70 NOTULAE ENTOMOLOGICAE 61, 1981 Notulae Entomologicae 6I :71-89, 198l
Distribution of bumblebees (Hymenoptera, Apidae: Bombus and Psithyrus) in eastern Fennoscandiar) Antti Pekkarinen, Ilkka Teräs, Juha Viramo & J"hani Paatela Abstract
Purrlnr^-Erv, ANrrr, TrinÄs, ILKKÄ, Vrn.e.uo, JuH.l & P,tarrle, JunaNr: Distribution of bumblebees (Hymenoptera, Apidae: Bombus and Psithyrus) in eastern Fennoscandia. Notulae Entomologicae 61:71-89. 1981. The distribution of -26 bumblebee and B cuckoo bumblebee species of Fin- land, and northwestern USSR are mapped according to the European UTM grid system. The species are grouped on the basis of their distribution patterns in the area studied. The limits of many species correlate with the isotherms of effective temperature sums and eight species have their northern or southern limit in a biogeographical transition zone running from the head of the Gulf of Bothnia to the southeast, B. syluarum and B. subterra¡teus have greatly Lettland, Salaspils, 1917-06-07' Ar¡n. 4. Aedoeagus von Monochroa simþIicella Lg, & 2., extended their range since the 1930s through the Karelian Isthmus to south- ern Finland. The effect of warming up of the climate and other ecological Kirja-arvostelu Bokrecens¡on factors on the distribution area are treated. The distribution of cuckoo bumble- - bees and their bumblebee hosts as well as the relative abundances of different bumblebee species in some areas of northern Europe compared. Press, New are Rocrsruw, M. (Ed.) 1978: Biochemistry of Insects' - Academic Authors' addresses: Antti Pekkarinen and Ilkka Teräs, Departrnent oÍ Zoo- York, San Francisco, London.649 sid',215 figs',60 tab. Pris: USD 29.50. Ìogy, University of Helsinki, P. Rautatiekatu 13, SF-00100 Helsinki 10, Fil- land; Juha Viramo, Oulanka Biological Station, University of Oulu, Torikatu Insekternas yttre struktur och utseende har svarsämnena hos steklar sammanfattas av M. S' 7, SF-90100 Oulu 10, Finland; Juhani Paatela, Agricultural Research Centre, länge studerats, främst ur systematisk synvinkel. Blum, som även tar upp vilka olika föreningar SF-31600 Jokioinen, Finland. Titiar man istället inuti en insekt hittar man som används som försvarssubstanser och varför, och hur insekterna undviker att förgifta sig själva. R. beskriver olika typer av D. O'Brien lntroduction insecticiders verkan med avseende på funktion (1979) has elucidated some taxonomic såsorn nerv-, muskel- och metaboliska gifter. The first accounts of bumblebee spe- and evolutionary problems of Fenno- Avgiftningsprocessen, dvs hur skadliga insekti- cies in Finland were given by NvreN- scandian bumblebees. The distribution cider omvandlas till för insekten ofarliga pro- hos insekter. DER (1848,1852). (1889) of bumblebee species is consequently G. M. Chippendale diskuterar kolhydraterna dukter diskuteras av W. C. Dauterman och E. S¡Hre¡nc genitiken och evolutio- published a catalogue of the occurrence now relatively well known in Fenno- med avseende på deras funktioner som närings- Hodgson. Den kemiska ,the nen sammanfattas av F. J. Ayala, som beskri- of. Bombus and Psithyr?rr species in the scandia. However, knowledge of ver olika biokemiska metoder för att mäta den biogeographical provinces of Finland the distribution in eastern Fennoscan- genetiska variationen inom arter och släktskap dia is far from complete and more 'de- mellan arter. and later (1902) a general review of Agosin som bl.a. tar upp struktur proteiner, Illustrationerna är klara och tydliga. Varje Bombus species in Finland. ErrvrNc tailed information on the distribution .i"y^.r, cytokromerna (c, bs och P-450) och kapitel är försett med en lista över allmän (1960) has made a revision of Finnish of these biologically and economically bioluminiscens. P. S, Chen beskriver protein- och a.vancerad litteratur. species and later (1968) a more com- important animals is needed. In the på syntes och lagring Till bokens nackdelar hör att titlar på upp- syntesen med avseende prehensive account distribution course of this study we have especially av proteiner, reglering av syntesen och muta- satserna ej finns i litteraturlistorna. Att Ayala of their tioner. De olika kemiska föreningarna i cuti- endast har givit de engelska namnen på de and food plants. In her thorough taxo- tried to map out the distribution lirnits culan tas upp av A. G. Richards. Kemin bak- organismer han tar upp (med undantag av nomic and faunistic monograph on of the s,pecies which do not occur om och de olika funktionerna hos biokromerna Drosoþhila) är synd eftersom latinska namn Scandinavian bumblebees LørnN throughout Fennoscandia and also to diskuteras av A. E. Needham. Biokemin bakom brukar underlätta läsandet av böcker som ska evaluate possible changes in the limits nå utanför den engelskspråkiga världen. (1973) has also treated Finnish fau,na de olika hotmonerna och tillväxtregulatorerna during the last few decades. Further- behandlas av L. M. Riddiford och J. W. Tru- En lämpligare sammanfa.ttning av biokemin to some extent. Pnxr¿.nINaN & T¡nÄs man. I två kapitel behandlar N. Weaver den hos insekterna till detta låga pris det får man (1977) have reviewed the bi,ology, idis- more, we have ,treated some zoogeo- kemiska kontrollen inom och mellan arter, där nog leta länge och väl efter. För alla som stu- tribution and identification of Finnish graphical aspects and abundance of f symbios derar entomologi borde denna bok yara en olika typer av eromonet respektive species (in Finnish) and Pnrx¡nrNnr.r the species. mellan myror och bladlöss beskrivs. Olika me- välkommen handbok, som ökar förståelsen för toder för att insamla och detektera feromoner kemin inuti den lilla insekten. tas upp av W. L. Roelofs, som även diskuterar 1) Dedicated to the late Mr. Erkki Valkeila in recognition of the great entomologist and his olika typer av feromon-fällor. De kemiska för- Ulf Carlberg magnificent work on Finnish Aculeata fauna. 