Scandinavian species of the genus Psithyrus Lepeletier ...

ASTRID I-OKENScandinavian species of the genusPsithyrus Lepeletier(Hymenoptera: Apidae)Entomologica ScandinavicaSupplement No. 23

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ENT. SCAND. SUPPL .23(1984)MethodsTaxonomyKeys to the genera of Bombinae and to thespecies of European Psithyrus are provided. Thepapers of Popov (1931) and Pittioni (1939) wereconsulted. The nomenclature of Bombus followsLoken (1973).Scandinavian species of Psithyrus Lepeletier 3follow Loken (1973) except that regional divisionfollows NU (1977). Danish and Finnish distributionsindicated on the maps are from Hammer(1970) and Pekkarinen et al. (1981).If not otherwise mentioned the general distributionof the individual species is taken from Popov(1931), Panfilov (1957), Reinig (1971), Tkalcfl(I974b) and Rasmont (1983).M orpholo gical terms, abbreviationsThe terminology follows Loken (1973). Abbreviationsand additional terms:Avtz in female, Ar-r: in male: Individual segmentsof antennae.Bt: Basitarsus.Genal furrow: Border between vertex and uppergena (Fig . 4).Str-e in female, Str-z in male: Individual gastralsternites.Tr-o in female,T p7 in male: Individual gastral tergites.Upper gena: Part of gena outside genal furrow andupper part of eye.MeasurementsThe following measurements were taken inboth sexes: (a) Rl (Fig. 1); (b) Iw (Fig.3); (c) Hw- head width; (d) hamuli number; (e) Lh (Fig. 5);(f) Dh (Fie s); (e) Ar; (h) A$ (i) A'; (j) or (Fig.+); (k) Oe (Fig. 4). The measurements of a-f presentedby number (N), mean, * standard deviationand range, and are given in mm. Thosemarked g-k are expressed in micrometer units asonly the proportions g:h:i and j:k are used.Interalar width is chosen as an indicator of bodysize and the following categories were used:Female: Body large, mean Iw >5.00 mm; bodyof medium size Iw 4.60-5.00 mrn; body small, Iw3.70 mm; body ofmedium size, mean Iw 3.55-3.70 mm; body small,mean Iw

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ENT. SCAND. SUPPL. 23 (1984) Scandinavian species of Psithyrus Lepeletier 57AFigs. 6-15. - 6. P. bohemicus (Seidl) Q; A = right mandible, B = ocellar-orbital field, C : callosities of sternite6.- T.P.vestalis(GeoffroyinFourcroy)9;A:ocellar-orbitalfield,B:callositiesofstemite6.- S.P.rupestris(Fabricius)Q,callositiesofsternite6.- 9.P.barbutellus(Kirby)Q,callositiesofsternite6.- l0.P.campestris(Panzer) Q, callositiesofsternite6.- 11. P. sylvestris Lepeletier Q;A: rightmandible,B = tergite6,C = callositiesofsternite6.-12.P.norvegicusSp.SchneiderQ;A:tergite6,B=callositiesofsternite6.- L3.P.flavidus (Eversmann) Q , callosities of sternite 6. - 14. P. quadricolorLepeletier t ; callosities of sternite 6. - 15.Fernaldaepsithyrus t; lateral view of tip of gaster.fringe hardly or not exceeding their greatest width.T6 keeled (Fig. l2A). Callosities of St6 as in Fig.I2BP. (Fernaldaepsithyrus)norvegicus Sp. Schneider p. 36Hind tibia and Bt with longest hairs in posteriorfringe markedly longer than their greatest width. T6inconspicuously or not keeled. Callosities as inFigs. 13-14 ....... 99. Callosities of St6 as in Fig. L3.T4 yellow-haired, T5black-haired P.(Fernaldaepsithyrus) flavidus (Eversmann) p. 28Callosities of St6 as in Fig. 14. T:-s ferruginoushaired,often with an admixture of whitish hairsP. (Fernaldaepsithyrus) quadricolor Lepeletier p. 23MALES1. Malar space shorter than 3lq the distal width. Stz-aand genitalia as in Figs. 16-17; penis valve nothooked beneath .......... Ashtonipsithyrus Frison 2Malar space longer than 3lq the distal width. Stz-sand genitalia as in Figs. 18-24; penis valve hookedbeneath 32. A.3 hardly or not longer than ,A.5. Hind tibia and Btwith longest hairs in posterior fringe at least as longas their greatest width. Stz-s and genitalia as in Fig.16.{{ P. (Ashtonipsithyrw) bohemicru (Seidl) p. 8,A.3 markedly shorter than As. Hind tibia and Bt withlongest hairs in posterior fringe shorter than their grea-

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ENT. SCAND. SUPPL .23(1984) Scandinavian species of PsithyrusLepeletier21A22A 23B,22823C24B,24CK/ffiGWG22DW24EG21CW23A 22C 22E 23DFigs.2l-24. Fernaldaepsithyras Frison; sternite 6, right half of male sternite 7-8 and genitalia. - )il . P. quadricolorLJpeletierd;A:steiniteT,B:sternite3,6:pdsteriorpartofgenitalia.- 22.P.norvegrcasSp.Schneiderd;A^:sterniteT,B=stemitei3,6=genitalia,D-E:posteiiorpartofgenitalia.- 23.P:sy.lvestrisl-epeletierd;A: sternite6, B = sterniteT, C =-sterniteS, p:lenitalis,B: posteriorpartofgenitalia.- 24. P' flavidus(Eversmann)d;A:sternite6,B=sterniteT,C:sterniteS,D-E=posteriorpartofgenitalia.or not exceeding their greatest width. Stz-s and genitaliaas in Fig. 18A{. Coat shaggyP. (Allopsithyrus) barbutellus (Kirby) p. 126. ,A.3 slightly shorter than ,A.5. Hind Bt parallel-sided, ffiorethan 4x longer than wide. Stz-s and genitalia as inFig. 19A-C . T+4with reddish hairsi; ;;;k;;d ;;#"{',":# Pr#{;'{:'Jl1:ffi:? Xoits greatest width. Stz-s and genitalia as in Fig. 20A{.T +4without reddish hairsP. (Metapsithyrw Popov) campestris (Panzer) p. 187 . Stu distally broadly rounded. St7-6 and genitalia as inFig. 21A{. T+ with an admixture of yellowish, whitishor pinkish-orange hairs, Ts-, with femrginous hairs .P. ( Fernaldaepsithyrus) quadricolor Lepeletier p. 23St6 as in Figs . 23A, 24A. Stz-s and genitalia as inFigs . 22-24. T q-s otherwise coloured 88. ,A.3 markedly shorter than As. Ao- tzZx their width.Genal furrow distinct. Stz-s and genitalia as in Fig.22A_E P.(Fernaldaepsithyrus) norvegicas Sp. Schneider p. 36,{3 hardly or not shorter than As. Ao-13 shorter than

