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Osmia (Melanosmia) steinmanni sp. n., a new bee species from the Swiss Alps (Hymenoptera, Apoidea, Megachilidae) PDF

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Revue suisse de Zoologie 109 (4): 803-812; décembre 2002 Osmia {Melanosmia) steinmanni sp. n., a new bee species from the Swiss Alps (Hymenoptera, Apoidea, Megachilidae) MÜLLER Andreas InstituteofPlantSciences,AppliedEntomology, Swiss Federal Institute ofTechnology (ETH), Clausiusstrasse 25/NW, CH-8092 Zürich, Switzerland. E-mail: [email protected] Osmia {Melanosmia)steinmannisp. n., a newbee species from the Swiss Alps (Hymenoptera, Apoidea, Megachilidae). - Osmia {Melanosmia) steinmanni sp. n. is described, illustrated and compared to other European species ofthe subgenus Melanosmia. It is morphologically most closely re- lated to O. svenssoni Tkalcu, 1983 hitherto recorded only from northern Sweden. O. steinmanni is known from two localities in eastern Switzerland situated in the subalpine and alpine zone oftheAlps. It was found on stony slopes exposed to the south-east. Known pollen sources are Hippocrepis comosa and Lotus corniculatus (both Fabaceae). Key-words: Megachilidae - Osmia -Melanosmia - taxonomy - new species - Switzerland. INTRODUCTION Within the scope ofthe preparatoryworkforthefourth volumeofthe Swissbee fauna (Amiet et al., in preparation) seven 9 9 of an unknown Osmia species (Hymenoptera,Apoidea, Megachilidae) werefound in theentomological collections of the ETH Zürich and ofthe University ofZürich. All specimens had been collected at m the Ebenalp (1500 above sea level) situated in the Alpstein massif in eastern Switzerland on 10th June 1907. An excursion to the Ebenalp nearly one century later, on 1st June 2002, resulted in the discovery of several 9 9 and ofone cT ofthis enig- matic Osmia. The inspection ofall museum and private bee collections in Switzerland revealed one further â which had also been collected in the Swiss Alps near Juf m (Avers, Graubünden, 2300 above sea level) on 19th June 1994. On closer morphological inspection these specimens turned out to belong to a hithertounknownbee species. Osmiasteinmannisp. n. is atypical representativeofthe Holarctic subgenus Melanosmia Schmiedeknecht, 1885. In the Nearctic region Mela- nosmia is the major subgenus of Osmia with 91 described species (Hurd, 1979; Michener, 2000). Seventeen species are known in the Palaearctic (Zanden, 1988), ten species occur in Europe (Tkalcû, 1983). Manuscript accepted24.10.2002 804 A. MÜLLER SYSTEMATIC PART Osmia (Melanosmia) steinmanni sp. n. Figs 1-7 Material Holotype: 9 Switzerland, Appenzell Innerrhoden, Wasserauen, Ebenalp, 749/238, , 1500m, 1.6.2002, leg. FelixAmiet, coll. ETHZürich. Paratypes: 59 9, let, same locality and date as holotype, leg. FelixAmiet, Mike Herr- mann and Andreas Müller, coll. ETH Zürich (19, le?), coll. Muséum d'histoire naturelle Genève(1 9),coll.FelixAmiet(19),coll. MikeHerrmann(19)andcoll.AndreasMüller(19); 79 9, same locality as holotype, 10.6.1907, coll. ETH Zürich; lo", Switzerland, Graubünden, Juf-Stallerberg, 2300 m, 19.6.1994, leg. Bernhard Merz, coll. FelixAmiet. Etymology Thenew speciesisdedicatedtotheexcellent bee specialistDrErwinSteinmann (Chur) who is as fond ofthe mountains as "his" bee seems to be. Diagnosis O. steinmanni sp. n. belongs to the subgenus Melanosmia Schmiedeknecht, 1885 described in detail by Tkalcû (1983). In both sexes it is morphologically most similar to O. svenssoni Tkalcü, 1983 hitherto known only from northern Sweden (Tkalcû, 1983), the 9 can easily be confounded with O. inermis (Zetterstedt, 1838) as well (see below). The diagnosis is given in the form of keys and tables. The keys including all European Melanosmia species except O. bulgarica Friese, 1922 (see discussion) lead to O. steinmanni and its morphologically most similar relatives which are then