The Drosophilidae (Diptera) of Estonia

2006·0.08 MB·

by  Stefan Escher

©EntomologicaFennica.3March2006
The Drosophilidae (Diptera) of Estonia
StefanA.Escher,JohanEkenstedt,KaupoElbergandAnssiSaura*
Escher, S. A, Ekenstedt, J., Elberg, K. & Saura, A. 2006: The Drosophilidae
(Diptera)ofEstonia.—Entomol.Fennica17:13–20.
Untilrecently,nopublishedrecordsondrosophilidsfromtheBalticcountriesex-
isted,exceptforanunpublishedspecieslistforEstonia.Wemadetwocollecting
tripsthroughthethreeBalticcountriesinthesummerof2000.Thefirsttripwas
madeinearlysummertogetspringspeciesandthesecondinlateAugusttoget
the fungivorous species. We used baiting methods for collecting Drosophila,
whiletheearlierresultswereobtainede.g.usingnetsweeping.Ingeneral,the
drosophilid fauna of Estonia resembles the well-known fauna of the Nordic
countries.ThesinglemostinterestingresultwasthatChymomyzaamoena,are-
centAmericaninvaderofcentralEurope,wasfound.Anotherinterestingfinding
wastherelativerarityofD.subobscuraandthevirtualabsenceofD.virilisgroup
species.
S.A.Escher,J.Ekenstedt&A.Saura,DepartmentofMolecularBiology,Umeå
University, SE-901 87 Umeå, Sweden; correspondent author’s e-mail:
[email protected]
K.Elberg,InstituteofZoologyandHydrobiology,UniversityofTartu,Vanemuise
46,EE-51014Tartu,Estonia
Received8March2005,accepted2May2005
1.Introduction DrosophilabiologytookplaceinNorthAmerica:
Morgan and his students, with Sturtevant and
The genus Drosophila is by far the best known Dobzhansky,pioneer-workedonthebiologyand
groupofinsects.DrosophilamelanogasterMei- evolution of these small flies. At that time it
genisamodelorganismofthegenomeproject. seemedthattheRussianswerecatchingup,with
The sheer weight of the ever accumulating Chetverikov,DubininandTimofeeff-Ressovsky
knowledgehaseffectivelypreventedarevisionof layingthegroundworkofevolutionarygenetics.
an obviously paraphyletic genus, held together History took, however, another way. We have
withD.melanogaster.TherestofthefamilyDro- data on drosophilids from Finland (Hackman
sophilidae is held together largely through this 1954),theSt.Petersburgarea(Shtakelberg1930,
paraphyly(Bächlietal.2004). Hackman 1957) and Denmark (Frydenberg
TheDrosophilastory beganinnorthernEu- 1956),andwehaverecentlycompletedanover-
rope,withFabriciusdescribingMuscafunebrisin view of a North European drosophilid fauna
Denmark in 1787. Fallén, working in Sweden, (Bächlietal.2004).Theimportanceofthisfam-
transferred this species to a new genus, Droso- ily in all biology makes publishing a review of
phila,in1823.Thisgenuswaslatersplitintosev- Estonian drosophilids timely. To give an exam-
eral others, with Hackman, in Finland, being a ple,whileD.subobscuraoccursintheStPeters-
specialist on Scaptomyza. The breakthrough in burgareaandallthewaytoMoscow(Sauraetal.
14 Escheretal. (cid:127) ENTOMOL.FENNICAVol.17
1998) and Kazakhstan (Gornostaev 1993) and philidfaunaoftheNordiccountries(e.g.Hack-
overtherestofEurope,theattemptstodescribe man1954,Frydenberg1956).Thesubarcticand
thesituationintheBalticcountrieshavejustbeen alpinespecieswere,ofcourse,missing.
guesses(e.g.Krimbas1993). WefoundsomeChymomyzaamoena.Thisis
The drosophilid fauna of Estonia is, in fact, arecentNorthAmericanimmigranttocentraland
wellknown.Kunberg(1981)haswrittenashort southernEurope(e.g.Bandetal.1999)Another
monograph,withrecordsonbiologyinEstonia. remarkable result was the almost complete ab-
Hisstudyhas,however,remainedanunpublished senceofspecimensofthevirilisgroup.Theriver-
thesis written in Estonian. The authors Escher, sidehabitatshostedmostly domesticspecies,in
Ekenstedt and Saura have collected additional particular D. melanogaster. D. subobscura was
material.Wemayalsomentionthatthespecimen alsoremarkablyscarce.Inhabitatswhereweex-
ofD.busckiithatwecollectedatagarbagecon- pectedit,wegotmostlyD.obscurainverylarge