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ENT. SCAND. SUPPL.23 (1984)DescriptionFEMALEBody of medium size. Coat rather shaggy.Measurements: N-20. Rl: 4.06 + 0.265 mm'3.55-4.45 mm; Iw: 4.79 + 0.316 mm, 4.2V5.30mm; Hw: 4.55 + 0.233mm, 4.151.90 mm; hamulinumber: 20.75 +1.4I0., 18-23; Lh: I.63 + 0.102mm, I.45-I.78 mm; Dh: 0.083 + 0.004 ffiffi,0.078-0-094 mm.Head. Malar space with close distribution offine punctures admixed with some coarser ones,distally fading into an almost impunctate, largesubtriangular area. Clypeus more sparsely andcoarsely punctate on disc than at base, distallywith a well defined median impunctate space.Upper gena with sparse to moderate distributionof punctures of varying size; genal furrow usuallyindistinct. OOb without accumulation of minutepunctures (Fig. 68). ,{3:Ao:As: 19.5: 11: 1,4,, Ao.njust longer than their distal width.Hind lrg.Bt about 2x longer than its greatestwidth. Longest hairs in posterior fringe of tibiaand Bt as least as long as the segment's greatestwidth.Gaster. T6 smooth, shiny, nearly impunctate onthe disc, closely punctured laterad and distad.Ridge-like callosities (Fig. 6C) distally morepointed and less separated than in P. vestalis (Fig.7B ); rather sparse - mainly fine - punctures on thedisc between the callosities.Colour pattern. Broad, pale to dull yellow collarextending aboutllq down the adjacent episternumand more or less projecting backwards below baseof wing. Extreme posterior fringe of scutellumusually with single yellow hairs concealed by blackones. Sparse pale yellow or whitish hairs in lateralpatches of T1 . Latero-posterial patches and extremedistal margin of T3 , T q, and lateral patchesof T5 whitish-haired. T6 laterally and distally, andSt6 outside and between callosities clothed withgolden-brown pubescence. Otherwise coat black.Yellow hairs paler than in P. vestalis.Variation. Pile of vertex encroached by yellow.Collar varying in width, reduced to lateral patchesor completely replaced by black hairs. Pile in crescent-shapedposterior fringe of scutellum rangingfrom all black to nearly all yellow. Slight variationin the amount of whitish hairs on T3 and Ts.Whitish hairs, particularly on T3 occationally replacedby yellow ones.Scandinavian species of Psithyrus Lepeletier 9MALEBody of medium size. Coat shaggy, long.Measurements. N - 20. Rl: 3.62 + 0.200 mm.3.05-3.85 mm; Iw: 3.62 + 0.195 mm, 3.25-3.90mm; Hw:4.13 + 0.204 mm ,3.6M.40 mm;hamulinumber: 17.15 + 0.988, 7Gl9; Lh: 1.32 + 0.079mm, 1.18-1.48 mm; Dh: 0.082 + 0.006 ffiffi,0.072-0.093 mm.Head. Malar space markedly longer than As,distally impunctate or nearly so, otherwise with ir-::ili':lL :r':x# 1lh 3: ?:'J; ff f T, * J I|spaced, shallow punctures of varying size; genalfurrow usually distinct. Or - 2x Oe or nearly so.OOb without accumulation of minute punctures(see Fig. 68). A3:Aa:As : 17:10.5:16, Ao-r3 about1.5 x their distal width; flagellum about 3 x longerthan scapus.Hind leg. Tlbia with a small almost hairless areaclose to distal margin; longest hairs in posteriorfringe up to 1.5x the segment's greatest width. Bt3.5x longer than its greatest width or nearly so;longest hairs in posterior fringe up to 2x Bt'sgreatest width.Gaster. Stz-s and genitalia as in Fig. 16A-C; thatpart of volsella visible from above about 1 .5 xlonger than gonostylus.Colour pattern. Pile of vertex with black andyellow intermixed. Collar laterally extendingabout l lq down episternum, hairs of posteriorfringe of scutellum and T1 yellow. Hairs of lateroposterialpatches of T z, T +-5 , lateral patches of T6,at least fringes of Sta-5 whitish. Otherwise coatblack.Variation. Pile of vertex ranges from all black topredominanty yellow. Yellow collar varying inwidth, laterally passing more than half way downepisternum, reduced to lateral patches or turningall black. Pile of scutellum ranging from all yellowto all black. Pile of particularly mid and hindbasitarsi occasionally pale yellowish-brown or tippedso. Whitish patches of T3 varling in size, occasionallywhitish hairs extending along extreme distalmargin. White hairs on T3 replaced by yellowones. Pile on T5 admixed with black hairs on thedisc. Pile on Stz-s ranging from all black to allwhitish.Melanism. Colour variations are too slight tochange the general colour pattern except forsporadically occurring pronounced melanic specimens.The darkest individuals are nearly black,having at most whitish hairs on T5. The tendency

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ENT. SCAND. SUPPL .23(1984) Scandinavian species of Psithyrus Lepeletier 11Fig.25. P. bohemicus (Seidl). The distribution in Fennoscandiand Denmark, Inset: The host B. lucorum(I-,)G;ken 1973) updated iSSf.'O = one or severalocalities; O : unrevised records (from literature or received inlitt.); X = inaccurate locality. Shaded areas refer to literature indications.

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ENT. SCAND. SUPPL.23 (1984)Scandinavian species of Psithyrus Lepeletierl37.4VI.78 mm; Dh: 0.082 + 0.004 mm, 0.075-0.087 mm.Head. Malar space about 3lq the distal width,about as long as A31a; integument more or lessstriate-rugose closely set with punctures of varyingsize fading into an almost impunctate narrow areaclose to distal margin. Main keel of mandible welldeveloped, anteriorly tapering, reaching marginor almost so. Labral furrow deep, broad; labraltubercles protuberant, blunt, the distal distancebetween them at least rlz labral width. Clypeusmore sparsely and markedly more coarselypunctate on disc than at base. Upper gena withrather dense, distinct, mainly coarse punctation;genal furrow absent or indistinct. Or about 2xOe. A3:Aa:A5 _ 19:lI:16, Ao-rz as long as theirdistal width.Hind lrg.Bt markedly narrower than adjoiningtibia, about 3x longer than its greatest width..Longest hairs in posterior fringe of tibia and Btequal to the segment's greatest width or justlonger.Gaster. Tip of gaster not much incurved. T+-swith rather close, mainly coarse puncturing. T6with close, rather uniform, mainly medium-sizedpunctation; usually with a weak, narrow impunctate,median longitudinal keel. Sto with flattenedsemi-circular callosities positioned at a distancefrom apex (Fig. 9).Colour pattern. Pile of vertex with an admixtureof yellow. Hairs of collar just passing margin ofepisternum, scutellum and variable part of T1 yellow.Hairs of Ta and lateral patches of T5 whitish.T6 and Sto outside the callosities with goldenbrownpubescence. Otherwise coat black.Variation. Pile of vertex ranging from predominantlyyellow to all black. Yellow collar more orless reduced. Pile of scutellum ranging from allyellow to all black except a few yellow hairs alongextreme posterior margin. Legs with variable admixtureof pale-coloured hairs, especially in thefringes.MALEBody of medium size. Coat rather shaggy.Measurements. N - 20. Rl: 3.44 + 0.178 mm.3.25-3.75 mm; Iw: 3.62 + 0.24I mm. 3.2H.05mm; Hw: 4.25 + 0. 163 mm,3.9M.50 mm; hamulinumber: 18.60 t 1.698, 17-23; Lh: 1.30 + 0.059mm, 7.2UI.44 mm; Dh: 0.074 t 0.007 mm.0.059-0.083 mm.Head. Malar space hardly as long as distalwidth, ?S long as A:*+; integument distally impunctateor almost SO, otherwise irregularlypunctured, punctures coarser centrally thanbasad. Upper gena with rather dense mainly shallow,coarse punctation;genal furrow absent, occasionallyindistinct. Or about 2x Oe. Hairs ofscapus about 2x width of segment; ,{3:Aa:45 -16:10.5:19, Ao-r: nearly 2x their distal width;flagellum about 3x longer than scapus.Hind lrg. Bt 3.75x longer than its greatestwidth. Tibia and Bt with longest hairs in posteriorfringe hardly or not exceeding their greatestwidth.Gaster. St6 with 2 mounded callosities. St74 andgenitalia as in Fig. 18A-C; penis valve hooked beneath.Colour pattern Pile of vertex, collar just passingmargin of episternum, scutellum and T1 yellow.Hairs of extreme distal margin of T3 ,Tq-s, lateralpatches of T6, Stz-s whitish. Otherwise coat black.Variation. Pile of vertex with variable admixtureof black. Yellow collar extending ll4 to ll2down episternum. Yellow hairs of scutellum moreor less reduced to crescent-shaped posteriorfringe.MelanismBoth sexes display a slight tendency towardsmelanism which is more pronounced in southernmostSweden (SK, OL) than further north.HostBombus hortorum (L.) (Ball l9l4; Cumber1949; Mulle r 1936; Postner 1952). B. hypnorum(L.) (Pouvreau 1973).Host species mentioned by Hoffer (1889) areomitted as the author confused P. barbutellus andP. sylvestris. P. barbutellus observed establishedin a colony of B. pratorum (L.) and B. ruderariusrespectively (Pouvreau 1973) are a misprint(Pouvreau in litt.).Flight seasonSouthern Sweden: Hibernated female: 9 May -2 August;female progeny: ? -3 September;male:lJuly-5October.

