Scandinavian species of the genus Psithyrus Lepeletier ... species of the genus Psithyrus Lepeletier (Hymenoptera: Apidae) Entomologica Scandinavica Supplement No. 23 tts€-s0r0 NSSI

ASTRID I-OKEN

Scandinavian species of the genusPsithyrus Lepeletier(Hymenoptera: Apidae)

Entomologica Scandinavica Supplement No. 23

tts€-s0r0 NSSI

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Scandinavian species of the genu s Psithyru, Lepeletie. '* W*(Hymenoptera: Apidae)

ASTRIDLAKEN

IntroductionThe genus Psithyrus Lepeletier comprises about

45 species described from the palaearctic, nearc-

tic, oriental and neotropical regions. The bulk of

the species is confined to the palaearctic region

where 30 taxa at present have specific status (e.g.

Popov l93l; Thalcfl 1961 ,,I974a). The greatest di-

versity is in the east part of the region; only 10

species occur in Europe (Reinig 1981), 9 of which

are found in Scandinavia, i.e. Norway and Swe-

den.The species in this genus, the cuckoo bumble

bees, are social parasites depending on the social

bumble bees , Bombus Latreille for their exis-

tence. They havb been recorded from sea level to

subalpine/subarctic regions but in general they do

not occur at the peripheries of the ranges of their

potential hosts.Variations in climate, structure and topography

cause great ecological differences throughout

Scandinavia which extends from 55"21'N to

L6ken, A.: Scandinavian species of the genus Psithyrus Lepeletier (Hymenoptera: Apidae).

Ent. scand. Suppl .23: I45. Lund, Sweden23 December 1984. ISSN 0105-3574.

Keys, synonymy, bibliographies, descriptions, distributions and host relationglri.ns are given

for-the-g ScanOinavian speiies of. Psithyrus Lepeletier,viz. P. bohemicus (Seidl), P. vestalis

(Geoffroy in Fourcroyf, P. barbutellus (Kirby) , P. rupestr,'t-(F3pricius) , P. campes.tris(Punr".), p. quadricolor Lepeletier, P. sylvestris Lepeletrer,, P. flavi.du1(Eversm.unQ and P.

iorvegicns Sp. Schneider. The only European species not occurring in Scandinavia, P. maxil'

losus (Kug), is included in the keys for the sake of completeness. Lectotypes are designated

for P. quadricolor meridionalis Richards and P. norvegicus.Key words: Hymenoptera, Apidae, Psithyrus, taxonomy, fauna, host selection, Scandinavia.

A. Lgken, Univ. of Oslo, Dept. of Bio l . , Div. of ZooL, P.O.Box 1050, Bl indern, N-0316 Oslo

3. Norway.

71"I1'N (NU 1977). As the inquilines are mar-

kedly less abundant than their industrious hosts,

information on distribution, geographic variation,

ecology, etc. of individual Psithyrus spp. requires

repeated observations in a variety of habitats.

The present work is an effort to provide a basis

for further studies of Psithyrus. Owing to the cur-

rent interest in this host/parasite relationship reli-

able references to the European host species are

included, i.e. references to observations of bum-

ble bee colonies usurped by the Psithyrus spp. in

question.

Historical reviewIn dividing the Scandinavian bumble bees into

two sections Dahlbom (1832) unintentionally

separated the social bumble bees from the in-

quilines. Lepeletier (1832) subsequently claimed

that the taxa listed in Dahlbom's divisio Z:da con-

Ent scalrd

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ENT. SCAND. SUPPL .23 (1984)

Methods

Taxonomy

Keys to the genera of Bombinae and to the

species of European Psithyrus are provided. The

papers of Popov (1931) and Pittioni (1939) were

consulted. The nomenclature of Bombus follows

Loken (1973).

M orpholo gical terms, abbreviations

The terminology follows Loken (1973). Ab-breviations and additional terms:Avtz in female, Ar-r: in male: Individual seg-

ments of antennae.Bt: Basitarsus.Genal furrow: Border between vertex and uppergena (Fig . 4).Str-e in female, Str-z in male: Individual gastral

sternites.Tr-o in female,T p7 in male: Individual gastral ter-gites.Upper gena: Part of gena outside genal furrow and

upper part of eye.

Measurements

The following measurements were taken in

both 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 pre-

sented by number (N), mean, * standard devia-

tion and range, and are given in mm. Those

marked g-k are expressed in micrometer units as

only the proport ions g:h: i and j :k are used.

Interalar width is chosen as an indicator of body

size and the following categories were used:

Female: Body large, mean Iw >5.00 mm; body

of medium size Iw 4.60-5.00 mrn; body small, Iw

<4.60 mm.Male: Body large, mean Iw >3.70 mm; body of

medium size, mean Iw 3.55-3.70 mm; body small ,

mean Iw <3.55 mm.If not otherwise mentioned the measurements

refer to populations in SE Norway and S Sweden.

Distribution

The presentation of maps, list of localities and

other references to the occurrence of the species

Scandinavian species of Psithyrus Lepeletier 3

follow Loken (1973) except that regional division

follows NU (1977). Danish and Finnish distribu-

tions indicated on the maps are from Hammer(1970) and Pekkarinen et al. (1981).

If not otherwise mentioned the general distribu-

tion of the individual species is taken from Popov(1931), Panfilov (1957), Reinig (1971), Tkalcfl( I974b) and Rasmont (1983).

BiologyThe life cycle of Psithyr,zs is broadly similar to

that of. Bombus (Alford 1975). Being an obligate

social parasite, however, Psithyrus is unable to

build up a colony of its own. The females lack pol-

len collecting apparatus and wax producing

glands; they cannot produce a worker caste and

usurp a bumble bee colony to raise the next gener-

ation (Pouvreau 1973; Alford 1975). The female

inquiline emerges from hibernation later then her

host (Postner 1952; Reinig 1973; Husband et al.

1980). She has great success in invading a nest

when the colony is not too small but has a suffi-

cient number of workers to rear her brood; if too

populous she may lose in fights with the workers

(Pouvreau 1973; Alford 1975).

PhenologyThe flight activity of hibernating females is gen-

erally confined to late spring - early summer; such

females are occasionally observed later in the sea-

son, however (Alfken I9I3; Pittioni & Schmidt

1942; Wagn er 1971). The young females are rarely

recorded in the field. They emerge approximately

at the same time as the males (Husband et al.

1980) and are occasionally observed feeding in

flowers during the flight period of the males

(Alfken l9l3; Pittioni & Schmidt 1942; Wagner

reTr).

ClassificationThe family Apidae is restricted in recent classifi-

cation to the subfamily Apinae, Meliponinae and

Bombinae (Winston & Michen er 1977 ). Itt the ho-

larctic region the family is represented by Apinae:

Apis L. and Bombinae: Bombus Latreille and

Psithyrus Lepeletier.

