94 A-I), more or lessforming the same contours. Starting from thecoast of Norw&y, the boundaries run close togetherand then spread fan-like across CentralSweden before joining to enter a southern andnorthern group framing the Gulf of Bothnia;further east most of them converge into Russia.This characteristic undulating belt of northernborder areas, comprising the northern limits of anumber of terrestrial animals (Lindroth 1956)and plants (cf. Hult6n 1950, the maps) is a densionzone due to a complex of ecological factors,of which climatic ones play an important role.

ur N o09 of lseerluou pepJoceJ z(pluecs pu?drlunoc eql go ped lsoruursglnos oql q Euprnrcodpreu uepe/(S q eUqA\ 'N "99 flrueu ol rllroufrlunoc eql Jo lulod lsoruuJeglnos eql tuo4lseoc eueJlxe eql of peprJlseJ sr femrolq uIselceds oql 'elqe>lrerueJ sl (V V6'Z,g'sEI.{) tantol-s?nct'gJo uollnqlJlslp u€rl€urpuecs punfsrp egl'eJerl popnlcw eJe sgtwnq 'g pue wnnqts'g'snu&ota 'tI 'snauDtatqns 'g 's44satta1 'g 'utnJ-ocsnta'g '(C-yV6'EIC) Dtuqrog Io ItnC to puauaqnos ar,lt Eulsson Uuil uaqttou tlUM sapadg'll Jo rluou Eurssud Jo eruqlog JoJInC Jo pue urerllnos eql Eurssorc slFull uJequouerll Jo uol4sod eql o1 Eurproccu sdnorEqns oa\lolq pepplp ere serceds eql ',te/ruoN Jo lsuoclse/( pu" qlnos erueJlxe etll Euole Eurpuelxe'serceds eql Jo eruos tfq peleJlsuouep dpeelcsr JelJJ"q I€Jnleu e s€ eEuer ulelunoru eqlsnuDnlat 'g : J :(snrcuqeg) sngotapnt 'g :g :(dqrry) snuDuryln? 'g:V Jostprll ureqlrou eEe.re^€ oql'(srucuqeg)'€6 'EIdut dl 00e dE @l 0uillo-T'Y z6l

Studies on Scandinavian Bumble Bees 193Fig. 94. The average northern limits of A: B. muscorum (Linnaeus); Bt B. tenestris (Linnaeus); C: B. subterrazeas(Linnaeus);D: B. magnus Vogt; E: B. sylvarum (Linnaeus); F: B. humilis Illiger; G: B. Iapidarius (Linnaeus);H: B. ruderarius (Miiller); I: B. distinguendus Morawitz; J: B. soroeensis (Fabricius).Uppland. This atlantic type of distribution isemphasized by the subspecific splitting of theScandinavian species (Fig. 82) of which B. m.Iiepetterseni is adapted to the atlantic climate onthe Norwegian west coast. However, owing toa few relic occurrences north of the indicatedboundary (Fig. 94 A), that of Finland:Kilpisjiirvifar northLe:situated in the mountain region asas 69" N, B. muscorum apparentlyproceeded further towards the north during morefavourable climatic conditions in earlier postglacialperiod than at present. Outside Fennoscandiathe species occurs in the British Islesand ranges throughout the European continentnortheast to almost the Polar Circle in Russiaand with relic occurrences north to 69" N.Further east it occurs in Kazakhstan, NorthernTien Shan, Southern Siberia, Northern Mongolia

'1e>1dege>leT ol lsee seEuer pue 'erssng ul N "09ol ls€eglJou edornE lnoq8nor{} spue}xe erpuecs-ouueC eprslno smto.tapwg'uorlelndod qsruurCet{t q}ll\ pe}couuoo sr serceds eq} eJerIA\ 'ue11oq-rroN ol ]seeqpou uope^\S lnoq8norgl pelnqrr]-slp sl lI 'lseoc uJelse^\qlnos pue qlnos erueJlxeeqt pue spuel^\ol uJe]seeg]nos eq] o] pe]3rr]seJsr ,(emro51 uI (ff V6 'Lg 'sEIC) snrtunpn.t 'g'snsecneJ o1 de4rnl tuorg EurEueJ eoJe eqlur leruoJo oql se]er]euod pue ersrunl ur sJn3coreloeroru'eE1onlI eql ol ls€o pue N "09 olrlgou Eurqceer erssn)I ur 'edornE lnoqEnor{}Eut8uer erpuecsouuoC eprslno 'serceds ueoueJ-relrpetr tr e sl snuoptdo1 '6. 'Bluqlog Jo JInD oql Jopue urerlgou oql ur uorlelndod qsruurC oql olpe{ull sr 1(ou ,{q uorlelndod r{srpe^\S oq} req}erl^rurepecun sr 1l 'e'r 'suorleErlsenur ]uercgJnsur olenp Jo leer sr ue]loqrroN pu€ ue]]oqre]sg^ ursnlerr1 eql rer.{}eqla uor}senb e sr }! snq; 'pueluetu-reEuy ul N ,08 "t9 .(1reeu ol lseeggou po^JesqoJeJ os sr serceds eql uepe^\S uJ'N "I9 ol qgoulseoc erueJlxe eql Euole spuelxe JeloeJoru pues,(e1en luecefpe eql Jo eruos sel€Jleued'spuu1-A\ol uJelseeqlnos et{l uI pelnqlJ}slp Sl{ezuro51ur (C V6 '59 'sElg) sntnptdol'g 'uorlnqlrlslpJo ed,& l€luourluoc e elug JoruJoJ eeJql oqlrlcrrll\l Jo dnorEqns s1ql ol Euoleq slsuaaons 'gpue snpuan&u11s1p'g 'snutonpwg 'sn1top1do1'g'zl^'sercods rnog'0-Ct6 'EI.{) otut1tog Io finCIo tlltou Sutuun"t Utttl untlt,tou Llttu sanadg'erloEuolN ureqlroN pue 1e4,(eg ol lseeEurqceeJ 'uorlnqrJlsrp crlersv Jeprl\ e oleq olul\oDtnl sr sltwnq 'g reJ os leql ldecxe uorlnq-lJlslp uerseJng erues eql lsowle ener,I sercedsqlog 'uailoqntsoAur N ,v9 lnoq€ ol quoupeprocer eruqlofl Jo JInc Jo ls€oc eql Euole 'E'e'qlrouJer1UnJ ,(pe4reur seEuer JeuJoJ eql leqlldecxe wnn^1,$ 'g Jo l€r{} e{11 rer,I]€r sl (C V6'6L'sEIC) sllfanll'g Jo urelled Ieuorlnqlrlslp eql'ueJI ur lerueJe eq] puesnsecneJ 'de>1rn; tuor; ul\ou>[ sr ]I 'erEeeur sruorleuJoJul crlersv eql lnq 'lell17 ur sJncoo lI's[€rn ureqlnos eqt o] lsee pue 'erssng A\N ulN .09 ol tllrou'sue>11e9 oqt 'dlell 'utedg A\Nruo4 se8uer luourluoc aql uo pue selsl t{slllJgeql q sJn3co elpue3souued oplslno wn,tortlts 'g'puelurg lsoluuJelseeqlnos ur pepJoseJ st lI 'puel-Eurs1g11 ul N "29 ,llreou ol lseoqgou pelnqlrlslp,{leprtr uepe,,!\S ur sr 'lseoc uJeqlnos eueJlxeeql pue spuell\ol ]soruuJo]seeqlnos er{l o} Eul-qceer ,temro51 ul '(g V6'69's8lg) wnn^l,ts 'g'snsecne]pue .(e>1rna tuo4 u^\oDI osl€ sr ]I 'eqoEuolNuJeqUoN ol ]see pue erssnU l€rlueJ lnoqEnoJql'sue>11egeql 'f,IelI 'ecuerg ol qlnos lueurluoc eqllnoqEnorel 'puelEug uJerllnos ur sJncco €rpuecs-ouueC eprslno serceds eql'lseoc qlnos eql uopeprocer .{16urs,{po sl ll pueprC w epq^\ 'puel-ddn ur ,0€.09 ol lseeqgou polngrrlslp ,(1eplmuepems ul sl snaua,ntatqns 'g 'pepEllse.Lul ,(1roodueoq eêq,{1e1e1 qJlqa seoJs ur Eurrrncco '^(rlunoceql Jo t"l;ed lsoruuJelseeqlnos eql ol ut\eJpqlrltdltueredde sr uorlelndod eql 'slsrxe [rls ]l JI ]€g]pepnlcuoc sr lI 'ursuocun sr ,(emro51 ur sercedssrrll Jo uollnqlrtslp eqt (C V6 '26 'sEl$ snauo.t-Dtqns 'g Jo lueluleeJl eqt ur pouorlueu sv'uelse>lJnl ur lerueJe eq] Eurgceerpue uelsq>leze>{ o} }see snsecne3'edornguJer{}nos pue 'ecpgyIer}ueJ lnoqEnor{} lsel\-r{uoN 'spue1s1 ttmue3'se1s1r{sl}lJfl ur Eul"uncco'uorlnqlJlslp ueeu€JJelrpeuoJng epltvr e 6um,eq€rleurpuecs eprslno 'ssceds ueeueJJelrpetr [ esr'serlrl€collIpele8rlselur IIoA\ e]rnb ,(lsnovr-erd epnlcq ,{1ged qcFlls 'seeJe repJoq uJeqlJoueql ruoJJ spuu lueJeJ ,(1uo Qte eJeql'sercedsenrsuedxe ue sr 'puepllrlsgD ul ,0€ "09 lnoqe olls€orluou pelnqlrlslp ,{leprzn uepe/Ks ur pu€ prolJ-olso Jo suoJrlue eql ul popJocer.'{po femro5l ur'(g V6'Lg 'sEtg) slusatnt'g 'spuel^\ol urolseo-qlnos eql ol peuguoc ,{emro51 uI lnq 'uepemSuJelseg ur qpou JeJ JeqleJ Eutqceet'erteurpuecsur uorlnqlJlslp Jo ed,(t Ielueurluoc e lueserddnorEqns sql ur serceds Eurutetuer JnoJ eql'(nsue;) Eulqf, pue errnqcuel4 ol lsee elsv urseeJe peJetlms ur 'JeloeJour 'pue 'eeg clllefl er.{}ol rllJou edornE [eJ]ueJ lnoqEnoJq]'seeuer,{4eql 's,{eu4r6 eqt spuel}ocs ur pepJoceJ reJ ossr erpuecsouueg eprslno pue puelurC Jo gedlsoruuJeqlnos eql ruoJJ uA\oDI sr 1I 'eruqlogJo JInD Jo lseos erll uo N "Eg 01 lseequou€rAeurpueos uJeqlnos 1r1 ecueJJncco esredstpe pue 'N "0L ol rlgou lseoc uerEezluoN eqlEuole uorlnqrJlslp snonurluoc € sel"clpul JeJ osuorleruroJul'(Atseel lv V6 'SS 'sElg) snaSow'gserceds Iucrlrrc eql sr uorlnqlrlslp Jo ed,{torluelle ue Eurneq ,{ltueredde setceds JeLI}ouv'snseone3 eql pue 'snsecnecsuuJl 'rte>1rn;ruo4 ul\ou>I osle sr pue 'etrnqcuery ol lseeua4o7 'Y v6r