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ENT. SCAND. SUPPL.23 (1e84) Scandinavian species of Psithyrus Lepeletier 15Fig.26.P.barbutellus(Kirby).Inset:ThehostB.hortorw (L.)updated1983.LegendasinFig.25.

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ENT. SCAND. SUPPL.23 (1984) Scandinavian species of Psithyrus Lepeletier 17Fig.27. P. rupestris (Fabricius). Inset: The host B. lapidarizs (L.), updated 1983. Legend as in Fig. 25.

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ENT. SCAND. SUPPL .23(1984)Scandinavian species of Psithyrus Lepeletier 19Apis rosiella Kirby,1802:331, No. 85. Type locality: England:E. Suffolk: Barham. Lectotype d KCL! (Yarrow1e68).Bombus rosiellus (Kirby); Dahlbom L832.Apathus rosiellus (Kirby); Thomson 1870, 1872.Apis leana Kirby ,1802:333, No. 86. Type locality: England:E. Suffolk: Barham. Holotype C KCL! (Yarrow1e68).Apis francisanaKirby, 1802:334, No. 87. Typ. locality:England: E. Suffolk: Barham. Holotype d KCt-! (Yarrow1968).Nomenclatural and taxonomical remarksWahlberg (1855b) argued that the descriptionof P. campestris by Dahlbom (1832) applied tocampestris Panzer while the appurtinant figuremay refer to the male of P. svaveolens Wahlberg -P. quadricolor Lepeletier. The figure is in myopinion too diffuse for a sperific division ofDahlbom's campeslris which should be confinedto that of.Panzer onlvJFirst S candinav ian recordsNorwav: HOy: Bergen (Lie-Petters en 1902).Sweden: SK, SM, VG (Dahlbom 1832). Recordsfrom Lapland (Dahlbom 1832) were revisedto P. flavidus (Thomson 1872).DescriptionFEMALEBody of medium size. Coat rather even, gastermarkedly sparser coated than in the remainingEuropean species.Measurements. Norway: (AAy-HOy): N :20.Rl: 3.94 + 0.159 mm,3.5H.20 mm; Iw: 4.82 !0.26I mm , 4.45-5.15 mm; Hw: 4.64 +0.166 mm,4.154.90 mm; hamuli numb et:22.25 + L916, 18-26; Lh: I.57 t 0.090 mm, 1.4V1.70 mm; Dh:0.074 t 0.004 mm, 0.067{).082 mm.Head. Malar space nearly as long as distalwidth, slightly shorter than Az*+; integumentsmooth with irregular distribution of punctures ofvarying size distally fading into a well defined minutelypunctured area. Main keel of mandibleusually broad, reaching aboutzlt down to margin,broadly abrupted distad. Labral furrow deep,broad, widened towards the base; labral tuberclesdistally pointed , feebly raised, the distal distancebetween them exceeding tltlabral width. Clypeuswith dense, coarse punctation except for a medianimpunctate area close to distal margin. Uppergena with scattered minute punctures betweenmainly shallow coarse ones; genal furrow absent,occasionally inconspicuous. Or shorter than 2xOe. A3:Aa:As : 77.5:10.5:15 , Ao-nhardly lcingerthan their distal width.Hind lrg. Tibia with longest hairs in posteriorfringe slightly or not longer than its greatest width.Bt markedly narrower than adjoining part of tibia,hardly 3 x longer than its greatest width, longesthairs in posterior fringe hardly exceeding 0.5x itswidth.Gaster. Tip of gaster not much incurved.Tqashiny; closer, finer punctured medially thanlaterad and distad. Terminal of St6 slightly produced;the callosities convex distally pointed, &lmostreaching apex and separated by a narrow,deep furrow (Fig. 10).Colour pattern Pile of vertex with slight admixtureof yellow. Hairs of broad collar, extendingabout ll4 down episternum, scutellum, lateroposterialpatches of T. and large patches of To-tyellow. Interalar-band laterally with varying admixtureof yellow hairs, posteriorly more or lessV-shaped. T6 and St6 with pale yellow to goldenbrownpubescence. Yellow pile on thorax usuallypaler than that of gaster. Otherwise coat black.Variation. Pile of vertex all black. Pile on collar,scutellum individually ranging from all black to allyellow, rarely hairs on thorax all yellow except fora few black hairs on the disc. Yellow hairs on gastermore or less replaced by black ones. Completelymelanic specimens occur.MALEBody of medium size. Coat shaggy.Measurements. Norway (AAy - HOy): N :20.Rl: 3.48 + 0.162 mm,3.2V3.80 mm; Iw:3.64 +0.244 mm ,3.25L 10 mm; Hw:4.11 + 0.121ffiffi,3.9M.30 mm;hamuli number: 19.85 + 1.226, 18-22; Lh: I.42 + 0.069 mm, 1.30-1.50 mm; Dh:0.075 + 0.004 mm, 0.069-0.083 mm.Head. Malar space hardly shorter than distalwidth, 3S long as A31+ or almost so; integumentfeebly alutaceous with irregular distribution ofmainly minute to moderate punctures distally fadinginto a fine-punctured or impunctate area.

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ENT. SCAND. SUPPL.23 ( 1984) Scandinavian species of Psithyrus Lepeletier 21Fig. 28. P. campestris (Panzer). Inset: The host P. pascuorum (Scopoli) updated 1983. The hatching refers to the distributionof subspecies. Legend as in Fig. 25.

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ENT. SCAND. SUPPL .23(1e84)overlap between the species, occasionally impedereliable identification of European specimens. Anexception is P. quadricolor where the variability inthe diagnostic characters interferes in a lesser degreewith the remaining species , a.E it is recognizedby the shape of St6 and details in the colourpattern (.f. below). Reliable determination ofcritical males of P. sylvestris, P. flavidus and P.norvegicus is, however, obtained by using the classificationfunctions derived from a step- wise discriminantanalysis routine based on morphometricvariables (Lgken & Framstad 1983). In thatpaper, however, there are some few errors: Page48 column 2: K/E everywhere to be read N/E. E(527.56870) column P. sylvestris read 527 .32467.Table I: I(7.138) column P. flavidus read 1.738.Page 49 column 1: E(I67 .99313) column P. norvegicusread 167 .88313.Psithyrus (Fernaldaepsithyrus) quadricolorLepeletierFigs. 14,21,29.Psithyrus quadricolor var. quadricolor Lepeletier deSaint Fargeau 1832:376,, 1841 :428. Type locality: ThePyrenees. Type not traced.Psithyrus quadricolor Lepeletier; Richards 1928.Apathus quadricolor (Lepeletier); Thomson 1870 (partim)Bombus frutetorum: Zetterstedt nec Panzet 1801, thed.Psithyrus globosus Eversmann, 7852. Type locality_:USSR: Oienburg, Spaskoje. Type (?holotype) ?MAL!; Lie Pettersen 1901, 1905 , 1907; Muchardt 1904;Aurivillius 1903; Strand 1904; Sp. Schneider 1918' 'Ander 1965; Kullenberg et al . I97U Bringe r 1973.Apathus globosus (Eversmann); Thomson 1872; Siebke1880; Strand 1898b.Psithyrus quadricolor globosus Eversmann; Richards1928; Meidell 1934; Ander 1963P s ithy rus s v av e o I e ns W ahlberg 1855 a .132.T y_pe local i ty :Sweden: Ostergotland. Lectotype ? ZMLI. (Ander1967); Wahlberg 1855b.Psithyrus quadricolor var. intercalaris Mtill et, 1913.Psithyrus meridionalis Richards, 1928:351. Type locality:Yugoslavia: Styria. Lectotype ? TRM!, hereby designated(cf. below).Psithyrus quadricolor meridionalis Richards; Tkalcfl7962.TaxonomicalremarksP. quadricolor is divided into 5 subspecies inEurope, viz. P. q. quadricolor,Richards, P. q. globosus Eversmann,P. q. rossicusPopov and P. q. meridionalisP. q. arvernicusRichards. TheScandinavian species of Psithyrus Lepeletier 23designations refer to variations in the colour patternof the coat and were based on a few specimens(Richards 1928; Popov l93I). P. q. meridionalis iswell defined (cf. below). The subspecific status ofthe remaining taxa is questionable (Tkal cfr' 1962).I find, however, it is most adequate to retain thesubspecific divisions above until further materialfrom populations throughout the range of thisspecies is studied.P. quadricolor meridionalis RichardsP. globosus auctt., nec Eversmann.D esignation of lectotyp eP. meridionalis was described from a Q and 12C C of which the Q and one of the C d are in coll.TRM. The ? is hereby designated as lectotypeand has the following labels: (1) "Styria, TragopObert"; (2) "E. Graeffe"; (3) Ps. meridionalisn.sp ? type" [handwritten]; (4) "lectotype P.meridionalis Richardt ? design. A. Loken 1984" .The specimen is in good condition except that leftAt-r2., 3last tarsal segments of the right hind legand 2 last tarsal segments of the left hind leg arelost. Colour pattern: Pile on more than posterior0.5x of T, , T* and fringes of Str-r ferruginous.Coat otherwise black.A male paralectotype designated as allolectotypehas labels as follows: (I-2) As for the lectotype;(3) "Ps. meridionalis C type" [handwritten];(4) "allolectotype P. meridionalis RichardsC design. A. Loken 1984". The specimen is ingood condition except that the 2 last tarsal segmentsof the left hind leg are lost. The sculptureagrees with that of P. quadricolor . Colour pattern:Pile on posterior half of T3 , T +-t, single hairs infringes of mid l.g, most hairs in posterior fringe ofhind tibia and Bt, and fringes of sternites ferruginous.Coat otherwise black.Psithyrus quadricolor globosus EversmannFig.29.First S candinav ian recordsI"lorway: On: Dovre, SFi: Jostedal (Wahlbergr 85sb).Sweden: Localities not mentioned (Kriech-