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uaTQ-l putsv v

ENT. SCAND. SUPPL. 23 (1984) Scandinavian species of Psithyrus Lepeletier 5

Figs. 6-15. - 6. P. bohemicus (Seidl) Q; A = right mandible, B = ocellar-orbital field, C : callosities of sternite6.- T.P.vestal is(GeoffroyinFourcroy)9;A:ocel lar-orbitalf ield,B:cal losit iesofstemite6.- S.P.rupestr is(Fabricius)Q,cal losit iesofsternite6.- 9.P.barbutel lus(Kirby)Q,cal losit iesofsternite6.- l0.P.campestr is(Panzer) Q, cal losit iesofsternite6.- 11. P. sylvestr is Lepelet ier Q;A: r ightmandible,B = tergite6,C = cal-los i t ieso fs te rn i te6 . - 12 .P.norveg icusSp.Schne iderQ;A: te rg i te6 ,B=ca l los i t ieso fs te rn i te6 . - 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.I2B P. (Fer-naldaepsithyrus) norvegicus Sp. Schneider p. 36Hind tibia and Bt with longest hairs in posteriorfringe markedly longer than their greatest width. T6inconspicuously or not keeled. Callosit ies as inF igs . 13 -14 . . . . . . . 9

9. Callosit ies 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 ferruginous-haired, often with an admixture of whitish hairsP. (Fernaldaepsithyrus) quadricolor Lepeletier p. 23

MALES

1. Malar space shorter than 3lq the distal width. Stz-aand genitalia as in Figs. 16-17; penis valve nothooked beneath . . . . . . . . . . Ashtonipsi thyrus Fr ison 2Malar space longer than 3lq the distal width. Stz-sand genitalia as in Figs. 18-24; penis valve hookedbeneath 3

2. 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-

7 A

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ENT. SCAND. SUPPL .23 (1984) Scandinavian species of Psithyrus

Figs.2l-24. Fernaldaepsithyras Frison; sternite 6, right half of male sternite 7-8 and genitalia. - )il . P. quadricolorLJpelet ierd;A:steiniteT,B:sternite3,6:pdsteriorpartofgenital ia.- 22.P.norvegrcasSp.Schneiderd;A^:sterniteT,B=stemitei3,6=genital ia,D-E:postei iorpartofgenital ia.- 23.P:sy. lvestr isl-epelet ierd;A: sternite6, B = sterniteT, C =-sterniteS, p: lenital is,B: posteriorpartofgenital ia.- 24. P' f lavidus(Eversmann)d ;A:s te rn i te6 ,B=stern i teT,C:s te rn i teS,D-E=poster io rpar to fgen i ta l ia .

Lepelet ier

K/ffiGWGW24E

G

W

or not exceeding their greatest width. Stz-s and genitaliaas in Fig. 18A{. Coat shaggy

P. (Allopsithyrus) barbutellus (Kirby) p. 126. ,A.3 slightly shorter than ,A.5. Hind Bt parallel-sided, ffio-

re than 4x longer than wide. Stz-s and genitalia as inFig. 19A-C . T+4with reddish hairs

i; ;;;k;;d ;;#"{',":# Pr#{;'{:'Jl1:ffi:? X ;' oits greatest width. Stz-s and genitalia as in Fig. 20A{.T +4without reddish hairs

P. (Metapsithyrw Popov) campestris (Panzer) p. 18

7 . 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 8

8. ,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)

Description

FEMALE

Body 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; hamul inumber: 20.75 +1.4I0. , 18-23; Lh: I .63 + 0.102mm, I .45- I .78 mm; Dh: 0.083 + 0.004 f f i f f i ,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 i ts 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 ex-treme distal 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 cres-cent-shaped posterior 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 re-placed by yellow ones.

Scandinavian species of Psithyrus Lepelet ier 9

MALE

Body 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 .90m m ; H w : 4 . 1 3 + 0 . 2 0 4 m m , 3 . 6 M . 4 0 m m ; h a m u l inumber : 17 .15 + 0 .988 , 7G l9 ; Lh : 1 .32 + 0 .079mm, 1.18-1.48 mm; Dh: 0.082 + 0.006 f f i f f i ,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 posteriorfr inge up to 1.5x the segment's greatest width. Bt3.5x longer than i ts 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 latero-posterial patches 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 tip-ped so. Whitish patches of T3 varling in size, occa-sionally whitish hairs extending along extreme dis-tal margin. 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 allwhit ish.

Melanism. Colour variations are too slight tochange the general colour pattern except forsporadically occurring pronounced melanic speci-mens. 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 11

Fig.25. P. bohemicus (Seidl). The distribution in Fennoscandia and Denmark, Inset: The host B. lucorum(I-,)

G;ken 1973) updated iSSf.'O = one or several localities; O : unrevised records (from literature or received inlitt.); X = inaccurate locality. Shaded areas refer to literature indications.

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7.4VI.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 just

longer.Gaster. Tip of gaster not much incurved. T+-s

with rather close, mainly coarse puncturing. T6with close, rather uniform, mainly medium-sizedpunctation; usually with a weak, narrow im-punctate, median longitudinal keel. Sto with flat-tened semi-circular callosities positioned at a dis-tance from 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 yel-low. Hairs of Ta and lateral patches of T5 whitish.T6 and Sto outside the callosities with golden-brown pubescence. Otherwise coat black.

Variation. Pile of vertex ranging from predo-minantly yellow 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 ad-mixture of pale-coloured hairs, especially in thefringes.

MALE

Body 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; hamul inumber: 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.

Scandinavian species of Psithyrus Lepeletier l3

Head. Malar space hardly as long as distalwidth, ?S long as A:*+; integument distal ly im-punctate or almost SO, otherwise irregularlypunctured, punctures coarser centrally thanbasad. Upper gena with rather dense mainly shal-low, coarse punctat ion;genal furrow absent, occa-sional ly indist inct. Or about 2x Oe. Hairs ofscapus about 2x width of segment; ,{3:Aa:45 -

16:10.5:19, Ao-r : near ly 2x thei r d is ta l 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 be-neath.

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 admix-ture of black. Yellow collar extending ll4 to ll2down episternum. Yellow hairs of scutellum moreor less reduced to crescent-shaped posteriorfringe.

Melanism

Both sexes display a slight tendency towardsmelanism which is more pronounced in south-ernmost Sweden (SK, OL) than further north.

Host

Bombus 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 l i t t .) .

Flight season

Southern Sweden: Hibernated female: 9 May -

2 August;female progeny: ? -3 September;male:l J u l y - 5 O c t o b e r .

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Fig.26.P.barbutel lus(Kirby).Inset:ThehostB.hortorw (L.)updated1983.LegendasinFig.25.

