ASTRID I-OKEN
Scandinavian species of the genus Psithyrus Lepeletier
(Hymenoptera: Apidae)
Entomologica Scandinavica Supplement No. 23
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Scandinavian species of the genu s Psithyru, Lepeletie. '* W*
(Hymenoptera: Apidae)
ASTRIDLAKEN
Introduction
The 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- d e n .
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 . L g k e n , U n i v . o f O s l o , D e p t . o f B i o l . , D i v . o f Z o o L , P . O . B o x 1 0 5 0 , B l i n d e r n , N - 0 3 1 6 O s l o
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 review
In 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-
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E N T . S C A N D . S U P P L . 2 3 ( 1 9 8 4 )
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 upper gena (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 proportions 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 . 6 0 m m .
Male: Body large, mean Iw >3.70 mm; body of medium size, mean Iw 3.55-3.70 mm; body small, m e a n I w < 3 . 5 5 m m .
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).
Biology
The 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).
Phenology
The 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).
Classification
The 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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= rlc 'rlnuuq
lere}el eou?lsrp Jo rlnrueH ruoq ecuulsrp pulq tqSu snlleJo ruoq r{-I iEur/l| snlleco lqErr JBeurl : lelrdrccoerd ol punoduoc o] 'a8prr 'S : JO'efe uee/rueq ecuulsrp
: eO eOO ,plelJ : 'eere elelcundrul - qOO 'puuq peJnlJund
Iulrqro-rullero :
'morrn;1eue3 : oo
Jc lpeeq snt{,t\trs4
Jo ued raddn ;o
^\ern lprcuC
'qlplly\ 'V
rulurelur 'ecuds Jplu[u - esJeASueJJ 'E Jo uorlces A\I lxeroql :
p-c 'qlpl,r\ -
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.I
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uer8urq .g-I .sEIC
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 sternite
6.- 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 = cal-
l o s i t i e s o f s t e r n i t e 6 . - 1 2 . P . n o r v e g i c u s S p . S c h n e i d e r Q ; A : t e r g i t e 6 , B = c a l l o s i t i e s o f s t e r n i t e 6 . - L 3 . 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. 36 Hind tibia and Bt with longest hairs in posterior fringe markedly longer than their greatest width. T6 inconspicuously or not keeled. Callosities as in
F i g s . 1 3 - 1 4 . . . 9
9. Callosities of St6 as in Fig. L3.T4 yellow-haired, T5
black-haired P.
(Fernaldaepsithyrus) flavidus (Eversmann) p. 28 Callosities of St6 as in Fig. 14. T:-s ferruginous- haired, often with an admixture of whitish hairs P. (Fernaldaepsithyrus) quadricolor Lepeletier p. 23
MALES
1. Malar space shorter than 3lq the distal width. Stz-a and genitalia as in Figs. 16-17; penis valve not
h o o k e d b e n e a t h . . . A s h t o n i p s i t h y r u s F r i s o n 2
Malar space longer than 3lq the distal width. Stz-s and genitalia as in Figs. 18-24; penis valve hooked
beneath 3
2. A.3 hardly or not longer than ,A.5. Hind tibia and Bt
with longest hairs in posterior fringe at least as long as 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 with longest hairs in posterior fringe shorter than their grea-
7 A
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srrur.{ lse8uol qlyt\ lg
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l'l .) qe rr uJ 8
: C'8alrur€ls:
g'relrurels = V:,9 (dq4S)m11asnqn_q.d.gtr -.Bllellue8 =
g,gegurels =
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'n
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ua>101 plttsv 9
E N T . S C A N D . S U P P L . 2 3 ( 1 9 8 4 ) Scandinavian species of Psithyrus
Figs.2l-24. Fernaldaepsithyras Frison; sternite 6, right half of male sternite 7-8 and genitalia. - )il . P. quadricolor
LJpeletierd;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
( E v e r s m a n n ) d ; A : s t e r n i t e 6 , B = s t e r n i t e T , C : s t e r n i t e S , D - E = p o s t e r i o r p a r t o f g e n i t a l i a . Lepeletier
ffi K/
W G
W 24E G
G
W
or not exceeding their greatest width. Stz-s and genitalia
as in Fig. 18A{. Coat shaggy
P. (Allopsithyrus) barbutellus (Kirby) p. 12
6. ,A.3 slightly shorter than ,A.5. Hind Bt parallel-sided, ffio-
re than 4x longer than wide. Stz-s and genitalia as in Fig. 19A-C . T+4with reddish hairs
i; ;;;k;;d ;;#"{',":# Pr#{;'{:'Jl1:ffi:? X ;' o
its 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 in
Fig. 21A{. T+ with an admixture of yellowish, whitish
or pinkish-orange hairs, Ts-, with femrginous hairs . P. ( Fernaldaepsithyrus) quadricolor Lepeletier p. 23 St6 as in Figs . 23A, 24A. Stz-s and genitalia as in
Figs . 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
24B,
22A 23B,
21A
23C 24C
228
22D
21C
23A 22C 22E 23D
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E N T . S C A N D . S U P P L . 2 3 ( 1 9 8 4 ) Description
FEMALE
Body of medium size. Coat rather shaggy.