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ENT. SCAND. SUPPL.23 (1984)Scandinavian species of Psithyrus Lepeletier 25subspecies of host and parasite, respectively (cf.below).P. quadricolor ursuping a colony of B. pratorum(L.) (Pouvreau 1973) is a misprint (Pouvreau inlitt.). B. ruderarirzs (Miiller) listed as a host (Pekkarinenet al. 1981) refers to an observation by A.Nordman of a Q flying into a nest of this Bombussp. (Pekkarinen in litt.).Flight seasonSouthern Sweden: Hibernating female: 4 May -12 August; female progeny: ? 13 September;male: 9June-9October.DistributionScandinavia (Fig.29). Confined to nemoral andboreonemoral zones. In Sweden also occurring inthe southern boreal zone.In Norway more often observed at sea level inluxuriant biotopes along the western coast than inthe southeastern lowlands. In Sweden widely distributednorth to "limes norrlandicus" (NU 1977)and further north occasionally recorded togetherwith B. soroeensis throughout the southern borealzone. A single record in central boreal zone (VB:Degerfors NRS) indicates a scattered distributionnortheast to at least about 64"N.Scarce. A total of 135 Norwegian and about 230Swedish specimens were examined.Vdstra Vram, Ovedskloster. HA: Halmstad NRS,Hdsslov, Snostorp NRS. - SM: FAgelfors. Grdnnaberget,Horeda NRS, Korsbergp, Markaryd, Ryssby,Sjotorp by Alvesta, Vdstervik, Osterkorsbeqg_a_GNMZML. - OL: Bengtstorp, Borgholm, Boda NRS ZrylU,Gdrdslosa, Hogsium-Rilla, Rod, Vickleby. OC:Borsjon NRS, Brunneby GNM ZML,, Linkoping,.Vadstena.- VG: Grhstorp NRS, Toreboda NRS. - NA: Arboga,Orebro NRS. - SO: Larslund NRS, NorrkopingNRS ZMA, Nykoping NRS, Runtuna NRS, Trosa,Tuna NRS, Tyreso. - UP: Hdvero NRS, Ingaro, Nacka,Stockholm, Uppsala. - VR: Alster NRS, Arvika, Eda,Forsvik SE Saffle, Grums NRS, Aris. - DR: BrunnsvikNRS, Ludvika NRS, Mora NRS, Oxberg NRS. - GA:Ockelbo NRS. - HA: Delsbo. - VB: Degerfors S Vindelnd 26.VIII.1935 Forsslund leg. NRS.Correction. A record for this species in Lapland(Friese 1902, referring to Wahlberg 1855b) is incorrectand due to linguistic misunderstanding.The total distributionof the subspecies P. quadricolorglobosus Eversmann. Fennoscandia - Denmark- The Baltic - European USSR.Sparse informationabout the Danish population(Hammer1970; a few dark coloured specimensexamined by me) indicate that the subspeciesmay intergrade with P. q. meridionalis inDenmark; in SE Europe it intergrades with P. q.rossicus (Popov 1931).World distribution of P. quadricolor Lepeletier.In Europe restricted to mountain islets in Spain,the Pyrenees, France, northern Italy, the Alps,northern Balkans and Slovakia; north of the Alpswidely distributed north to Fennoscandia and toabout 62" in European USSR. Discontinously occurringfrom Turkey NE to Transbaykal.The present information about distribution ofthe species indicates a division into two geographicallysegregated groups similar to that of the hostB. soroeensis (LOken 1973). P. q. meridionalis occurswithin the range of the red-tailed B. s. proteus(Richards 1928; Tkalcfl 1969;Beinig 1973) and theislet distribution of the remaining subspecies iswithin the distribution of the white-tailed B. s.soroeensis.List of localities. l'{orway. @: Moss: JelOy ZMA. -Oslo: ZMO. - On: Vestre Slidre: Valdres ZMO; Dovre:Dovrefjell d Boheman leg. 1.832 NRS. - TEi: Lunde:Vommestol. AAy: Tvedestrand: Nes Verk TRM.AAi: Bygland: AustadZldO.- Ry: Strand: Tau; Fister:Fister; Tysvrer: Nedstrand. Ri: Forsand: Le"rang; Ardal:Eskeland, Ardal; Hjelmeland; Suldal: Aimjodlo byMostol 700 m. - HOy: Bergen: ZMB TRM, DolvikZ}y'.A, Fantoft, Hop, Minde, Sanddalen, SkipanesZ}dA. HOi: Ullensvang: DjOnno, IJtne; Ulvik: Hjeltnes.- SFy: Gloppen: Lotsberg. SFi: Stryn, Flo. - MRy:Orskog:Skjodje ZMO. MRi: Norddal: Valldal. Psithyrus (Fernaldaepsithyrus) sylvestrisUnrevised records: SFi : Luster: Jostedalen(Wahlberg 1855b). HOy: Askoy, islands in Feiefjord Lepeletierand HOi: Voss (Lie Pettersen 1901, 1905). Ry: Eigersund:Heskestad, Ri: Suldal: Jelsa, HOi: Odda and SFi:Figs. I1,23, 30.Balestrand: Balholm (Meidell L934).Correction. A female from VAy: Kristiansand: RandesundZMO (Richards 1928) is revised to P. campes-Fargeau , 1832:377, 1841 :429. Type locality: Describ_edPsithyrus quadricolor var. sylvestris Lepeletier de St.tris.on specimens from the Pyrenees and Paris. Type:2 d dSweden SK: Andrarum, Brosarp, Helsingborg, Hiickeberga,Jordholmen ESE Malmo ZMA,, Klostersig_el, types (Kellner-Pillaut in litt.)labelled "ex coll Spinola scat'Il712" SCT! might be syn-Kiseberga NRS, Lund, Nytorp, Sandhammaren NRS, Apathus quadricolor auctt. nec P. q.quadricolorTorekov, Torna-Hdllestad, Vallf,ker, Vitemolla, Vomb, Lepeletier; Thomson 1870 (partim).