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ENT. SCAND. SUPPL.23 (1984) Scandinavian species of Psithyrus Lepeletier 17

Fig.27. P. rupestris (Fabricius). Inset: The host B. lapidarizs (L.), updated 1983. Legend as in Fig. 25.

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8Z , OZ.OI

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nodo4 sru{qqsdn4ay4 snue8qns

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9PP9N '-IWZ YWZ STINS SNNO

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EN uorelsol '11WZ seuEuerl5 ,SUN Bunlun5 'SIIN punslexo 'SUN Eurdol,{p ,SUN glrgry ,SUNpunlsJu-I 'seuEuerlg

N oru81e11 ,SIIN punslupuurg 'SUN e>p,{1to_g :eS - .SUN orqaro ,8iWZ BSollaINBrols 'SSNS dro1u1 'SUN eEoqry tu..r .gu-tg ;;-Sod-rV :yN - 'SuN uueq8reqaln 'SuN pelsr.uorts 165relso{pAs '-IWZ

SUN petsruorls ,SUN pelsreêN .SUNpunsualsêH'suN pelsaqqerc,suN puBrlsllBô8:Og - 'SUN uueldralsg '-IWZ

SUN epoqerol ,SUN

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(ruor) x '-Iddns 'cNVJS 'JNa !

'SuN euuv t:! SUN unre8ulu ,ywz SdN Erequg 'vwz

sgNS SurdoluoN 'suN elelolN ,-lwz awz

SUN Eurdolull 'SUN persolleH ,SdN Eru11 ,ldb ,iU1.fuolsrog 'nWZ -IWZ hlNO Ereqsuerog -SUN nfnlolnf71,15 erlsu^lv :C_e -'1INZ-IWZ SUN oqEurlugL,-IWZSUN ,{qsl4 'eEirgrs 'St{N 1epr981c9ug ,lWZ

SdtorqreT 'ef,-I 'e1s[o1 '-IWZ

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SUN e8urlddeg ,SUNn1gU'3uo11_'oruseg 'run1e sgusred ,-IWZ

SUN eErCI{d-".W 'lg-I 'Eu1dg; '-IWZ SUt t esollse; ,-IINZ

Sl{N r,unrs-Epg

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SUN ruloq8-rog 'dro1s18ueg 'SfgJ tunrsln8lv :lO - .SUN orueu-rg1'SdN sBuBrJ'urueqsre{so'SUN oiq,i5l,porJsllnH '!)wz -Iwz suN wNc euuurc 'slIN plsla 1Y\Ndrotsuurg'SUN erenlSreg :WS -'dnrey,ruNg rgsiuq:vH - '{lôllgH :-Ig - 'suN snqy ,-Iwz

sdN pe}s^ 'pun-I E qruo1 'SUN olôISlUA ,SUtrt glsp1n ,e11oue

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VWZ SUN grulul4tr 'SUN dnrapol ,-IWZ gWZ VWZWNC pun-I'eEuu1e1cu1 'Eraqe11n;1 ,ueEesrelsol)'gnl 'SUN relleH sôH '-IWZ

vWZ EroqEursleH ,Eues-l_oEgg 'a8urllefg '-IWZ

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E1ryzdnrepuog 'SUN er{u>lreg '-IWZ gWZ ereg ;,(qseg'Sl{N d:o1s1e1ry 'runrurpuy 'dreuly :)S .uapats's4sadwn 'd ol pesrêr seA\ (nsost puer15) L6gr.xl.t

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t0t ropreuqcs 'dS) ue8reg :{OH '(tOOt 'I06I uesrel}od-elT) sgrellenj:ue8reg j{Off pue ddepl - :,{U '(.1qndun

tiaptelnt)Iupuery

'6V* puu lupuerv :,iVV jsprorar pesrêruflVWZ tSoA 'lloo xe 'Ee1 erreg L06I'IIA'0I

p sauedllS :ueEreg :{OH - '1EWZ'3e1 11eplohtr It6I 'II4

6 ntpnW :raEuenels puu 1E6I'IIA'tI 66E euEg:uu8g :,{U - 'upuurls-olN :euofl iarel :[,O{aigN :Ei- 'ueruruur6l :OWZ:e>pre8ulg :OB - 'OWZ preErnul:1a5 iOWZ :uorC :uO - '511{OS :reruuH :OWZ IupO -ros :uolBpo :sEH - 'owz -Iwz gwz vwz nuJNV1S -IWg :olso - 'urelln :runr*g i11eqe11on :sV:)tV - 'vWZ ssoIAI 'vWZ fgpt:ssow lT,re18nC :apgul>1t,rrBg :fosuO :tgrcl.vt>l :[,ota{gr) i,{opueg erpuos ',{o{ru 'laytel1:relp^H :g 'tnrwoN 'saurp)ot

lo ts!.l

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'serceds

snquog luBurr,uop aql sr ,{11eco1 sntnprdn\ : geJaql\ seeJe ueqJn Jo palellllnJ ur ldecxe oJJeJS

'(mo1aq ]sll eql ';c) sreef

0g tsel aql pe^rosqo ueaq lou ser4 eurlrnbur eql sesJsrxe IIIIS ,{ernro51 Jo }seoJ ure}se^r eq} 3uo1e uo!}-elndod eql roqleq^\ ureuerun sr lI 'N.29

tnoqe ot

ua4al plttsv 8I


Apis rosiella Kirby,1802:331, No. 85. Type locality: En-gland: 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: Eng-land: E. Suffolk: Barham. Holotype C KCL! (Yarrow1e68).Apis francisanaKirby, 1802:334, No. 87. Typ. locality:England: E. Suffolk: Barham. Holotype d KCt-! (Yar-row 1968).

Nomenclatural and taxonomical remarks

Wahlberg (1855b) argued that the description

of P. campestris by Dahlbom (1832) applied to

campestris Panzer while the appurtinant figure

may refer to the male of P. svaveolens Wahlberg -

P. quadricolor Lepeletier. The figure is in my

opinion too diffuse for a sperific division of

Dahlbom's campeslris which should be confinedto that of.Panzer onlv

J

First S candinav ian records

Norwav: HOy: Bergen (Lie-Petters en 1902).

Sweden: SK, SM, VG (Dahlbom 1832). Re-

cords from Lapland (Dahlbom 1832) were revised

to P. flavidus (Thomson 1872).

Description

FEMALE

Body of medium size. Coat rather even, gaster

markedly sparser coated than in the remaining

European species.Measurements. Norway: (AAy-HOy): N :20.

R l : 3 .94 + 0 .159 mm,3 .5H .20 mm; Iw : 4 .82 !