61, 1981 Ef- AL., BUMBLEBEES IN E. FENNOSCANDIA tc 72 NOTULAE ENTOMOLOGICAE A. PEKKARINEN
ed (referred to in the text). The number of (4. Saarinen); Ofr: Kuhmo (J. Paatela); Kon: (cf' specimens inspected is about 3B 170 Jalguba (4. Günther). Scarce. Table 1). Recoràs from the eastern Fennoscandian B. lucorum (Linnaeus, 1761) (MaP 2) areas of the Soviet lJnion are old. There are Distributed throughout E Fennoscandia. eas which Very abundant, sca nd World biotopes of Lapland .'Records ently conspecific wit 'sPorad'ic. cimens regarded as B. þratoru ø (Linnaeus, 1761) (Map 11) Most of the records have been made by the light males oÍ B. Iucorum Distributed thloughout E Fennoscandia. A closely allied sPecies, Very abundant. s), occurs on the SE coast 60oN (LørrN 1973:55), B. taþidarius (Linnaeus, 1758) (Map 12) where it is locally abundant. B. terrestris has The noithern limit of distribution runs from not been found in E Fennoscandia, but its the head of the Gulf of Bothnia to the south- occurrence in SW Finland is possible. The east. The northernmost records: Oá; Oulu, Ç, specimens from S Finland, which G. Kruseman has determined as B. terrestris (Er,rvrrc 19t60), are B. lucorum specimens (cf. LøxrN 1973:56). (Map 3) järvi, (E. Q, Finland. No Sw Fi ,19+2-07-15 A. Merisuo). B. alpinus (Linnaeus, 1758) (Map 13) mountain (Map 4) Distribuied over most of the arctic Bombus The records of very rare species (only a few B. sþoradicus Nylander, 1B4B area of N Fennoscandia. The uurÍlenü 6.9 1.6 Arbsent from Äland and perhaps from ex- and tundra records from the area) and the northernmost southernmost records: Li: Ivalo, several (W' sor oe ensxs 4.5 2.9 2.9 Lapland and N Kola. The southe¡n- Ç, or southernmost records of more common spe- treme N Hibiny (LørrN 19'73:98). lucorum 11.6 9.7 19.4 most records: Ab: Pargas (Ingelius), Karja- He1lén) ; Lirn: Quite the Finnish scarce. sþoradicus t.2 2.0 O & RAATI- lohja (U. Saalas); Nr Espoo (8, Poppius), 3.1 ry terrest, Ns 0.0 . The names Porvoo (8. Suomalainen). The northernmost B. polaris C,urtis, 1835 (8. arcticus Kitby, 0.1 cullumanus iterature are records: le.' Palojoensuu 758:3'3 (P. Niemelä); 1821) (Map 14) 0.3 0.8 2.5 Very eingulatus given in parentheses. li: Karigasniemi 771:46 (8. Valkeila). Distribution about the same as that of B. al- hyþnorum 3.J 4.3 6.8 scarce in S Finland, abundant in tr and N Fin- distribution t2.6 þinus,bwt the southern limit of the jonellus 9.7 5.8 Abbreviations land. The records in SW Finland are old and (to about 6BoN). 7.4 : èxtends farther in Finland laþþonicus 7.9 3.6 C central the species is nowadays absent from wide areas southernmost records : l¿: Pallastunturi 7.9 7.6 : The þratorunl 9.3 E east, eastern of SW and S Finland. 7ú6':37 (J. Kaisila); .Li; Kivipää 759:52 (8. laþidarius J.¿ 10.8 6.8 N: north, northern B. semenouiellas Skorikov, 1910 (Map 5) alpinus t.2 0.4 0.4 NW : northwest, northwestern 0. 6 Only one record. Kl; Parikkala 682:63, $, Quite scarce. þolaris 0.3 0.4 S: south, southern 1.9 , 2 1964-07-2+ (R. Elfving) (ErrvINc 1965). b alte atus SE = southeast, southeastern B. balteatus Dahlbom, 18,32 (MaP 15) hyþerboreus 0.2 0.2 0. ¿ SW : southwest, southwestern B. cingulatus Wahlberg, 1854 (Map 6') 'fhe southern limit of the distribution in consobrinus ,i 0.9 1. i W = west, western Absent from S Finland and possibly from Finland runs from about 67o30'N in W Ifr to 4.4 see back hortorum 1 0.0 11.5 For abbreviations of the provinces the the western coast of the Gulf of Bothnia and about 65o15'N in E Ok. The southernmost re- ruderatus 0.5 cover. N Kola. The southernmost records: Tb: Yüta- cords: lk: Kittilä (R. Krogerus) ; Ok: Suomus- humilis t.4 +.5 1.3 saari (8. Valkeila) ; Sá; Rantasalmi (E. Pulliai- salmi Ruhtinassalmi (O. Sorsakoski). Locally muscorufn 2.9 1.5 0.4 Distribution of lhe species nen); Kl: Sortavala (O. Hulkkonen), Ruskeala abundant. 10.8 tl.7 þascuorunT 15.4 Bombus wurllenü Radoszkowski, 1B5g (8. (E. Rantalainen). Scarce, more abundant in E 1.1 ¿,¿ 2.4 B. hyþerboreus Schönherr, 1809 ruderarius mastrucatus Gerstaecker, 18,69') and N Finland. syluarurn t.2 +.7 0.8 (Map 16) B. hyþnorum (Linnaeus, 1758) (Map 7) aeteranus t.+ ¿.o The following records from the northern- found also 0.6 43 ¿.1 Clearly absent only from N Kola; dístinguendus in Ä.land (cf. Errvruc 1960:36). Very abund- subterraneus 0.1 2.2 1.3 ant. 27700 3376'2 B. jonellus (Kirby, 1802) (Map B) n 28900 Distributed throughout E Fennoscandia. Very abundant, rather scarce in S Finland, but Psithyrus land, especially in more abundant in the southern archipelago. 