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ENT. SCAND. SUPPL .23(1984)Scandinavian species of Psithyrus Lepeletier 27Psithyrus quadricolor auctt. nec Lepeletier; Sp. a median longitudinal line usually forming an indistinctlyinterrupted keel (Fig. 118). Sto with nar-Schneider 1898, 1909, 1918; Friese 1902; Aurivillius1903; Strand 1904; Soot-Ryen 1925; Bengtsson 1931.Apathus sylvestris (Lepeletier); Thomson 1872; Siebkerow angled callosities (Fig. 11C).1880; Strand 1898b.Colour pattern Pile of vertex with a variable admixtureof yellow. Broad yellow collar laterallyPsithyrus silvestrus Lepeletier; Richards L928; Meidell1934; Barendrecht l94l;Tjeder 1954; Ander 1963,,1965; covering adjacent border of episternum. LateralLoken 1966b; Kullenberg et al. 1970; Bringer 1973;Lundberg & Svensson 1977; Svensson 1980.patches of T3 more or less extending inwards alongextreme distal margin and T4 with white hairs. T6and St6 between and outside callosities covered bypale ferruginous to pale brownish pubescence.Taxonomical remarksCoat otherwise black.P. sylvestris var. citrinus Schmiedeknecht, P. s. Variation. Pile of vertex ranging from all yellowvar. albicans Richards and P. s. var. atrithorax to all black. Pile on T3 ranging from all white toRichards one or several of which mentioned by rarely all black. Fringes of legs, particularly Bt,Sp. Schneider (1898, 1909), Richards (1928), pale yellowish or tipped so.Bahrendrecht (1941) are infrasubspecific formsoccurring occasionally throughout Scandinavia.MALEFirst S candinav ian recor dsNorway: Tromso (Sp. Schneider 1898).Sweden: Distributed north to Lapland (Thomson1872).DescriptionFEMALEBody small. Coat shaggy.Measurement. N _ 25. Rl: 3.56 mm t 0.129,3.30-3.80 mm; Iw: 4.47 + 0.209 mm, 4.2V5.00mm; Hw : 4.23 t 0. 123 mm, 4.0M.40 mm; hamulinumbe r: 21 .52 + 1 . 53 L, 19-24; Lh: 1 . 48 + 0. 081 ,I.23-1.60 mm; Dh: 0.072 t 0.004 mm, 0.067-0.081 mm.Head. Malar space usually just longer thanA:*o; integument smooth with irregular, closepunctures of varying sizes anteriorly fading into alarge almost impunctate area. Main keel of mandiblewell developed, reaching margin or almost so,tapering distally (Fig. 11A). Upper gena with distinct,usually dense mixture of fine and coarsepunctures; genal furrow indistinct, occasionallyabsent. Surface of scapus smooth, shiny, rathersparsely haired. A::Aa:As : 17:10.5:14, Ae-r z 7slong as their width.Hind leg. Tibia with longest hairs of posteriorfringe about 1 .5 x its greatest width ; Bt withlongest hairs about as long as its greatest width.Gaster. Punctation of T5 as that of Tz-q orslightly sparser. T6 closely punctured except alongBody small. Coat shaggy.Measurements. N _ 50. Rl: 3.23 + 0.148 ffiffi,2.80-3.55 mm, Iw: 3.51 + 0.471 mm, 3.00-3.90mm; Hw :3.92 + 0.139 mm ,3.551.30 mm; hamulinumber: 18.30 + 1.389, 15-21; Lh: I.24 + 0.073mm, 1.10-1.38 mm; Dh:0.072 + 0.006 mm,0.06-0.09 mm.Head. Malar space about as long as distal widthequal to A3a + or just longer; integument smoothwith rather sparse, fine and medium-sized punctationdistally fading into an almost impunctate ateaof various size. Upper gena usually with close, distinctand mainly coarse punctation; genal furrowindistinct. Scapus shiny, smooth rather sparselyhaired. A3:Aa:A5 _ 16:lI:17, Ao-r: about 1.5xtheir distal width.Hind lrg. Tibia with longest hairs in posteriorfringe about 1.5x its greatest width; Bt's longesthairs almost 2 x its greatest width.Gaster. St6 distally sligthly protuberant, truncate,more or less bilobed (Fig. 23A). Stz-a andgenitalia as in Fig .238-F-; inner basal extensionsof gonocoxite usually slender, longer than wide;gonostylus varying in shape (Fig. 23D-E), innermargin concave, usually about as long wide andabout as long as that part of volsella visible fromabove (Fig . 23D).Colour pattern Pile of vertex with varying admixtureof yellow. Broad yellow collar laterallycovering upper part of episternum. Tl with varyingadmixture of whitish and yellow hairs. Hairs ofT z-q and lateral patches of T5 whitish, hairs of T6-7predominantly ferruginous or whitish. Otherwisecoat black.

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ENT. SCAND. SUPPL.23 (1e84) Scandinavian species of Psithyrus Lepeletier 29Fig. 30. P. sylvestrisLepeletier. Inset: The host P. pratorum (L.), updated 1983. Legend as in Fig. 25.

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ENT. SCAND. SUPPL.23 (1984) ScandinavianDF GHFig.3l. P. flavidus (Eversmann). Variations in the colour pattern ofthe Fennoscandian populations, see the text. A-G:males,H-I:females.Black:black,o:yellow,o:black,verticallines:ferruginous.and melanics (Fig. 31F-G). The pie chart demonstratesa pronounced tendency towardsmelanism throughout the Scandinavian peninsula,intermediates and melanic specimens everywherebeing in majority. The examined Finnish males indicatea slight decrease in melanism to the east.However, more Finnish males and representativematerial from the USSR are needed to evaluatethe proportion of melanism east of Scandinavia.Melanism occurs at least as far east as Tobolsk inwestern Siberia (Popov 1931; Pittioni 1942).ConclusionThe rather stable colour pattern of the femalesargues against subdividing the Scandinavianpopulation of P. flavidus. It belongs to the nominatesubspecies as indicated by Popov (1931).Host?Bombus jonellus (Kirby) (Brinck 1951). TwoP. flavidus Q Q and one B. jonellus Q were foundin a colony of B. ionellus located in an old rodent's

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ENT. SCAND. SUppL. 23 (1984) Scandinavian species of. Psithyrus Lepeletier 33Table.I. Proportion and degree of melanism in P. /avldus (Eversmann). Area = province or district thereof (cf. Fig.JJ I.Number of individualsTotalPer centAreaTvp.formintermedeatesmel.formnumbertyp.formintermedeatesmel.formNorway:OsOnBvHOiSFiTRiFO13101424T93775l69343031428 1 326 1268 11 0 07 1 424 1713 15507354707268703715443014t715Sweden:DRJAVBNBAsLYPILUTOFinland:KbokobLKLeLiUSSR:Petsamo43130052t7t9114461852731132353157t992l9104932235603477883811171201533 t229 101 7 6r22 256 05 7 9t2 17r20 14t76 11[ , 936 3914 2923 2679 238 6 325t68210652650925136882586444623725680294950825873229373015060Total t45 4t8 361, 924nest (Sweden: LU: Virihaure 8.VII .1944 ZI$-4L).The colony was not analyzed and the presence ofa B. jonellus ! makes it uncertain whether it wasusurped by P. flavidus or the inquilines were temporarilyresting in the nest. The 2 specimens arethe only records of P. flavidus observed in a bumblebee colony.Bombus lapponicus s.l. ( - B. lapponicus (Fabricius)and B. monticola (Smith)) has been consideredas host of P. flavidus (Richards 1928; Pittioni1942, 1943; Tkalcfl 1969; Lundberg &Svensson 1977; etc.) based on the fact that B. lapponicuss.l. and the inquiline are boreo-alpinespecies locally observed in the same areas. Pittioni(1942) in recognizing that P. flavidus has a widerdistribution than B. lapponicus s.l. in Finland,proposed B. cingulatus Wahlberg as a secondhost. Pekkarinen et al. (1981) confirmed that in EFennoscandia P. flavidus occurs markedly furthersouth than B. lapponicus s.l.; they suggested thewidespread B. jonellus might be the only host ofthis inquiline in Finland. The present study revealsthat P. flavidus has a wider distribution in Scandinaviathan that of of B. lapponicus s.l. or B. cingulatus(cf. below).Until more colonies invaded by P. flavidus aterecorded it remains question whether B. ionellusis preferred host (cf. p . 34). The inquiline resemblesB. jonellus in colour pattern and is widely distributedwithin the range of this Bombus species(cf. below).,.Difference in colour pattern and inecological valence may, however, not prevent thespecies from attacking colonies of B. lapponicuss.l. or B. cingulatus. The 3 Bombus speciessuggested as hosts of this inquiline are all belongingto the subgenus Pyrobombus Dalla Torre.