0 .26 I mm , 4 .45-5 .15 mm; Hw: 4 .64 +0.166 mm,

4.154.90 mm; hamul i 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 distal

width, slightly shorter than Az*+; integument

smooth with irregular distribution of punctures of

varying size distally fading into a well defined mi-

nutely punctured area. Main keel of mandible

usually broad, reaching aboutzlt down to margin,

broadly abrupted distad. Labral furrow deep,

broad, widened towards the base; labral tubercles

distally pointed , feebly raised, the distal distance


between them exceeding tltlabral width. Clypeus

with dense, coarse punctation except for a median

impunctate area close to distal margin. Uppergena with scattered minute punctures between

mainly shallow coarse ones; genal furrow absent,

occasionally inconspicuous. Or shorter than 2x

Oe. A3:Aa:As : 77.5:10.5:15 , Ao-nhardly lc inger

than their distal width.Hind lrg. Tibia with longest hairs in posterior

fringe slightly or not longer than its greatest width.

Bt markedly narrower than adjoining part of tibia,

hardly 3 x longer than its greatest width, longest

hairs in posterior fringe hardly exceeding 0.5x itswidth.

Gaster. Tip of gaster not much incurved.Tqa

shiny; closer, finer punctured medially than

laterad and distad. Terminal of St6 slightly pro-

duced; the callosities convex distally pointed, &l-

most reaching apex and separated by a narrow,

deep furrow (Fig. 10).

Colour pattern Pile of vertex with slight admix-

ture of yellow. Hairs of broad collar, extending

about ll4 down episternum, scutellum, latero-

posterial patches of T. and large patches of To-t

yellow. Interalar-band laterally with varying ad-

mixture of yellow hairs, posteriorly more or less

V-shaped. T6 and St6 with pale yellow to golden-

brown pubescence. Yellow pile on thorax usually

paler than that of gaster. Otherwise coat black.

Variation. Pile of vertex all black. Pile on collar,

scutellum individually ranging from all black to all

yellow, rarely hairs on thorax all yellow except for

a few black hairs on the disc. Yellow hairs on gas-

ter more or less replaced by black ones. Com-

pletely melanic specimens occur.

MALE

Body of medium size. Coat shaggy.

Measurements. Norway (AAy - HOy): N :20.

R l : 3 .48 + 0 .162 mm,3 .2V3 .80 mm; Iw :3 .64 +

0.244 mm ,3 .25L 10 mm; Hw:4 .11 + 0 .121f f i f f i ,

3 .9M.30 mm;hamul i 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 distal

width, 3S long as A31+ or almost so; integument

feebly alutaceous with irregular distribution of

mainly minute to moderate punctures distally fad-

ing into a fine-punctured or impunctate area.

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1EWZ {qleC ,Ereqe>1gg , g;WZôlsJeg 'dresorg ',(qeuunJg 'SUN drelualg 'drolsuro[g'-IWZ

11WZ ra[qe11lg 'runrerpuy :)S 'uapaws'(zrd'p)

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lprgEeplo4 (retsol) es,(1 :sO :,iOH 'uelupallels'uagfsepne5 'es1e1 :luplns lEur1e61 :puusroC :ru .figp-ul-I :reEuenelg lsurl 'Oqeltetg :seupuu5 leseN :lupsefglaErssol4l-:orurl lddepl :ddep1 iloruy :lepu{os llupsenl,-tatrlijtg :lbry 'pun1 :f111 'etg iptirdoqra"H 'rvi :rurer{re[g :loru :pun-I :fU - 'elg ":petsoqe8eg

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to ts!7'pauruexe eJeA\ suerurJeds qsrpat\S 0Sg puu

uurEamroN 0rI lnoqe Jo Iulol V 'ecrers reqlell

'GZ'8tg) tanton)-sod 'g lsoq ureru eql Jo aEuer eql puu uepa^\S

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lnoqB ol r.llrou uoll-nqlJlsrp lulseoJ peJelleJs lnq snonurluoJ 3 ales-lpq ,{eur (qeqpuorl :lIS) quou requnJ prorerelEurs V

'N"29 lnoqe o1 prolyolso relno tuoq ]seoluJalsel\ puu uJeqlnos eql Euole sadololq luerJnxnl'peralleqs ul Eurrrncco ,(eruoN ul sl sarcads eql

'uepa1Y\s ul euoz IBeJoq uJeqlnos e{}olul Eurlerlaued ,{11uco1 'seuoz

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ISOH

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ua>1Q'l putsv 0z

ENT. SCAND. SUPPL.23 ( 1984) Scandinavian species of Psithyrus Lepeletier 21

Fig. 28. P. campestris (Panzer). Inset: The host P. pascuorum (Scopoli) updated 1983. The hatching refers to the dis-tribution of subspecies. Legend as in Fig. 25.

qclq^\ Jo IeJeAes 'seJnlBoJ

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Jo rnoror eqr ur srr'lep or uorrrppe ur r"rffi:rtill-orelJs lereâs fq peqsrn8urlsrp aJe saletueJ aql

'(tEOt) nodo4i pue (SZOD spreqrlg ,(q pe$rrelc,{1ureu sely\ rlrp{1y\

uorsnJuoJ ,{Fee qcnu pesnBJ 'lanelroq 'serceds

eql uao^\taq sdrqsuolleler esolJ eql'snua8qnspeurJep IIer!\ e solnlrlsuoJ snt,(.qnsdaoppuni

sTtouat lo)ltuouoxoJ

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sarcads uvttt)urpuns

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lsotule sn1,(lsouoE puo,{eq elles-lo^ :VZ-yZ 'sBIC ur su ellellueE puu 8-tts 'ra$ng

'tllplA\

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lnoqe unlleEBIC 'eO x Tuvtll reuor{s tO'poaH

gTvw

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' pe^JnJ,i1qeeJ,(lrouetsod' qlpl/!\

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'eo xzueql rouoqs ro 'qlpyn lerqsl

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'SuN ellH :yc - 'SuN ufqru4rA ,StgJurow 'sfgJ

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;YN - 'SdN pnrellB>ls 'gwz

slIN >lrolsou :sc - 'slIN^qrel]A :og - 'nwz eprBtsre^ 'nwz -lwz uurepogerlse1 'nwz

sdN ep^g{s 'wNc ere{s '-Iwz urglw'SUN drotue[qH 'SUN EurdolluC 'SUN ofsunly 'nWZsgsEurly :gA - 'uure4 '11WZ relsol) uler1 'euelspen'gWZ otullglJ 'g.WZ Surdoryepgs 'vWZ