Studies on Scandinavian Bumble Bees 195It is also known from Tunisia, Turkey and theCaucasus.As mentioned in the treatment of B. distinguendus(Figs. 91, 94 I) the population on thesouthwestern coast of Norway might have disappeared.The recent investigations indicate adistribution limited to Southeastern Norway andto the inner part of Trondheirnsfiord, the latterapparently connected to Sweden in Jiimtland.The Swedish population is otherwise widelydistributed throughout the country, but onlysporadically recorded in Lapland. Outside Fennoscandiait occurs in the British Isles, in Centraland NorthernEurope, in fact almost reachesthe tundra in Russia. Further east it rangesthroughout Northern Kazakhstan, Altai, Westernand Southern Siberia, reaches Sakhalin,and occurs in Tien Shan. B. distinguendus atleast in Europe has a more northern distributionthan the remaining species hitherto mentionedand might be a Siberian fauna element.It is with doubt that the last species, viz.B. soroeensrs (Figs. 51, 94 J) is included in thisgroup. It has expanded wider and further norththan the remaining southern species; in fact itis dispersely distributed north to 70" N on theNorwegian coast. It occurs throughout Swedenyet nowhere reaches the high mountainous region.The species ranges throughout Finland exceptin the northeastern part. B. soroeensis has expandedwider and further north than previouslynoted (Reinig 1939; Erlandsson 1950), and mustbe considered as a dynamic southern species nolonger being peculiar to the southern Scandinavianfauna. Outside Fennoscandia, it occurs inEngland, Scotland and on the continent, is continuouslydistributed throughout Central Europeeast to Northern Mongolia. It has additionallya complex of disjunct ranges in Southern Europe,Northern Africa, Turkey, and the Caucasusfrom where it penetrates the eremial south ofCaspian Sea. It is remarkable that the specieshas expanded north to 70" N in Norway.Eastern speciesThree species, viz. B. sporadicus, B. cingulatusand B. consobrinus are termed 'eastern' accordingto their occurrence in Scandinavia and theirtotal area of distribution. They all have theirmain distribution in regio coniferina (Fig. 2),and thus are broadly characterized by theirabsence from Southern Scandinavia, the highmountainous regions, and the extreme westernand northern coast (Fig. 95). They are northernEurosiberian species, representing the taiga speciesof the Siberian elements.B. sporadicus (Figs. 56,95 C) is locally frequentand widely distributed in the forest east of themountain chain in Southern Norway, reachesthe coast in the wooded areas in Sor-Trcindelagand Nord-Trondelag and is, moreover, scantilyrecorded north as far as the Polar Circle. Thespecies in Sweden is widely distributed throughoutthe forest from the northern paft of Dalslandand northwards. Outside Scandinavia it occursthroughoutthe taiga east to the Pacific coastand at least in Finland, extending north to 70o N.It ranges south to Altai and Northern Mongoliaand is distributed along the coast from NorthernManchuria to Kamchatka.B. cingulatus (Figs. 60, 95 B), locally few innumber, is in Norway distributed in the southeasternforest and adjacent subalpine zone, andscantily occurs further north to 70" N. In Swedenit is dispersely observed from Dalarne and northwardsthroughout the country. The species isconfined to the northern and eastern part ofFinland and further east the distribution is morenortherly than that of ^8. sporadiczs. The southernlimit is in Siberia running north of Lake Baykal,and its occurrence along the Pacific coast rangesfrom the area east of Amur north to Anadyr.The Scandinavian distributional pattern ofB. consobrinus (Figs. 73, 95 A) accords withthat of the main food source Aconitum septentrionale,both of which are distributed in coniferforest and adjacent subalpine zone. In Norwoy,the species ranges between 59o and 68" 30' N,and locally reaches the inner end of some westernfiords and the extreme coast north of Trondheimsfiord.The Swedish population ranges fromHiirjedalen to Lapland (Lu. Lpm.) and extendseast to the Gulf of Bothnia in Medelpad. B.consobrinus has a disjunct North-European distributionrelated to that of Aconitum. OutsideScandinavia it is confined to Northern Karelia

ur uo4nqlrlslp epl/I\ pu" osuop e seg selJedseql ',{eluoN uI '(0S 'EIc) rutagftnm 'g 'zlt',(1uoserceds euo ttq polueserder sr 'erpuecsouueC Jolsua ecuosqe sll pu€ slêulpuscs uI uollnqlJl-slp .(1re1sen e ,{q pez'relc€Jerlc 'dnor8 sFIIsalcads un$anl'?{}€IICtus) puuuqetples 'uede1 'euFIC }suoq}roN 'lellv ruor;u1rrou>[ sr pue N .09 Jo qlnos eEre] uerreqls or{llnoqEnorqt seEuer eoq elqtunq oql ts"o JeqUnC'ued €[oX Jo ]seoc ls€e oq] uo pepJoceJ ueae sl ll! erssng ur plro11 - ^\ocsol I perEurusl Jorlgou seeJe psleredes IeJeAes ol pue puelurd ur'repu€IdN snctpo,tods'g :3 :Ereqqe76sn7o7n&4c 'g : g i uoqlqe q snuqqosu@ 'g : V Jo uorlnqrrlsrp rrerpuetsormed egl roJ sllull e€erele eql'96 'EtguaToT'Y 96r

Studies on Scandinavian Bumble Beest97conifer forest and adjacent subalpine zone northto nearly 68" N, and also occurs along thesouthern and western coast. In Sweden, it rangesfrom Dalsland north to Lapland (Lu. Lpm.)and occurs scattered east to the Gulf of Bothnia.B. wurfleini is a boreo-alpine species. With theexception of a single record in Finland, it is inthe north restricted to Scandinavia. The southerndistributional area displays disjunct occurrencesconfined to mountain ranges and adjacent forestsin Central and Southern Europe, Turkey, theCaucasus, and the Urals.Arctic and subarctic speciesThe fauna in alpine/ arctic regions of Scandinaviais distinguished by five tundra elementsthree of which, viz. B. hyperboreus) B. arcticusand B. balteatus, are the only holarctic speciesin Scandinavia while the remaining two species,viz. B. alpinus and B. lapponicus, have a palearcticarcto-alpine distribution. The members ofthis group have a disjunct distribution relatedto the southern and northern part of the mountainchain (Fig. 96). The most arctic species isthe rather rare B. hyperboreus (Figs. 71, 96 A)with the southern area of distribution restrictedto scattered localities ranging from Jotunheimento Dovrefiell in Norway and a local mountainousoccurrence in Hiirjedalen in Sweden. The northerndistributional area is confined to severallocalities in the mountainous Lapland and northernmostNorway.It is a circumpolar specieswith islet occurrences in the arctic fringe of bothhemispheres, inclusive Novaya Zemlya andGreenland.B. arcticus, in Fennoscandia and Kola pen.represented by the subspecies B. A. diabolicus(Figs. 68, 96 B), has a disperse distribution inthe mountainous massif in Southern Norwayand adjacent mountains in Sweden, i.e. in Hiirjedalenand Jtimtland. The northern distributionarea refers to a scattered occurrence along thenorthern part of the mountain chain with awider distribution in northernmost Norway. B. Q.diabolicus extends east throughout NorthernFinland to Kola pen. and ,8. arcticus is other-wise distributed in Arctic Siberia, Novaya Zemlya,Kamchatka, Arctic America and Greenland.With exception of a rather narrow gap in thearea separating the southern and the northernpart of the mountain chain, B. balteatus (Figs.70, 96 D) is widely distributed in regio alpinalarctica and regio subalpinaf subarctica from thesouthern elevation of the mountain range to thenorthern coast. It is also recorded locally inadjacent spruce forest and at sea level in theinner part of the western fjords. It is distributedthrbughout Northern Finland, occurs in NorthernSiberia, Kamchatka, Alaska, western partof Arctic Canada, along the mountain range inthe Rocky Mountain States, and has a relicoccurrence in Mongolia.In Scandinavia, the arcto-alpine B. alpinus(Figs. 67, 96 C) possesses a clearly disjunct distributionrelated to the southern and the northernpart of the mountain chain, and in NorthernNorwayhas expanded to the extreme coast.It is a European species, in the north restrictedto Fennoscandia and the Kola pen., and in thesouth to the Alps.The distributional pattern of the arcto-alpineB. lapponicus (Figs. 63, 96 E), the dominantbumble bee in the mountains, is noteworthy.The species is continuously distributed in regioalptnalorctica and regio subalpine f subarcticathroughout Scandinavia and frequently observedin adjacent forest as well. In Norway, it locallyreaches the extreme western and northern coast,in Sweden approaching the Gulf of Bottfiriaand moreover singly recorded as far southeastas in Uppland. Owing to its wide distribution,there is no clear gap corresponding to the hiatusbetween the southern and northern part of themountain chain. The species in Finland is confinedto the northern part of the country andfurther east it extends throughout northern Russiaand Siberia to Bering Strait, and Kamchatka.The southern area of distribution is related toScotland, Western England, and disjunct rangesin the mountains of Central and SouthernEurope. Relic occurrences ate moreover recordedin Altai, Pamirs, and Tien Shan. B.lapponicus has a wider distribution than theremaining Scandinavian tundra elements and isoften mentioned as a subarctic species.

