The and fauna of the Ridti archipelago in Lake Milaren, Sweden (, )

CONEN E. NILSSON

Nilsson, G. E.: The wasp and bee fauna of the Ridd archipelago in Lake Miilaren, Sweden (Hymenoptera, Aculeata). IInventering av gaddstekelfaunan i Riddarkipelagens nat-trrreservat i Malaren (Hymenoptera, Aculeata).1 - Ent. Tidskr. ll2:'79-92. UmeA, Sweden 1991. ISSN 0013-886x.

The aculeate fauna (excluding Formicidae, s.str- , and ) was surveyed in the Ridd arghipei-ago nature reserve (19 km2) and the small adjacent mainland area Solbacken (0.5 km2). The-Ridi, archipelago is characterized by a mosaic of deciduous forests (including some of nearly primeval type), coniferous forests, wet and d-ry meadows, grazing ground. grazing marshes, and parkland. During 1982 - 1990. a total of I 139 specimens iere c-ollected ii the alchipelago and more than 2000 specimens at Solbacken. 222 (45 Vo of the Swedish fauna) arelepresented in the material ( I 50 species from the archipelago). Of the species found, 35 occur on red data lists from Britain and/or Central Eurgpe.. The wood-neiting fauna was the richest. Wood-nesting sphecid were_represented by.57 species (72 4o of the wood-nesting Swedish faunaj, a figure that outnumbe-rs most Swedish piovinces (including SkAne. Oland and Gotland) and equ_als_the lqTlgr of Danish. spec-ies. bf-wood-nesting eumenid wasps and megachilid , 63Vo and 8lo/o, respectively, of all Swedish wood-iesting species were found. The richness of the wood-nesting aculeate fauna is probably related to the wealth of dead trees and fallen logs, the rich flora of flowe-ring plants, ttui exceptionally mild and dry local climate, and a continuous influence from ireighbouring populationi. Five species are reported as new to Sweden, and l3 new province records are given.

G. E. Nilsson, Department of Zoophysiology,IJppsala University, P-O. Box 560, S-751 22 Uppsala, Sweden.

Introduction course of this study, it soon became clear that the RidO archipelago contains one of the richest fau- and social wasps and bees re- In general, solitary nas of wood and stem nesting aculeates in nort- types of nesting habits: ( I those present two main ) hem Europe. who dig their nest in sand or soil, and (2) those who nest in cavities in other materials, mainly At least in northem Europe, wood and plant stems. Study area these two ecological groups include similar num- bers of species distributed among most major fa- The Rid6 archipelago is a nature reserve situated milies. in the western part of Lake Miilaren (59'30' N, , the most diverse aculeate family in 16'40' E). The main part belongs to the province Sweden, has 77 species in the first group and 19 Viistmanland (VS), and a smaller part belongs to in the second. The many aculeate species that are Siidermanland (SO). It consists of 160 islands parasites on other aculeates are generally rather ranging in size from 7.5 kmz (Ridttn) to less than host-specific, and, thus, can be referred to the 100 mz (Fig. 1). The total area is about 19 km2. same ecological groups as their hosts' The area, which has been surveyed botanically All recent Swedish regional surveys ofaculeate by Kers (1978), contains a mosaic of deciduous Hymenoptera have dealt with sandy areas domina- forests, coniferous forests, wet and dry meadows, ted by soil nesting species. The present survey is grazing ground, grazing marshes, and parkland. the first study of an area with very few sandy The deciduous forests are rich in oak, elm, lime, biotopes but rich in woodland, notably old, more ash, maple and alder. Probably due to the difficul- or less natural, deciduous forests. During the ties of transporting logs to the mainland, parts of 80 Gδ・′α″E./V′ ′ssο ′ ♪ ６

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$8. l. Map of the Ridti archipelago. A = Flaten; B-E = Ridctn; F-H = Aggarrin; I = Lilla Aggarcin; J = Hrigholmsskiir; K=Svinholmen; L-Hemmingsskiir; M=AggarciLAngholme; N=Grisslingen;O=Kurci-Nysslingen; P= Ekholmen; Q = Sundbyholmsdn; R = Nysslingen; S = Solbacken.

Karta tjver Ridd-arkipelagen med anviind omridesindelning.

the forests are in a nearly primeval state. There is a/ri (Fischer v. Waldheim) (Lundberg 1963). The a long history of pasturage (cattle) on the larger Lepidoptera fauna is also rich. For example, Ri- islands (Fig. l: areas B-E, G-H, M, and Q), and diin has the only known population of Ethmia the flora of open land is rich. The only noteworthy dodecea (Haworth) in Fennoscandia (Palm 1989), sandy biotope is an old sand pit (ca 1000 m2; on a beautiful oecophorid moth living on Lithosper- Flaten (Fig. l: area A). mum fficinale. Some notable aculeates from the The climate of westem Lake Miilaren is excep- Ridti archipelago were commented in Nilsson tional for its latitude, the mean summer tempera- ( r986). ture being among the highest (17"C in July) and the precipitation among the lowest in Sweden (Angstrdm I974). Methods The deciduous forests of the Ridti archipelago All families of aculeate Hymenoptera were sur- hold a very rich wood-living beetle fauna inclu- veyed except (Formicidae), social bees (Api- ding such rare species as Masosa curtulionide.s dae s.str.) and two families of solitary bees - And- (L.), Chlonphorus herhsti (Brahm), and Diterco renidae and Halictidae. The latter two families The u'asp and bee Jauna oJ'the Ridi arc.hipelago 8l contain only sand and/or soil nesting species. Of becomes even more impressive. After 9 years of the studied families, 489 species are known to collecting, the list of sphecid species from the occur in Sweden. Ridci archipelago (incl. Solbacken) contains the About 55 full-day collecting trips were made same number of wood-nesting species as has ever to the archipelago. Most specimens were collected been recorded from Denmark (57 species accord- with a hand-net at sites which appeared to be good ing to Lomholdr 1916 and Pulawski 1984). Den- habitats for aculeates, like logs, thatch, flowers mark is situated 250 - 600 km S of the archipelago. and sun-exposed foliage. Ca lO Vo of the material is more than 2000 times larger, has a direct conti- were collected in yellow pan traps and trap-nests. nental connection, and, ofcourse, contains a larger Apart from the Ridtj archipelago nature reserve variety of biotopes. The comparatively high abun- (Fig. 1: areas A-R), also Solbacken (Fig. l: area dance of aculeate species in the Ridtj archipelago S) was included in the investigation. Solbacken is is not due to an addition of northern elements. situated at the mainland shore (VS), only 500 m None of the species found in the archipelago has from the nearest island in the archipelago. I have its southemmost Swedish record in VS, whereas a summer cottage at Solbacken, which has allo- l3 species have their northemmost known Swe- wed an intensive collecting activity there since dish occurrence in VS. Indeed, all species found 1982. At Solbacken I have used netting, pan traps, in the archipelago are also known from continen- trap-nests and Malaise traps. This area includes tal Europe. wet and dry meadows, and more or less natural Although all biotopes found at Solbacken ap- deciduous forest, although not as old as some of pear to exist also in the Ridd archipelago (but not the deciduous forest in the Ridij archipelago. In vice versa), the intensive study of Solbacken has this paper, I only present those species from Sol- yielded several aculeate species not found in the backen that have not been recorded from the Ridii archipelago (Tab. 2). These species should be ex- archipelago. pected to occur also in the archipelago. For each species listed in Tabs I and 2, the Abbreviations used for Swedish provinces (from south to north): SK=SkAne, BL=Blekinge, HA=Halland, principal nesting habit is stated. However, the SM=SmAIand, OL=Oland, GO=Gotland. GS=Gotska choice of nest substrate is not always constant for Sandrin, oc=Ostergdtland, .VG=Viistergitland, BO= a certain species. For example, some usually Bohusliin, DS=Dalsland, NA=Niirke, SO=S0derman- ground-nesting species may occasionally nest in land, UP=Uppland, VS=Viistmanland, VR=Viirmland, hollow plant stems, decayed wood or empty snail DR=Dalama. .GA=Giistrikland. HS=Hiilsin&land.. ME= Medelpad. HA=Hiirjedalen. JA=Jiimtland, AI)=Alger- shells. This is true for species like Agenioideus manland, VB=Viisterbotten, NB=Norrbotten. AS=Asele cinctellus, c'onc'innus and A. nig,errimus lappmark, LY=Lycksele lappmark, PI=Pite lappmark, (Oehlke & Wolf 1987). Moreover, some chrysidid LU=Lule lappmark, TO=Tome lappmark. The provin- cial division of Sweden is shown on a map in each species that usually parasitize on wood-nesting volume of the series Fauna Entomologica Scandinavica. aculeates may sometimes choose a ground living wasp as host. In Sweden, the SE provinces have the richest Results wasp and bee fauna (e.g. Wolf 1969, Erlandsson During the period 1982-1990, 1020 specimens 1971, Lomholdt 1974, 1976, Erlandsson et al. were collected in the Ridti archipelaeo. In addi- 1988, Svensson et al. 1990, Janzon etal. 1991). tion, Nor6n (1988) collected I l9 specimens of the In Tab. 3, the eumenid, sphecid and megachilid presently studied families. These are included in faunas of the Ridri archipelago have been com- Tab. l. The total number of species collected in pared to those of the 10 south-eastem provinces. the archipelago is 150 (Tab. l). In addition, 72 Especially of the wood-nesting species, Tab. 3 species have been found within 200 m from the clearly shows that the Rid