Measurements: N-20. Rl: 4.06 + 0.265 mm'
3 . 5 5 - 4 . 4 5 m m ; I w : 4 . 7 9 + 0 . 3 1 6 m m , 4 . 2 V 5 . 3 0
m m ; H w : 4 . 5 5 + 0 . 2 3 3 m m , 4 . 1 5 1 . 9 0 m m ; h a m u l i
n u m b e r : 2 0 . 7 5 + 1 . 4 I 0 . , 1 8 - 2 3 ; L h : I . 6 3 + 0 . 1 0 2 m m , I . 4 5 - I . 7 8 m m ; D h : 0 . 0 8 3 + 0 . 0 0 4 f f i f f i , 0.078-0-094 mm.
Head. Malar space with close distribution of fine punctures admixed with some coarser ones, distally fading into an almost impunctate, large subtriangular area. Clypeus more sparsely and coarsely punctate on disc than at base, distally with a well defined median impunctate space.
Upper gena with sparse to moderate distribution of punctures of varying size; genal furrow usually indistinct. OOb without accumulation of minute
punctures (Fig. 68). ,{3:Ao:As : 19.5: 11: 1,4,, Ao.n
just longer than their distal width.
Hind lrg.Bt about 2x longer than its greatest width. Longest hairs in posterior fringe of tibia and Bt as least as long as the segment's greatest width.
Gaster. T6 smooth, shiny, nearly impunctate on the disc, closely punctured laterad and distad.
Ridge-like callosities (Fig. 6C) distally more
pointed and less separated than in P. vestalis (Fig.
7B ); rather sparse - mainly fine - punctures on the disc between the callosities.
Colour pattern. Broad, pale to dull yellow collar extending aboutllq down the adjacent episternum and more or less projecting backwards below base of wing. Extreme posterior fringe of scutellum
usually with single yellow hairs concealed by black
ones. Sparse pale yellow or whitish hairs in lateral patches of T1 . Latero-posterial patches and ex- treme distal margin of T3 , T q, and lateral patches of T5 whitish-haired. T6 laterally and distally, and St6 outside and between callosities clothed with
golden-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 patches or completely replaced by black hairs. Pile in cres- cent-shaped posterior fringe of scutellum ranging from all black to nearly all yellow. Slight variation in the amount of whitish hairs on T3 and Ts.
Whitish hairs, particularly on T3 occationally re- placed by yellow ones.
Scandinavian species of Psithyrus Lepeletier 9
M A L E
Body of medium size. Coat shaggy, long.
Measurements. N - 20. Rl: 3.62 + 0.200 mm.
3 . 0 5 - 3 . 8 5 m m ; I w : 3 . 6 2 + 0 . 1 9 5 m m , 3 . 2 5 - 3 . 9 0
m m ; H w : 4 . 1 3 + 0 . 2 0 4 m m , 3 . 6 M . 4 0 m m ; h a m u l i
n u m b e r : 1 7 . 1 5 + 0 . 9 8 8 , 7 G l 9 ; L h : 1 . 3 2 + 0 . 0 7 9
m m , 1 . 1 8 - 1 . 4 8 m m ; D h : 0 . 0 8 2 + 0 . 0 0 6 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; genal furrow usually distinct. Or - 2x Oe or nearly so.
OOb without accumulation of minute punctures
( s e e F i g . 6 8 ) . A 3 : A a : A s : 1 7 : 1 0 . 5 : 1 6 , A o - r 3 a b o u t
1.5 x their distal width; flagellum about 3 x longer than scapus.
Hind leg. Tlbia with a small almost hairless area close to distal margin; longest hairs in posterior
fringe up to 1.5x the segment's greatest width. Bt
3.5x longer than its greatest width or nearly so;
longest hairs in posterior fringe up to 2x Bt's greatest width.
Gaster. Stz-s and genitalia as in Fig. 16A-C; that
part of volsella visible from above about 1 .5 x longer than gonostylus.
Colour pattern. Pile of vertex with black and yellow intermixed. Collar laterally extending about l lq down episternum, hairs of posterior fringe 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 coat black.