erpuecsouueC eprslno ere (99 'EIC) tantonrsod .gpue '(SL 'Elg) unrcuoq 't[ '(Vg 'Slg) wnrctotd'g '.{e4rn; ur pepJoceJ ueeq seq pue .rnury'lellv 5€lregls urer,Ilnos pue urelselA 'edornEurequoN pu" 'se1s1 IerlueJ qslllrg ur pelnq-lrlslp sr lI 'puelecl ur Euurncco serceds u(poeqt sl (29 'Elg) sry1auof 'g 'N .01, Jo r{Uou pueou1d1ol&at q luenberg osle ere tantoptd .g puesnyauof 'g ,{1uo loutdloqns o8at ol Ielel ees tuor;se8uer deullrop ureqlnos q uorlnqr4slp IecrUeArleql'N"oL }s€el le ol qlnos I'uoJJ pue }se^\ol lffie tuorg 'o'r 'elnsurued uurA?urpu€cs eJrluoetlt lnoqEnorg] pelnqrrlslp ,{leprm er€ qcrq^\ Jo[11u'utntoncsod 'g pue ownto1rcq 'g ,tunto1otd'g 'snpauof 'g 'utntoud{q 'g 'tantocn1 .g .zfl'sorcods xls Jo Ielol e sesuduroc dnorE sF{Io!ôulpuo?S Tnoq&nonfl panqlustp rQaplu salcadg.(snlcpqeg)sncyoddol 'g :g Jo lltull ureqlnos erIJ 'ruogLqeq snpaqDq .g :q !(sneeuql) snuldlo .g :3 ifqtyltr mcu'tro 'g ig:rrequoqcs snanqndrbl 'g. :V Jo uoqnqrJlsrp uerl€rrprrersorrueC eql roJ silrurl e8erele eql '96 .ErguaryoT'v 86I

Studies on Scandinavian Bumble Beesr99widely distributed in British Isles, throughoutthe European continent, and eastwards at leastto Lake Baykal. They are known from Turkey.B. pratorum and B. hortorum moreover reachthe eremial in Iran and the latter also occursin Algeria. B. lucorum (Fig. 54) has a wideEurasian distribution, outside Fennoscandia extendingthroughout Europe and ranging throughoutSiberia and adjacent southern countries eastto Manchuria, Sakhalin, and Kamchatka. It isknown from Northern Turkey, Pamirs, and Tibet.The ubiquitousspecies comprise a group ofminor zoogeographical interest except for thelatter species, viz. B. hypnorum (Fig. 61), whichshould be commented upon. Though widelydistributed throughout Fennoscandia, it displaysa great affinity to the forest. This is clearlyillustrated by its scattered occurrence in NorthernNorway, everywhere being related to theconifer forest (cf. Figs. 6l and 2), and by thefact that the species is less frequent along thewest coast than inland and not at all recordedin Gotland or the Aland islands. B. hypnorumis a Siberian fauna element, expanding the taigafar throughout Central Europe until the Channelconstitutes the barrier to the British Isles. It iswidely distributed throughoutSiberia and onthe Pacific coast ranges from Japan north toAnadyr. The species is treated by Reinig (1965,p. 131) as a sub-boreal taiga element.bulk of the Scandinavian Bombus spp. and comprisesthe fourteen southern species in additionto the six ubiquitous ones. A decline in numberof species and a change in fauna elements ongoing north is to be expected, but as many as14 species occur in northernmost Norway, i.e.the counties Troms and Finnmark, and reflectthe fairly favourable living conditions that farnorth. In addition to the ubiquitous species andthe southern B. soroeensis, the five tundra elements,viz. B. hyperboreus, B. arcticus, B. balteattts,B. alpinus, B. lapponicus, and the taigaspecies B. sporadicus and B. cingulatus, ate represented.Flowever, the fauna in regio arcticais limited to the five tundra elements besidesthe ubiquitous B. jonellus and B. pratorum, acomposition corresponding to that of regioalpina in Southern Norway.Of particular interest is the zonation along thewest coast of Southern Norway being included inregio quercina (Fig. 2). A total of twelve speciesis represented, viz. all the ubiquitous species inaddition to B. muscoruffi, B. magnus, B. lapidarius,B. soroeensis, B. wurfleini and B. lapponicus.With exception of the two latter, they are southernspecies and defend the rather awkward useof regio quercina applied to the west coast.However, occurrence of the tundra element B.lapponicus and the fact that B. muscorum isrepresented by the atlantic subspecies B. nt.liepetterseni clearly distinguish this northern partof regio quercina from the more typical southernfauna in the remaining part of this region.REMARKS ON THE SCANDINAVIANFAUNAThe zoogeographical survey above articulatesthe mixture of elements constituting the compositionof the Scandinavian bumble bee fauna.It reflects the great variability in ecologicalconditions, the contrast in topography and climate,and the position of the peninsula,extending frorn 55o20' to 7loll' N and formingthe northwest corner of the Eurasiancontinent. Not only the various fauna elementsbut also the number of species constitute thefaunal composition. A total of twenty speciesrepresented in Southern Sweden constitutes theIMMIGRATIONROUTES OFSCANDINAVIAN BUMBLE BEESThe quatern ary glaciations of Northeast Europeexterminated the flora and fauna of Scandinavia,and, in the geological time scale, plants andanimals now distributed throughout the peninsulaare young immigrants. Thus it should beeasier to sllrvey the natural history of Scandinaviathan that of other parts of the worldthe main problem being related to the biologicaleffect of the last glaciation (Wtirm), timing,

eql'serceds eurdle-olJre pue eurdle-oerog (g)'[IeA\ sB lsee pue qlnos tuo4 uer^surpuecspopelul seq tan.ton)snd'g pelngrJlslp ,{1epnroqt 'e1et {ue 1y 'lecr}og1od,{q Jer.{teJ uorlseEEnssF{} e{etu pueprC Jo }s€e suor}elndod }noqeuorleuJoJul Jo {cel'JeAoA\oH 'suor}ceJrp e}rsoddour suorlerJel l€urlc 8urlueserd suorlelndod ,(qper1oeoJ uoeq ener{ ue{} plnoc s€eJe EurperEJelureql'lseequou eql tuoJJ ,{emro51 }soruuJequouEurqceer snuotqtlws 'd 'g pue 'eruqlofl Jo JIncpunoJ lseo uro4 snuna.t"tods 'd 'g 'qlnos ruo4erneurpuecs Eurpenur satcotoptllod 'd 'g :selnorledrcurrd eerql tuo4 uorselur IerculE-lsod lseEEnsol uoseeJ entE lqEFu gcSIA\ 'puelureur uerneu-rpuecs eql ol peuguoJ Eureq sercedsqns eerqlsu fueur se sel€crput 'suorlerJel leurlc uo peseq'utntoncsod'gJo uorlenlene cgrcedsqns'lseeeql tuo4 elnsurued oql peqouer tqE[ elod-eueEtlllA\ uorlelndod reglou€ elrr4^\ q}nos tuo4 erleu-rpuecs pepelur en€r1 l{Erul >lrep elod-eueE q}rznuorlelndod e WLll pelseE8ns sl 1I 'erleurpuecs uJe-LIuoN uI polJosqo 11e ]ou eJe sueurrceds crueloruolF{r!\ ,(eAro51 uJoq}nos q lusrueletu 01 ^(cuepuelpocunouord e s,(eldsrp serceds eql'unto|,tot1 'gJo lueruteen eql ut peuorluetu sv 'tuntoncsod 'gpue unnt"toq 'g sdeqred ere suorldecxg'suorlel-ndod Iecrtuepr se peleerl ere pue 'peqsrnEull-sp ,{gectdfloueqd }ou lereueE ur ete suorlelnd-od qsruurC pue uerl€urpuecs eql'suorlelndodqsruur{ pue qsryea\S eq} ueeaUeq de? + e eqilps .(eur ereql 'smnp1do1 'g EulpreEeg 'eruqlogJo JInC eqt punor uepeas ol relo Eurpeerdssuorlelndod qsruurC ,(q letu eJea\ suorlelndodeql 'uelloqrroN ol lseor4gou Eurqceordde uO'Wflos eql tuor; erleurpueJs pepelq }srg serceds, esoql 'utntoncsnd 'g pue wnnryo7'g 'tan,toto,td'g 'sn11auo['g 'utntoud,fil 'g 'tantocnl 'g '4L'sercods snolrnblqn eql dlleuorllppu pue (f-C fO'EIC) srsuaaotos 'g pue snpuanSutlstp 'g 'snrto"t-apwg ,smnptdol .g .zll €ruqlog Jo JInC Jotluou Euruunr slprll urorllrou eql qlla sercedspecuelpe ..{peqpou eqt sureluoc dnorEqns sFII'puelurc JeAo ls€e tUoJJ pue {Je(uuoc JeAo qlnosruo4'zIL 'suorlcerrp oJoru Jo ol[l ruo4 erleu-rpuecs pepelur oler{ qcFllvr serceds'e'r '(6V61tllorpulf) uollerEurull .elqnop, rlll^\ sercedg (7)'uorlcnpoJlur eJoqcodorqlue Ielueproceol enp eq l{Erur prooer qsrpe^\S elEurs eq}se '1n;tqnop sr 'tan"totuod 'g 'z!L 'serceds qluele Jo uorseluJ '(g-V V6 'sEtg) slllwnt! 'g pueuntull,ft 'g 'snauo"tntqns 'g 's1"t7sailal'g 'snuSotu'g'utntocsnut'Userceds uJoqlnos pecueApe,(lreqUou eql pu€ (t6 'EIg) snun,tata^'g'sn1oapnt't[ 'snuoutnnugsercods qsrpel\s-q]nos oueJ]-xe eql 'zIL 'serceds ouru Jo lelol e sesudurocdnorEqns egl'qlnos Jer,{UnJ Euruunr Jo eruqtogJo JInD eql Jo puo uroqtnos oql Eurssorc stprlluJequou eql gllzn 'e'r 'erêurpuecs uJerllnos olpeuguoc uortnqlrlslp luecer qllÂ sercedg (t):seuoEeleJeerql olq pep$lpqns eq uec ereq pepnlcurserceds eql'uepel\S uJer,Ilnos ol {reruuo6l JeAodlureur 'qlnos ruo4 erleurpueos Eurpeurur serc-eds seceJqrue sluerElulull uJeqlnos tuJe] eqlstuo.t8rtatur unqruos'sreldeqc snornerd ur dn petutuns uorler,uJoJullecqderEoeEooz uo peseq sr 'mo1eq pol€crpur.(1e.r.r1eluel selnoJ uorlerEprurl ledrcurrd ;o ^/ylorleql'eur dq pelndsrp oJe/( slcrleJ lercelErelulJo serJoeql'snultqosaoJ'g pue'sn)ntoddnl'g'rutagftnu 'g Jo ]uetu]eoJ] eq] uI'r(1uo slueru-ele lercelE-lsod fq pepunoduroJ sr €uneJ luecoreqt l€t{} se}uolpul 'dds snqwog uelleulpumseql Jo urelled Ieuorlnqlrlslp eqt'n11e1 eqtol sV 'pesserls ueeq eleq ,fto1sq IeuneJ pueuorlnqlrlslp sll EurureloE sturelc 1ec6o1oce eql'uorleEllselur Jer1UnJ JoJ srseq e se elJes ol'ddssnEaog uerleurpuecs Jo uorlnqlJtslp pue 6suoll-errel crdr(loueqd 'rtruouoxel uo slceJ epduocol ueoq seq >Irozlr lueserd eql Jo edocs eql'GgAI qlorpulD pessncslp ueeqserl '^(1uo .poued ser{rq reEunoa, oqt qEnorqllelrlrns ep€cep lsed eql Eurrnq'(Zget'6V61rlloJpulT 'JJ) ,{1e,rr1cedser slstEoloeE pue sreqderE-olq ,{q pe}nJeJ pue 'perno,re; 'pelseEEns ueeqeûg z(eznro51 go lseoc }so1v\ oql uo seEnJer eeJJ-ecl ur urJn1\ lnoqEnoJql IeAIAJnS uo selJoeql'uorselurlercep-lsod ,{q peureldxe eq }ouuecsl€ruru€ pue s1ue1d go reqlunu e Jo uol]nqlr]slpluoceJ l€r{} ?ceJ oqt o} Eutrvro 'slo,tllJns Ielc-eIE JoJ serlrlrqrssod Uel Jo €rAeulpueJs peJeloc,t1e1e1druoJ uorlercelE 1se1 oql JeqleqlA o1 se ueslrseq uorlsenb eql odrnluec lsel eql Jo pue eqlecurs 'uorleJErurur lercelE-1sod yo uollceJlp pueua4o7'v 002