Tab. l. List of species found in the Rid6 archipelago. Areas (A-R) in which the specie.s were found refer to Fig. l. Inds. denotes the number of specimens collected; c means common and that additional individuals were observed. The months when each species were collected are listed. The nesting habitat is described as: (builder) builds nest with clay or wood mixed with saliva, and is relatively free in its choice of nesting site (thus, excl. Vespa u'abro): (Col) paiasitc on Coleoptera larvae; (ground) nests in sand or soil; (Hom) parasite on Homoptera Auchenorrhyncha; (Lep) parasite on Lepidoptera larvae; (stems) nests in hollow plant stems, e.g. Rubus or Ph.ragmites; (wood) nests in more or less decayed wood; (wo/st) nests in either wood or stems; (p) the species parasitizes on aculeates with the nesting habits given. Distribution gives the southem- and northemmost province records in Sweden and the total number of the 30 Swedish provinces from which records are known; (new) new to Sweden. The following families were not found (no. of Swedish species within parenthesis): Embolemidae (l), (3), (3), and (l).

Family and Species Inds Month Nesting Distribution

Bethylidae ( l3) Ｆ 晦 Ｃ SK― VS(7) Bethylus nilidls (Thomson) Ｇ 呻 Ｃ SK― LU(24) B. fuscicornis (Jurine) Ｇ Ｃ ‐ ｏ VS(6) La e I i u s fe mo ralis (Frirster) SK― Chrysididae (46) Omalus auratus (L.) AE 2 67 wo/st(p) SK― LU(20) O. r'iolaceus (Scopoli) C 1 7 wood(p) SK― GA(5) C h ry s u ro ilrinsuta Gerstiicker G 3 6 wood(p) SM― LY(10) viridula L. M 1 7 ground(p) SK― GA(15) C. mediata Linsenmaier EM 2 ← 7 wo/gr(p) SK― NB(17) C. pseudohrevitarsls Linsenmaier F 1 6 wood(p) SM―JA(9) C. impressa Schenck AEGHMNP 36 ← 7 wood(p) SK― TO(21) C. angustula Schenck EGN 11 ← 8 wood(p) SK― LY(23) C. longula Abeille de Perrin EGM 8 ← 7 wood(p) SK― LY(20) C. equest'is Dahlbom CP 2 7 wood(p) SM― HS(5) SK― LY(25) cyanea (L.) A―R 3c “ 7 wood(p) (22) “ ４ ８ F Unclear Anteon exiguum Haupt １ ８ Hom Lonchodryinus ruficornis (Dalman) F Hom SK― TO(7)

Sapygidae (3) ６ Sopyga c'lavicornis (L.) D ６ wood(p) SK― H撃 (16) S. qui nquepunc'tata (F.) C wood(p) SK― GA(9)

(63) Pompilidae １ ７ SK― VB(19) Priocnemis exaltata (F.) G ８ ↑ ground SK― HS(15) D ipo gon subi nte rme dius (Magr.) EFGIJMN １ ６ wood SK― PI(18) D. bifasciatus (F.) E ９ ７ wood (Scopoli) SK― GA(14) Auplopus corbonarius GM ７ ８ builder (Wesmael) SK― VB(20) Arachnospila arceps AN １ ７ ground p (Zetterstedt) SK― LU(16) A. fu mi e nni s P ９ ↑ ground (Schiridte) A― SK― VB(19) A. spissa Q ４ ７ ground c' i (Spinola) SK― LY(16) Age nioide us nc te ll us AJM ４ ７ ground i i (Dahlbom) SK― LU(18) Anopl us conc nnus A １ ↑ ground ge (Scopoli) SK― LY(19) A. ni rri mus AGP ２ ７ ground Episyron rufipes (L.) AP ground SK― VB(17)

Eumenidae (36) ６ Odynerus spinipes (L.) CG ground SK― NB(17) Gymnomerus laevipes (Shuckard) ７ Ｓ ｔ ｅ ｍ ｓ ssp. scandinavicas (Saussure) SK― HR(17) Q ６ Ｗ ¨ Anc i stroc' e rus q uadratu s (Panzer) G ↑ SK― ME(18) Ｗ ｏ ／ ｓ ｔ A. trifasc iat us (Mtiller) CEGP “ SK― LU(25) Ｗ ｏ ｏ ｄ A. parietinus (L.) AEFGMNP ６ SK― TO(20) Ｗ ｏ 。 ｄ A. antilope (Panzer) E ↑ SK― LY(17) Ｗ ｏ 。 ｄ A. ot' ive ntr i s (Wesmael) AGMN ↑ SK― LU(24) Ｗ ｏ 。 ｄ Symmorphus c rassi cor n i s (Panzer) EF ↑ SK― HS(12) Ｗ ｏ ｏ ｄ S. gracilis (Brull6) ABCGM SK― LY(11)