34.4 result. There are s bohemicus caught in Le: Kilpisjärvi, 2 196'7- 11.6 specimen in B. Iaþþonicus (Fabricius, 1793) (Map 9) have been ç ruþestrNs the (J. Paatela). Only four workers 6.4 Lrxono'rH 1945:1 The southern limit of distribution extends 07-19 & H. camþestris have been recorded from Scandinavia (LørnN barbutellus 2.0 this species from t to about 65oN in Finland. The southernmost 17.9 records: Oå: Hailuoto Vesanniitty 722:39, 1973 rt7). flauidus (Fabricius, 1776) (Map 1) $, 7.8 B. soroeensis 1966-0,7-11 (8. Valkeila), Pudasjärvi, several globosu.s northern limit of the distribution runs B. consobrinus Dahlbom, 1832 (Map 17) .1. J The (C. Brander) Kuusamo, several re- norûe gLcus Bothnia to the Þ ; Ks; The distribution is associated with the main 16.5 from the head of the Gulf of cords. Abundant. sylue stris southeast. The northernmost records: Lk: Ko- food plant Aconitum seþtentrionale Koelle lari, Þ, 19'46-0'7-22 (P. Niemelä); Oå; O-unas-- B. monticola (Smith, 1849) sensu Svrns- (Lørnn 1961, 1973124). In Finland the largest n 4408 vaara (8. Kanervo), Kemi Karihaara 730:3'8 soN 1979 (Map 10,) stands of Aconi.tum arci Kb: Tohmajärvi Piilo- BUMBLEBEES IN E. FENNOSCANDIA 74 NOTULAE ENTOMOLOGICAE 61, 1981 A. PEKKARINEN ET AL., 75 vaara 6,90:67 and Kitee Papinniemi 6BB:6'5' B. þascuorum (Scopoli, 1793ì (Map 21) gasniemi; Oa: Wasa, but these ¡ecords are un- Distribution patterns These are also the only localities of B. conso- Clearly absent only from N Ii and N Kola. certain. The species has evidently immigrated brinus in Finland and the species occurs re- The northernmost records in Finland: Le: Kil' into Finland; through the Karelian Isthmus 'Ihe Bom.bus and Psithyrus favna of. pisjärvi abundant; Li: Jvalo 761:52 (E. during the last few decades. gularly in these places. Other records: KIr area, E Fennoscandia comprises 34 species Sortavala area (several collectors), Harlu (J. Valkeila), 762:52 (A'. Albrecht). Subspecific Kaisila, P. Niemelä), Ruskeala (8. Rantalai- delimitation in Fennoscandia, see LørrN (1973) Psithyrus bohemicus (Seidl, 1B'3'7) (26 Bombus and 8 Psithyrus species). nen), Impilahti (several collectors) ; Kon: and P¡rx¡rnrwlN (1979). (Map 27) This is aboutthe half of all the species (4. Günther); Iu: Kusomen (W. The northern limit of the distribution in 0/o Jalguba (Map 22) found in Europe and about l0 of the Hellén) Lp: Ponoj (W. Hellén). Often abun- B. ruderarius (Müller, 1776') Finland extends tc about 6'8o40/N. The north- ; runs world figure. The corresponding dant in its habitat. The species has been more The northern limit of the distribution ernmost records: I¿.' Ounastunturit, Q, 19r51- scarce in Tohmajärvi Piilovaara during the from the head of the Gulf of Bothnia to the 07-24 (J. Kaisila) ; Li: Ivalo 761:5'2, ó, 1979- number in Scandinavia is 38 (29 Born- records: Oulu last few years than in the 1960s. southeast. The northernmost Ob; 08-12 (8. Sirjola). The most abundant Psithy- bus and 9 Psitlzyrus). The Scandina- 1890 ( Elfving); Kó: Nurmes (4. about K. røs species in S Finland. vian species which have not been found B. hortorum (Linnaeus, 176,1 (Map 18) Saarinen) ; Koz; Karhumäki (J Carpelan) ) zøurflenü Throughout E Fennoscandia to 6B'o45'N in Quite scarce, locally abundant in S Finland. P. ruþestris (Fabricius, 1793) (Map 28) in E Fennoscandia arei B. Finland; the information is too scanty for the The northern limit of the continuous distri- Radoszkowski, B. terrestris (L.), B. cul- B. sylaarurn (Linnaeus, 1761) (Map 23) determination of the northern limit of the dis- bution in Finland extends to about 6,2o30'N. lumanus (Kirby), B. ruderatus (Fabri- tribution in Kola, USSR. The northernmost The northern limit of the distribution ¡uns The northernmost records: Oa: Tjock (P. Nie- from about 61o15'N in SW Finland to about cius), B. (Panzer) and P. aes- records: .Lk: Muonio 754:36 (E. Valkeila) ; melä) Kb; Tohmajärvi (4. Pekkarinen); iso- þomorum 62o45'N in SE Finland. The westernmost and ; (Geoffroy). These a Tl.' Karesuando (Sweden) (LørnN 19'78); Li: lated record from Ks: Kuusamo Jyrkänkoski talis all have records: z4ä: Turku Ruissalo 6'71 : Ivalo 761:52 (several collectors); Ips: Pum- northernmost 733:61, 1979-09-06 (J. Viramo). Scarce, southern distribution in Scandinavia, q, (4. Pekkarinen) Ta: Lem- ó, manki, Þ, 1930-07-12, Yläluostari,9, 1930- n, ß75-08-18 ; locally abundant in southernmost Finland, often except B. zøurflenü, which is a boreo- päälä 681:31, Þ, 1977-0'8-14 (M. Raekunnas) ; 07-15, Peuravuono, Þ, 1930-07-21 (O. Hulk- in cultivated and urban biotopes. alpine species (.f LørnN 1973) (a abundant, local, favours culti- kbr Kontiolahti 696:64,9, 1978-06-03 (M. . konen). Quite 696:70, vated habitats. Raekunnas), Ilomantsi Mekrijärvi .