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SUN fqsh 'eEugls 'ellls 'ej'sto1 'uueqelull) 'lulIEgH'esue11 ( epreg 'punsorgC 'le

lgrg 'aplC 'S)IN erunpreC

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IA erpqs 'nWZ fqss,{g 'urueqsru{sb 'SUNpunsrur.ule) 'St{N rer.ule) 'SUN 8urdo1uo1 'unr;8un1

BIg:htrs 'osrol "(qeuuog:'Ig 'relsol>lspeô

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awz spunlêd'u1oq,{qsg 'dro1,(5q ',(qeso51 'dru133n urroN 'gruluhtr'-IWZ

AWZ pun-I 'urutuol 'gwz paq,(q3un[-I 'pgrop-ul-1 'dro1s11e;1 'e8urppaqry,i;1 'Sraqelln) 'e^gl) 'ue8-EsJelsol) '{l^l) 'ulerurul 'JggH (ruloqolssgH 'e8reqal

(rso r) tz'-Iddns 'cNVf s 'JNauaTa'I pursv zz

ENT. SCAND. SUPPL .23 (1e84)

overlap between the species, occasionally impedereliable identification of European specimens. Anexception is P. quadricolor where the variability in

the diagnostic characters interferes in a lesser de-gree with the remaining species , a.E it is recog-nized by 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 clas-

sification functions derived from a step- wise dis-

criminant analysis 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. f lavidus read 1.738.

Page 49 column 1: E(I67 .99313) column P. nor-

vegicus read 167 .88313.

Psithyrus (Fernaldaepsithyrus) quadricolorLepeletierF igs . 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 (par-t im)Bombus frutetorum: Zetterstedt nec Panzet 1801, the

d .Psithyrus globosus Eversmann, 7852. Type locality_:USSR: Oienburg, Spaskoje. Type (?holotype) ?MAL!; Lie Pettersen 1901, 1905 , 1907; Muchardt 1904;Aurivi l l ius 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 1 85 5 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 local-i ty: Yugoslavia: Styria. Lectotype ? TRM!, hereby de-signated (cf . below).Psithyrus quadricolor meridionalis Richards; Tkalcfl7962.

Taxonomical remarks

P. quadricolor is divided into 5 subspecies in

Europe, viz. P. q. quadricolor, P. q. arvernicus

Richards, P. q. globosus Eversmann, P. q. ros-

sicus Popov and P. q. meridionalis Richards. The


designations refer to variations in the colour pat-

tern of 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 Richards

P. globosus auctt., nec Eversmann.

D esignation of lectotyp e

P. meridionalis was described from a Q and 12

C C of which the Q and one of the C d are in coll.

TRM. The ? is hereby designated as lectotype

and has the following labels: (1) "Styria, Tragop

Obert"; (2) "E. Graeffe"; (3) Ps. meridional is

n.sp ? type" [handwritten] ; (4) "lectotype P.

meridionalis Richardt ? design. A. Loken 1984" .

The specimen is in good condition except that left

At-r2., 3last tarsal segments of the right hind leg

and 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 allolec-

totype has labels as follows: (I-2) As for the lec-

totype; (3) "Ps. meridionalis C type" [handwrit-ten]; (4) "allolectotype P. meridionalis Richards

C design. A. Loken 1984". The specimen is in

good condition except that the 2 last tarsal seg-

ments of the left hind leg are lost. The sculpture

agrees with that of P. quadricolor . Colour pattern:

Pile on posterior half of T3 , T +-t, single hairs in

fringes of mid l.g, most hairs in posterior fringe of

hind tibia and Bt, and fringes of sternites ferrugin-

ous. Coat otherwise black.

Psithyrus quadricolor globosus Eversmann

Fig.29.

First S candinav ian records

I"lorway: On: Dovre, SFi: Jostedal (Wahlberg

r 85sb).Sweden: Localities not mentioned (Kriech-

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subspecies of host and parasite, respectively (cf .

below).P. quadricolor ursuping a colony of B. pratorum

(L.) (Pouvreau 1973) is a misprint (Pouvreau in

litt.). B. ruderarirzs (Miil ler) l isted as a host (Pek-

karinen et al. 1981) refers to an observation by A.

Nordman of a Q flying into a nest of this Bombus

sp. (Pekkarinen in l i t t .) .

Flight season

Southern Sweden: Hibernating female: 4 May -

12 August; female progeny: ? 13 September;

ma le : 9June -9Oc tobe r .

Distribution

Scandinavia (Fig .29). Confined to nemoral and

boreonemoral zones. In Sweden also occurring in

the southern boreal zone.In Norway more often observed at sea level in

luxuriant biotopes along the western coast than in

the southeastern lowlands. In Sweden widely dis-

tributed north to "limes norrlandicus" (NU 1977)

and further north occasionally recorded together

with B. soroeensis throughout the southern boreal

zone. A single record in central boreal zone (VB:

Degerfors NRS) indicates a scattered distribution

northeast to at least about 64"N.Scarce. A total of 135 Norwegian and about 230

Swedish specimens were examined.

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: Hjelt-nes. - SFy: Gloppen: Lotsberg. SFi: Stryn, Flo. - MRy:Orskog:Skjodje ZMO. MRi: Norddal: Valldal.

Unrevised records: SFi : Luster: Jostedalen(Wahlberg 1855b). HOy: Askoy, islands in Feiefjordand HOi: Voss (Lie Pettersen 1901, 1905). Ry: Eiger-sund: Heskestad, Ri: Suldal: Jelsa, HOi: Odda and SFi:Balestrand: Balholm (Meidell L934).

Correction. A female from VAy: Kristiansand: Ran-desund ZMO (Richards 1928) is revised to P. campes-tris.

Sweden SK: Andrarum, Brosarp, Helsingborg, Hiic-keberga, Jordholmen ESE Malmo ZMA,, Klostersig_el,Kiseberga NRS, Lund, Nytorp, Sandhammaren NRS,Torekov, Torna-Hdllestad, Vallf,ker, Vitemolla, Vomb,


Vdstra Vram, Ovedskloster . HA: Halmstad NRS,Hdsslov, Snostorp NRS. - SM: FAgelfors. Grdn-naberget, Horeda NRS, Korsbergp, Markaryd, Ryssby,Sjotorp by Alvesta, Vdstervik, Osterkorsbeqg_a _GNMZML. - OL: Bengtstorp, Borgholm, Boda NRS ZrylU,Gdrdslosa, Hogsium-Ril la, Rod, Vickleby. OC:Borsjon NRS, Brunneby GNM ZML,, Linkoping,.Vads-tena. - VG: Grhstorp NRS, Toreboda NRS. - NA: Ar-boga, 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 Saff le, Grums NRS, Aris. - DR: BrunnsvikNRS, Ludvika NRS, Mora NRS, Oxberg NRS. - GA:Ockelbo NRS. - HA: Delsbo. - VB: Degerfors S Vin-deln d 26.VI I I .1935 Forsslund leg. NRS.