com- pared in tabularform. The morphological terminology follows Micheneretal. (1994). The opinion that O. svenssoni and O. uncinata Gerstäcker, 1869 are syno- nymous (Schwarz etal., 1996) is notfollowed here. The examinationofthe type speci- mens of O. svenssoni corroborates the view of Tkalcü (1983) that O. svenssoni and O. uncinata are valid species morphologically clearly separated from each other. Female 1 Propodeum shiny, propodeal triangle nearly completely polished or at least along sides with shiny area. Body length at least 11 mm. Osmia alticola Benoist, 1922; Osmia maritima Friese, 1885; Osmia xanthomelana (Kirby, 1802). SeeTkalcu (1983) andHaeseler(1999) for distinctive characters. 1* Propodeum and propodeal triangle completely shagreened and dull. Body length at most 10 mm 2 2 Declining basal portion of first tergite shiny, at most superficially sha- greened here and there. Osmia hyperborea Tkalcu, 1983; Osmiaparie- tina Curtis, 1828; Osmiapilicornis Smith, 1846; Osmia uncinata Ger- stäcker, 1869. See Tkalcû (1983) and Haeseler (1999) for distinctive characters. 2* Declining basal portion of first tergite densely shagreened, only with silkylustre. Osmiainermis (Zetterstedt, 1838); OsmiasvenssoniTkalcü, 1983; Osmia steinmanni sp. n.. See Tab. 1 for distinctive characters. OSMIA (MELANOSMIA)STEINMANNISP. N. 805 Table 1: Distinctivecharacters of 9 ofOsmia steinmannisp. n., OsmiasvenssoniTkalcû, 1983 and Osmia biennis (Zetterstedt, 1838). Osmiasteinmannisp. n. OsmiasvenssoniTkalcû, 1983 Osmia inermis (Zetterstedt,1838) (n= 13) (n = 1. paratype) (n= 10) SwissAlps(Ebenalp.Juf) NorthernSweden (Abisko) Northernandcentral Europe, North (Tkalcü, 1983). America(Tkalcû, 1983;Warncke, 1988). Secondmandibulartoothasyme- Secondmandibulartoothsym- Secondmandibulartoothbroadand tricallytriangular,separatedfrom metricallytriangular,broadly long, littleprominent, notclearly thirdtoothbyacute-angledand separatedfromboth firstand separatedfromfirsttooth; indenta- broadindentation,fromfirst thirdtoothbyacute-angledand tionbetween secondandthirdtooth toothbyshallowandrounded broadindentation (Fig. lb). verynarrow, acute-angled(Fig. lc). indentation (Fig. la). Mandibleatitsinferiorouter AsinOsmiasteinmanni. Mandibleatits inferioroutermar- marginnearly straight,without ginwithprominent, asymmetrically tooth(Fig. 2a). triangulartooth (clearlyvisible only when mandibleisopened) (Fig. 2b). Ocelli lessdistantlyseparated AsinOsmiasteinmanni. Ocelli moredistantly separated fromeachother,distance fromeachother,distancebetween betweenmiddleandlateralocel- middleandlateralocellusabout lusabouttwothirdofdiameterof equal todiameterofmiddleocellus. middleocellus. Nervulus(cu-v)offorewing Asin Osmiasteinmanni. Nervulus(cu-v)offorewing postfurcal.i.e.runninginto interstitial, i.e. runningexactly into cubital veindistallyfromorifice orificeofbasal vein. ofbasalvein. Tibialspineofmiddle leglonger, AsinOsmiasteinmanni. Tibial spineofmiddleleg shorter, narrowerandweaklybent(Fig. broaderandmore triangular(Fig. 3a). 3b). Pilosityofvertex, scutum, Pilosityofvertex, scutum, Asin Osmiasvenssoni. propodeumandfirsttergite propodeumandfirsttergite whitishtoyellowish-white. brightlyyellowish-brown. Pilosityofscutuminitsanterior Pilosityofscutumandscutellum Asin Osmiasvenssoni. fourfifthwhitishtoyellowish- uniformlybrightly-yellowish white withsingleintermixed brownwithoutblackishhairs. blackishhairs,thatofbackmost partofscutumandofscutellum brightlyyellowish-brown. Pilosityofmesepisternum Pilosityofmesepisternum Pilosityofmesepisternuminits uniformlywhitetoyellowish- predominantlyblackish,onlyon upperpartbrightlyyellowish- white,in some specimenswith itsmostanterioranduppermost brown, initslowerpartwhitish. singleintermixedblackishhairs. partonnarrowzonebrightly yellowish-brown. Marginalzoneofsecondand Marginal zoneofsecondand Marginalzoneofsecond andthird thirdtergite impressedonits thirdtergite