S. bdasciatus (L.) ∈ Ｗ ｏ ｏ ｄ (mutinensis auct.) ACEFGM SK― LU(20) The wasp and bee fauna of the Ridd archipelago 83

Family and Species Arca Inds Month Nesting Distribution

“ Ｗ ／ ｓ ｔ S. de bi I itatus (Saussure) ABEFGMN0 39 NA― GA(6) “ Ｗ “ S. connexus (Curtis) CEFG 22 HA― DR(6) (Saussure) S. a I lobrogus ９ ７ wo/st SK― TO(23) (bifast'iatus auct.) CEGMN １ ７ wood SK―VB(12) Euodynerus notatus (Jurine) M ６ “ (F.) wood SK― VB(17) E. quadrdasciatus GMP １ ７ SK― LU(19) Eumenes pedunculatus (Panzer) E ０ “ builder (Panzer) ABEGM SK― HS(16) E. <'oronatus ７ “ builder (P AEGP wood SK― VS(6) D i sc oe I i u s zo na I i s anzer) (13) ｃ “ wood SK― GA(12) Vespa L. CEFG ２ “ SK― TO(28) Dol i chove spula nedia (Retzius) EF ６ builder D. norv+'egica (F.\ E 1 “ builder SK― TO(19) D. saxonic'a (F.) ADEGMP 9 ６ builder SK―TO(29) Vespula vulgaris (L.) E 1 ７ builder SK―TO(27) V. germanica (F.) F 1 “ builder SK― LY(19) V. rufa (L.) CEHM 5 ６ builder SK― TO(29) V. austriaca (Panzer) E 1 builder(p) SM― LU(18) Sphecidae (156) Ammophilo sabulosa (L.) O 1 7 ground SK― TO(26) P emphredon lugubris (F.) CGP 41 6-7 wood SK― TO(24) P. montanus Dahlbom G 2 7-8 wood SK― LU(24) P. inornatus Say CFGP 21 7 wood SK―LΨ(20) P. lethifer Shuckard D 1 “6 wood SK‐GA(12) C e rat op h o r u s mo r i o (v.d.Linden) G 2 ←7 wood SM― VB(12) C. t'lypealis (Thomson) G 3 6 wood SK― Vs(5) P a s so I oe c' u s e re mi ta Kohl G 1 6 wo/st SM―JA(12) P. corniger Shuckard CEFGMN 20 ← 8 wo/st SK― HS(17) P. insignis (v.d.Linden) CGNP 7 ← 7 wo/st SK― DR(15) P. sing,ularis Dahlbom CEFGHM 11 ← 7 wo/st SK― VB(16) Sti gmus salstyi Morawitz CEGMP 22 ← 8 wo/st SK― Hs(11) Spilomena enslin i Bltithgen G 1 7 STEMS SM―GA(8) S. troglodytes (v.d.Linden) EGM 7 8 wood SK― HS(13) S. beara Bliithgen C 1 “7 “ wo/st |⊃ M i mume sa da hlbomi (Wesmael) CGP 4 “ wood 獣 M. beaumonti (van Lith) CGPQR 10 “ wood OG―G本(5) 13 wo/st SK―`職Gネ P s e nu I u s fu st' i penn is (Dahlbom) GMNP ７ (14) P. pallipes (Panzer) ACEGI 8 ７ wo/st Ar go goryte s mystac eus (L.) CH 4 ７ ground :奎 求やB 2 ground SK― ME(14) G oryt e s lat i <' i nc t us (Lepeletier) C ７ Harpactus tumidus (Panzer) N 1 “ ground SK―VB(16) Nysson spinosus (Forster) CE 4 ７ ground(p) SK― ME(17) Astata boops (Schrank) C 1 “ ground SK― GA(17) N itela spinolae Latreille EFG 4 “ wo/st SK― DR(12) SK― Jハ Try poxy I o n c I av i c e r um Lepeletier EFG 5 “ wo/st (14) T. attenuatum Smith CG 6 ．７８ stems SK― GA(14) T. figulus (L.) CGMNP 12 ７ wo/st SK― DR(8) T. minus de Beaumont CFGP 12 “ wo/st SK― VB(14) Oxybelus unig,lumis (L.) ACD 7 “ ground SK― NB(24) Ec'temnius cephalote s (Olivier) EFG 8 “ wood SK― GA(14) E. cavifi'ons (Thomson) ACEGIP 23 “ wood SK― GA(15) E. rufi cor ni s (Zetterstedt) FG 2 ７ wood SK― TO(25) E. lapidarius (Panzer) B 1 “ wood SK― VB(20) E. continuus (F.) CEFM 11 “ wood SK― TO(25) E. tlives (Lepeletier & Brull6) CDE 7 ６ wood SK― GA(15) 2 SK― GA(7) Le sti ca c I ypea ta (Schreber) G ７ wood SK― DR(15) Entomo gnat hu s breli.s (v.d.Linden) CDG 4 ７ ground albilabris (F.) C 1 ．７８ ground SK― NB(21) clatipes (L.) CI 3 “ wo/st SK― VB(19) R. coardltum (Scopoli) AEFGHM 31 ７ wo/st SK― LU(19) R. ni gri nunt (Kiesenwetter) C 1 stcms SK― VS(5) 84 Gδ′ ENゴなsο ″ `″

Family and Species Inds. Month Nesting Distribution ６ ７ C rossoc e q u (F.\ C ground SK― GA(13) rus uadri mac lat us ５ “ podagricus (v.d.Linden) ABEFGMR wood SK― VS(10) C. ４ “ (Dahlbom) EG OG,UP,VS C. congener ３ “ wood (Shuckard) EFGMP wood SK― GA(9) C. walkeri ９ “ (Lepeletier wood GA(9) C. annulipes & Brulld) CEGM ３ “ sK― OG― DR(6) C. heydeni Kohl G ２ ６ wood (Dahlbom) BL― VB(9) C. barbipes G ４ ６ wood (L.) SK― C. leucostomus CG ９ “ wood To(26) (Shuckard) CEFG wood SK― C. cetratus １ ６ GA(12) me gacephah.rs (Rossi) G C. ７ “ wood vagabundus (Panzer) wood :鷺 C. GMP １ ６ NttB (F.) E wood SK― LU(20) C. dimidiatus １ ７ C. binotatus Lepeletier & Brulld G wood HA― VS(6)

Colletidae (25) ７ C ol lete s davlesanls Smith MN “ ground SK― NB(23) H ylaeus communis Nylander A―Q 37c ７ wo/st SK― TO(27) H. angustatus (Schenck) D l ↑ wo/st SK― ME(19) H. c'onfusus Nylander ACDFGHMQ 20 ７ wo/st SK― VB(23) H. gibbus Saunders D 1 ７ wo/st SK― ME(16) H. difformis (Eversmann) CM 8 ７ wo/st SK― DR(12) H. rinki (Gorski) CQ 2 stems BL― JA(14)