Variation. Pile of vertex ranges from all black to predominanty yellow. Yellow collar varying in width, laterally passing more than half way down episternum, reduced to lateral patches or turning all black. Pile of scutellum ranging from all yellow to all black. Pile of particularly mid and hind basitarsi 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 yellow ones. Pile on T5 admixed with black hairs on the disc. Pile on Stz-s ranging from all black to all whitish.
Melanism. Colour variations are too slight to change the general colour pattern except for sporadically occurring pronounced melanic speci- mens. The darkest individuals are nearly black, having at most whitish hairs on T5. The tendency
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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 in
litt.); X = inaccurate locality. Shaded areas refer to literature indications.
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7 . 4 V I . 7 8 m m ; D h : 0 . 0 8 2 + 0 . 0 0 4 m m , 0 . 0 7 5 - 0 . 0 8 7 m m .
Head. Malar space about 3lq the distal width, about as long as A31a; integument more or less
striate-rugose closely set with punctures of varying
size fading into an almost impunctate narrow area close to distal margin. Main keel of mandible well developed, anteriorly tapering, reaching margin or almost so. Labral furrow deep, broad; labral tubercles protuberant, blunt, the distal distance between them at least rlz labral width. Clypeus more sparsely and markedly more coarsely punctate on disc than at base. Upper gena with rather dense, distinct, mainly coarse punctation;
genal furrow absent or indistinct. Or about 2x Oe. A3:Aa:A5 _ 19:lI:16, Ao-rz as long as their distal width.
Hind lrg.Bt markedly narrower than adjoining tibia, about 3x longer than its greatest width.
.Longest hairs in posterior fringe of tibia and Bt equal to the segment's greatest width or just longer.
Gaster. Tip of gaster not much incurved. T+-s with rather close, mainly coarse puncturing. T6 with close, rather uniform, mainly medium-sized punctation; 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 admixture of yellow. Hairs of collar just passing margin of episternum, 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 or less reduced. Pile of scutellum ranging from all yellow to all black except a few yellow hairs along extreme posterior margin. Legs with variable ad- mixture of pale-coloured hairs, especially in the fringes.
MALE
Body of medium size. Coat rather shaggy.
Measurements. N - 20. Rl: 3.44 + 0.178 mm.
3 . 2 5 - 3 . 7 5 m m ; I w : 3 . 6 2 + 0 . 2 4 I m m . 3 . 2 H . 0 5 m m ; H w : 4 . 2 5 + 0 . 1 6 3 m m , 3 . 9 M . 5 0 m m ; h a m u l i n u m b e r : 1 8 . 6 0 t 1 . 6 9 8 , 1 7 - 2 3 ; L h : 1 . 3 0 + 0 . 0 5 9 m m , 7 . 2 U I . 4 4 m m ; D h : 0 . 0 7 4 t 0 . 0 0 7 m m .
0.059-0.083 mm.
Scandinavian species of Psithyrus Lepeletier l3
Head. Malar space hardly as long as distal width, ?S long as A:*+; integument distally im- punctate or almost SO, otherwise irregularly punctured, punctures coarser centrally than basad. Upper gena with rather dense mainly shal-
low, coarse punctation;genal furrow absent, occa-
sionally indistinct. Or about 2x Oe. Hairs of scapus about 2x width of segment; ,{3:Aa:45 -
1 6 : 1 0 . 5 : 1 9 , A o - r : n e a r l y 2 x t h e i r d i s t a l w i d t h ;
flagellum about 3x longer than scapus.
Hind lrg. Bt 3.75x longer than its greatest width. Tibia and Bt with longest hairs in posterior fringe hardly or not exceeding their greatest width.
Gaster. St6 with 2 mounded callosities. St74 and
genitalia as in Fig. 18A-C; penis valve hooked be- neath.
Colour pattern Pile of vertex, collar just passing margin of episternum, scutellum and T1 yellow.
Hairs of extreme distal margin of T3 ,Tq-s, lateral patches of T6, Stz-s whitish. Otherwise coat black.
Variation. Pile of vertex with variable admix- ture of black. Yellow collar extending ll4 to ll2 down episternum. Yellow hairs of scutellum more or less reduced to crescent-shaped posterior fringe.
Melanism
Both sexes display a slight tendency towards melanism which is more pronounced in south- ernmost Sweden (SK, OL) than further north.
Host
Bombus hortorum (L.) (Ball l9l4; Cumber 1949; Mulle r 1936; Postner 1952). B. hypnorum (L.) (Pouvreau 1973).
Host species mentioned by Hoffer (1889) are omitted as the author confused P. barbutellus and P. sylvestris. P. barbutellus observed established in a colony of B. pratorum (L.) and B. ruderarius respectively (Pouvreau 1973) are a misprint (Pouvreau in litt.).
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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