Ç. P. camþestris (Panzer, 1800) (Map 29) single uncertain record of B. uurflenü 1978-06-0'9 and Ç, 19179"06-11 (8. Ranta). The northern limit of the distribution in from E Finland is excluded from this (Map 19') The species has immigrated to Finland through B. humilis Illiger, 1806 Finland extends to about 63oN. The northern- study, see p. 72). The E Fennoscandian The northern limit of the distribution runs the Karelian Isthmus during the last 40 years most record: ,9å: Nilsiä (K. Levander). Scarce. from about 6'1oN in SW Finland to about 620 (cf. NIrunr.:i 1947 and, the progress of the species absent from Scandinavia are 30'N in SE Finland. The northernmost record: distribution in Map 23). Scarce, locally abund- P. barbutellus (Kirby, 1802) (Map 30) B. þatagiatus and B. semenoaiellus, Kb: Liperi, Þ, 1961-08.26 (R. Blfving). ant in SE Finland. The northern limit of the distribution runs both of which have an eastern distri- 60o30'N Finland about Scarce, very local, (Fabricius, (Map from about in SW to B. ueteranus 1793) 2a) 62oN in SE Finland. The northernmost records: bution. The northern limit of the distribution runs B. muscorum (Linnaeus, 1758) (Map 20) S¿.' Rantasalmi (4. Vesterlund) Kl: Suistamo maps the from the headt of the Gulf of Bothnia to the ; Distribution often reflect Scattered occurrence in the archipelago of (E. Hammarström). Very scarce. collectìng activity in different areas, SW Finland, on the coast of Oa: Vaasa, (8. southeast. The northernmost records: Oä; Oulu Ç, 1978-0i-09 (J. Viramo) Så: Ii- Lingonblad) ; Ob; and. White Sea: Onega (out- l2l:42, 9, ; P. llaaidus Eversman, lB52 (lissonurus to,o. Maps 35 and 36 reveal superficial side E Fennoscandia). Also ¡ecorded inland: salmi 705:50, several Þ. 1980 (E. Pankakoski); Thomson, 1Bi2) (Map 31) collecting activity in wide and relative- (P. Ha- Kb: Ilomantsi, 1865-07-13 (E. Grönvik) (un- Kå; Nurmes 705:60, ó, 1979-08-10 Absent from wide areas of S Finland, but ly continuous areas of S Finland ar¡d certain record) ; Ok: Yaala Säräisniemi 714:48'' kala). Rather scarce. the southern limit of the distribution is un- (J. certain. The southernmost records: O¿; Kris- Lapland. The highest number of species Þ, 1980-07-08 Viramo) ; Ob: Yaala làwi- B. distinguendzs Morawitz, 18'69 kylä, V, 1979-01-28, Pudasjärvi Korvenkylä tinestad; Z¿: Keuruu (S. Sulkava) ; Kl: Pa- is recorded from the grids on the south- (Map 25) (E. several 725:51,2 Þ, 1980-07-09, Syväoja 724:48, ë, in rikkala Valkeila), So¡tavala area, ern coast of Finland in the Helsinki The northern limit of the distribution specimens (E. Rantalainen); Kol: Mäkrätjärvi 1980-08-13 (J. Viramo), Ylitornio. ?. 1946- 68o30'N' Z¿ to area and in southeastern Finland in Finland runs from about in S and Vaaseni (P. Niemelä). The most abundant 08-03 (P. Niemelä); Ia: Kilpisjärvi, 2 Ç, 19'39 the southeast (to about 65015'N in E Finland). thorough these local- Psithyrus species in N Finland. the Parikkala area where (8. Kivirikko). The specimens in The northernmost records : Iz.' Karesuvanto, ities have a dark coat on the episternum and collections have been made. This num- 2 (LørnN 19'78); Lk: Muonio, Þ, 1946- P. globosus Eversman, 1852 (Map 32) vente¡ and no distinct local differences can be Ç, ber (23) is about 68 0/o of all the species 0B-03 (P. Niemelä) Ofrr Suomussalmi Ruhti- The northern limit of the distribution runs seen in the colour of the coat. The Finnish ; nassalmi, 2 (O. Sorsakoski) ; Ko¿; Karhu- from about 61oN in SW Finland to about 6,3'oN in the whole of E Fennoscandia, A re- specimens greatly resemble the specimens from 9, mäki (J. Carpelan). Quite scarce. Finland and Kon. The northernmost re- latively high number (16) has been re- Norway, which have been named as a subspe- in E cords: ,9å: Kuopio (4. Vesterlund., R. Elfving) cies, B. m. lieþetterseni, by LøxnN (19'73:146), B. subterraneus (Linnaeus, 1758) ; corded from the Kuusamo area, where Kon: Tiuti.a and Tulvoja (8. Scarce. and also resemble the specimens from NW (Map 26) Poppius). several southern and northern species Uusikirkko, 193,8- Russia ("f. Popov 1930 and Pnrxlnrw¡¡t Recorded from: fk: ?, P. noruegicus Sparre Schneider, 1918 occur. 06-06 (P. Niemelä) N: the rural district of 1979). The specimens ftom Ik: Muolaa, Ç, ; (Map 33) We have made intensive collections 1936-07-26 and 2 Ç, 1938-08-11 (P. Niemelä) Helsinki 668:39, about 30 specimens, 1959-0'7- The northern limit of the distribution in re- in the marginal areas of the distribu- have a yellow episternum and venter and they 31 ...08-15 (M. Markkula), later found Finland is uncertain; found in several localities Helsinki are identical with the nominate form occurring gularly and locally abundant in the up to 63oN and some specimens in Ks; Ou- tion of different species, especially in on the SW coast of Norway, S Sweden and C àrea; Kl: Parikkala 682:63. 