Studies on Scandinavian Bumble Bees 201boreo-alpine B. wurfleini and the arcto-alpineB. alpinus and the subspecies ,8. lapponicus scan'dinavicus refer to this subgroup. It seems likelythat they lived at the edge of the ice during thelast glaciation and when the ice retreated theyfollowed northwards, i.e. they are southern postglacialimmigrants. The earlier faunal historyof these species is not commented upon here.Eastern immigrantsThe term eastern immigrants of Lindroth(1949) is used for all species in post-glacial timeinvading Scandinavia east of the Baltic Sea.Thus the species mentioned as southern immigrantsunder the heading 'double' immigration,ate eastern immigrants as regards the invasionover Finland. Otherwise eastern immigrants referto the tundra elements,B. hyperboreus, B. arcticus,B. balteatus and B. l. lapponicus (Fig. 96 A-B,D-E) and to the taiga elements, viz. B. sporadicus,B. ctngulatus and B. consobrinus (Fig. 95). Inthe treatment of B. balteatus, attention is directedto a possible double immigration, based on thegreat phenotypical variability including a pronouncedtendency to melanism in Southern Norwayyet mentioned only as a speculativetheory. F{owever, double immigration of B. lapponicus,represented by two subspecies (Fig. 63),is obvious. The nominate subspecies is a northeasternimmigrant while .8. l. scandinavicus is asouthern invader and consequently is includedabove as a southern immigrant.B. consobrinus (Fies. 73, 95 C) is the onlypuzzling member of the Scandinavian bumblebeesowing to its recent disjunct distribution inFennoscandia, presenting a wide gap in Finland.Clinal variations along an immigrationroute extending from Eastern Asia, throughoutthe taiga in Siberia, and west into Scandinavia,is mentioned in the treatment of the species.The very close morphological and phenotypicalrelationship between the separated populationsin Scandinavia, Finland, and adjacent part ofLJ.S.S.R., emphasize their subspecific identityand gives reason to surmise a former continuousdistribution in post -glacial time.ACKNOWLEDGMENTSI am under heavy obligation to a number ofpersons, institutions and supporting agenciesthat have made this study possible.I owe the late Ove Meidell deep gratitude forintroducing me to apidological research. Stimulatingdiscussions shortly before his early deathin 1942 and access to his numerous unpublishednotes, some of which are included here, havecontributed greatly to my general knowledge ofthe subject.I am very much obliged to Dr. G. Kruseman,Professor O W. Richards and Dr. I. H.I{. Yarrowfor profitable discussions on taxonomic andnomenclatural problems, for confirming my identificationsof some abercant specimens, and otherwiseproviding me with information about typesand classical collections. As the work was writtendirectly in English, Professor Richards andDr. Yarrowkindly corrected the language inthe major part of the manuscript. The latter alsogenerously allowed me to refer to his unpublishedrecords whenever they contributed to the worlddistribution of Bombus spp. treated here.I am especially grateful to Dr. K. Ander forhis never-failing interest in this study. His effortsto provide me with detailed information aboutold and recent collections, with permission toinclude his unpublished records, greatly helpedto complete the Swedish part of this work.I have profited much by communication withthe following professors and doctores regardingcertain phyto- and zoogeographical aspects and/or locating of types, interpretations of labelsin classical collections, etc.: A. M. Antonova,Moscow; R. Y. Berg, Oslo; R. Delmas, Monpellier,France; S. Erlandsson, Stockholm; K-H.Forsslund, Stockholm; H. Kauri, Bergen; thelate N. Knaben, Oslo; C. H. Lindroth, Lund;H. E. Milliron, Ottawa; H. Olsen, Bergen;D. V. Panfilov, Moscow; B. Petersen, Copenhagen;T. Soot-Ryofl, Oslo; A. Strand, Oslo;B. Tjeder, Lund; B. Tkalcu, Prague.Professor K. Fregri kindly made a few pollenanalyses.I am most grateful to Mrs. Inger MeidellTambs-Lyche for her untiring efforts as assistantin the continuous and tedious field work

luenbesqns oql'uepoAtg ,(rlunoc oruoq sFI ruor;'dds snqwog plel uel peqrJcsep ogrlr,sneeuullqlll\ EuruurEeq 'ue,rF sl zlrerleJ lecrJolsFl v.I(uepezrg puefemrop - ) erneurpuecs q snquog snuoE eql Josrsdou^(s e lueserd ol sr ,(pn1s sFIt Jo urle eqlAIIVruWNS'peqs[qnd ueeq eêqJeleu sdeqred plnoA\ 'setull I€JeAes peldnuelurueeq sBrI suosueJ snorJBA JoJ r{clql\,>llozvr luo-serd eql lueueEernocue sq 'ldrrcsnuerulnoqll1\eql perro.rdulr e^€q qclq^\ rusrcrlrJc pue ,ecrzrpe'suorssnoslp Eurlelnulls JoJ rJne) .H JossoJoJd olseo8 epnlrl€rE lserecurs ,(u lseel lou lng lseT's{u€rl} InJo}eJE ,(ru sserdxe I suol}n}r}sureseql oI'pgrsEurDISJoC eErlede{suolrluoullyseEroSl ,{q pue ueEreg go ,(lrsrenpn oq} ruo4spunJ ,{q JoJ pled ,(1ged oJo^&\'e1ep go uor}ecg-lpoc ol peleleJ sesuodxe pue 'suorlnlllsur u8rerogur dels dppuoC sEuu4rsro;sElDleftA S/Vdq pegoddns ,(gercueug se^\ {ro^\ pleg eql'EreqsEg1\ IulN'srl tr pu€ ue,{oqcg 1uu,Cl sslntr'uespnu;F{epuesol{ ppquEeg 'srtr tr'uesqoceg'H lpelq'srl tr'sueErl uellg sslntr'oqsEneH uef'JtrAI 'uesserlg slly'srtr ['ueslueJefl reEul.sJI tr o]pue 'seusletutuo6l lesls sslntr ol enp ,{llercedse eJequeql ^(ur 'ecuelsrsse lmruqce] snorJel JoC'pepoc,{11enperE erezuepp eql se pelnduoc pue peqcund lerreletu eqlJo t"l:ed uelEervuoN erll EurlteE ur dleq luerledJoJ 'ueEreg odlrsrenp,O oql te eJlueJ relndurocoql Jo ro8eueul 'pueslolC ') ol pelqepur rue I'uopuoT'(.fio1sr11 1ern1e5f tunesntrAI qsrlrJfl '.urouea'H 'H 'IiEroqelog 'runesn141 €{srro}sr,Irn}eNErogelog 'u?ple1A 'Hipueplg 'ueurlueurgH6elre{le1 'E ! ue8uquedo3 'urnesn141 4srEo1oo7'uesre1e4'g pue uexnl .-I .s i{lutpg repefu .J,: oQy orurepe{V oqy 's}[n{S '11 iureqpuo{ '1oos-nIN'qe>1s1es srege{sueprn eErleEuo;'ueslrenrg'E iolsg'runrlolercqe-I >1srEo1oo7'uossu€rlof-ques 'V! pe"lEupe'I 'ecuercg Jo ,(ulepecyoeleJeulouod 'V elel eqt lelesddn 'rEo1o1edtxg1Jo.J ueuorlnlllsq 'uosspuuersso 'C i olsg'urnes-nIAI 'reurureqelllT 'V 4srEo1oo7 pue Ern1e51')I'T I punl 'ueuorlnlrlsuJ e>lsrEol ooz'uossJepuv'Ff pue qlorpurf 'Ff 'C ielesddn .uossrel .H: oqy'srsue4Jnl 'AIun 'ulncrE olooz unesntrAJ'seredllqef ') 'E ! urlreg 'runesnlN soqcs6olooT'r1 etflolooT gerzeds 'J elnlrlsul .uuer,us81ugy'E! uleprelsurv 'urnesntr41 seqcstE olgoz .ueur-esnr) 'g !'C'C[ uofuryse1\ .runesntrAlI€uol]eN'S',O'ureqruoJ)'A'Xiulloq4cols,urnesnws{lg">lsrJo]sil1Jn]eN 'uossle4 'I 'd pue Jepuufief;tr'E elel eqt ielesddn ueuor]n]l]sq e4srEol ooz'uossuol 'S ! reEuenelg ournesnl4l reEueurulg .ues-reE1o11'H i r4uls1e11 '1eesntr41e>lslEol ooz .uouu-€{{ed 'V pue uetu{3uH 'A\ ! euuern .urnesn141seqcsrJolsrrlJnleN'reqcsrg'tr^[ !puuprg,ordny'EupglE 'U elel eql ! osurorl 'urnesn141 osruor1'uosuBrlslJqf 'giqetn'ueEo1 ..,rlcqcJeeseulectEoloruolug 'V'61'S'n 'geqog .E .D : suoll-nlrlsur JIerIl ur ,(els ,(ul Euunp serl{rceJ qll,t\otu peprlord pue ,(pn1s sFIl ol lueulUed suorlcel-[oc eunuuxe o] otu pe]]plred osrly\Jeq]o Jo ueoluo peceld ,{lsnoreueE oq,!\ ser}rJoq}n€ Eurmolloyeqt ol pepuelxe sr uor]€rcerdde eJecurs .(tr41'plo^lso 'f T'C \mtEg 'I .ueprgEep7 .S'rnelen 'd 'pueleuual;- 'V 'eqc,{1-sqtuel .H'H 'guepuos 'A'lpung 'E 'reurrolg .d 6uelolsT'E 'e11e,re1o5'O 'uertg-]oos 'H ,,(oq1og .I'lepulelos 'd 'elpueg 'D 'ueprenr{11'1 ,uesJelled'U 'V 'uelng 'I 'urreqrg 'ru .V .uessp51T'H 'dxe lnocs Jo ropeel oqloues1er51 .I .VT'qcta1A1 'V '1snqep1,(6 'lI-Sry 'lS 'd A O'unllpl6 'g'guuo1'V'punT 'I'rqnT .C .C ,sno1'C 'J '4rnro1 'a 'ues rc-I'V 'efieqEuel .1 .ure1s-JeA) 'n 'rrr,{r) 'H 'reEnr;1 'C 'd 'uespnu)Ir{ep-uesoll 'U 'uesuel eAeJC '-I'uesuel 'g .)T V''Ie le soruef 'C ',(qesn1tr'f 'oqsEne11 .f .IA'ues>1e1lng'f o'Ie le e{spoC 'C e}e[ eq],11on-srefg 'O 'u?plrC 'C 'gsplelC 'f T y 'EreqllefC.V'uelepplelC 'f 'pleqcneC 'y 'ues{lrg '4 'uessellg'v 'eplg 'V 'selEnoq 'W 'f 'sousJetutuo( .S'pue11eq 'I 'erceq '1 'uesuerlsrJrl3 'E 'luqr(rg .I'>lcurrg 'd 'punl4rsfg 'U ') 'uosueg .U eluleql pue 'f 'e{{efl'V 'repuy ') 'uesgly 'A '[ep-uolv 'E : peleErlsonur esrl\reqlo lou seere ruor;'fprcd'1eue1eru qlrl\ etu perlddns ,(lsnoreueE oqzlrsroleroqelloc Eurznollog et{t {ueq} o} r{sl,!\ I'J1e 'se1ou Euldr(1 pue Eut8ueJJe 'sueurrcedsSurrederd fq elqelreê fiepre6'O pueqsnr1 req Jolerreleru reô-tJel eqt epetu osle oqs 'peuueldf11eut8rro ue{} IerJe}etu qcJeeseJ re8rel ,{1qere-prsuoc e q Eurllnsot'19-L561 sree.( eql Eurrnpua4o7 'v z0z