Correction. A record for this species in Lapland(Friese 1902, referring to Wahlberg 1855b) is incorrectand due to l inguistic misunderstanding.

The total distribution of the subspecies P. quad-

ricolor globosus Eversmann. Fennoscandia - Den-

mark - The Bal t ic - European USSR.

Sparse information about the Danish popula-

t ion (Hammer 1970; a few dark coloured speci-

mens examined by me) indicate that the sub-

species may intergrade with P. q. meridionalis in

Denmark; 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 Alps

widely distributed north to Fennoscandia and to

about 62" in European USSR. Discontinously oc-

curring from Turkey NE to Transbaykal.

The present information about distribution of

the species indicates a division into two geographi-

cally segregated groups similar to that of the host

B. soroeensis (LOken 1973). P. q. meridionalis oc-

curs within the range of the red-tailed B. s. proteus(Richards 1928; Tkalcfl 1969;Beinig 1973) and the

islet distribution of the remaining subspecies is

within the distribution of the white-tailed B. s.

soroeensis.

Psithyrus (Fernaldaepsithyrus) sylvestrisLepeletierFigs. I1,23, 30.

Psithyrus quadricolor var. sylvestris Lepeletier de St.Fargeau , 1832:377, 1841 :429. Type locality: Describ_edon specimens from the Pyrenees and Paris. Type:2 d dlabelled "ex coll Spinola scat' I l712" SCT! might be syn-types (Kellner-Pillaut in litt.)Apathus quadricolor auctt. nec P. q.quadricolorLepeletier; Thomson 1870 (partim).

alqrssod

ftsot) tz

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.-IddNS 'CINVJS 'INEua4a7 putsv gz


Psithyrus quadricolor auctt. nec Lepeletier; Sp.Schneider 1898, 1909, 1918; Friese 1902; Aurivil l ius1903; Strand 1904; Soot-Ryen 1925; Bengtsson 1931.Apathus sylvestris (Lepeletier); Thomson 1872; Siebke1880; Strand 1898b.Psithyrus silvestrus Lepeletier; Richards L928; Meidell1934; Barendrecht l94l;Tjeder 1954; Ander 1963,,1965;Loken 1966b; Kullenberg et al. 1970; Bringer 1973;Lundberg & Svensson 1977; Svensson 1980.

Taxonomical remarks

P. sylvestris var. citrinus Schmiedeknecht, P. s.var. albicans Richards and P. s. var. atrithoraxRichards one or several of which mentioned by

Sp. Schneider (1898, 1909), Richards (1928),

Bahrendrecht (1941) are infrasubspecific forms

occurring occasionally throughout Scandinavia.

First S candinav ian recor ds

Norway: Tromso (Sp. Schneider 1898).Sweden: Distributed north to Lapland (Thom-

son 1872).

Description

FEMALEBody small. Coat shaggy.

Measurement. N _ 25. Rl: 3.56 mm t 0.129,

3.30-3.80 mm; Iw: 4.47 + 0.209 mm, 4.2V5.00

mm; Hw : 4.23 t 0. 123 mm, 4.0M.40 mm; hamuli

numbe 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 than

A:*o; integument smooth with irregular, closepunctures of varying sizes anteriorly fading into a

large almost impunctate area. Main keel of mandi-

ble well developed, reaching margin or almost so,

tapering distally (Fig. 11A). Upper gena with dis-

tinct, usually dense mixture of fine and coarse

punctures; genal furrow indistinct, occasionally

absent. Surface of scapus smooth, shiny, rather

sparsely hai red. A: :Aa:As : 17:10.5:14, Ae-r z 7s

long as their width.Hind leg. Tibia with longest hairs of posterior

fringe about 1 .5 x its greatest width ; Bt with

longest hairs about as long as its greatest width.

Gaster. Punctation of T5 as that of Tz-q or

slightly sparser. T6 closely punctured except along


a median longitudinal l ine usually forming an in-distinctly interrupted keel (Fig. 118). Sto with nar-

row angled cal losit ies (Fig. 11C).Colour pattern Pile of vertex with a variable ad-

mixture of yellow. Broad yellow collar laterally

covering adjacent border of episternum. Lateralpatches of T3 more or less extending inwards alongextreme distal margin and T4 with white hairs. T6

and St6 between and outside callosities covered bypale ferruginous to pale brownish pubescence.

Coat otherwise black.Variation. Pile of vertex ranging from all yellow

to all black. Pile on T3 ranging from all white to

rarely all black. Fringes of legs, particularly Bt,

pale yellowish or tipped so.

MALE

Body small. Coat shaggy.Measurements. N _ 50. Rl: 3.23 + 0.148 ffiff i,

2 .80-3 .55 mm, Iw: 3 .51 + 0 .471 mm, 3 .00-3 .90

mm; Hw :3 .92 + 0 .139 mm ,3 .551.30 mm; hamul i

number : 18 .30 + 1 .389, 15-21 ; Lh : I .24 + 0 .073

mm, 1 .10-1 .38 mm; Dh:0 .072 + 0 .006 mm,0.06-

0 .09 mm.Head. Malar space about as long as distal width

equal to A3a + or just longer; integument smooth

with rather sparse, fine and medium-sized puncta-

tion distally fading into an almost impunctate atea

of various size. Upper gena usually with close, dis-

tinct and mainly coarse punctation; genal furrow

indistinct. Scapus shiny, smooth rather sparsely

haired. A3:Aa:A5 _ 16: l I :17, Ao-r : about 1.5x

their distal width.Hind lrg. Tibia with longest hairs in posterior

fr inge about 1.5x i ts greatest width; Bt 's longest

hairs almost 2 x its greatest width.Gaster. St6 distally sligthly protuberant, trun-

cate, more or less bilobed (Fig. 23A). Stz-a and

genitalia as in Fig .238-F-; inner basal extensions

of gonocoxite usually slender, longer than wide;

gonostylus varying in shape (Fig . 23D-E), inner

margin concave, usually about as long wide and

about as long as that part of volsella visible from

above (Fig . 23D).Colour pattern Pile of vertex with varying ad-

mixture of yellow. Broad yellow collar laterally

covering upper part of episternum. Tl with vary-

ing admixture of whitish and yellow hairs. Hairs of

T z-q and lateral patches of T5 whitish, hairs of T6-7

predominantly ferruginous or whitish. Otherwise

coat black.

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ua4@'l plttsv gz

ENT. SCAND. SUPPL.23 (1e84) Scandinavian species of Psithyrus Lepeletier 29

Fig. 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) Scandinavian

Fig.3l. P. flavidus (Eversmann). Variations in the colour pattern ofthe Fennoscandian populations, see the text. A-G:males ,H- I : females .B lack :b lack ,o :ye l low,o :b lack ,ver t i ca l l ines : fe r rug inous .