Melittidae (8) ７ M e I i tta hae morr hoida I i s (F.) D 1 ７ ground SK― VR(15) Mat'ropis labiata (F.) CDEG 7 ground SK― GA(16) (51) ７ Stelis breviuscula (Nylander) N 2 “ wood(p) SK― MF(13) S. ornatula (Klug) EG 3 “ wood(p) SK― GA(13) Heriades trunt'orum (L.) CFGNQ ll “ wood SK― ME(17) C helostoma florisomne (L.) CEG 21 “ wood SK― ME(14) C. campanularum (Kirby) CGMN 9 ６ wood SK― VB(20) Osmia pilicorni.s Smith C 1 ６ wood SK― VS(7) O. uncinata Gerstiicker C 5 ６ wood SK― TO(16) O. parietina Curtis G 1 “ wood SM―TO(18) O. leaiana (Kirby) CEM 10 ６ wood O. caerulescens (L.) C 1 “ wood :鷺えヽ3 H opl it is tube rc ulata (Nylander) CFG 10 “ wood SK― NB(16) H. clovit'entris Thomson CDEGKMQ 12 “ stems SK― LY(22) H. leucomelana (Kirby) GQ 4 “ stems SK― G4(10) M e gac hi le I i ni (Kirby) BCEFMP 12 SM― I seca ．７８ wood GA(9) M. <'e ntun<'ularis (L.) CFM 5 “ wo/st SK― HS(15) M. alpit'ola Alfken CDGMQ 12 ．７８ wood SK― VB(14) M. wrsicolor Smith CDGM 6 “ wo/st SK― VB(14) M. u' i I I u g hbi e I la (Kirby) CP 6 ６ wood SK― VB(22) M. circumcinc'ta (Kirby) C 1 “ ground SK― LU(22) M. nigriventris Schenck GQ 8 ６ wood hitli um p u nctat um Latreille G 1 ７ ground C oe I i r tt r r uJc :x c n.s Lepeletier 4 1' BC “ wood(p) :燃 |』 C. elongata Lepeletier G 2 ６ wood(p) SK― VS(13) C. inermis (Kirby) G 3 wood(p) SK― NB(16) Anthophoridae (44) “ Anthophora .furcatu (Panzer) BCEFG ６ wood SK― VB(19) Nomada hititlu Thomson E ５ ground(p) SK― LU(13) N. laucolththalnra (Kirby) FG ６ ground(p) SK― VB(12) N. murshumcllu (Kirby) CD ６ ground(p) SK― qハ (14) N..fluloguttata (Kirby) G ７ ground(p) SK JA(17) N. .lubriciunu (L.) CD ７ ground(p) SK― VS(8) Eltu tl us va ri c gatus (L.) C ground(p) SK― DR(8) The v'asp and bee fauna rf the Ridd archipelago 85 pelago contains more wood-nesting sphecid spe- Threatened species in the Ridii archipelago cies than the Baltic islands Oland and Gotland, as Unfortunately, the red data list for aculeates in well as Skflne, the southemmost Swedish pro- Sweden (Andersson et al. 1987) is still rudimen- vince. These three provinces have all been relati- tary (except for ants). Hence, I will here refer to vely thoroughly investigated with regard to acu- the more comprehensive lists issued for Britain leates. (Shirt 1987), Germany (Blab et al. 1984), Austria The fauna of wood-nesting megachilid bees in (Gepp 1984), and Baden-Wtirttemberg (Westrich the Rid6 archipelago is also very rich, only out- & Schmidt 1984, Westrich 1989). Those for Ba- numbered by 4 Swedish provinces, whereas the den-Wi.irttemberg are especially valuable because wood-nesting eumenid fauna is somewhat less im- detailed background data have been published pressive when compared to that of whole provin- (Schmidt 1979-1983, Westrich 1989). ces (Tab. 3). Tab. 4 lists the 35 species from the Ridci archi- The following 5 species are here reported as pelago (incl. Solbacken) that are included on Eu- new to Sweden: Aphelopus querceus, A. nigri- ropean lists over threatened species. The high ceps, A. serrotus, Anteon tripartitum, and Spilo- number of endangered species shows that the Ridci mena beata (dryinids determined by M. Olmi). archipelago is well worth its protection also in an Symmorphus debilitatus is reported from SO for intemational perspective. Although envi- the first time. The following 12 species are new ronmental destruction is somewhat less severe in to YS'. Emholemus ruddi, Goniozus claripennis, Fennoscandia than in the more densely populated Bethylus Aphelopus melaleucus, An- fusc'ic'ornis, parts of Europe, one may assume that the wasps teon A. pubicot"ne, A. A. flavicorne, fulviventre, and bees that are threatened in other parts of Eu- jurineanum, Mesodryinus niger, Spilomena ex- rope are potentially endangered also in Fenno- pectata, c'lypeata, and Rhopalum nigri- scandia. As almost all aculeates in Fennoscandia num. exist near their northem distributional limits, they should be especially vulnerable to environmental Discussion change. Thus, the thermophilic nature of most aculeates will put additional demands on the A combination of several favourable factors are micro-climate and the degree of sun exposure of seemingly responsible for the high diversity of their nesting sites in Fennoscandia as compared wood-nesting aculeates in the Ridd archipelago. to continental Europe. A key factor must be the mosaic of small forests with many dead and dying trees (deciduous and coniferous) mixed with grazing grounds, marshes Comments on selet'ted species and other sites rich in herbs. This mosaic provides a wealth of both nesting substrates and nutritional Symmorphus debilitatus is here recorded from SO resources. Pollen and nectar are the main energy for the first time. It appears to be very rare within sources for adult aculeates and for the larvae of its entire distributional range (Bltithgen l96l), bees. Another essential factor is probably the ex- and has a scattered occurrence in northem and ceptionally warm climate in the western part of central continental Europe (Cumming 1989). In Lake Miilaren, with hot summers, comparatively the Ridd archipelago, it was found nesting in large mild winters, and low precipitation. Finally, the numbers in eaves of thatched roofs (Phragmites) presence of several similar areas (notably Tidti, on old cottages and barns. I have also seen it Strijmsholm, and Angsci) within 20 km from the nesting in beetle-tunnels in logs of old deciduous archipelago is likely to be of importance. Modem trees, especially oak. It also nests in maple (L.-A. theories on metapopulations predict that the pre- Janzon in litt.) The only other sites where I sence of many small populations within a limited have found debilitatus are Tidti in VS, and Bi- geographic area will improve the chance for survi- skops-Amd and Kungshamn (S of Uppsala) in UP, val of a species during periods of unfavourable i.e. areas still rich in old deciduous trees. Needless conditions (Gilpin 1987). Thus, in the long run, to say, thatched cottages, and deciduous forests neighbouring populations may very well be essen- rich in dead and dying trees, are decreasing in tial for preserving a diverse fauna in the Rido number or area. S. debilitatus has been found to archipelago. prey on larvae of a small oak leaf-mining moth 86 Gdran E. Nilsson

Tab. 2. List of species found at Solbacken (area S) but hitherto not recorded from the Ridci archipelago Se legend to Tab. I for explanations.