Studies on Scandinavian Bumble Bees 203pioneer contributions to the knowledge of theScandinavian bumble bee fauna are briefly summarized.The topography and climate in Scandinaviaare then shortly commented upon, followedby a simple regional division based onecological conditions and expressed by treelimits.2. The main areas investigated by me duringthe years 1939-1969 with several years of intervalsare reviewed, and the museum and privatecollections under revision are listed. About 56,7A0specimens were examined altogether.3. Procedures related to the treatment ofbibliography, taxonomy and distribution of theScandinavian species are described.4. Classification is commented upon and thestrong homogeneity in the subgenera of Bombusis in particular emphasized.5. Keys to the Scandinavian species of Bombusare given.6. The main part of this work is devoted tothe study of the Scandinavian species, arrangedby subgenera. An attempt is made to providevalid designations and synonyms with informationabout the type and the typ e area. The treatmentof each species deals with the following items(some of which are for convenience assigned tothe actual subgenus or subspecies): List of synonymsalso including bibliographic references tothe Scandinavian fauna; first record in Norwayand Sweden respectively; nomenclatural andlortaxonomical remarks, e.g. listing named infrasubspecificforms; diagnostic descriptions (morphologicalcharacters, colour pattern , variations);distribution in Norway and Sweden respectively,showed by dot maps, and detailed list of localities;distribution in Finland and general occurrencethroughout the world; biology. In severalspecies, phenotypic variations, distribution, andbiology are subject to remarks or discussion.A total of 29 species, divided into 34 subspecies,is recognized, of which 25 species (28 subspecies)occur in Norway, while all the species (33 subspecies)are recorded in Sweden.The study of the subgenus Bombus s.s. supportsspecific recognition of the critical taxonB. magnus. The male of this species is, however,not distinguished from the male B. lucorotm.The subgenus is represented by four species, viz.B. lucorum, B. magnus, B. sporadicus and B.terrestrus of which the latter is new to Norway.The subgenus Pyrobombus is represented byfive species, of which B. lapponicus is divisibleinto two subspecies, viz. B. l. lapponicus andB. l. scandinavicus. As a result of a phenotypicstudy it is stated that the distribution of thenominate subspecies extends west to northeasternmostSweden and Norway. Regarding the rapidcycle of ,8. jonellus, B. lapponicus and B. pratorumthe seasonal length is noteworthy, lasting to thebeginning or end of September. In addition tolate-emerging queens lengthening the season, itis indicated that under favourable conditions ,8.lapponicus and B. pratorum possibly producetwo generations, as recently suggested for B.jonellus.A total of four species is distinguished in thearctic subgenus Alpinobombu.s. The conspecificityof the Canadian B. q. arcticus and the ScandinavianB. a. diabolicus ( - B. alpinifurmis) isevaluated and maintained, i.e. a specific rankingof the latter is refuted. The unstable colouring,including melaniSffi, of -8. balteatus is studied.It is concluded that the gre at range of variationsare not geographically segregated except forthose related to melanism. As to the latter, thefrequency of recorded melanics, being high inmountainous Southern Norway, is markedlyreduced on going north; a tendency to melanismis in fdct not observed at all in northernmostNorway. The theory of arctic species beingadapted to solitary living is moreover disputed.A great number of collected workers, and recordsof colonies, prove that B. alpinus, B. arcticusand B. balteatus regularly produce a workercaste. Regarding B. hyperboreus, the revisionreveals no reliable records of colonies in itsentire geographical range. Although few in number,workers have, however, been observed inScandinavia and further east. Recent observationsof B. hyperboreus usurping colonies ofB. arcticus in the western hemisphere and acolony of B. jonellus in Scandinavia indicatethat this high-arctic species is a facultative inquilinerather than (even temporarily) adaptedto solitary living.The subgenus Megabombus is represented bythree species. The information acquired about

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Studies on Scandinavian Bumble Bees 205\roo\r,o-ej'trta.lr2,r,/ atat'IItttI tl a a 'r IRURAL DISTRICTS AND CITIES IN NORWAYThe numbers below refer to official index number of the ruraldistricts and cities. The two first figures correspond to the indexnumber of the county and the two last to the index of the ruraldistrict or the city. As the two first figures are common for alldistricts within a county, only the two last are used in Figs. 92,98, to save space. The suffixes (i), (y), (n), (s), (v), (o) refer tothe inner, outer, northern, southern, western and eastern districtot the county cf. Fig. 99.S9--9!ttn'!Ef12o\r.'1 - a -'r-'.-"33'50km18 Nordland countytgOa Bod" (,1805 Narvik (o)1811 Bindal (y)1814 Bronnoy (y)1815 Veea (y)1816 Vevelstad (y)1818 Heroy (y)1820 Alstahaug (v)1822 Leirfjord (y)1824 Vefsn (i)1825 Grane (i)1826 Hattfjelldal (i)1827 Dsnna (y)1828 Nesna (y)1832 Hemnes (i)1833 Rana (i)1834 Lursy (y)1835 Trrena (y)1836 Rsdoy (y)1837 Melsy (y)1838 Gildeskil (y)1839 Beiarn (i)1840 Saltdal (i)1841 Fauske (i)1842 Skjerstad (i)1845 Ssrfold (o)1848 Steieen (s)1849 Hamaray (a)1850 Tysfjord (o)1851 Lsdingen (v)1852 Tjeldsund (s)1853 Evenes (s)1854 Ballangen (o)1855 Ankenes (s)' ," 1938 Lyngen (i)1856 Rsst (v) 1939 Storfjord (i)1857 Vrersy (v) 1940 K&fjord (i)1858 Moskenes (v) 1941 Skjervay (y)1860 Vestvigoy (v) 1942 Nordreisa (i)1865 Vflean (v) 1943 Kvenangen (i)1866 Hadsel (v)1867 Bo (v)1868 Oksnes (v)1870 Sortland (v)1871 Andsy (v)19 Troms county.1901 Harstad (y)1902 Tromso (y)1911 Kvafj'ord (v)1913 Skenland (i)1915 Bjarkoy (y)l9l7 lbestad (y)l9l9 Grataqgen (i)1921 Salangen (i)1922 Bardu (i)1924 Milselv (i)1925 Sarreisa (i)1926 Dyray (y)1927 Tranoy (y)1928 Torsken (y)1929 Berg (y),193'I Lenvik (y)1933 Balsfjord (i)1936 Karlssy (y)20 Finnmark county2OW Hammerfest (v)2002 Vardo (n):,2003 Vadso (n)'t011 Kautokeino (i)2Ol2 Alta (i)2Ot4 Loppa (v)2Ol5 Hasvik (v)20rc Soroysund (v)2017 I(valsund (v)2018 Missy (v)2Ol9 Nordkapp (v)2O2O Porsanger (n)2O2L Karasjok (i)2022 Lebesby (n)2023 Gamvik (n)2024 Berlevig (n)2025 Tana (n)2027 Nesseby (n)2028 Bitsfjord (n)2O3O Ssr-Varanger (o)Fig. gT.Division of Northern Norway. Names and limits dated to 1969. O-O.14. Bumble Bees.