D

HGF

and melanics (Fig. 31F-G). The pie chart de-monstrates a pronounced tendency towardsmelanism throughout the Scandinavian peninsula,intermediates and melanic specimens everywherebeing in majority. The examined Finnish males in-dicate a 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).

Conclusion

The rather stable colour pattern of the femalesargues against subdividing the Scandinavianpopulation of P. flavidus. It belongs to the nomi-nate subspecies 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

'SJIUUIatu

eleldurocur;o uollrodord : pelddns 'srrueleu eleldruor;o uotlrodord : :1oe1g 'JoeJeql lJlllsry ro acurnord e ulqlrn

spJocar peuruexeJo lequ.rnu plol eqt ol reJer sern8g aqa 'uslueloru Jo uotgodor4 '(uueusreirg) snyaoTf '4 '79'3rg

(ru0 t) fz'-Iddns 'cINVJS 'JNEua4@7 pulsv zt

ENT. SCAND. SUppL. 23 (1984) Scandinavian species of. Psithyrus Lepeletier 33

Table.I. Proportion and degree of melanism in P. /avldus (Eversmann). Area = province or district thereof (cf . Fig.J J I .

Number of individuals

inter-medeates

Total

number typ.form

inter- mel.medeates form

Per cent

AreaTvp.form

mel .form

3715443014t715

68029495082587322

937

30150

60

50735470726870

82106526509

251368

825864446237

25

8 1 326 1268 11 0 07 1 4

24 1713 15

33 t229 101 7 6

r22 256 0

5 7 9t2 17

r20 14t76 11

[ , 936 3914 2923 2679 238 6 3

34

303142

4T93775

l69

1310142

431

30052

t7t9

273

1 132353

157t9

2235

603

477

8838

Norway:OsOnBvHOiSFiTRiFO

Sweden:DRJAVBNBAsLYPILUTO

Finland:KbokobLKLeLi

USSR:Petsamo t625

1117

120

15

92l9

10493

11446

185

Total t45 4t8 361, 924

nest (Sweden: LU: Virihaure 8.VII .1944 ZI$-4L).The colony was not analyzed and the presence of

a B. jonellus ! makes it uncertain whether it was

usurped by P. flavidus or the inquilines were tem-porarily resting in the nest. The 2 specimens are

the only records of P. flavidus observed in a bum-

ble bee colony.Bombus lapponicus s.l. ( - B. lapponicus (Fab-

ricius) and B. monticola (Smith)) has been consi-

dered as host of P. flavidus (Richards 1928; Pit-

tioni 1942, 1943; Tkalcfl 1969; Lundberg &

Svensson 1977; etc.) based on the fact that B. lap-ponicus s.l. and the inquiline are boreo-alpinespecies locally observed in the same areas. Pittioni(1942) in recognizing that P. flavidus has a wider

distribution than B. lapponicus s.l. in Finland,proposed B. cingulatus Wahlberg as a second

host. Pekkarinen et al. (1981) confirmed that in E

Fennoscandia P. flavidus occurs markedly further

south than B. lapponicus s.l.; they suggested the

widespread B. jonellus might be the only host of

this inquiline in Finland. The present study reveals

that P. flavidus has a wider distribution in Scan-

dinavia than that of of B. lapponicus s.l. or B. cin-

gulatus (cf . below).Until more colonies invaded by P. flavidus ate

recorded it remains question whether B. ionellusis preferred host (cf. p . 34). The inquiline resem-bles B. jonellus in colour pattern and is widely dis-

tributed within the range of this Bombus species(cf . below).,.Difference in colour pattern and in

ecological valence may, however, not prevent the

species from attacking colonies of B. lapponicus

s.l. or B. cingulatus. The 3 Bombus species

suggested as hosts of this inquiline are all belong-

ing to the subgenus Pyrobombus Dalla Torre.

I

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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.

MALE

Coat rather even. Body large.Measurements. N - 40. Rl: 3.78 + 0.083 ffiff i,

3 . 35 -4 15 mm; IW:3 .70 + 0 .183 mm,3 .254 .05mm; Hw:4.22 + 0. l32mm, 3.8(H.50 mm; hamul inumber : 17 .58 + 1 .130,15 20 ; 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 bor-

der of 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 post-

erior fringe rarely longer than 1.5 x their greatestwidth.

Gaster. St6 distally slightly protuberant with ele-vated margin more often inconspicuously bilobed

than truncate. Stz-s and genitalia as in Fig. 22;

inner basal extension of gonocoxite broad, flat-tened, slightly or not longer than broad (Fig.

22C); gonostylus varying in shape (Fig. 22C-E),usually broad, longer than that part of volsella vis-ible from above, inner margin more or less con-cave and 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 cov-

ering adjacent border of episternum and extend-ing backwards 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 ad-mixture of black hairs. Hairs of T3-a more or less


yellow. Pile on T6-7 ranging from all whitish to allferruginous or black.

Host

Bombus hypnorum (L.) (Stoeckert 1954;Roseler 1972; Cederberg 1976; Mjelde in l i t t .) .

Flight season

Southern Sweden: Hibernating female: 3 May-1 1 September; female progeny: ? 5 August;male:25 june - 15 September.

Distribution

Scandinavia (Fig . 34). Sparsely distributedthroughout the peninsula north to almost 70'N.Occurring from sea level to timberline; rarely ob-served in 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: Kon-nerud; 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; Ul-lensvang: Fresvik, Lofthus, Ringoyi Ulvik: Hjeltnes,Ulvik. - SFi: Balestrand: Flesje; Luster: Leirdola, Ok-leviki. - 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, Hel-singborg NRS, Horby, Kulla-Otorp(-Kulla-Gun-narstorp), Kullaberg, Lund ZM^, Skiilderviken NRS,Strandbaden, Vegeholm NRS, Ahus NRS. - BL: Ron-neby ZMB, 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,

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Table IL Host recordsfor European Psithyrus.N: numberofBozbus colonies usurped byPsitlryrus: + = 5-10co-lonies, ** : > 10 colonies, 0 : number ofcolonies recorded in Scandinavia, x : nonscandinavian species.