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ENT. SCAND. SUPPL.23 (1984) Scandinavian species of Psithyrus Lepeletier 35Fig.33. P. flavidas (Eversmann). Inset: The supposed host B. jonellus (Kirby) updated 1983. Legend as in Fig. 25.

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ENT. SCAND. SUPPL.23 (1984)Variation. Hairs on T1 black. Lunate part of T3with black hairs varying in size, occasionally hairsof T3 all white. Ts with small whitish patcheslaterad. Pale hairs on gaster occasionally moreyellow than whitish.MALECoat rather even. Body large.Measurements. N - 40. Rl: 3.78 + 0.083 ffiffi,3.35-415 mm; IW:3.70 + 0.183 mm,3.254.05mm; Hw:4.22 + 0.l32mm, 3.8(H.50 mm; hamulinumber: 17.58 + 1.130,1520; Lh: 1.30 + 0.071,l.13-I.46 mm; Dh: 0.080 + 0.006 mm, 0.068-0.092 mm.Head. Malar space slightly longer than wide,about as long as A3aa; integument smooth withuniform moderate distribution of mainly minutepunctures becoming sparser in front of inferiorlower margin of eye and close to distal margin.Upper gena with irregular, shallow punctureswhich are particularly coarse and close along borderof eye; genal furrow distinct. Scapus coarselypunctured, densely haired. ,A'3:Aa:,A5 :16:12.5:20, Ao-r: 1.75x their distal width.Hind leg.Tibia and Bt with longest hairs in posteriorfringe rarely longer than 1.5 x their greatestwidth.Gaster. St6 distally slightly protuberant with elevatedmargin more often inconspicuously bilobedthan truncate. Stz-s and genitalia as in Fig. 22;inner basal extension of gonocoxite broad, flattened,slightly or not longer than broad (Fig.22C); gonostylus varying in shape (Fig. 22C-E),usually broad, longer than that part of volsella visiblefrom above, inner margin more or less concaveand coarsely punctate (Fig.22C); inner basalextension always with tuft of long fluffy hairs;basal part of volsella with markedly denser fringesthan the remaining species of the subgenus.Colour pattern. Pile of vertex laterally more orless extending along preoccipetal ridge, collar coveringadjacent border of episternum and extendingbackwards below tegula, yellow. Distal fringeof scutellum with slight admixture of whitish orpale yellow hairs, but more or less concealed byblack hairs. Hairs of T1 , extreme posterior marginof Tz, Tz-+, To-t and fringes of St+--e, whitish.Otherwise coat black.Variation. Hairs of T1 yellow or with varying admixtureof black hairs. Hairs of T3-a more or lessScandinavian species of Psithyrus Lepeletier 37yellow. Pile on T6-7 ranging from all whitish to allferruginous or black.HostBombus hypnorum (L.) (Stoeckert 1954;Roseler 1972; Cederberg 1976; Mjelde in litt.).Flight seasonSouthern Sweden: Hibernating female: 3 May-1 1 September; female progeny: ? 5 August;male:25 june - 15 September.DistributionScandinavia (Fig . 34). Sparsely distributedthroughout the peninsula north to almost 70'N.Occurring from sea level to timberline; rarely observedin subalpine/subarctic biotopes. Rare orabsent along the extreme west coast of Norwaywhere the frequency of the host B. hypnorum ismarkedly lower than inland.A total of 110 Norwegian and about 190Swedish specimens were examined.List of localities. I,,lorway. A. Onsoy: Rgrvik; Moss:Jel6y ZMA; Hobol: Tomter. - AK: Asker; Brerum:Burudvann; Holand: Skogen; Nittedal: Hakadal kirke.Oslo: TRM ZMB. - HEn: Rendal: Solbakken ZMO. -Os: Sondre Land: Enger MJEN; Nordre Land: Dokka;Gjovik: Biri ; @yer: Tret"ten-Losna, Oyer. - Bv: Nore &Uvdal: SkjOnne kirke; Al: ZMO. - VE: Skoger: Konnerud;Notteroy: Teie. - TEy: Lunde: Vommestdl. TEi:Kviteseid: Kviteseid; Tinn: VAer. - AAy: Grimstad:Dommesmoen. AAi: Bygland : L4ndal; Valle: Kvestad,Rygnestad. - VAi: Hregeland. - HOy: Os: Stendalsetra;Bergen: Fantoft, Ytre Arna. HOi: Rgldal: ROldal; Ullensvang:Fresvik, Lofthus, Ringoyi Ulvik: Hjeltnes,Ulvik. - SFi: Balestrand: Flesje; Luster: Leirdola, Okleviki.- STi: Oppdal: Engan; Trondheim. - NTi:Merflker: Vardetun; Hegra: Hegra st.; Snf,sa: Hegge. -TRy: Troms@ TRM. TRi: Bardu: Stromsmo TRM;Milselv: Bjerkeng TRM, Nordmo TRM.(Jnrevised records (Varley, in litt.): SFi: Balestrand:Balholm, Mundal.Sweden SK. Alnarp, Brunneby, Fulltofta ZMA, HelsingborgNRS, Horby, Kulla-Otorp(-Kulla-Gunnarstorp),Kullaberg, Lund ZM^, Skiilderviken NRS,Strandbaden, Vegeholm NRS, Ahus NRS. - BL: RonnebyZMB, Sjoarp NRS. - HA: Enslov, Steninge. - SM:Aneboda, Grdnna, Horeda I'{RS, Jonkoping NSS,Ljungby Zl/.A, Osterkorsberga ZM\, ZMU. - OL:Borgholm NRS, Morbylinga NRS. - OG: Borensberg.- VG: Hdndene NRS, Kinnekulle E Lidkoping, StenumE Skara NRS, Skatis E Gothenburg. ZMA, UtbyniisNRS. BO: Brastad. Sdmstad NRS. - SO: Enhorna NRS,

'SZ'8lC ul su pua8al 'gg6y pelepdn '('1) untoud,fi1 'g tsoq oqJ :losul repreuqrg'dg sntSauou 'd 'tt'ElJ*'r (rso ) tz '-Iddns'ctNvls'rNE ua4@j putsv 8t