2 ç, 1962-07-09 lanka 737:6'A,737:61, 1977-78 (J. Viramo). Ob and Kr N Finland. We therefore (R. Elfving); 7a: 676:36, 1968- Europe, The subspecific status of dark speci- Janakkala ?, Scarce. suppose that our maps approximately mens in Finland is uncertain and a comparative 07-01 and later in the Hämeenlinna area some revision of the colour forms in N Burope is specimens in different years (E. Valkeila); Sa: P. syluestris (Lepeletier, 1833) (Map 34) reveal the boundaries of the distribu- necessary. Usually scarce, found abundantly in Lappeenranta 6,771:561, 2 ç, 19'77-07-27 (8. The northern limit of the distribution ex- tion in Finland, despite the existence Oå; Hailuoto 721:39'. Very local, restricted to Valkeila) ; Ka: Anjalankoski Myllykoski 674: tends to about 6,6o30'N. The northernmost re- of superficially investigated areas. The (M. Sarl¡nnc cords: Oó: Rovaniemi (E. Rantalainen); Ks; damp, mainly coastal meadows rich in flowers, +8, 9, 19'77-06-12 Viitasaari). records from :the Soviet part of B l-en- also on the fields with sv/ampy ground and with ( 1BB9) has notified the species from the follow- Oulanka, several specimens 1977-80 (J. Vi- Kan- ramo). Abutrdant. noscandia are sporadic except for the fireweed (Eþilobium angustifoliurn L.). ing localities: Iftr Isthmus Karelicus; ^S¿: A. PEKKARINEN ET AL., BUMBLEBEDS E. FENNOSOANDIA 77 76 NOTULAE ENTOMOLOGICAE 61, 1'981 IN provinces of Ih, Kl and Kol, but the and 'therefore an ambient temperature ?åble 2. The percentåge dlstrlbubton oi Bombus lndlvlduaLs counted ln some lleld studles ln Fennoscandla information on this area is al'so useful obviously plays an important role in and northern Germany, for some conclusions concerning the many biotic and abiotic factors de- limits of the distribution. termining the distribution of bumble- ó @ ó r oÔ a o The species can be grouped accord- bees. The temperature of the whole rts ôô F. ..ro ol do ô^ x@ z ø/oo â tstã ôts in ing to their recent distribution in E season is important to bumblebees and Ôts d Ørt tsroo õ tsHôz cdÉ ciH c^ CJ c6t Fennoscandia as follows: therefore isotherms describing thermal OH doe Eoø olo cd
3 B, PATACIATUS 5 B. SE1IENOVIELLUS I B. LAPPONICUS 10 B. IIONTICOLA
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4 B. SPORADICUS I B. JONELLUS 11 B. PRATORUM 12 B. LAPIDARIUS
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15 B. BALTEATUS 16 f. HvpnRgonrus
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a a a a a a t0 NOTULAE ENTON{OLOGICAE 6I, I9B1 A. PEKKARINEN ET AL., BUMBLEBEES IN E. FENNOSCANDIA 8l
24 ''t IIORTORUI\f 22 B. RUDERARIUS B. VETENANUS 17 B. CONSOBRINUS I B.
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25 B. DISTINGUENDUS 26 B. STJBTERRANEUS
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19 B. HUMILIS 23 R- SYLVARUM
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B. PASCUORUM 27 P - BOHEMICIJS 28 P. RUPESTRIS 20 B. MUSCORUM 21
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a a \\ a aa a n a a a a -È a a a a a a a a a a a a a a a a t a a a a - .{. PEKKARINEN ET AL., BUMBLEBEES IN E. FENNOSCANDIA 82 NOTULAE ENTOMOLOCIC,{E 61, l98l 83
35 36 29 P. CAMPESTRIS 30 P. BARBUTELLUS n tr t-25 5 E 26-50 E T 76 -9 e a t foo a a a
P. GLOBOSUS 31 P. FLAVIDUS 32
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3z' 37 /u b- I I r\1 I ,gorn Maps 1-34. The dlstrlbutloo of Bonbu! and ^ff Pslbhyrus gpecler 1n eaaten¡ Fennoscandla,
Map 35. the numbers of Bombw and prlthyRs specles recorded 1n eastern Fennorcandla
accordlng to the European UTM grld ryrten.
Map 36, The numbera of Bonbus ånd prlthyrur
specles recorded (Map 35) as a fercentage of the probable total oumberr 1n llfferent
33 P. NORVEGICUS 34 P. SYLVESTRIS UTM squares.
Map 37. Effectlve tenperature sun! ln Flntând o. and northeastern Scandlnavla 1n 192Ì{950
(after LAAIßoNEN 1979).
zlo
{ ru a a a -f a a -Y
A. PEKKARINEN ET AL., BUMBLEBEES E. FENNOSCANDIA 34 NOTULAE DNTOMOLOGICAE 6I, I98I IN 85
has recently been recorded as far north with the distribution in Karelia, where as Abisko (68"22'N) (LørnN 1978). The the range of Aconitum extends as far scattered distribution of B. muscorurn as Onega on the coast of the White Sea. is possibly due to a wider post-glacial Thus the distribution of B. consobrinus distribution and some isolated popula- in Scandinavia perhaps originates from oo oo ddoo o tions have perhaps become extinct in one or more glacial refuge populations I I I I I tØt tØ I I I I I Ø Scandinavia during the last few de- (cf. PexxrnrNan 1979). The distribu- È!È ggg È 5=A AtA J cades (Løxrrv 1973). The dark form of tion of the group 3 species in Fenno- É Ê iÈÈ ËdJ Ètttttttttt B. muscorurn in Scandinavia and Fin- scandia is associated with the area of ÞÞ Ð Þ land is restrioted mainly coastal coniferous forests (regio coniferina) as ¡ rì rll JJ J É ttltttttttt ¡ to a d oø aaøø ûûø habitats and the abundant local occur- (1973:195) has emphasized in ø o oo ooqo ooo oo o LørrN È ¡¡ J¡ ÈtÈÈ ÈÊË ¡ rar rìl r r I r J rence on the eastern coast of the head connection with B. sþoradicus and B. o o ooo o oo o ¡ r I r r I r r ËËJ r r Ë r r Jd r r r r t r r r r r r r r È I I I of the Gulf of Bothnia and eastwa¡ds cingulatus. These species are represen- inland is worthy of mention. tatives of the Palaearc'tic taiga element J r q r r I I I ¡È r il r r I d 14 I I I È I I I I I Ë I I I I I I I É Ê ÉÉEE É EEE E É É and they are absent from wide areas of ¡ ridd t ildi t il i The biotopes of B. laþþonicus in J t¡lr r r¡ËìÈlÈrÉÉËË¡ r I lJ¡ I I I I Èl I I I I I I S Scandinavia and the western and jl Scandinavia and Lapland are mainly g mottntain heaths and meadows (.f. northern Atlantic coast. ooo o ooo o 9^ o Êqo o LøxnN 1973:76). The record of B. Iaþ- ç6 v tt rvr rxr r rx r I r r¡4 The Finnish climate has become !