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Studies on Scandinavian Bumble Bees 207RURAL DISTRICTS ANDThe numbers below refer to officiolCITIES IN NORWAYindex number of the rurol districts ond cities.The iwo first figures correspond to the index number of the county ond the two lostto the index of the rurol district or the city. As the two first figures ore commonfor cf l districts within o county, only the two lost ore used in Figs. 97, gg, to sovespo.e. The suffices (i), (y), (n), G), (u), (O) refer to the inner,outer, northern,soutliern, western ond eostem district ofthe counly cf Fig, 99.0l Ostfold county0l0l Holden0102 Sorpsborg0103 FredrikstodOlM Moss0l I I Hvoler0l l 3 Borge0l 14 Vorteig01 l5 Skieberg0l l8 Aremork0l l9 Morker0l2l Rgmskog0122 Trggstod0123 Spydeberg0124 AskimCl25 Eidsberg0127 Skiptvet0128 Rokkestod0130 Tune0l3l Rolvsgy0133 Kr6kergy0134 Onsgy0135 R6de0136 Rysse0137 V6ler0138 Hobgl02 Akershus counfy021I Vestby02r3 skioztl At.0215 Frogn0216 Nesodden0217 Oppeg6rd0219 Bcerum0220 Asker0221 Aurskog-Hglond0226 Sgrum0227 Fet0228 Relingen0229 Enebokk0230 Lgenskog0231 Skedsmo0233 Nittedol0234 Gierdrum0235 Ullensoker0236 Nes0237 Eidsvoll0238 Nonnestod0239 Hurdol0301 OsloO4 Hednnrk county0401 Homor (s)0402 Kongsvinger (s)0412 Ringoker (s)0414 Vons (s)0415 Lgten (s)0417 Stonge (s)O4l8 Nord-Odol (s)0419 Sor-Odol (s)0420 Eidskos (s)0423 Grue G)0425 Asnes (s)M26 V6ler (s)0427 Elverum (s)0428 Trysil (n)O4zS Amot (n)0430 Stor-Elvdol (n)0432 Rendolen (n)0434 Engerdol (n)0435 Tolsq-Os (n)0437 Tynset (n)0438 Alvdol (n)0439 Folldol (n)05 Opplond county0501 Lillehommer (s)0502 Giovik (s)051 I Dovre (n)0512 Lesio (n)0513 Ski6k (n)0514 Lom (n)0515 Vdsd (n)0517 Sel (n)0518 Fron (n)0520 Rinsebu (s)0521 @yer (s)0522 Gousdol (s)0528 Qstre Toien (s)0529 Vesire Toten (s)0532 Jevnoker (s)0533 Lunner (s)0534 Gron (s)0536 Sgndre Lond G)0538 Nordre Lond (s)0540 Sgr-Aurdol (s)0541 Etnedol (s)0542 Nord-Aurdol (s)0543 Vestre Slidre (n)0544 Qystre Slidre (n)0515 Vong (n)06 Buskerud county0601 Rinserike (9)0602 Dommen (g)0604 Kongsberg (g)0615 Fl6 (v)0616 Nes (v)0617 Gol (v)0618 Hemsedol (v)ootr At 1u;0620 Hol (v)0621 Sisdol (v)0622 Krgdsherod (9)0623 Modum (p)0624 Qvre Eiker (9)0625 Nedre Eiker (g)0626 Lier (9)0627 Rpyken (9)0628 Hurum (9)0631 Flesbers (9)0632 Rollos (v)0633 Nore og Uvdol (v)07 Vestfold county0702 Holmestrond0703 Horten0705 Tgnsberg0706 Sondefiord0707 Lorvik0708 Stovem071I Svelvik0713 SondeQ714 Hof0716 V6le0717 Borre0718 Rormes0719 Andebu0720 Srokke0721 Sem0722 Ngttergy0723 figne0725 Tiglling0726 Brunlones0727 Hedrvm0728 larcrrl08 Telemork county0805 Porsgrunn (y)0806 Skien (y)0807 Notodden (i)081 I Si I ion (y)0814 Bomble (y)0815 Kroserg (y)0817 Drongedol (y)0819 Nome (y)0821 Bp (i)0822 Souherod (i)0826 Tinn (i)0827 Hio*dol (i)0828 Seliord (i)0829 Kviteseid (i)0830 Nissedol (i)0831 Fyresdol (i)0833 Tokke (i)0834 Vinie (i)09 Aust-Agder counly0901 Risgr (y)0903 Arendol (y)09Ol Grimtod (y)09ll Gierstod (y)0912 Ves6rchei (y)0914 fvedestrond (y)0918 l/olond (y)0919 Frolond (y)0920 @yestod (y)0921 Tromgy (y)0922 Hisgy Q)0923 Fiere (y)0924 Londvik (y)0926 Lillesond (y)0928 Birkenes (y)0929 Amli (i)0935 lvelond (y)0937 Evie og Hornnes (i)'0938 Byslond (i)0940 Volle (i)0%l Bykle (i)l0 Vest-Agder countyl00l Kristionsond (y)1002 A/ondol (y)1003 Forsund (y)1004 Flekkefiord (y)l014 Venneslo (y)l0l7 Songdolen (y)l0l8 59sne (y)l02l Momordol (y)1026 Aserol (i)1027 Audnedol (y)1029 Lindesnes (y)1032 Lynedol (y)1034 Hegebostod (i)1037 Kvinesdol (i)1046 Sirdol (i)I I Rogolond countyll0l Eigersund (y)I 102 Sondnes (y)I 103 Stovonger (y)I 106 Hougesund (y)llllSokndol (y)lll2 Lund (y)ll14 Bierkreim (y)ll19 Hd (y)ll20 Klepp (y)l12l Time (y)I122 Giesdol (y)I124 soto (y)I 127 Rondoberg (y)I129 Forsond (i)I I30 Strond (y)I133 Hielmelond (i)I 134 Suldol (i)I 135 Soudo (i)ll4l Finngy (y)I 142 Rennespy (y)I144 Kvihgy (y)1145 Bokn (y)I146 TpverI149 Kormgy (y)ll5l Uhiro (y)I154 Vindofiord (y)l2 Hordolond county12ll Etne (i)l214 @len (y)12l6 Sveio (y)l2l9 Bgmlo (y)l22l Stord1222 Fitior (y)1223 Tysnes (y)1224 Kvinnhercd (i)1227 )mdol (il1228 Oddo (i)1239 Ullensvong (i)1233 Ulvik (i)1234 Gronvin (i)1235 Voss (i).1238 Kvom (i)'1241 Fuso (y)1242 Somnonger (y)1243 Os (y)1244 Austevoll (y)1245 Sund (y)1246 Fiell (y)1247 Ask|y (y)1248 Loksevds (y)1249 Fona (y)1250 Amo (y)l25l Voksdol (y)1252 Modolen (y)1253 Ostergy (y)tzss Asone (y)1256 Melond (y)1259 @ysorden(y)1260 Rodgy (y)1263 Lindas (y)1264 Austrheim (y)1265 Fedie (y)1266 lvlosfiorden (y)l30l Bergen (y)14 Sogn og Fiordonecountyl40l Floro (y)l4ll Gulen (y)t4t2 stluna (y)14l3 Hyllestod (y)1416 Hgyonger (y)14r7 vik (i)14l8 Bolestrond (i)14l9 Leikonger (i)1420 Sogndol (i)l42l Aurlcnd (i)1422 Lcrdol (i)t424 Araol (i)1426 Luster (i)1428 Askvoll (y)1429 Fioler (y)1430 Goulor (y)l43l Jglster (y)1432 Fgrde (y)1433 Noustdol (y)1438 Bremonger (y)1439 V6ss9y (y)l.t4l Selie (y)1443 Eid (y)1445 Gloppen (y)1,148 Stryn (i)l5 r\&re og Romdolcountyl50l Alesund (y)'1502 lvlolde (y)1503 Kristionsund (y)l5l I Vonylven (y)l5l4 Sonde (y)l515 Her6y (y)l5l6 Ulstein (y)l517 Horeid (y)l5l9 Voldo (y)1520 @rcro (i)1524 Norddol (i)1525 Stron& (i)1527 @rckos (y)1528 Sykkylven (y)1532 Giske (y)1534 Horom (y)J535 Vestnes (y)I539 Rounn (i)1543 Nesser (i)1545 Midsund (y)1546 Sondgy'(y)1547 Aukro (y)1548 Fr

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Studies on Scandinavian Bumble Bees209Abbreviations of countiesNorvegiaAA :Aust-Agder O :AK : Akershus (incl. Oslo) R :B : Buskerud SF :F : Finnmark ST :HE : Hedmark TE :HO : Hordaland (incl. Bergen) TR :MR : Msre og Romsdal VA :N : Nordland VE :NT :Nord-Trondelag O :The suffixes (small letters) mean:i : indre (inner) n : nordre (northern) vy : ytre (outer) s : Soro (southern) oOpplandRogalandSogn og FjordaneSsr-TrsndelagTelemarkTromsVest-AgderVestfoldOstfold: vestre (western): OStre (eastern)Fennoscandia orientalisAb : Regio aboensis LkAl : Alandia LmIk : Isthmus karelicus LpKa : Karelia australis LpsKb : Karelia borealis LtKk : Karelia keretina LvKl : Karelia ladogensis NKol : Karelia olenetsensis OaKon : Karelia onegensis ObKpoc : Karelia pomorica occidentalisOkKpor : Karelia pomorica orientalis OmKs : Kuusamo SaKton : Karelia transonegensis SbLe : Lapponia enontekiensis StLi : Lapponia inarensis TaLim : Lapponia Imandrae Tb: Lapponia kemensis: Lapponia murmanica: Lapponia ponojensis: Lapponia petsamoensis: Lapponia tulomensis: Lapponia Varsugae: Nylandia: Ostrobottnia australis: Ostrobottnia borealis: Ostrobottnia kajanensis: Ostrobottnia media: Savonia australis: Savonia borealis: Satakunta: Tavastia australis: Tavastia borealisSueciaBl. : Blekinge Jmt. : JdmtlandBoh. : Bohusliin Lu.Lpm. : Lule LappmarkDlr. : Dalarne Ly.Lpm. : Lycksele LappmarkDlsl. : Dalsland Med. : MedelpadG.Sand. : Gotska Sandon Nb. : NorrbottenGstr. : Gdstrikland Nrk. : NiirkeGtl. : Gotland P.Lpm. : Pite LappmarkHall. : Halland Sdm. : SodermanlandHls. : Hdlsingland Sk. : Skine (Scania)Hrj. : Hdrjedalen Sm. : SmilandLapland (Lpl.) is divided into five Lappmarks.T.Lpm. : Torne LappmarkUpl. : UpplandVb. : VdsterbottenVg. : VdstergotlandVrm. : VirmlandVstm. : VdstmanlandAng. : AngermanlandAs.Lpm. : Asele LappmarkOg. : OstergotlandOt. : OlandI : JyllandDania2 : the main islands3 : Bornholm