ENT. SCAND. SUPPL.23 (1984) Scandinavian species of Psithyrus Lepeletier 39Table IL Host recordsfor European Psithyrus.N: numberofBozbus colonies usurped byPsitlryrus: + = 5-10colonies,** : > 10 colonies, 0 : number ofcolonies recorded in Scandinavia, x : nonscandinavian species.Social parasiteP. bohemicusP. vestalisP. barbutellusP.rupestrisP. campestrisP. quadricolorP. sylvestrisP. flavidusP. norvegicusHostB. (s. str.) lucorumB. (s. str.) terrestrisB. ( Megabombus) hortorumB. (Pyrobombus) hypnorumB. ( Melanobombus) lapidariusB. ( Melanobombus) sichelissp. alticola"Thor ac ob o mb us ) p as cuo rumThoracob omb us ) sy lv arumThor ac ob o mb us ) p as cuo rumThoracob omb us ) humilisRhodobombus) pomorumPyrobombus) pratorumS or o e ensib omb us ) s or o eensisPyrobombus) pratorumPyrobombus) jonellusPyrobombus) jonellusPyrobombus) hypnorumN++++-r1+ +(1)222++T11aJ+ +(2)1(? 1)+ (3)Tyreso NRS, Uttran NRS, Valla NRS. - UP: BjorklingeNRS, Ingaro, Fogdo NRS, Stockholm NRS ZML,,Svartsjo, Uppsala, Vallentuna NRS, Yxlan. VR: Arvika.- DR: Bingsjo NRS, Brunnsvik NRS, Falun NRS,Horndal CBJS, Leksand TJTS, Ludvika NRS, MoraCBJS, Ore iilv TJTS, Orsa, Sandsjo, Siljansborg NRS,Sjurberg NRS, Smedbacken, Tiillberg NRS. - GA: HilleNRS, Ockelbo. HA: Bergvik, Fredriksfors NRS,Soderhamn NRS. - JA: Griddede NRS, Hallen, HAllandNRS, Jormlien, Mattmar, Snasahogarna NRS, AreZMIJ. - AN: Junsele ZMB, Kniisjo 65 km NWOrnskoldsvik Zl/rB, SE Asele Z}dB. - VB: SkellefteAZMB, Tdftefl ZMIJ. - NB: Bjorkfors ZMB, Kalix, Ojtbyn.- LY: Sorsele: Ammarnds, Ndckrostjiirn. - LU:Messaure ZMB ZMU, Nattavaaraby ZMB, Njunjes,Vuollerim ZMB. - TO: Abisko BML, DjupvikenLHAS.Unrevised records (Ander, in litt.): SK: Osby. - HA:Akulla NE Varberg. - DS: Bengtsfors, Lingbron. - DB'Vikarebyn.World distribution. Eurosiberian. Central andnorthern Europe - Siberia - Kamchatka - Transbaykal- Northern Mongolia. The only Psithyrussp. recorded in Japan (Ito & Tadauchi 1981).The range of P. norvegicus is within that of B.hypnorum, a Siberian fauna element (LOkenre73).The host-parasite relationship Bombus- PsitiyrusThe host species of the individual Psithyrusspecies mentioned above and the frequency of ursupedbumble bee nests (calculated from the figuresof the host species references) have beencompiled in Tab. II. Each Psithyrus sp. seems toprefer a certain Bombus sp. commonly occurringin Europe. Some species are more specific in theirchoice of host than others.Scent is important for nest-seeking Psithyrusfemales (Slad en I9l2; Pouvreau 1973). RecentlyCederberg ( 1979, 1983) found that P. rupestris followstrail pheromone laid by B. lapidarius, thepreferred host. Further experiments demonstratedthat when given a choice, female P.rupestris at first reacted strongly to trailpheromone laid by P. lapidarius but after a fewminutes they reacted as well to trail pheromoneproduced by Bombus spp. representing 5 differentsubgenera and left that laid by B. lapidarius. P.rupestris occasionally usurps other Bombu.s spp.than the preferred host (Tab. II).According to Reinig (1981) 53 Bombus speciesoccur in Europe (29 in Scandinavia (LOken1973)), of which 14 species have been observedvictimized by Psithyrus (Tab. II). A few moreBombus species repeatedly listed as host specieswithout referring to ursuped colonies (Reinig1935; Pittioni 1942; Panfil ov 1957; Pouvreau 1973;etc.) are not included in the table. It is, however,likely that other Bombus spp. occasionally are occupiedby Psithyrus. P. bohemicus, P. vestalis, P.norvegicus and, possibly, P. quadricolor may be

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ENT. SCAND. SUPPL.23 (r984)Scandinavian species of Psithyrus Lepeletier4lperiuntur. ..(new species added by Geoffroy) . 544pp.Paris.Franklin, H. J .1911. New North American Bombidae. -Trans.Am.ent. Soc . 37 :157-168.Latr. and'Psithyrus Lep. males and localization ofsite of production of the secretion . - Zoon, suppl.1:23-30.Lepeletier, A. de Saint-Fargeau. 1t132. Observations surFrid€n, F., Eskilsson , L. & Bingefors, S. 1962. I'ouvrage intituld: " Bombi Scandinaviae MonographicdTractati, etc.., d Gustav Dahlbom. LondoniBumblebees and red clover pollination in CentralSweden. I. Int. symposium on pollination 1960. - Gothorum, 1832; "..... Ann.Soc.ent. FranceMeddn.Sver. Froodlare-forbund 7: 17-26.Friese, H. 1902.Die arktischen Hymenopteren, mit Ausschlussder Tenthrediniden. - Fauna arct. 2:439--l:36G382.1841. Histoire Naturelle des insectes. Hym€nopteres.I. Genre Psithyrus. 2:424 437 .498.Frison, T. H. 1927. A contribution to our knowledge ofthe relationship of the Bremidae of America north ofLie-Pettersen, O. J. 1901. Bidrag til kundskaben omVestlandets Bombus- & Psithyrus-arter. - BergensMus.Arb. 1900, 3:1-19.Mexico (Hymenoptera) Trans.Am.ent.Soc. - 1902. Biologische Beobachtungen an norwegischen53:5l-78.Hummeln. - Bergen Mus.Arb. 1901, 6:1-10.Haeseler, V. 1970. Zur Wirtswahl des Psithyrus rupestris - 1905. Entomologisk bidrag til skjrergaardsfaunaen i(Fabr.) (Hymenoptera: Apidae) Faunist. - det vestlige Norge. - Bergen Mus.Arb. 1904,11:1-Okol.Mitt .3:29G298.Hammer, K. 1970. Arternes biologi og kendetegn. InHammer, K. & Holm, Sv. N.: Danske humlebier og-25.1907. Neue Beitrdge zur Biologie d. norwegischenHummeln. - Bergen Mus.Arb. 1906 ,9:142.snyltehumler. - Natur og Museum I4(2-!:aA3.Hasselrot, T. 1960. Studies on Swedish bumblebees.(Genus Bombus Latr.). - Opusc.ent.Suppl. 17:l-Loken , A. I966a. Notes on Fabrician speciesof BombusLatr. and Psithyrus Lep., with designation of lectotypes(Hym., Apidae). - Ent.Meddr. 34:199-206.192.- I966b . Ekskursjonsberetning. Insekter og- 1962. Humlor och bin i froodlingen. Vira vanligastehumlor. - HushAlln.szillsk.Forb.Smiskr. 51 :8-13.arachnoider samlet under det 13. Nordiske Entomologmotetsekskursjon til Flflm (SFi: Aurland)Hoffer, E. 1889. Die Schmarotzerhummeln Steiermarks.Lebensgeschichte und Beschreibung derselben.l3-t6386.august 1965. - Norsk Ent.Tidsskr. L3:371-- Mitt.Naturw.Ver.Steiermarks (1888) 25:82- - 1973. Studies on Scandinavian bumble bees158.Hoppner, H. 1901. Beitrige zur Bienenfauna der(Hymenoptera, Apidae). - Norsk ent.Tidsskr .20:1-218.Ltineburger Heide. - Jber.Ver.Naturk. Unterweser - 1978. Notes on the Scandinavian fauna of social1900:922.Aculeates (Hy-., Vespidae and Apidae s.s.). -,ii Husband, R. W., Fischer, R. L. & Porter, T.W. 1980. Norw.J. Ent . 25 :165-170'Description and biology of bumblebees (Hymenoptera:- & Framstad, E. 1983. Contribution to the taxonomyApidae) in Michigan. - Great Lakes Entomol.13:225-239.of. Fernaldaepsithyrus (Hy-.,tomol. Fennica 42:46-50.Apidae). - Acta En-Illiger, J. C. W. 1806. William Kirbys Familien derbienenartigen Insecten mit Zusatzen, Nachweisungenu. Bemerkungen. - Magazin ftir Insektenkunde5:28-175.Ito, M. & Tadauchi, O. 1981. The occurrence ofPsithyrus norvegicus in Kyushu, Southern Japan. -Kontyu 49:715.Jansson, A. 1925. Die Insekten-, Myriapodq.n- undIsopodenfauna der Gotska Sandon. 182pp. Orebro.Lundberg, H. & Svensson, B. G. 1977. Altitudinal distributionof Psithyrus Lep. in relation to that of theirhosts Bombus Latr. in a subalpine/alpine area.Proc. 8th Int.Congr.I.U.S.S.I. 8:305-306.Lundblad, O . 1924. Nigra ord om aculeat' och tubuliferfaunani Lycksele lappmark. - Ent.Tidskr. 45:17-28.May , J . 1937. Bionomie rodO Bombus Latr. a PsithyrusLep. (1) - Casopis Cs.Spol.Ent. 34:115-118.1959. Cmeldci v. CSR jejich bionomie, chow a hospoddisky- 1935. Supplement till Die lnsekten-, Myriapodenundvyznam. 170pp. Praha.Isopodenfauna der Gotska Sandon Meidell, O. 1934. Bier og hu^mler i Rogaland (Apidae,Ent.Tidskr. 56:52-87.Hy* ) - Stavanger Mus.Arsh . 43(1932-33):85-131.Kirby, W. 7802. Monographia Apum Angliae.2.387pp. M6czin, L. 1977. Das Nest von Bombus alticolaIpswichKriechbaumer (Hymenoptera: Apidae). - ActaKriechbaumer, J. 1854. Beitriige zur Kenntnis deutscherBienen. - Linnaean Ent. 9:170.-188.Kruseman, G. 1959. Enkele zoogeografische opmerkingenover de hommelfauna van Zweden (Hym. ,Apid.) . - Ent. Ber. 19:49-51 .Biol. Szeged . 23:133-138.Muchardt, H. 1904. Bidrag til humlornas och snylthumlornasutbredelse. - Ent.Tidskr. 15:204.Mi.iller, M. 1913. Beitrdge ztrr Kenntnis unserer Hummeln.- Arch.Naturgesch . A. 79:17UI23.Kullenberg, B. 7973. Field experiments with chemical - 1936. Psithyrus barbutellus Kirby und P. maxillosussexual attractants on Aculeate Hymenoptera males. Klug. (Apid., Hy-.). Mitt.Deutsch.Ent.Ges.II. - Zoon, suppl .l:3142.(193s) 6:73-76.- ,Bergstrom, G. & Stiillberg-Stenhagen, S. 1970. Vol- Nerdn, C. H. 1892. Entomologiska sommarstudier. -atile components of the cephalic marking secretion Ent.Tidskr. 13:97-116.of male bumble bees. - Acta Chem.Scand. 24:1481- Newman, E. 1835. Attempted division of British insects1483. into natural orders. - Ent.Mag. (1834) 2:379431., Bringer, B., Carlberg, B. & Cederberg, B. N(ordisk)U(tredningsserie).I9TT.NaturgeografiskRel9T3.ObservationsonthescentmarkingbyBombusgionindelning av Norden. B 1977 No 34 130pp.