õ g È in Hailuoto (a large island on DC o o þonicus 6ø o warmer during the last hundred years. ø rV the eastern coast of the Gulf of Both- According to HuNo (1978) rthe increase oç c c nia) implies that coastal meadows are c o o 3 in the mean annual temperature cul- }1 I I I I I I I I I I I I I I I I I lyl I lt also suitable biotopes for this species. çc cc q ç ç cc c c cç minated around the 1940s when it was cç o o o oo Some single records from the lowlands 99 , € , ,9 , , , ,9 , , , €€ , ry 1.5"C higher than 50 years earlier in oÐ ÀÀÀÀ ' à4Å' ii ' 4Å dd ÀÅÅÅ oooo ooo oo oo oo oo ¿¿¿ooo oooo of C Sweden (Løruw 1973:77) in'dicate od y!¿l
Local and habitat differences in the The material reveals a great change this century has possibly aiided the ex- Nest parasite Host tension of B. subterr&neu's. No distinct abundance of different species are large in the abundance of B. lucorum, B. la- (e.g. RnrNrc 1972). Thus the material and B. distinguendrns, On the extensions of the ranges are noticeable P. bohemicus B.Iucorum þidarius for other species. Old northern records P. ruþestris B. Iapidarius in Table 2 reflects the composition of a other hand, the abundances of B. hor- from about 1890 concerning B. soroeen- P. camþestris ? (8. humilis, B. þascuorum) local fauna very imperfectly and only toru,m, B. þascuorum and B. subterra- (about P. barbutellus ? (8. hortorum) tentative comparisons are possible be- neus seem to be more stable. The great sis and B. aeteranus in Kuopio jonellus) P. syluestris ? (8. þratorum, B. tween different areas of Fennoscandia and stable abundance of B. subterra- 63oN) a B. jonellus P. flauidus spe- and B. P. norae gicus B. hyþnorurn and N Germany. B. lucorum, B. þrato- neus is worthy of note, since this 65"N) r P. globosus B. ruderarius rum, B. laþidarius, B. hortorum and B. cies is a newcomer to Finland. 8., lu- of these þascuorum are among the most abund- corltnx and B. Iaþidarius (workers) are species short tongue and during the last 90 years. However, The greatest conformity of distribu- ant species in S Finland, N Germany the with a and England (Table and INT. BEE thus they are not efficient foragers on ro-. äf the new notihetn records of tions occurs between P. bohemicus and 2 RES. ASSOC. 1980). B. lucorum and red clover. Variation in the abundances these species mentioned above may be B. lucorunt, P. ruþestris and B. Iaþida- are most abundant of other nectar flowers is one possible due to a recent extension. B. terrestris rius, P. camþestris and B. humilis, P' B. þascuorurn the species the material from S Finland, expl,anation for their abundance fluct- has recently extended its range north- and B. jonellus, and P. globo- in flaaidus but B, lucorum is distinctly rnore scarce uations on red clover (.f. TnnÄs wards in Scandinavia sus amd B. rucJerarius. The isolated re- ir: Germany, where the closely re- 1976a). 194), as have some soli cords of B. Iaþidariu.t an'd P. ruþestris N lated species B. terrestris is the mos,t Modern agriculture and urbanisa- the beewolf ( Philanthus in Kuusamo confirm their host-parasite abundant species. In N Europe, B. ter- tion have presumably changed the hab- in Finland (NIaunrÄ 1947, VÀxure relation. B.laþþonicus is often assumed restris is more anthropochorous spe- and abundance bumblebees. 1953 and VIrrnsa.tnr 1975). to be the host of P. but the a itats of flavidus, cies (Awonn 1963, Regrerttably, we have very exact range P. extends much than B. lucorum little of flauidus H¡,rs¡r,nn 1972 and PsrrÄ,nrx¡N 1979: information concerning the human in- There is a notable correspondence furthelto the south than that of B.Iaþ- and these species occur fluence on these useful animals and between the northern limits of many ar,.d B. jonellus is the most 9) do not þonicus habitat long-term studies this subject are southern Bornbus species and summer probable host of P. the abundantly in the same on flaaidus.,on (WacNnn l97l). In Fennoscandia important. isotherms both in Scandinavia and Fin- basis of their habitats and distributions. N B. hyþnoru,rn and B. cingulatus, and B. land (cf. the limits of ranges in Lørnru is also worth noting that B. jonellus Acknouledgernents. We are greatly obliged It alþinus and B. (in alpine and 1973 and fig. 14 in LaersoNnN 1979). is scarce in the wide area of SW Fin- þolaris to the late Mr. Erkki Valkeila for many valu- B. ue\eranus is an exception. This spe- absent. arctic areas) are closely related species, able records and observations on bumblebees. land from which P. Tlaaidus is We are also grateful to the following persons a time in S but no distinct habitat differentation cies has occurred for long for delivering records, collecting material or Scandinavia, but only in extreme SW has been observed between the mem- Abundance other assistance: A. Albrecht, Alli Brander, T. Sweden. The absence of the species bers of. these species pairs. Brander, Y. Haila. H. Hippa, E. Kaarnama, from other parts of S Scandinavia is The