notimpressedand tergiteonly slightlyimpressed, wholewidth,clearlyseparated notseparatedfromdisc. weakly separatedfromdisc. fromdisc. Punctationoftergite2medio- Punctationoftergite2medio- Asin Osmiasteinmanni. basallycoarser, interspacesas basallyfiner, interspacesupto largeasthesizeofonetomaxi- thesizeofthreepunctures. mallytwopunctures. Punctationofcentral areaof Punctation ofcentral areaof As in Osmiasteinmanni scutumjustbehindlongitudinal scutumjustbehindlongitudinal suturemoredispersed,inter- suturedenser,interspaces maxi- spacesuptothesizeoftwoto mallyas largeasthesizeofone threepunctures. puncture. Longesthairsonoutermarginof Longesthairson outermarginof As in Osmiasvenssoni. tibiaofhindleglongerthan tibiaofhindlegaslongasmaxi- maximal widthoftibia. mal width oftibia. 806 A- MÜLLER Male 1 Outer margin of gonoforceps preapically with semicircular widening, gonoforceps here nearly twice as broad as at its narrowest width. Body length at least 11 mm. Osmiaalticola Benoist, 1922; OsmiamaritimaFriese, 1885; Osmiaxan- thomelana (Kirby, 1802), See Tkalcû (1983) and Haeseler (1999) for distinctive characters. 1* Outer margin of gonoforceps preapically without widening or only weakly widened, gonoforceps here at most little broader than at its mm narrowest width. Body length at most 10 2 2 Fourth sternite with normal hairs, without hooked bristles. Osmia inermis (Zetterstedt, 1838); OsmiahyperbofeaTkalcû, 1983. See Tkalcû (1983) and Haeseler (1999) for distinctive characters. 2* Fourth sternitewithhookedbristlesbothalongapicalmarginandonpre- marginal area; along apical margin the bristles are oriented horizontally whereas on the premarginai area they are directed increasingly vertically ... 3 3 Underside ofantennal segments only microscopically haired. Osmia parietina Curtis, 1828; Osmia uncinata Gerstäcker, 1869. See Tkalcû (1983) and Haeseler (1999) fordistinctive characters. 3* Underside ofantennal segments with conspicuousbristles whichareone fourth as long as diameter ofantenna to as long as diameter ofantenna .... 4 4 Bristles on underside of antennal segments as long as diameter of an- tenna. Projecting middle part of sixth sternite narrow, densely covered with conspicuously knobbed hairs. Osmiapilicornis Smith, 1846. 4* Bristles on underside of antennal segments about one fourth as long as diameter of antenna. Projecting part of sixth sternite broad, without knobbed hairs. Osmia svenssoni Tkalcu, 1983; Osmia steinmanni sp. n.. See Tab. 2 for distinctive characters. Table 2: Distinctive characters of S ofOsmia steinmanni sp. n. and Osmia svenssoni Tkalcû, 1983. Osmiasteinmannisp. n. OsmiasvenssoniTkalcu, 1983 (n=2) (n= 1, holotype) Vertex(seenfromthefront) weaklyascending, out- Vertex (seenfromthefront)morestronglyascend- lineofheadroundish (Fig.5a). ing,outlineofheadmorequadrangular(Fig. 5b). Thirdsternitemoredeeplyrecessed,emargination Thirdsternite lessdeeplyrecessed,emargination aboutonethirdasdeepasbroad(Fig. 6a). aboutonefifthasdeepasbroad(Fig. 6b). Projectingmiddlepartofsixthsterniteshorter,less Projectingmiddlepartofsixth sternite longer,more thanhalfaslongasbroad(Fig. 6a). thanhalfaslongasbroad(Fig. 6b). Outermarginofgonoforcepspreapicallynot Outermarginofgonoforcepspreapicallydistinctly widened,gonoforcepshereasbroadasmorebasally widened,gonoforcepsherebroaderthanmore (Fig. 7a). basally(Fig. 7b). Marginalzoneofsecond,thirdandfourthtergite Marginalzoneofsecond,thirdandfourthtergite stronglyimpressedonitswholewidth,polishedto weakly impressed,densely shagreenedatleaston only superficiallyshagreened. itsbasalhalf. Pilosityofsecondtofourthtergiteyellowish-white. Pilosityofsecondtofourthtergitebrightly yellowish-brown. OSMIA (MELAN0SM1A)STEINMANNISP. N. 807 ^^XuaJXD^t^ 6a 6b Figs 1-7 1:Rightmandibleofthe 2: aOsmiasteinmannisp.n.,bOsmiasvenssoniTkalcû, 1983,c Osmia inermis(Zetterstedt, 1838). - 2: Inferioroutermargin ofrightmandibleofthe 2: aOsmia stein- mannisp. n.,b Osmia inermis(Zetterstedt, 1838). - 3: Tibial spine ofright middle legofthe 2: aOsmiasteinmanni sp. n., b Osmia inermis (Zetterstedt, 1838). -4: S ofOsmia steinmanni sp. n.: aRight antenna from the front, b Right basitarsus ofhind leg, cApical margin ofsixth and seventh tergite. - 5: Head of the 6 seen from the front: a Osmia steinmanni sp. n., b Osmia svenssoni Tkalcû, 1983. - 6: Apical margin of third and sixth sternite of the 6: a Osmia steinmannisp.n.,bOsmiasvenssoniTkalcû, 1983.