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ENT. SCAND. SUPPL.23 (1984) Scandinavian species of Psithyrus Lepeletier 43Forh.Videnskselsk.Krist. 1903,8:1-8.in Zetterstedt, J. W.: Insecta lapponica descripta.Svensson, B. G. 1974. Bin (Hymenoptera, Apoidea) 1139columns. Voss, Lipsiae (Leipzig 1838-1840).fflngade i fiillor vid Messaure ekologiska station, Zimsen, E. 1964. The type material of I. C. Fabricius.Lule Lappmark, med anmiirkning om humlornas 656pp. Munksgaard, Copenhagen.fenoloei. - Ent.Tidskr. 95:122-124.- 1979. Patrolling behaviour of bumble bee males in asubalpine/alpine area, Swedish Lapland. - Zoon7 :67-94.- 1980. Species-isolating mechanisms in male bumble Indexbees (Hymenoptera, Apidae). - Abstr.Upps.Diss.-Fac. Sci. 549 :142 .Valid names in Roman type. Synonyms,Thomson, C. G. 1870. Oversigt af Sveriges humlor. - homonyms, infrasubspecific forms in italics. InOpusc. ent. 2:251-26I.Psithyr,rzs only those pages referring to description- 1872. Hymenoptera Scandinaviae. 2. 286pp. Berling,or taxonomic status are indexed .In Bombus onlyLundae (Lund).Tjeder, B. 1954. Ostskinska insekter. II. - Opusc.ent. those pages referring to host species are indexed.19:205-209.Tkalcfl, B. 1967. Zur Hummelfauna der UmgebungKuku-Nors (Hymenoptera, Bombinae) psithvrusCas.Cdske Spol.ent. 58:344-379 .- 1962. Bourdofis capturds par M. Ed. Klinzigen 1960et196l. - Bull.Soc.ent. Mulhouse 1962:44 48.- 1969. Ergebnisse der Albanien-Expedition 1961 desdeutschen entomologischen Institutes. 78 Beitrag.Hymenoptera: Apidae IV (Bombinae). - Beitr.Ent.19 :887-916.- 1974a. Eine Hummel-Ausbeute aus dem Nepal-Himalaya (Insecta, Hymenoptera, Apoidea, Bombinae).- Senckenbergiana biol. 55:31I-349.- 7974b . Ergebnisse der 1 . und 2. mongolischtschechoslowakischenentomologisch-botanischenExpedition in der Mongolei. Nr. 29:Hymenoptera,Apoidea, Bombinae. Acta faun.ent.Mus.Nat.Pragae 15:25-57 .Van Honk, C., Roseler, P.-F., Velthuis, H. &MalotauX, M. 1981. The conquest of a Bombus terrestriscolony by a Psithyrus vestalis female. -Apidologie 12:5747 .Wagner, R. 1971. Die Verdnderung der HummelfaunaCuxhavens in diesem Jahrhundert. Der Versucheiner Deutung. - Ent.Mitt.ZooL Mus. Hamburg 4.No.75.Wahlberg, P. F. 1855a. Iakttagelser ofver de nordiskahumlorna. - Forh.Skand.naturf. 6. mote, Stockholm1851 . Bilagan C:23V233 .- 1855b. Iakttagelser o.ch anmdrkningar ofver de nordiskahumlorna. - Ofvers.K.VetenskAkad.Handl.Ir:r99-211. (1854).Wahlgren, E. 1915. Det oltindska alvarets djurvdrld. I. -Ark.ZooL 9:1-135.- t9t7-1918. Det oldndska alvarets djurvdrld. II.Ark.ZooL 11(I):l-129.Winston, M. L. & Michener, C. D.1977. Dual origin ofhighly social behaviour among bees (Hymenoptera/Apidae/Phylogeny) . Proc. Natn. Acad. Sci. USA74:1135-1I37 .Wollin, H. 1975. Kring nornans biologi. - Fauna ochFlora 70:89-98.Yarrow, I. H. H. 1968. Kirby's species of British bees;designation of holotypes and selection of lectotypes.Part I. Introduction and the species of Apis Linnaeusnow included in the genera Bombus Latreille andPsithyrus Lepeletier. - Proc.R.ent.Soc.Lond. (B)37:9-15.Zetterstedt, J. W. 1838. Bombus Fabr. columns 473474aestivalis Panzeraestivalis Dahlbom nec Panzeralbicans Richardsalpium Richardsalbinella KirbyAllopsithyrus Popovamo enus SchmiedeknechtApathus Newmanarenaria Panzerarvernicrzs RichardsAshtonipsithyrus Frisonashtoni (Cresson)atricolor Richardsatrithorax RichardsWl#if.,#N:. :.' .",\ni";::):,ffll"?n::::::citrinus Schmiedeknechtdistinctus P€rezFernaldaepsithyrus Frisonfernaldae Franklinflavidus Eversmannflavidus (Eversmann)francisana Kirbyfrutetorum Panzerfruteto rum : Zetterstedt nec P anzerglobosus Eversmannintercalaris Miillerlato fas ciatus Pittionileana Kirbylissonurus Thomsonmaxillosus (Klug) ...Metapsithyrus Popovmeridionalis Richards ..nemorum Fabriciusnorvegicus Sp. Schneiderobscurus SeidlPsithyrus Lepeletier (s. str.) ..quadricolor Lepeletierquadricolor auctt. nec Lepeletierrosiella Kirby1082728t4t28T4142388282728121281818t22782222282819t423232328t92812182383614l42325t9

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ENT. SCAND. SUPPL .23(1984) Scandinavian species of Psithyrus Lepeletier 45Fig. 35. Division of Fennoscandia and Denmark. For abbreviations of provinces, see p.44.Manus received December 1983Journal and subscription to reprints of particular groups to be ordered from:Publishing House of the Swedish Research Councils,P.O. Box 6710, S-113 85 Stockholm, Sweden.