evaluation of abundance of the No comprehensive study has been I. M. Kerzhner (Leningrad), K. Koivulehto, pro- and made of the temporal variation in the M. Koponen, S. Koponen, Mervi Kunelius, K. an interesting zoogeographical species is a complicated problem Laaksonen, (Oslo), Pamilo, abundance of different bumblebee spe- Astrid Løken P, blem. in this context we can only compare S. Panelius, E. Pankakoski, Yu. A. Pesenko the relative abundances between dif- cies in N Europe. Yearly fluctuations (Leningrad), E. Pulliainen, M. Raekunnas, E. Knowledge of the hosts of different ferent species. The material in Table I are obviously great and vary in differ- Ranta, H. Silfverberg, E. Sirjola, S. Sulkava, Psithyru,s species is incomplete and ìs mostly based on public collections, ent species. The following observations B. G. Svensson (Uppsala), J. Terhivuo, P. were made on red clover the same Utrio and Sirkka-Liisa Varvio-Aho. many reports indicate that some Psi- and, of course, forms no random in This study has been aided by a grant from thyrus species possibly have several locality in Helsinki (668:39) in 1966 the Finnish-Soviet Commission for Scientific hosts and that areal differences in the (07-22,07-26), 1967 (08-04) and 1980 and Technical Co-operation. host specificity are possible, too. There- (07 -22, 07-31) (values in percentages): fore, only some tentative concl'usions References of 1966 1967 19,80 can be drawn from the distribution Arrono, D.V. 1975: Bumblebees. Davis- Psithyrus species compared with their cies in Finland and Fennoscandia. The B, soroeensis 0.3 Poynter, London, 352 pp. - potential hosts. The mosrt probable species distributed practically all over lucorum 34.1 - 7 0.6 ANonn, K. 1963: Om humlor och snylthumlor. hosts of Psithyrus species in E Fenno- the Fennoscandia (8. lucorum, B. pra- Iaþidarius 13.7 t7.1 49.6 Notiser om gaddsteklar (Hym.) 3. hortorum 6.2 9. I J.J - scandia are, according to the literature torum, B. hortorum and B. 4 Opuscula Entomol. 2B:189-195. þøscuorum) þascuorum 6.7 06 4.2 Cæ¡nnsnnc, B. 1976: Snylthumlan Psithyrus (Hervrrvrnn & Holu 1970, Pouvnrau are among the most abundant species ruderarius 1.8 1.1 3.1 norvegicus Sp. Schn., boparasit hos hus- 1973, Ar¡onn 1975, C¡¡annnnc 1976 in the materials of the three Fenno- sylaarum 0.3 0.3 humlan Bombus hypnorum L. (Hym., etc.), the observations of the late Erkki scandian countries. These species are aeteranus 4.1 2.2 2.5 Apidae). Entomol. Tidskr. 97:90 distinguendus 2.9 14.9 0.6 - own observations, as also the most abundant species in rthe Valkeila and our subtcrraneu.ç 30.2 47.0 25.5 Etrvrxc,-91. R. 1960: Die Hummeln und, Schma- follows (very uncertain hosts are in random materials from the several ¡otzerhummeln Finnlands. Fauna Fen- parentheses): areas in N Europe (Table 2). n 9Bl 181 353 nica 10:1-43. - PEKKARINEN AL., BUMBLEBEES IN É. FENNOSCANDIA 88 NOTULAE ENTOMOLOGICAE 6'1, 1981 A. ET 89
Proc. Int. Congr. and P. monticola (Smith, 1849) (Hy- (EH) mesipistiäisfaunasta. Ann. En- --"- 1965: Bombus semenoviellus Skor' ra, Apidae). llth - (Hym., Apoidea) in Finnland. gefunden. Entomol. 1:59'8-603.- menoptera, Âpoidea, Bombinae). En- tomol. Fennici 19:186-190. Notulae Entomol. 45:101-104' 19,73: Studies on Scandinavian bumble tomol. Scand. lO:275-296. - Vrrres.r.anr, M. 1975: Notes on the occurrence --"- -1968: Die Bienen FinnÌands, Fauna -o- bees (Hymenoptera, A.pidae). Norsk Tnnris, I. 1976a: Bumblebees, Bombus Latr. of Philanthus triangulum F. (HYm., Fennica 21:l-69. - Entomol. Tidsskr. 20:l-218. - (Hymenoptera, Apidae), on red clover Sphecoidea, Philanthidae) in Finland. *"- fauna in South Savo, Finland, Ann. Agric. Ann. Entomol. Fennici 41 :79. Fnrns, M. 1949: Den nordiska utbredningen av 1978: Notes on the Scandinavian Fenniae 15:116-127 , - Vorrro,- P. 1956: The biological zonation of Lactuata alpina, Aconitum septentrio- of social Aculeates (Hym., Vespidae and Apidae s.s.). Norwegian J. En- 1976'b: Flower visits of bumblebees, Finland as reflected in zootaxonomy. naJe, Ranunculus platanifolius and Po- - -u- (Hymenoptera, Apidae), Ann. Zool. Soc. "Vanamo" 1B'(3):1- Acta Phyto- tomol.25':165 169. Bombus Latr. lygonatum ve¡ticillatum. during one summer. Ann. Zool. Fen- geogr. Suecica 24:1-80.- Nrnurr.Ä, P. l9+7: Bräitä Karjalan Kannakselta käsin maahamme parhaillaan leviäviä nici 13:200-232. WecNnn,-36. R. 1971 : Die Veränderung der I{,tnsnrnn, V. 1972: Anthropogene Biotope mehiläislajeja (Hy-.). Ann. Ento- Txarcu, B. 1967: Sur deux espèces de Bour- Hummel-Fauna Cuxhavens in diesem (Kahlschlag, Kiesgrube, Stadtgärten) mol. Fennici 13 I75-179,- dons décrites par William Nylander (Hy- Jahrhundert. Der Versuch einer Deu- als Refugien für Insekten, untersucht am Nrr-eNonn, W. 1B4B: Adnotationes in exposi- menoptera, Apoidea, Bombus). Bull. tung. Entomol. Mitt. Zool. Mus. Beispiel der Hymenoptera Aculeata. - Hamburg- 4 (75) :207-232. - tionem monographicam Apum borea- Soc. Entomol. Mulhouse 1967:41-58. Zool. Jahrb. Syst. 9'9:133-212. lium. Not. Sällsk. Fauna Flora Fen- Varrnrr.L, E. 1953: Tietoja Etelä-Hämeen Iìer,xxa, O., RlarrrarnEN, M. & Vnn,,.srr, J' nica Förhandl- . l:164-282. 