-1:6 genitaliaindorsalview: aOsmiastein- manni sp. n., b Osmia svenssoniTkalcû, 1983. 808 A- MULLER Description Female Body length 8-10 mm, fore wing length 7.0 - 7.7 mm (0 = 7.3 mm, n = 12). Body colour dark brown to black without metallic sheen. Head: Head seenfromthefrontaboutas longasbroad. Distancebetweenlateral ocellus and margin of vertex about three times as long as ocellar diameter. Mandible four-toothed; second tooth asymmetrically triangular, separated from third tooth by an acute-angled and broad indentation, from first tooth by a shallow and rounded inden- tation (similar to O. uncinata, see Tkalcü, 1983) (Fig. la). Inferior outer margin of mandible nearly straight, without tooth (Fig. 2a). Punctation of vertex, genal area, supraantennal area, supraclypeal area and clypeus (apart from the polished marginal area) very dense, interspaces much smaller than the size of one puncture. Paraocular area less densely punctate, interspaces up to the size of one puncture. Pilosity of clypeus predominantly blackish with single intermixed white hairs, pilosity of the remaining head whitish to yellowish-white with many intermixedblackish hairs. Hairs on apical halfofclypeus dense obstructing the view onto the polished marginal areaof the clypeus (as in O. inermis and O. svenssoni). Thorax: Punctation ofscutum (except in centre), scutellum and mesepisternum verydense, interspaces much smallerthanthe sizeofonepuncture. Centralareaofscu- tumjustbehind longitudinal suture more dispersely punctate, interspaces uptothe size oftwo tothree punctures. Metanotum, propodeal triangle and propodeum densely sha- greened, dull. Pilosity ofbackmostpartofscutumandofscutellumbrightly yellowish- brown clearly contrasting with the remaining pilosity ofthe thorax which is whitish to yellowish-white. Scutum and in some specimens also mesepisternum with single intermixed blockish hairs. Veins and stigma of wings dark brown to black. Nervulus (cu-v) offore wing running into cubital vein distally from orifice ofbasal vein ("post- furcal"). Tibial spine of fore and middle leg long, narrow and weakly bent (Fig. 3a). Tibial spurs ofhind legblack. Erect pilosity offemurofall legs predominantly whitish with intermixed blackish hairs, that of tibia and tarsus predominantly blackish with intermixed white hairs. Longest hairs on outer margin of tibia and basitarsus of hind leg longer than maximal width oftibia and basitarsus respectively. Abdomen: Declining basal portion offirst tergite densely shagreened, only with silky lustre (as in O. inermis and O. svenssoni, see Tkalcu, 1983). Punctation of disc of second to fourth tergite basally dense (interspaces mostly of about the size of one puncture), distally more dispersed (interspaces upto the sizeofthree punctures). Inter- spaces on second tergite nearly polished, on third tergite superficially shagreened and on fourth tergite distinctly shagreened. Marginal zone of second and third tergite impressed on its whole width, clearly separated from disc, shagreened and dispersely punctate. Pilosityon firsttergite whitishto yellowish-white, on secondandthirdtergite yellowish-brown with many intermixed blackish hairs, on fourth and fifth tergite predominantly blackish with whitish hairs along the marginal zone. Sixth tergite covered with appressed white hairs. Longest hairs on second to fourth tergite about as long as the last three