Family and Species Month Nesting Distribution

Embolemidae E mbole mus rzrldii Westwood unknown sK-ME(7) G oniozus c' lari pe nnis (Fdrster) 6-8 Lep sK-AN(13)

Chrysididae ２ ７ (F.) Omalus aeneus ２ “ wood(p) sM-ME(7) H ground(p) edychrum niemeluei Linsenmaier １ ７ sK-GA(13) h s i b i t' o I o ground(p) OL,UP,VS C ry s r Lepeletier １ ５７ ． wood(p) C. fulgida L. １ ６ sK-AN( l8) C. mediadentata Linsenmaier ２ “ wood(p) sK-LY(16) C. schencki Linsenmaier wood(p) sK-LY( l3)

Dryinidae ３

Ap he lopus me la le uc' us (Dalman) ７ 5-6 Hom sK-LU(4)

A. querceus Olmi １ G8 Hom VS(new) '7 A. nigric'eps Kieffer ４ Hom VS(new)

A. serratus Richards ２ 6-7 Hom VS(new) (Dalman) Anteon flavicot'ne ８ 6-7 Hom vG,vs(2) (Dalman) A. pubicorne １ 6-8 Hom sK-TO(7)

A. tripartitum Kieffer １ 5 Hom VS(new)

A. gaullei Kieffer ０ 8 Hom sK-vs(4) (Haliday) A. fu lvive nt re ５ 6-7 Hom UP,VS jurineanum A. Latreille １ 6-7 Hom sK-vs(4) M e sodry i nus niger (Kieffer) 7 Hom VS.VR Sapyga similis (F.) 5-6 wood(p) sM-LU(14)

Mutillidae ９ “ ground Myrmosa atra Panzer ３ “ sK-vB(12) Smicromyrme rurtpes (F.) ground sK-HS( r7)

Pompilidae ７ “ P rioc nemis pe (Harris) ground rturbator １ ７ sK-LU(18) P. pusilla ground Schiddte ０ ７８ sK-vR(l l) schioedtei ． ground P. Haupt ６ ７８ sK-HS( r6) ． ground l P. cordivalvato Haupt ９ ７８ SK-NB( l ) (F.) ． ground P. hyalinato ２ “ sK-GA(ls) P. ground fennica Haupt ２ ７ sK-vB( l3) C al iadur gus iate I I us (Spinola) ground fasc ７ “ sK-GA(13) varie gatus (L.) wood １ ６ sK-GA(r6) A ra c h no s p i I r (Haupt) ground SK-HS( I 3 a ufa １ ５ ) Anoplius viaticus (L.) ground SK-AN( I9)

Eumenidae ６ ．５８ An<' i stroc e ni grit'orn i s (Curtis) wood rus ４ “ SK-LY(15) Symmorphus murarius (L.) wood sK-cA(8) Sphecidae “ Ｓ Ｋ Ｖ Ｂ ＜ ‐ ８ ＞ ・ Dol ic' u I (Spinola) hurus corni<' us 20 ％ ground Ｓ Ｋ Ｎ Ｂ ＜ ‐ ８ ＞ ・ P e m p h re do n /lrgens Dahlbom 30 “ wood Ｓ Ｍ 」 Ｕ ＜ ‐ ６ ＞ ・ D t i i odo n u s me d us Dahlbom t2 ground Ｓ Ｋ Ｖ Ｓ ＜ ‐ ３ ＞ ７ ・ P assaloec grac i I i (Curtis) us s I ７ wo/st Ｓ Ｋ 」 Ｕ ＜ ‐ ６ ＞ P. Dahlbom I ・ turionum ５７ wood Ｓ Ｍ 」 Ｙ ＜ ‐ 。 ＞ P. boreolis Dahlbom 8 ． ・ ７ wood Ｓ Ｋ Ｖ Ｂ ＜ ‐ ６ ＞ ・ Sp i lome na diffe re ns Bliithgen 2 ７ wood Ｓ Ｋ Ｇ Ａ ¨ ＜ ‐ ‐ ＞ ・ S. e.rpectata Valkeila I “ stems ¨ P se nu I us conc o (Dahlbom) lor t4 ７ wo/st Ｏ 」 Ｇ Ａ ＜ ９ ＞ P. schentki (Toumier) 2 ・ ７８ wood Ｓ Ｋ Ｍ Ｅ ＜ ２ ０ ＞ Cerceris arenoria (L.) 6 ． ground ・ ７ Ｓ Ｋ Ｌ Ｕ ＜ ‐ ５ ＞ ・ C. q uad r ifa s <' i a t a (P anzer) 3 ground The wasp and bee fauna of the Ridd archipelago 87

Family and Species Inds. Month Nesting Distribution ７ ground Ar go goryt e s far3er (Shuckard) ７ sK-LU(22) (F.) ground SK-NB(18) G o ry t e s q ua dr ifasc iotus ．７８ N ysson trimaculatus (Rossi) ７ ground(p) sK-qA04) N. dimidiatus Jurine ．７８ ground(p) SK-AN( l2) Mellinus arvensis (L.) ６ ground sK-ME(20) Astata pinguis (Dahlbom) ７ ground sK-LU(20) ground sK-vR(14) T ac hy s p h e.r oh s c' u r i p e nni s (Schenck) ．７８ T. pompiliformrs (Panzer) ７ ground sK-NB(21) wo/st sK-vB(ls) N i t e la borea lis Valkeila ．７８ C rabro c ribrat'lzs (L.) “ ground sK-vB(23) rubit'ola (Duf. & Perris) ．５７ stems sM-ME(10) E. guttatus (v.d.Linden) “ wood sK-vB(15) E. boreali s (Zetterstedt) “ wood sK-TO(27) Crossoc'erus ovalis Lep. & Brull6 ７ ground sK-ro(18) C. pusillus Lepeletier & Brull6 ground sK-vB(17)

Colletidae ４ ７ ８ ． wo/st SK-JA(18) Hylaeus brevicornis Nylander ３ “ wo/st sK-TO(27) H. annulatus (L.) ９ “ H. hyolinatus Smith wo/st sK-AN(20) Megachilidae Anthidium manicatum (L.) 2 7 ground sK-JA(19) S t e I i s punc t ulatissirra (Kirby) I 7 wood(p) SK-HS(16) Osmia rufa (L.) .J 5 wood SK-GA(13) '7 M e gachi I e lapponic a Thomson I wood SM-TO(10) C oe I io.ry s I u nc e o lat a Nylander I 6 wood(p) sM-LY(7) Anthophoridae ７ ground Ant hop hor a q uadr i mac ul at a (Panzer) 価 SK-AS(23) N omada obscura Zetterstedt ７ ground(p) sK-LU(14) N . r obe rj e ot i a na P anzer ground(p) sK-LU(2r)