1967: Modes of balance in the PolY' apum borealium, (L.) 18,52: Revisio synoptica morphism of Philaenus spumarius -u- comparatis speciebus Europae mediae. (Ilomoptera). Ann. Acad. Scient. - Not. Sällsk. Fauna Flora Fennica Fennicae (A IV) 107:1-16. -Förhandl. 2:224-286. Selostus Referat 19'70: Danske hum- Heunnn, K. & Hor.v, S. N. PrxranrunN, A. 1979 : Morphometric, colour Iebier og snyltehumler. Natur og Mu- Kimalaisten levinneisyys ltä-Fennoskandiassa Humlornas utbredning i östfennoskandien - and enzyme variation in bumblebees seum 14:1-22. (Hymenoptera, Apidae, Bombus) in Fen- ja loiskimalaislajin Utbredningen för våra 26 arter humlor och Ilra.ru, J. I97l: European invertebráte survey, noscandia and Denmark. Acta Zool, 26,:n kimalais- kahdeksan har karterats med UTM rutsys- Instructions for record,ers. Abbots Fennica 158:1-60. - levinneisyys on kartoitettu UTM Grid -ruutu- B snylthumlor - järjestelmän ja Neuvosto- temet för Finland och nordvästra Sovjetunio- Ripton, 23 pp. 19'81: Some aspects to the morphology mukaisesti Suomessa --o- käyttäen nen. Arterna har grupperats utgående från de- Huxnvnuuo, O. & R¡erIxatllrN, M. 1971: and specific status of Bombus lapponicus liiton luoteisosissa. Lajit on ryhmitelty The recording of localities of biological perusteena niiden levinneisyyttä tutkimusalueel- ras utbredning i det undersökta området. Flere (Fabricius) and B. monticola Smith följer isotermerna för finds in Finland. Ann. Entomol. Fen- (Hymenoptera, Apidae). Bntomol, la. Useiden lajien levinneisyysraja seuraa tehois- arters utbredningsgräns ja kahdeksan den effektiva värmesumman, och B arter har ntci 37 (la): 9-12.- Scand. (in press). - ten lämpötilasummien isotermejä HrrNo, R. 1978: Climatic changes in Finland lajin pohjois- tai etelàraja on eliömaantieteelli- sin nord- eller sydgräns på den biogeografiska Prxr.lnrNnN, A. & TnnÄs, I. 1971: Suomen ki- joka av Bott- during the last hundred years. Fen- ja sellä vaihettumisvyöhykkeellä, kulkee Poh- övergångszonen, som löper från bottnen - malaisista. loiskimalaisista. Luon- janlahden pohjukasta kaakkoon. Bombus syltta- niska Viken mot sydost. Botnbus sylaarum och nia 150:3-13. non Tutkija EI I-24, - INT. BEE RBS. ASSOC. 1980: Atlas of the rum ja B. subterraneus ovat levittäytyneet 1930- B. subterraneus har spritt sig över Karelska Nä- Porov, V. V. 1930: Note on Agrobombus luvulta lähtien Karjalan kannaksen kautta set till södra Finland sedan 19'30-talet. Vidare bumblebees of the British Isles. Cam- smithianus White (Hymenoptera, Bom- bridge, 32 pp, - Etelä-Suomeen. Kirjoituksessa tarkastellaan li- diskuteras effekten av temperaturstegring och bidae). Entomol. Oboz. 24:95-99. ja muiden några andra miljöfaktorer på humlornas ut- 1979: Climatic - säksi ilmaston lämpenemisen eräiden JÄnvrxnw, O. & VÄrsÄNEN, R. Posrxnn, M. 19'51: Biologisch-ökologische Un- ympäristötekijöiden vaikutusta kimalaisten le- bredning, snylthumlors och deras värdarters ut- changes and competi- changes, habitat tersuchungen an Hummeln und ihren loiskimalaisten ja niiden isäntien bredning jämförs och humlearternas relativa dynamics geographical overlap (A) vinneisyyteen, tion: of Nestern. Veröff. Mus. Bremen levinneisyyttä sekä kimalaislajien suhteellis- abundans på vissa områden i Nordeuropa pre- pairs of congeneric bird species - in two 2:45-86. muutamilla alueilla Pohjois-Euroo- senteras. in Finland. Oikos 33:261-271. ta runsautta - Pouvna.lu, A. 1973: Les ennemis des bourdons. passa. Kersrr.l, J. 1962: Immigration und Expansion I. Apidologie 4:103-148. den der Lepidopteren in Finnland in Pur.r.r.r.tunN,- E. 1979: The Bombus and Psithy- 1869-1960. Acta Entomol. Jahren - rus (Hymenoptera, Apidae) fauna of the Fennica lB:l-452. Värriötunturi fell area in eastern Fin- Lrr,rxsoNnN, K. 1979: Effective temperature nish Forest Lapland. Notulae Ento- sums and durations of the vegetative mol. 59:159-162. - period in Fennoscandia ( 19 2 1-19'50 ) , Rar,rr,,l, 8., Lunoilnnc, H. & T¡nÄs, I' 1981: Fennia 157 :17 t-197 . Patterns of'resource utilization in two Lruonoru,- C. H. 1945: Die Fennoskandischen Fennoscandian bumblebee communities. Carabidae. Eine tiergeographische Stu- C)ikos 36,:1-1 1. die. I. Göteborgs Kungl. Vetenskaps RnrNrc,- W. F. 19,72: Ökologische Studien an Vitterhets-samhälles- Handl. 6 B 4(1): mittel- und südosteuropäischen Hum- 1-709. meln (Bombus Latr., 1802) (Hym., Api- 1949: Die Fennoskandischen Carabidae. dae). Mitt. Münchner Entomol. Ges. -"- Eine tiergeographische Studie. IIL 60:1-56.- Götebolgs Kungl. Vetenskaps Vitter-- Slrrlnnnc, J. 1BB9: Catalogus praecursorius hets-samhälles Handl. 6 B 4(3') :1-911. Ilymenopterorum Anthophilorum Fen- Lurvornnc, H. 1975: The interrelations be- niae. Meddel. Soc. Fauna Flora Fen- tween Vaccinium species and, Bombus nica -l5:167-177. (Hymenoptera, Apidae) in an arctic 1902: Yleiskatsaus Suomen Bombusla- environment. Suppl. Bull. Technique -o- jeihin. Luonnon Ystävä 6:1BB.-193. Apicole 2:191-197.- SvnussoN, B. G.- 1979: Pyrobombus lapponicus Lørnrv, A. 1961: Bombus consobrinus Dahlb., auct., in Europe recognized as two spe- an oligolectic bumble bee (Hymenopte- cies: P. lapponicus (Fabricius, 1793)