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Studies on Scandinavian Bumble Bees2rlen mindre kiind art i vflr fauna. Ent. Tidskr. 71,20-26.Erlandsson, S. 1951. Drag ur vera humlors och binsutbredning. Friltbiologen 5, 36-39.Erlandsson, S. 1953. Bombus agrorum F. ssp. gotlandicusn. ssp. (Hym.). Ent. Tidskr. 74, 76-80.Erlandsson, S. 1960. Notes on Hymenoptera. I. Investigationsof the bee-fauna in south-eastern Sweden.Ent. Tidskr. 81, 123-130.Esmark, L. 1930. Zoologiske bemprkninger paa enreise i Norge (1832). Norsk ent. Tidsskr.2,318-329.Fabricius, J. C. 1775. Systema Entomologiae.832 pp.,Flensbyrgi et Lipsiae (Flensburg).Fabricius, J. C. 1776. Genera insectorum. 310 pp.,Chilonii (Kiel).Fabricius, J. C. I 787 . Mantissa insectorum. 1 . xx + 348pp., Hafniae (K. havn).Fabricius, J. C. 1793. Entomologia Systematico. 2.i-viii+519 pp., Hafniae (K. havn).Fabricius, J. C. 1798. Entomologiae SystematicAe.Suppl. 572 pp., Hafniae (K. havn).exp. 1900-1903 und einiger anderen arktischen Ausbeuten.Mem. Acad. Sci. Sr. Petersb. ( Sci, meth.,phys., nat.) 18, l-17.Friese, H. 1909. Neue Varietiiten von Bombus (Hym.).Dt. ent. Z.pp. 673-676.Friese, H. 1911. Neue Varietiiten von Bombus (Hym.)U-I[, 456-457, 57I-572. Zur Synonymie der Bombus-Artenp. 684. Dt. ent. Z.Friese, H. 1923. Hymenoptera. Apidae, pp. 3-9 inNorwegian Expedition to Novaya Zemlya 1921. Reportof the Scientffic Results. l, No. 14. publ. byVidenskapsselskapet i Kristiania (Oslo).Friese, H. 1935. Apiden aus Nordost-Gronland. Skr.Svalbard Ishavel No. 65, 1-10.Friese, H. & Wagner, F. 1909. Zoologische Studienan Hummeln. I. Die Hummeln der deutschen Fauna.Zool. Jb. Abt Sys/. , 0k, Geogr. 29, l_ 104 (1910).Friese, H. & Wagner, F. 1912. Zoologische Studienan Hummeln. II. Die Hummeln der Arktis, desHochgebirges und der Steppe. Zool. Jb. suppl. IS, I,t55-2r0.aFaester,K. 1959. In Dinemark getroffene Arten von Frison, T. H. 1927. A contribution to our knowledgeBombus und Psithyrus (Hym., Apidae). Ent. Meddr. of the relationship of the Bremidae of America north29, 152-153.of Mexico (Hymenoptera). Trons. Am. ent. Soc. 53,Forsius, R. 1925. Uber Bombus smithianus White in 51-78.Finland. MedCn. Soc. Fauna Flora fenn. 49, (1922- Fuglir, T. 1963.'Ndd lrerer naken kvinne i spinne'1923), 9-1 1.Naturen 87,3I1.Forsslund, K.-H. 1929. En tur til Vimhusfjrillen. Gaunitz, D. 1929. Gadd-och guldsteklar frin Sorsele,Sveriges Natur 20, 94-100.Lycksele lappmark. Ent. Tidskr. 50, 196-202.Forsslund, K.-H. 1951. Insektfynd pi fjrillet Hiirjehflgna.Norsk ent. Tidsskr. 8, 196-202.221-227.Gaunitz, D. 1931. Gaddstekelnotiser. Ent. Tidskr, 52,Fourcroy, A. F. 1785. Entomologia Parisiensis, sive Gaunitz, D. 1938. Smiirre meddelanden och notiser.catalogus Insectorum quae in agro Parisiensis reperiuntur. . . (new species added by Geoffroy). 544 pp., Gerstaecker, A. 1869. Beitrige zur niiheren KenntnisTre gynandromorfa bin. Ent. Tidskr. 59, 246-249.Paris (not seen).einiger Bienen-Gattungen. Stettin. ent. Ztg. 30,139-Free, J. B. & Butler, C. G. 1959. Bumblebees. 208 pp. 184, 315-367 .Collins, London.Grosvenor, M. B. 1963. National Geographic Atlas ofFrid6n, F. 1967. Fortsatta undersokningar rorande the World. 300 pp. Wash. D. C.insektpollination. Meddn. Sver. Frdodlfcirb. No. 8, Gurr, L. 1964. The distribution of bumblebees in the65-109.south island of New ZeaIand. N. Z. JI. Sci. 7, 625-Frid6n, F., Eskilsson, L. & Bingefors, S. 1962. Bumblebeesand red clover pollination in Central Sweden. ".Hammer, K. & Holm, Sv. N. 1970. Danske642.I. Int. symposium on pollination,{humble-196A. MedCn. Sver. bier og snyltehumler. Natur og Museum, /4 No. 2-3,Frtiodlareftrb. No. 7, 17-26.3-22.Fries, M. 1949. Den nordiska utbredningen av Lactucaalpina, Aconitum septentrionale, Ranunculus K. k. naturhistorischen Hofmuseums. Annln. naturlt.Handlirsch, A. 1888. Die Hummelsammlung desplatanifolius och Polygonatum verticillatum. Acta Mus. Wien 3,209-250.Phytogeographica Suecica. 24, 1-80.Harris, M. 1776. An Exposition of English Insects.Friese, H. 1902. Die arktischen Hymenopteren, mit 165 pp., London.Ausschluss der Tenthrediniden. Fauna arct. 2, 439- Hasselroto T. B. 1960. Studies on Swedish bumblebees.498.(Genus Bombus Latr.). Opusc. ent. suppl. 17, l-192.Friese, H. 1903. Neue Bombus-Arten aus der neotropischenRegion (Hym.). Z. syst. Hymenopt. Dip-Yhra vanligaste humlor. Hushdlln. scillsk. Ftirb. Smd-Hasselrot, T. B. 1962. Humlor och bin i froodlingen.terol. 3,253-255.skr. No. 5108-11.Friese, H. 1904. Uber die Hummelleben im arktischen Hell6n, W. 1933. Hymenoptera acaleata aus Finnmarkenin Norwegen. Notul. ent. 13, 4l-43.Gebiete. (Hym.). Allg. Z. Ent. 9,409-414.Friese, H. 1905. Neue oder wenig bekannte Hummeln Hobbs, G. A. 1964. Ecology of species of Bombusdes russischen Reiches (Hymenoptera). Esheg.-zool. Latr. (Hymenoptera: Apidae) in Southern Alberta.Mus. 9 (1904), 507-523.I. Subgenus Alpinobombus Skor. Can. Ent. 96,Friese, H. 1908. Uber die Bienen der russischen Polar- 1465-1470.

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Studies on Scandinavian Bumble BeesZl3Vestlandets Bombus- & Psityrus-arter. Bergens Mus.Arb. 1900 No. 3, l-lg.Lie-Pettersen, O. J. 1902. Biologische Beobachtungenan norwegischen Hummeln. Bergens Mus. Arb. 1901No. 6, 1-10.Lie-Pettersen, O. J. 1905. Entomologiske bidrag tilskjrergaardsfaunaen i det vestlige Norge. BergensMus. Arb. 1904 No. ll,l-25.Lie-Pettersen, O. J. 1907. Neue Beitriige nJr Biologieder norwegischen Hummeln. Bergens Mus. Arb. 1906No. 9,l-42.Lindroth, C. H. 1.949. Die fennoskandischen Carabidae.Eine tiergeographische Studie. 3 . Gbgs. K. Vet.Vitt. Samh. Handl. 6. F. serien B 4 (3), l-911.Lindroth, C. Ff. D56. Movements and changes ofarea at the climatic limit of terrestrial animal species.pp. 226-230 in: Zoological Papers in Honour ofBertil Hanstrdm's Sixty-Fifth Birthday November20th, 1956. Berling, Lund.Lindroth, C. H. 1962. Om istidsrefugier i Skandinavien.Sveriges Natur Arsb. 1962, 119-136.Lindroth, C. H. 1969. The theory of glacial refugia inScandinavia. Comments on present opinions. Notul.ent. 59, 178-192.LinnaeuS, C. 1758. Systema Naturre.I. 10th ed.824pp.,Holmia.Linnaeus, C. 1761. Fauna svecica. 2nd ed. 578 pp.,Stockholmiae.Linnaeus, C. l790.In Gmelin, J. F.: Systema Naturae.13th ed. I ,2225-3020, Lipsiae (Leipzig).Laken, A. 1949. Bumble bees in relation to Aconitumseptentrionale in Central Norway (Oyer). Nytt Mag.Naturv. 87, l-60.Loken, A. 1950. Bumble bees in relation to Aconitumseptentrionale in Western Norway (Eidfjord). Norskent. Tidsskr. 8, 1-16 (1951).Loken, A. 1954. Observations of bumble bee activityduring the solar eclipse June 30, 1954. Univ. BergenArb. 1954. Nat. rekke No. 13, l-6 (1955).Laken, A. 1958a. Pollination studies in apple orchardsof Western Norway. Xth Int. Congr. Ent. 1956. 4,961-965.Laken, A. 1958b. Bombus sylvarumv. nigrescensP6reznew to Norway (Hymenoptera, Apidae). Norsk ent.Tidsskr. 10, 236-238.Laken, A. 1960. Preliminary notes on Norwegianspecies of Bombus (Hymenoptera, Apidae). Norskent. Tidsskr. I I, 107-110.Loken, A. 1961a. Observations on Norwegian bumblebee nests (Hymenoptera, Apidae, Bombus). Norskent. Tidsskr. I I, 255-268.Laken, A. 1961b. Bombus consobrinus Dahlb., anoligolectic bumble bee (Hymenoptera, Apidae).XIth Int. Congr. Ent. 1960. l, 598-603.Laken, A. 1962. Occurrence and foraging behaviourof bumble-bees visiting Trifolium pratense L. inNorway. Proc. I. Int. Symposium on pollination.Meddn. Sver. Frriodl. Frirb. No. 7, 52-59.Laken, A. 1963. The neotype of Bombus alpinus(Linnaeus, 1758). Opusc. ent. 28,187-188.Loken, A. 1964. Bumble bees from Austria (Hymenoptera,Apidae). Norsk ent. Tidsskr. 12, 246-250.Laken, A. 7966a. Notes on Fabrician species ofBombus Latr. and Psithyrus Lep., with designationsof lectotypes (Hym., Apidae). Ent. Meddr. 34, 199-206.Loken, A. 1966b. The neotype of Bombus consobrinusDahlbom 1832. (Hym.). Opusc. ent. 31, 203-204.Loken, A. 1966c. Ekskursjonsberetning. Insekter ogarachnoider samlet under det 13. Nordiske Entomologmotetsekskursjon til Flflm (SFi: Aurland)13-16. august 1965. Norsk ent. Tidsskr. 13,371-386.Lundblad, O. 1924. Nflgra ord om aculeat- och tubuliferfaunani Lycksele lappmark. Ent. Tidskr. 45t7-28.May, J. 1959. amehc; y iSR jejich bionomie, chow ahospodaf sk! vyznam. I70 pp., Praha.Mayr, E. 1969. Principles of Systematic Zoology. 428Fp., McGraw-Hill, New York.Meidell, O. 1934a. Bier og humler i Rogaland (Apidae,Hym.). Stavanger Mus. Arsh.43 (1932-3j),85-133.Meidello O. 1934b. Fra dagliglivet i et homlebol. Naturen58, 85-95, 108-116.Meidell, O. 1946. Notes on the pollination of Melampyrumpratense and the 'honeystealing' of humblebeesand bees. Bergens Mus. Arb. 1944. Nat. rekkeNo. ll,l-12.Meidell, O. 1968. Bombus jonellus (Kirby) (Hym.,Apidae) has two generations in a season. Norsk ent.Tidsskr. 14, 3l-32.Michener, C. D. 1944. Comparative external morphology,phylogotry, and a classification of the bees(Hymenoptera). Bull. Am. Mus. Nat. Hist. 82, l5l-326.Milliron, H. E. 1960. Recognition of bumblebee typespecimens, with notes on some dubious names(Hymenoptera: Apidae). Bull. Brooklyn ent. Soc. 5587-99.Milliron, If. E. l96L Revised classification of thebumblebees - a synopsis (Hymenoptera: Apidae).J. Kans. ent. Soc. 34,49-61.Milliron, H. E. & Oliver, D. R. 1966. Bumblebeesfrom Northern Ellesmere Island, with observations onusurpation by Megabombus hyperboreus (Schonh).(Hymenoptera: Apidae). Can. Ent. 98,207-2 3.Morawitz, F. 1869. Die Bienen des Gouvernmentsvon St. Petersburg. Trudy russk. dnt. Obshch 6,27-71 .Morawitz, F. 1873. Die Bienen Daghestans. Trudyrussk. dnt. Obshch. 10, 129-189.Morawitz, F. 1880. Ein Beitrag zur Bienenfauna Mittel-Asiens.Bull. Acad. Sci. St. Petersb. 26,337-389(: Melanges Biol. 10, 443-518).Morawitz, F. 1881. Die russischen Bombus-Arten inder Sammlung der Kaiserlichen Academie der Wissenschaften,St. Petersb. Bull. Acad.,Sci. S/. Petersb.27,213-265 (: Melanges Biol. 11,69-144).Morawitz, F. 1888. Hymenoptera aculeata nova.Trudy russk. dnt. Obshch. 22,224-302.