lTist'heria complanella (Hiibner)l in France logs in the Ridci archipelago and in some other (Bliithgen 1961). However, Van Lith (1964), who sites rich in deciduous forest and close to the wes- gave a detailed account of the nesting of debilita- tem part of Lake Miilaren. Unlike S. debilitatus ('o,',- /rzs in the thatch of an old bam in Ulvenhout (Hol- and S. allobrogus, I have hitherto not found land), found that it can use several species of Mic- ,?exus nesting in thatch. However, Jorgensen rolepidoptera as larval provision. Nevertheless, if (1942) found (:onnexus nesting in a thatched roof debilitatus is relatively fastidious in its choice of in northem Sjaelland, Denmark. He observed that prey and nesting site, this may contribute to its ('onne-yus preyed upon leaf mining larvae of Zeu' rarity, and dehilitatus should probably be regarded gophora subspinosa (Fabricius) (Col., Chrysome- as endangered or vulnerable in Sweden. Indeed, lidae) and Caloptilia stigmatella (Fabricius) debilitatus is regarded as potentially endangered (Lep., Gracillariidae) in leafs of Populus tremula. in northem Germany (Haeseler 1978). On Rid

near barkless logs of oak, probably in search for Spilomena beata is here recorded from Sweden a suitable nesting substrate. S. murarius nests in for the first time. The single male collected was beetle tunnels in wood, in thatch and in holes in found at Viisteriing on Ridcin, l.VII.90, on a stone walls, and uses larvae of Chrysomela populi thatched roofofa barn. The species is known from L. (Col., Chrysomelidae) that lives on Populus Finland, south and central Europe, and England tremula. as larval provision. (Dollfuss 1986, Vikberg 1986). Due to its small Discoelius zonalis is abundant in at least 4 areas size (2.5-3.0 mm), it may have been overlooked. in the Ridti archipelago, all with deciduous forests However, as all congeners are of this small size. of primeval type and rich in lime-trees, viz. Flaten beata is probably considerably less abundant than (area A), Osterangen on Rid

Tab. 3. Species richness of Eumenidae, Sphecidae and Megachilidae in Sweden, in each of the south-eastem provinces, and in the Rid6 archipelago. Percentage values refer to the number of species recorded from Sweden.

Familv and Area Total no. No. of species nesting in: of species Wood/stems Ground/buildcr

Eumenidae Sweden 36( l00o/o) 27 (IOlc/,) 9(l0OVc) Rid6 archipetago Areas A-R l8(50olc) l5(56c/c) 3(33so) Areas A-S 2O(56ok) t7(63Ec) 3(33Vo) Swedish provinces Skine 28(7sEc) 22(8la/a) 6(66Ec) Blekinge t3(36ok) gt 11o/, \ 4(44o/o) Smfiland 29(8tok) 23(85Vc) 6(66Vo) Oland 29(8tEa) 23(85Ea) 6(66Ea) Gotland 22(6lVo) 17(63E{,) 5(557o) Ostergotland 26('724/o) 2t('78Ea) 5(55Vc) Niirke 2l(58o/o) l6(59Vo) 5(55Vc) Sddermanland 28(7sok) 22(8la/o) 6(66Vo) Uppland 31(86ok) 25(93oft,) 6(667o) Vzistmanland 25(69o/a) 2l(7\Vo) 4(44Vo) Sphecidae Sweden 1s6( 100%) 79(t00Eo) 77(t00ok) Ridri archipelago Area A-R 55(35o/o) 45(57o/a) 10(t3o/c) Area A-S 82(537o) 5702qa) 25(32Vo) Swedish provinces Skflne I l9('76ok) 55(70o/o 64(84Ea Blekinge 69(44o/o) 33(42o/a 36(41Eo Smflland I l8(7 6Vo) 62(78%, 56(73Vc OIand 112(727o) 5266Vc 60(78ch Gotland 96(620/o) 44(56c/c 53(69Ea Ostergcitland I 16(7 4o/o) 65(82Vc 5l(66Vc Niirke 91(58Vo) 48(6lVc 43(56Eo Scidermanland lO0(640/o) 54(68Vc 46(6oc/a Uppland I 16(7 4o/o) 67(85Vo 49(64Vc Vastmanland 99(63Vo) 63(80Vo 36(47Eo Megachilidae Sweden 5l(100o/o) 32(l00Vo l9( 1 00c/o) Rid6 archipelago Area A-R 24(477o) 22(697o) 2(1lqo) Area A-S 29(57o/o) 26(8lVo) 3(16%,) Swedish provinces SkAne 39(767o 25(78o/c) 14('7 4Eo) Blekinge 23(45o/o A@4qo) 9(47ok) SmAland 4t(80ok 30(94ok) 1l(58Vo) Oland 37(727o 22(697o) 15(79Vc) Gotland 32(634/o 22(69Va) t0(53Ea) Ostergdtland 43(84o/a 3t(97ok) t2(637c) Niirke t5(29ok I t(34Eo) 4(21Vc) Scidermanland 39(',767a 30(94Vo) 9(47a/c) Uppland 40('784/a 29(9tok) I t(58Vo) Viistmanland 32(63oh 26(8lVo) 6(32Vc)

as thermophilic, being common in the Mediterra- l.VII.90. R. nigrinum occurs mainly near fresh or nean area. brackish water, where it has been found nesting Rhopalum nig,rinum. The single female hitherto in Phragmires (Lomholdt 1976). This apparently collected in the archipelago was netted in the sporadic species is in Sweden known from 5 pro- Phrogmites zone at Tegelviken on Ridcin vinces, VS being the northemmost. 90 Gdran E. Nilsson

Tab. 4. Species of Aculeata from the Ridti archipelago in the Rid6 archipelago.. (4 9 23.VI.-g.VIII. (areas A-R) and Solbacken (area S) that are included on Lindsbo on Aggarcin and Osterdngen on Ridcin) European red data lists: (D) Germany, Blab et al. ( 1984)l were all flying near decayed deciduous wood wit- (BW) Baden-Wtirttemberg, Westrich & Schmidt (1984) & Westrich (1989); (GB) Great Britain, Shirt (1987); hin forests. I have also found c'ong,ener in a very (A) Austria, Gepp (1984). The fotlowing categories are similar biotope at Broholmen near Strijmsholm, used: (l) on the edge of extinction; (2) endangered, VS (l ? 27.Vll.g0). Hence, congener seems to (3) po- referred to as category 2-3 on continental lists; differ from most other Crossoc'erus species by tentially endangered or vulnerable, referred to as cate- gory 2 on the British list and category 4 on continental preferring a shady habitat, and it might be associa- lists; (-) family not covered by list, or species not recor- ted with natural or primeval deciduous forests. ded from area. u,alkeri is common in the Rido archipelago and at Solbacken but elsewhere rare. BW GB Species It nests in decayed wood (e.g. alder, oak and ash) ３ 一 near water and uses Ephemeroptera as larval pro- Chrysura hirsuto １ 一 Chrysis Julgida vision (Schmidt 1980). Due to water pollution and 一 Sopyga similis _, 一 the removal of dead wood near \tater, walkeri is 一 二 Arat hno spi La nnis 2 fumipe １ regarded as endangered ("stark gefdhrdet") in A rufct 22 Symmorphus murarius 2 一 Germany (Schmidt 1980, Blab et al. 1984, We- Odynerus spinipes 22 strich & Schmidt 1985). It is distributed from Eumenes pedunculatus 2 southem Fennoscandia and Central Europe east- Dist'oelius zonalis wards to Japan. Vespa crabro DoIichot,espula media Crossocerus heydeni is like C. cong,ener a