segments of the antenna, on first tergite sligthly longer. Abdominal scopa black. OSMIA (MELANOSMIA)STEINMANNISP. N. 809 Male Body length 9 mm, fore wing length 6.7 - 7.0 mm (0 = 6.85 mm, n = 2). Body colour dark brown to black; head, thorax and abdomen with faint blue green metallic sheen. Head: Head seen from the front about as long as broad (Fig. 5a). Distance betweenlateral ocellus andmargin ofvertex abouttwo andahalftothreetimes as long as ocellar diameter. Underside of antennal segments each with 10-20 white bristles about one fourth as long as antennal diameter (as in O. svenssoni, see Tkalcû, 1983) (Fig. 4a). Punctation ofvertex, genal area and face (apart from the polished marginal area ofclypeus) very dense, interspaces much smaller than the size of one puncture. Pilosity of head long and whitish, on clypeus, paraocular area and supraclypeal area very dense. Thorax: Punctation ofscutum, scutellum and mesepisternum very dense, inter- spaces much smallerthan the size ofone puncture. Metanotum, propodeal triangle and propodeum densely shagreened, dull. Pilosity of thorax whitish, on scutellum more yellowish but not as brightly yellowish-brown as in the 2 Veins and stigma ofwings . dark brown to black. Nervulus (cu-v) of fore wing running into cubital vein distally from orifice ofbasal vein ("postfurcal"). Tibial spurs ofhind leg black. Erect pilosity of all legs whitish. Basitarsus ofhind leg toothed, with slightly diverging sides in its basal two third (parallel in O. parietina, see Tkalcü, 1983) (Fig. 4b). Abdomen: Declining basal portion offirst tergite densely shagreened, only with silky lustre (as in O. inermis and O. svenssoni). Punctation ofdisc ofsecond to fourth tergite very dense, interspaces up to the size of one puncture but mostly smaller. Marginal zone of second to fourth tergite strongly impressed on its whole width, polished to only superficially shagreened, nearly impunctate. Pilosity on first, fifth and sixth tergite whitish, on second to fourth tergite yellowish-white. Longest hairs on second to fourth tergite little shorterthan the last two segments ofthe antenna, on first tergite distinctly longer. Apical margin of sixth tergite slightly notched in the middle, seventhtergitebilobed(Fig. 4c).Apical margin ofsecond sternite moreorless rounded (slightly emarginate in O. uncinata, see Tkalcû, 1983). Third sternite deeply recessed, the emargination about one third as deep as broad and densely beset with long yellow- ishhairs (Fig. 6a). Fourth sternite as in O. svenssoni (seeTkalcû, 1983): apical margin straighttovery shallowly emarginate, premarginai areablack, strongly shagreened and completely dull with hardly visible punctures (in contrast to O. parietina and O. unci- nata where the premarginai area is dark brown, less strongly shagreened and more shiny with more distinct punctures); apical margin and premarginai zone beset with longandhooked, conspicuouslyyellowish-redcolouredbristles (lighterin O.parietina and O. uncinata); along apical margin thebristles are oriented horizontally whereas on thepremarginai areathey are directed increasingly vertically. Fifth sternitebroadly but weakly emarginate. Projecting part of sixth sternite rectangular (more roundish in O. parietina and O. uncinata, see Tkalcû, 1983), less than halfas long as broad (Fig. 6a). Gonoforceps inflected in its apical part, preapically with atuft ofstiffand yellow- ish-red bristles both on the outer and the inner side (Fig. 7a); its outer margin pre- apically notwidened (weakly widened in O.parietina, seeTkalcû, 1983), gonoforceps 810 A.MÜLLER here as broad as more basally; proximal to the inflexion and seen from above sides of gonoforceps nearly parallel forsome distance (diverging towards base in O. uncinata). DISTRIBUTION, HABITATAND FLOWER PREFERENCES O. steinmanni is known only from two localities in eastern Switzerland: Ebenalp (Wasserauen, Appenzell