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Studies on Scandinavian Bumble Bees 215skinska stranddynernas insektfauna. Ent. Tidskr. 42,2L-40, 65-92.Roth, C. D. E. 1897. Bidrag till en bild af Skinesinsektfauna. Ent. Tidskr. 18, 127- 38.Sagemehl, M. 1882. Yerzeichnis der in Est-, Liv- undCurland bisher gefundenen Bienen. Arch. Naturk.Liv-, Ehst-, Kurlands, 2 Serie 8, 451-468.Sakagami, S. F. & Ishikawa, R. 1969. Note pr6liminairesur la r6partition g6ographique des bourdonsjaponais, avec descriptions et remarques sur quelquesformes nouvelles ou peu connues. .f. Fac. Sci. HokkaidoUniv. series VI, Zoology 17, 152-196.Sandhouse, G. A. 1943. The type species of the generaand subgenera of bees. Proc. U.S. natn. Mus. 92,519-619.Schenck, A. 1873. Uber einige streitige und zweifelhafteBienen-Arten. Dt. ent. Z. ( : Berl. ent. Z.) 17,243-259.Schenck, A. 1875. Aus der Bienen-Fauna Nassau's.Dt. ent. Z. ( : Berl. ent. Z.) 19, 321-332.Schmiedeknecht, O. 1878. Monographie der in Thiiringenvorkommenden Arten der Hymenopteren-Gattung Bombus. 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Om en i sommeren 1869 foretogenentomologisk reise gjennem Ringerike, Hallingdal ogValders. 7l pp., Gundersen, Christiania (Oslo).Siebke, H. 1873. Bidrag til Norges insektfauna. Beretningom en i Osterdalen foretagen reise i 1870. NyttMag. Naturv. 19, 39-1A2.Siebke, H. 1880. Enumeratio Insectorum Norvegicorum.Catalogum hymenopterorum continentem. Auctoredefuncto H. Siebke, ed. Schneider, J. Sparre, 5 (1).95 pp., Broegger, Christianiae (Oslo).Skorikov, A. S. 1908. Formes nouvelles des bourdons(Hymenoptera, Bombidae). (In Russian.) Russk. ent.Obozr. 7 (1907), I I 1-113.Skorikov, A. S. 1910. Nouvelles formes des bourdons(Hymenoptera, Bombidae). (In Russian.) Russk. ent.Obozr. 9 (1909), 409-413.Skorikov, A. S. l9ll. Bombus pratorum (L.) et sesformes (Hymenoptera, Bombidae). (In Russian.)Russk. ent. Obozr.11, 380-382.Skorikov, A. S. 1912. Bombus lapponicas (F.) et sesformes (?? et Hil (Hymenoptera, Bombidae). (InRussian.) Russk. ent. Obozr. 12,95-102.Skorikov, A. S. 1914a. Les formes nouvelles des bourdons(Hymenoptera, Bombidae). (In Russian.) Russk.ent . Obozr . 14, tl9-I29 .Skorikov, A. S. I9I4b. Hortobombus consobrinus(Dahlb.) et ses variations (Hymenoptera, Bombidae).(In Russian.) Russk. ent. Obozr. 14,283-286.Skorikov, A. S. I922a. Faunae Petropolitanae catalogus.(In Russian.) Petrograd agron. Inst. ent. Sta.Fauna Petrograd 2 No. 11, 1-51 (not seen).Skorikov, A. S. L922b. Les bourdons de la fauna

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Studies on Scandinavian Bumble Bees2t7INDEXValid names in roman types. Synonyffis, homonyms,in{rasubspecific forms, etc., in italics. Only thosepages referring to description or taxonomic status areindexed.aceruorum LinnaeusAdv ent ori bo mbus Skorikovadventor SkorikovAgrobombusVogt............ ......agrorum Fabricius ....agrorum (Fabricius)Alpigenobombus Skorikovalpigenus Morawitz....oalpiniformis Richards . . . . . . . . . . o . . . . . 95,Alpinobombus Skorikov . .. . . . ... . . . o. . .. . . .alpina Linnaeusalpinus (Linnaeus)apollineus Skorikov ............ .....apricaFabricius .. ............ ..,...apricus(Fabricius) ............arbustorum Fabricius ....ar b u s t o r um Zetterstedtarctica AcerbiarcticusDahlbom ....arcticusKirby .... .....arcticus Krtiger .... ...o.... ..r.arcticus (Quensel in Acerbi)arenicola ThomsonargillaceaScopoli o... ......argillaceus (Scopoli) ....atrocorbiculosus Vogt . . . . . . . . . . o . . . . . . . .auranticzs Dalla TorreautumnalisDahlbom ....autumnalis Fabricius . . . . . . . .autumnalis Zetterstedt . . . .balteatusDahlbom .....balticus Scholz . . . . . . . . . . . .barcaiVogt ........ 154,186137t37137154ts425251019496965664649083tl41541014011,417911813669t3917912469105t2s159beckwithellaKirby o. . . . . . o . . . . . . . . . o . . . 154bicolor Sparre Schneider . . . . . . . . . . . . 154, t66bimacularzs Skorikov .. o....... ....... o. ....bipunctatus Friese & Wagner . . . . . ^ . . . . . .8432Bombus Latreilles.s. .. q. ........ .... 37borealis Schmiedeknecht . . . .186brevigenaThomson ....oo.. ..... 25BremusJurine .o...... ..... 37burgllana Kirby o. o.. .. .......... ...........8384burrellanus (Kirby) . . . . . . . ..... . . . .burrellanusThomson ....56calidusErichson ...............o.... . 64.. 68calidus Erichson ........... 64Chromobombus Dalla Torrg ...o.. o.... o.....cingulatus Wahlberg ..... ........ o..........citrinus Schmiedeknecht... ......cognatu^y auctt.nec Stephen . . . . . . . . . . o . . . . . . .cognatusStephen ................. ......collarisHell6n ...o............ .o.......consobrinus Dahlbom . . . . . r, . . . . . . . . . . .cryptarum FabriciusCullumanobombus VogtcullumanaKirby ....cullumanus (Kirby) . . . . . . . .curtisella Kirby ....curtisellus (Kirby). .decoloraras AlfkenderhamellaKirby. ..............derhamellus (Kirby) ....4056565615415484169r69r0275t82diabolicusFriese.... ...101,dissidens Friesedistinguendus Morawitz. ....donovanellaKirby .... .... 83donovanellus (Kirby)84ephippiumDahlbom .... 83equestris auctt.nec Fabricius . . . . . . . . . . . . . 179equestris (Fabricius) ........ 180ericetorum Panzer .erlandssoni Kruseman64154eurynotusDalla Torrg .... .......... 1,36eurynotzs Kriechbaumer .... t36eurynotus vogtfidens(Harris) .......... .......r36t25fidus Harris ..... 83, 84flavicolorFriese .... 69flavoscutellarisTrautman ......... o.. 46floralis LinnaeusGmelin ..o........... 154formolo,su.t SkorikovforsterellaKirby ... ........ .75ts4fragransKirby... o......... 183fragrans(Kirby) .... ....... 183francillonella Kirby154frigidusFriese .... ........ ..... 64frisei Skorikov ....101gerstaeckeri Morawitzgmelini SkorikovgotlandicusErlandsson ........118105. 154, 169hafsahli VogthafsahlianusVogt....139139, 145hafsahloides Vogt 139, r40harrisella Kirby 125, 136harrisellus(Kirby) .............o........helveticzs Frieso ........ o...................12575helftranrzsSeidl ...............o........ 139hiemalis Friese........ ......... 64hirtus Vogto...r24HortobombusVogt ............ ......... il8hortorumLinnaeus ........................ o r24hortorum (Linnaeus) . . o . . . o o . . . . . . . . . . . 124humilisllliger.... .... ..........139hyperboreus Schonherr . . . . . o . . . . ... ll4hypnorum (Linnaeus) . .636316969

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A free drawing by G. Langhelle, from Skorikov 1922b.

Bumble Bees

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