Cerotophorus morio -1 wood-nesting species that is rare in Sweden and C. t'lypealis 3 in the rest of Europe (Lomholdt 1976, Schmidt Passaloecus eremita 3 1980). This boreoalpine species is regarded as po- Argogorytes fargeii 2 Crossocerus w'alkeri tentially endangered in Baden-Wi.irttemberg (We- C. congener 二 strich & Schmidt 1985). The specimens collected hevdeni C. １ in the Ridri archipelago (192d) were found on C. vagabundus decayed deciduous logs in a similar (or even the H. ang,ustatus 一 一 same) shady habitat as that of congener. H. dffirmis １

Stelis breviusculo Crossocerus binotatus is rare and has a scatte-

Osmia pilicornis ３ red occurrence in Sweden, north to the Ridci archi- O. uncinata ２ pelago. It is known to nest in old timber and logs. O. leaiana ３ The single female collected in the archipelago Megachile ligniseca ２

M. cirt'umcincta ３ (28.VII.85, near Lindsbo on Aggartin) was found M. alpicola ２ flying near decayed deciduous logs inside a deci- M. nigriventris １ forest. Coelioxys rufecens duous 一 Megachile ligniseca is rare, and has a scattered C. lanceolata ２

Nomada leucophthalma ３ occurrence in northem, central and eastem Eu- N. obscura ３ rope. This big leafcutting bee nests in dead wood. N. roberjeotiana ２ cossas (L.), and is regar- Epeolus variegatus e.g. in tunnels of Cossas ded as a forest species (Westrich 1989). Although ligniseca is generally very rare in Sweden, it is Crossocerus congener is one of the rarest spe- common in the Ridij archipelago, suggesting that cies of the genus in Sweden; only known from it is favoured by an abundance of dead deciduous OC, Up and VS. It has a sporadic occurrence in wood. It is regarded as potentially endangered in Europe, and it is regarded as potentially endan- Baden-Wiirttemberg (Westrich 1989), and, be- gered in Baden-Wtirttemberg (Westrich &. cause of the disappearance of natural forests, /ig- Schmidt 1985). It has been found nesting in wood nl.seca should probably be given the same status in Sweden and Germany (Kjellander 1954, in Sweden. Schmidt 1980) while Japanese individuals (ssp. Coelio.rys lanceolata is probably the rarest spe- fukuiensis Tsuneki) apparently nest in hollow cies of its genus in Europe (Westrich 1989). This plant stems (Tsuneki 1960). The specimens found boreoalpine species is very rare also in Fenno- The wasp and bee fauna of the Ridd archipelago 9l scandia (Erlandsson 1955, Rassi & VAis5nen Wiss. Berlin, Kl. f. Chem., Geol., Biol., l96l(2): l- 1987). It is known to parasitize on Megachile ni- 251. Cumming, J. M. 1989. Classification and evolution of griventris, a big leafcutting bee that occurs in the the eumenine wasp genus Symmorphus Wesmael Ridri archipelago. C. lanceolata is considered to (Hymenoptera: Vespidae). Mem. ent. Soc. Can. be on the edge of extinction in Germany, where 148: l-168. - it is only known from Bavaria (Blab et al. 1984), Dollfuss, H. 1986. Eine Revision der Gattung Spilomena Shuckard der westlichen und zentralen paliiarktischen possible given a and it is that it should be similar Region (Hymenoptera, Sphecidae). Ann. naturhist. status in Sweden. The single female collected in Mus. Wien. 88/89: 481-510. - the area (18-24.VI.89., Solbacken) was caught in Erlandsson, S. 1955. Die schwedischen Arten der Gat- a Malaise trap near some dry barkless sun-expo- tung Coelioxys Latr. Opusc. ent. 2O: 174-191. Erlandsson, S. 1971. Catalogus- Insectorum Sueciae sed oak logs. XIX. Hymenoptera: Aculeata. Ent. Tidskr. 92:87- 94. - Erlandsson, S., Janzon, L.-4. & Svensson, B. G. 1988. Concluding remarks Catalogus Insectorum Sueciae. Hymenoptera, Apoi- dea. 1. Colletidae and . Ent. Tidskr. 109: When compared to other areas, even entire provin- l6l-163. - ces or countries, the Ridci archipelago contains an Cepp. J. 1984. Rote Listen gefiihrdeter Tiere Oster- impressive number of wood- and stem-nesting reichs. Wien (Bundesministerium ftir Gesundheit und aculeate species. Even so, it is evident that there Umweltschutz). Gilpin, M. E. 1987. Spatial structure and population are still species in the area that remain to be found. vulnerability. In: Soul6, M. E. (ed.). Viable popula- For example, three sphecid wasps species, Spilo- tions for conservation.- pp. 125-139. Cambridge mena beata, Lestica clypeata and Rhopalum nigri- (Cambridge Univ. Press). num, and two megachilid bees, Anthidium mani- Haeseler, V. 1978. Flugzeit, Bliitenbesuch, Verbreitung und Hiiufigkeit der solitiiren Faltenwespen im Nord- catum and Megachile lapponica, were found in deutschen Tiefland (BRD) (: Eumeni- the Ridd archipelago or at Solbacken for the first dae). Schr. Naturw. Ver.- Schlesw. Holst. 48: time in 1990, i.e. during the 9th year of the survey. 63-131.- - Indeed, of the 222 aculeate species recorded in Haeseler. V. 1984. Mimumesa sibiricana R. Bohart, eine fiir die Bundesrepublik Deutschland neue Grabwespe, the Ridci archipelago and at Solbacken so far, 66 und weitere ftir Norddeutschland seltene Hautfltiger have been found only as single specimens. More- (Hymenoptera: Aculeata s. l.). Drosera 84: 103- over, although all biotopes at Solbacken occur in 1 16. - Wespen the Ridd archipelago, extensive collecting activity Haeseler, V. 1985. Zum Auftreten von und Bienen in einem abgestorbenen Birkenbestand im at Solbacken has resulted in 72 species hitherto Ostlichen Htigelland Schleswig-Holsteins (Hymenop- not recorded from the archipelago. Thus, there is tera: Aculeata). Faun.-rikol. Mitt. 5: 345-363. every reason to expect further interesting records Jacobs, H.-J. & Oehlke,- J. 1990. Beitriige zur Insekten- from the Ridci archipelago, provided that this el- fauna der DDR: Hymenoptera: Sphecidae. l. Nach- trag. Qeitr. Ent. Berlin 40: l2l-229. sewhere increasingly rare mixture of biotopes will Janzon. -L.-A.. Svensson, B. G. & Erlandsson, S. 1991. continue to exist within the archipelago also in the Catalogus Insectorum Sueciae. Hymenoptera, Apoi- future. dea. 3. Megachilidae, Anthophoridae and Apidae. Ent. Tidskr. l12: 93-99. - Acknowledgements. I thank Dr Massimo olmi (vi- Jprgensen, P. 1.942. Biological observations^on some vespides..- Ent' Meddr' 22:299-33s' ;;rd-i;;tii;; d;;;r-iri";-rll tte Jiyinios, Dr Lars -- solitarv neasirtimii-lppsala) for a.t.".rnining nro"i ofiirr.trysi- Kers, L. E. 1978. Ridti-arkipelagen i Miilaren. Naturin- ventering. Del' I .: och_ flora. Statens dia;; ;A rji b'unnui Steintroiiiiup?.ilu) for .orn16n6 _vegeration - on the manuscript. Kj:iilH:1',tl';!##,' ,i."T,l lll.ly,'.1flil'".,. Grabwespen aus Schweden nebst Beschreibung des Mlinnchen von Crabro conggner Dahlb. und- Crabro Kererences styrius Kohl (Hym., Sphecidae). Ent. Tidskr. 75: Andersson, H., Coulianos, C.-C., Ehngtrdm, 8., Harl- 238-244. - marstedi, O., ImUy, L., Janzon, L.-A., Lindelow, A. Lomholdt, O. 1974. Grundtraek af udbredelsen af gra- & Walddn, H. W. i987. Hotade evertebrater i Sverige. vehvepsene i Fennoskandien og Danmark (Hymenop- Ent. Tidskr. 108: 65-75. tera Aculeata, Sphecidae). Ent. Meddr. 42: 189- Blab,- J., Nowak, E. & Trautmann, W. 1984. Rote Liste 209. - der gef?ihrdeten Tiere und Pflanzen in der Bundesre- Lomholdt, O. 1976. The Sphecidae ofFennoscandia and publ'ik Deutschland. Greven (Kilda-Verlag). Denmark. Fauna ent. scand. 4: 1452. Bltthgen, P. 1961. Die Faltenwespen Mitteleuropa-s Lundberg, S.- 1963. Bidrag till ki.innedomen om svenska (Hfmenoptera, Abh. Dtsch. Akad. skalbaggar 7. Ent. Tidskr. tl4: 242-246. Diploptera). - - 92 Griran E. Nilsson