Innerrhoden) and Stallerberg (Juf, Graubünden). With 1500mand2300m above sealevelthesetwolocalities are atthe lowersubalpine and the lower alpine zone respectively. At the Ebenalp O. steinmanni was found on stony slopes exposed to the south- east. The vegetation cover varied between densely vegetated meadow-like areas to barely vegetated screes intermixed with groups of spruces. The most prominent plant in flower was Hippocrepis comosa (Fabaceae). Bee species accompanying O. stein- manni were Andrena intermedia Thomson, 1870, Andrena semilaevis Perez, 1903, Lasioglossum cupromicans (Perez, 1903), Osmia inermis (Zetterstedt, 1838), Osmia xanthomelana (Kirby, 18Ö2), Hoplitis tuberculata (Nylander, 1848) and Bombus mon- ticola Smith, 1849, all typical inhabitants ofupper montane and subalpine regions. The 9 $ ofO. steinmanni were observed at flowers ofH. comosa. The micro- scopic analysis ofthepollenloadsoffour 9 9 collected in 2002revealedthatthisplant species is an important pollen source. All loads contained pollen ofH. comosa. In ad- dition, Lotus corniculatus (Fabaceae) was recorded in all samples as well. DISCUSSION Owing to the detailed study of Tkalcû (1983) and the supplemental work of Haeseler (1999) the subgenus Melanosmia is well known in Europe. The discovery of anewMelanosmia species inthe well investigatedcentral Europeanbeefaunais asur- prise. The only unclear Melanosmia species in Europe is O. bulgarica described by Friese (1922) based on a single 9 from Macedonia. Warncke (1988) treats O. bulga- rica as a subspecies of O. inermis and mentions its occurrence in Macedonia, Greece and Italy. The type specimen obviously got lost (Tkalcû, 1983), but according to the original description O. bulgarica differs from all otherEuropeanMelanosmia species, including O. steinmanni, by its lack of black hairs on head, thorax and abdomen. Tkalcû (1983) mentions a Melanosmia 9 from Gran Sasso (Abruzzo, Italy) which might be eitherconspecific with O. bulgarica orrepresent a new species. Its declining basal portion ofthe first tergite being polished and its mandible bearing a tooth on the inferior outer margin exclude the possibility that this Italian specimen and O. stein- manni belong to the same species. Although all museum and private bee collections available in Switzerland were examined only three contained specimens ofO. steinmanni indicating that this species is extremely rare. However, its early flight period falling in late May, June and proba- bly early July in combination with its occurrence in subalpine and alpine areas might explain why this species was sorarely collected. Bee researchers collect bees athigher elevations preferably later in the season when the weather conditions are more favou- rable. Therefore, O. steinmanni is expected to occurnot only at additional localities in eastern Switzerland but also at otherplaces in the Swiss and probablyAustrian alps. OSMIA (MELANOSMIA)STEINMANNISP. N. 811 Melanosmia is especially well represented in montaineous regions (Michener, 2000). Two central European species, O. cuticola and O. inermis, are known to occur m beyond the timber line, the latter mounting up to 2800 (Haeseler, 1999; Schedi, 1982). O. steinmanni is a furtherMelanosmia species reaching the alpine zone. Based on morphological characters ofthe 9 (but not ofthe 6), O. inermis and O. steinmanni seem to be closely related. Most noteworthy is that both species were recorded together in the same habitat on the same day at the Ebenalp. Similarly, O. svenssoni, the closest known relative of O. steinmanni, coexists with O. inermis in northern Sweden (Tkalcu, 1983). Given this syntopic occurrence the question arises how O. inermis and O. steinmanni are ecologically separated in the Swiss Alps. The few data indicate that the flight period of O. steinmanni starts earlier than that of O. inermis.AttheEbenalp, twofreshlyemerged 6â butno $ ? ofO. inermiswerefound on 1st June 2002 whereas the only 6 of O. steinmanni collected that same day was heavily worn. However, most $ 2 of O. steinmanni recorded on this day were rather fresh indicating that there is an overlap rather than a clear separation in the flight periods of these two species. O. inermis is known to prefer flowers of Hippocrepis, Lotus and Onobrychis (all Fabaceae) for pollen collection (Westrich, 1990). The few pollen loads of O. steinmanni were found to consist of pollen from partly the same Fabaceae. Therefore, an ecological separation with respect to the preferred pollen sources seems rather improbable. The possibility exists that the two species differ in theirnesting sites ornest architecture. O. inermisis known tobuild itsbrood cells with masticated leaves under stones (Priesner, 1981; Schedi, 1982; Else & Edwards, 1996; own observation). Unfortunately, nests of O. steinmanni have not yet been recorded leaving the possibility ofdiffering nesting biologies an open question. ACKNOWLEDGEMENTS I thank Prof. Dr B. Svensson and Dr Mats Eriksson (Museum of Evolution, Uppsala) for loaning type specimens of O. svenssoni, Mr George Else (The Natural History Museum, London) for verifying the English morphological terminology and Dr Kathrin Tschudi (ETH, Zürich) for correcting the English. Comments of t)r Bernhard Merz (Muséum d'histoire naturelle, Genève), Dr Mike Herrmann (Konstanz), FelixAmiet (Solothurn), Prof. Dr S. Dorn (ETH, Zürich) and two anony- mous reviewers substantially improved the manuscript. REFERENCES Amiet, F., Herrmann, M., Müller, A. & Neumeyer, R. (in preparation). Fauna Helvetica, Apidae 4 (Megachilidae). Centre Suisse de Cartographie de la Faune, Schweizerische Entomologische Gesellschaft. Else,G.R. &Edwards, M. 1996. Observationson Osmia inermis(Zetterstedt)and O. uncinata Gerstäcker (Hym., Apidae) in the central Scottish Highlands. Entomologist's Monthly Magazine 132: 291-298. Friese, H. 1922. Eine Kriegsausbeute anApiden (Bienen) aus Makedonien. ZoologischesJahr- buch, Systematik46: 175-216. Haeseler,V. 1999.ZurKenntnisvonOsmiaalticolaBenoist 1922,OsmiamaritimaFriese 1885 sowie der für Mitteleuropa bislang unbekannten Osmia hyperborea Tkalcu 1983 (Apidae: Osmia (Melanosmia Schmiedeknecht 1885)). Entomofauna 20: 449-460. 812 A.MÜLLER Hurd, P. D. Jr. 1979. SuperfamilyApoidea (pp. 1741-2209). In: Krombein, K. V., Hurd, P. D. Jr., Smith, D. R. & Burks, B. D. (eds). Catalog ofHymenoptera in America north of Mexico,Vol. 2. SmithsonianInstitution Press, Washington. Michener, C. D. 2000. The Bees ofthe World. TheJohnsHopkins UniversityPress, Baltimore andLondon. Michener, C. D., McGinley, R. J. & Danforth, B. N. 1994. The bee genera of North and Central America (Hymenoptera: Apoidea). Smithsonian Institution Press, Washington andLondon. Priesner, E. 1981. Beobachtungen zur Nistbiologie der Alpen-Mauerbiene Osmia inermis (Hymenoptera: Apoidea: Megachilidae). Carinthia II 171/91: 349-356. Schedl, W. 1982. Über aculeate Hautflügler der zentralen Ötztaler Alpen (Tirol, Österreich) (Insecta: Hymenoptera). Berichte des naturhistorisch-medizinischen Vereins Innsbruck 69:95-117. Schwarz,M.,Gusenleitner,F.,Westrich,P. &Dathe, H. H. 1996. KatalogderBienenÖster- reichs, Deutschlands und derSchweiz. Entomofauna, Supplement 8. Tkalcu, B. 1983. Die europäischen Osmia-Arten derUntergattungMelanosmia (Hymenoptera, Apoidea, Megachilidae). Vestnik CeskoslovenskéSpolecnostizoologické47: 140-159. Warncke, K. 1988. Isolierte Bienenvorkommen auf dem Olymp in Griechenland (Hymeno- ptera, Apidae). LinzerbiologischeBeiträge 20: 83-117. Westrich, P. 1990. Die Wildbienen Baden-Württembergs. 2 volumes. Eugen Ulmer Verlag, Stuttgart. Zanden, G. Van Der 1988. Beitrag zur Systematik und Nomenklatur der palaearktischen Osmiini, mitAngaben überihre Verbreitung. ZoologischeMededelingen 62: 113-133.

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