Social parasite

P. bohemicusP. vestalisP. barbutellus

P.rupestris

P. campestris

P. quadricolorP. sylvestris

P. flavidusP. norvegicus

Host

B. (s. str.) lucorumB. (s. str.) terrestrisB. ( Megabombus) hortorumB. (Pyrobombus) hypnorumB. ( Melanobombus) lapidariusB. ( Melanobombus) sicheli

ssp. 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) hypnorum

N

+ ++ +-r

1+ + (1 )

222+ +T

11a

J

+ +(2)1

(? 1)+ (3)

Tyreso NRS, Uttran NRS, Valla NRS. - UP: Bjorkl ingeNRS, Ingaro, Fogdo NRS, Stockholm NRS ZML,,Svartsjo, Uppsala, Vallentuna NRS, Yxlan. VR: Ar-vika. - DR: Bingsjo NRS, Brunnsvik NRS, Falun NRS,Horndal CBJS, Leksand TJTS, Ludvika NRS, MoraCBJS, Ore i i lv TJTS, Orsa, Sandsjo, Sil jansborg NRS,Sjurberg NRS, Smedbacken, Tii l lberg NRS. - GA: Hil leNRS, 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, Ojt-byn. - LY: Sorsele: Ammarnds, Ndckrostj i irn. - LU:Messaure ZMB ZMU, Nattavaaraby ZMB, Njunjes,Vuollerim ZMB. - TO: Abisko BML, DjupvikenLHAS.Unrevised records (Ander, in l i t t .): SK: Osby. - HA:Akulla NE Varberg. - DS: Bengtsfors, Lingbron. - DB'Vikarebyn.

World distribution. Eurosiberian. Central and

northern Europe - Siberia - Kamchatka - Trans-

baykal - Northern Mongolia. The only Psithyrus

sp. recorded in Japan (Ito & Tadauchi 1981).

The range of P. norvegicus is within that of B.

hypnorum, a Siberian fauna element (LOken

re73).

The host-parasite relationship Bom-bus - Psitiyrus

The host species of the individual Psithyrusspecies mentioned above and the frequency of ur-

suped bumble bee nests (calculated from the fi-

gures of the host species references) have been

compiled 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). Recently

Cederberg ( 1979, 1983) found that P. rupestris fol-

lows trail pheromone laid by B. lapidarius, thepreferred host. Further experiments de-

monstrated that 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 species

occur in Europe (29 in Scandinavia (LOken

1973)), 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 oc-cupied by Psithyrus. P. bohemicus, P. vestalis, P.

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Forh.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 bumblebees (Hymenoptera, Apidae). - Abstr.Upps.Diss.-Fac. Sci. 549 :142 .

Thomson, C. G. 1870. Oversigt af Sveriges humlor. -

Opusc. ent . 2:251-26I .- 1872. Hymenoptera Scandinaviae. 2. 286pp. Berling,

Lundae (Lund).Tjeder, B. 1954. Ostskinska insekter. II . - Opusc.ent.

19:205-209.Tkalcfl, B. 1967. Zur Hummelfauna der Umgebung

Kuku-Nors (Hymenoptera, Bombinae) psithvrusCas.Cdske Spol.ent. 58:344-379 .

Index

Valid names in Roman type. Synonyms,homonyms, infrasubspecific forms in italics. InPsithyr,rzs only those pages referring to descriptionor taxonomic status are indexed .In Bombus onlythose pages referring to host species are indexed.

- 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.1 9 :887-916 .

- 1974a. Eine Hummel-Ausbeute aus dem Nepal-Himalaya (Insecta, Hymenoptera, Apoidea, Bom-binae). - Senckenbergiana biol. 55:31I-349.

- 797 4b . Ergebnisse der 1 . und 2. mongolisch-tschechoslowakischen entomologisch-botanischenExpedition in der Mongolei. Nr. 29:Hymenoptera,Apoidea, Bombinae. Acta faun.ent.Mus.Nat.Pragae 15:25-57 .

Van Honk, C. , Roseler , P.-F. , Vel thuis, H. &MalotauX, M. 1981. The conquest of a Bombus ter-restris colony by a Psithyrus vestalis female. -

Apidologie 12:5747 .Wagner, R. 1971. Die Verdnderung der Hummelfauna

Cuxhavens in diesem Jahrhundert. Der Versucheiner Deutung. - Ent .Mi t t .ZooL Mus. Hamburg 4.N o . 7 5 .

Wahlberg, P. F. 1855a. Iakttagelser ofver de nordiskahumlorna. - Forh.Skand.naturf. 6. mote, Stockholm185 1 . Bilagan C:23V233 .

- 1855b. Iakttagelser o.ch anmdrkningar ofver de nor-diska humlorna. - Ofvers.K.VetenskAkad.Handl.Ir:r99-211. (1854).

Wahlgren, E. 1915. Det olt indska 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 or ig in of

highly social behaviour among bees (Hymenoptera/Apidae/Phylogeny) . Proc. Natn. Acad. Sci. USA7 4:1135-1 I37 .

Woll in, H. 1975. Kring nornans biologi. - Fauna ochFlora 70:89-98.

Yarrow, I. H. H. 1968. Kirby's species of Brit ish bees;designation of holotypes and selection of lectotypes.Part I. Introduction and the species of Apis Linnaeusnow included in the genera Bombus Latrei l le andPsithyrus Lepeletier. - Proc.R.ent.Soc.Lond. (B)37:9-15.

Zetterstedt, J. W. 1838. Bombus Fabr. columns 473474

aestivalis Panzeraestivalis Dahlbom nec Panzeralbicans Richardsalpium Richardsalbinella KirbyAllopsithyrus Popovamo enus SchmiedeknechtApathus Newmanarenaria Panzerarvernicrzs RichardsAshtonipsithyrus Frisonashtoni (Cresson)atricolor Richardsatrithorax Richards

Wl#if.,#N:. : . . . . . ' . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .",\ni";::):,ffll"?n : : : : : : : : : : : : :: : : : : : : : : : : : : : : : : : : : :: :citrinus Schmiedeknechtdistinctus P€rezFernaldaepsithyrus Frisonfernaldae Franklin

flavidus Eversmannflavidus (Eversmann)

francisana Kirby

frutetorum Panzer

fruteto rum : Zetterstedt nec P anzerglobosus Eversmannintercalaris Miillerlato fas ciatus Pittionileana Kirbylissonurus Thomsonmaxi l losus (Klug) . . .Metapsithyrus Popovmeridionalis Richards ..nemorum Fabriciusnorvegicus Sp. Schneiderobscurus SeidlPsi thyrus Lepelet ier (s. s t r . ) . .quadricolor Lepeletierquadricolor auctt. nec Lepeletierrosiella Kirby

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ENT. SCAND. SUPPL .23 (1984) Scandinavian species of Psithyrus Lepeletier 45

Manus received December 1983

Journal 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.

Fig. 35. Division of Fennoscandia and Denmark. For abbreviations of provinces, see p.44.

Documents

Scandinavian species of the genus Psithyrus Lepeletier ... species of the genus Psithyrus Lepeletier (Hymenoptera: Apidae) Entomologica Scandinavica Supplement No. 23 tts€-s0r0 NSSI