Nilsson, G. E. 1986. Nya landskapsfynd av gaddsteklar, van Odynerus (Symmorphus) debilitatus (Sauss.; med en riversikt av de fennoskandiska artema i famil- (Hymenoptera aculeata, Eumenidae). Ent. Ber. jen Dryinidae. Ent. Tidskr. 107: 85-90. (Amst.) 24:256-262. - Nor6n, L. 1988. Rapport- frAn exkursion: Riddn, gadd- Vikberg, V. 1986. A checklist of aculeate Hymenoptera steklar. Graphosoma 3(2): ll-12. (Stencil.) of Finland (Hymenoptera, Aculeata). No- Oehlke, J.- & Woli H. 1987. Beitriige zur Insekten- tulae Ent. 66: 65-85. - Fauna der DDR: Hymenoptera Pompilidae. Westrich. P. 1989. Die Wildbienen Baden-Wtirttem- Beitr. Ent. Berlin 37: 279-390. - - bergs I-II. Stungart (Verlag Eugen Ulmer). Palm, E. 1989. Nordeuropas Prydevinger (Lepidoptera: Westrich, P. & Schmidt. K. 1985. Roten Liste der Stec- Oecopholidae). Danmarks Dyreliv 4, Kobenhavn. himmen Baden-Wi.irttembergs (Hymenoptera Acu- Pekkarinen, A. 1989. The homet (Vespa crabro L.) in leata ausser Chrysididae). Ver6ff. Naturschutz Finland and its changing northem limit in northwes- Landschaftspflege Bad.-Wtirtt.- 59160: 93-120. tem Europe. - Ent. Tidskr. I l0: 16l-164. Wolf, H. 1969. Catalogus Insectorum Sueciae XVII. Hy- Pulawski, W. J. 1984. The status of Trypoxylon figulus " menoptera: . Opusc. ent. 34: l2-16. (Linnaeus, 1758), medium de Beaumont, 1945, and Angstrdm. A. 19'74. Sveriges- Klimat. 188 sid. GLA, minus de Beaumont, 1945 (Hymenoptera, Sphecidae). Stockholm. Proc. Califomia Acad. Sci. 43: 123-140. Rassi,- P. & Viiisiinen, R. 1987. Threatened and plants in Finland. Helsinki. Schmidt, K. 1979. Materialien zur Aufstellung einer Ro- Sammanfattning (Grabwespen) ten Liste der Sphecidae Baden-Wtirt- Gaddstekelfaunan i Ridciarkipelagens naturreser- tembergs I. Philanthinae und Nyssoninae. Verdff. Naturschutz Landschaftspflege Bad. Wtirtt.- 49150: vat (SO och VS) samt Solbacken, ett angrdnsande 271-369. - strandomr6de i VS, har inventerats under 1982 - Schmidt, K. 1980. Materialien zur Aufstellung einer Ro- 1990. Sammanlagt har 222 arter insamlats, vilket ten Liste der Sphecidae (Grabwespen) Baden-Wtirt- utgcir 45 Vo av den svenska faunan av de studerade tembergs II. . Vertjff. Naturschutz gaddstekelfamiljema. gaddste- Landschaftspflege Bad.-Wiirtt.- 5 I 152: 309-398. Den vedlevande Schmidt, K. 1981 . Materialien zur Aufstellung einer Ro- kelfaunan visade sig vara extremt rik, sannolikt ten Liste der Sphecidae (Grabwespen) Baden-Wiirt- ocivertriiffad i Nordeuropa. Bl a pltriiffades 57 (ausser tembergs IIL Oxybelini, Larrinae Trypoxy- arter vedlevande rovsteklar (Sphecidae), vilket iir lon), Astatinae, Sphecinae und Ampulicinae. Verdff. Naturschutz Landschaftspflege Bad. - samma antal som har p6triiffats i Danmark, och Wiirtt. 53/54: 155-234. - mer an vad som iir kiint frAn t ex Sk6ne, Oland Schmidt, K. 1983. Materialien zur Aufstellung einer Ro- eller Gotland. Av vedlevande buksamlarbin (Me- ten Liste der Sphecidae (Grabwespen) Baden-Wtirt- gachilidae) patraffades 26 arter, vilket utgdr 8l 7o tembergs IV. Pemphredoninae und Trypoxylonini. - - Vertiff. Naturschutz Landschaftspflege Bad.-Wtirtt. av den svenska faunan. M6ngfalden av vedlev- 57 158: 2t9-304. ande gaddsteklar torde hdnga samman med den Shirt, D. B. 1987. British Red Data Books: 2. . rika fiirekomsten av murken ved i kombination pp. 402 Nature Conservancy Council. Londo"n. med en rik flora gynnad av bl a kreatursbete. Ett Svensson, B. G., Erlandsson, S. & Janzon, L.-A. 1990. Catalogus Insectorum Sueciae. Hymenoptera, Apoi- flertal av arterna iir troligen bundna till iidelldv- dea. 2. Andrenidae and Halictidae. Ent. Tidskr. skogen, vilken delvis dr urskogsartad. NAgra arter lll:47-52. - verkar vara mer eller mindre knutna till vasstak Tsuneki, K. 1960. Guide to the study of Japanese Hyme- pA gamla lador. Av de petreffade gaddstekelar- noptera 10.3. 7. Biology of the Japanese species of Crossocerus. Life Study (Fukui) 4(l). tema f