tainerattheharbourofTallinnhasfiguredgreatly numbers. This contrasts with observations from
inthestudyofidentifyinganovelsystemofgene southernFinlandincentralSweden.
regulation(Larssonetal.2001),anditschromo- AsforthefourspeciesoftheD.virilisgroup
someshavebeenusedtoillustratetheconceptof foundinSwedenandinFinland,D.lummeiand
epigeneticsinaspecialissueofthejournalSci- D. ezoana havevirtually disappeared fromFin-
ence(Riddihough&Pennisi2001).Allinforma- landandfromSweden,andalsoD.littoralisand
tion on Drosophila biology is valuable, and we D. montana have become uncommon. D. sub-
believethatthespecieslistofEstoniawillleadto obscurahaddisappearedfromthesitesinSwe-
furtherstudies. denandinFinlandwherewecollectedthespecies
in1994.Earlieritwasactivelyspreadingtowards
thenorthandhadbecomecommonandabundant
2.Materialandmethods (Saura1995).Currentlyitsdistributionseemsto
decrease. It is unclear what has happened with
Thematerialforthisreportwascollectedbythe these species. However, laboratory strains are
authorsinJuneandAugust2000.Allotherinfor- available from the period before the population
mationisderivedfromtheunpublishedthesisof crashandtherearealsoavastamountsofunpub-
Kunberg(1981).Heusedbothnetsweepingand lished enzyme allele frequency data from Nor-
baitstocollectDrosophila. way, Sweden and Finland from mid 1970’s.
Wecollectedfliesfromasmany biotopesas These may allow an evaluation of possible ge-
possible, including dust bins, mushrooms and netic consequences of the drastic decrease, e.g.
felledlogs,usingeithernetsweepingorsuctionin viacomparingallelefrequencies(andDNAs)be-
fermenting baits with rotting bananas and other foreandafterthecrash.
fruits.Dependinge.g.onweatherconditions,the
trapswerecheckedafteratleastonceanhour.Af-
ter preliminary examination of the flies in the 4.Anannotatedlistofspecies
field,theywerestoredinethanolandfurtherex-
aminedinthelab. 4.1.GenusAmiotaLoew,1862
Wemadeaspecialefforttogetspecimensof
thevirilisgroupbycollectingalongthebanksof Amiotaisaverylargegenus,with44Palaearctic
smallandlargerivers. species; most of which live in the eastern
Palearctic(Bächli&RochaPité1981).Thespe-
cies presented here belongs to the subgenus
3.Resultsanddiscussion AmiotaLoew,1862.
The results are given in the accompanying spe- A.alboguttata(Wahlberg,1839)
cieslist,wherecertainspeciesarehighlighted.In Kunberg (1981) reports this species from Uus-
general, there was nothing completely unex- Kasaritsa (Võru area). This is a widespread but
pected compared with the well-known droso- ratheruncommonspeciesinnorthernEurope.It
ENTOMOL.FENNICAVol.17 (cid:127) DrosophilidaeofEstonia 15
comes to fermenting baits, and lives among the 4.3.GenusDrosophilaFallén,1823
foliageoftrees.
4.3.1.SubgenusDorsilophaSturtevant,1942
4.2.GenusChymomyzaCzerny,1903 D.busckiiCoquillett,1901
A domestic species that breeds in decaying or-
C.caudatula,C.costataandC.fuscimanaarein- ganic material, in particular potatoes but also
dependentofhumanculture,i.e.“wild”species. mushrooms,etc.Comestofermentingbaits.Can
Allhavelarvaldiapause.Theycometoferment- be cultured on modified Drosophila medium.
ingbaitsandcanbeculturedonDrosophilame- Cosmopolitan.Wefoundthemamonggarbagein
dium.Theyareattractedtofreshly-cutsurfacesof wastebaskets in Narva, Otepää and Tallinn.
felled logs in early summer. They engage in Kunberg (1981) reported D. busckii from
lengthyandhighlyvisiblecourtshipbehaviorand ViljandiandTartu.
areeasilycollectedwithanexhaustor.
There is a monograph on niche shifts and 4.3.2.SubgenusDrosophilaFallén,1823
sympatryinthegenus(Band1996)andanother
on the diapause, cold-hardiness and circadian D.funebris(Fabricius,1787)
eclosionrhythm(Riihimaa1996). This species belongs to the funebris species
group. Kunberg (1981) noted that this wide-
C.amoena(Loew,1862) spread,domesticspeciesiscommonalloverEs-
Tartu, 13.VI.2000; a park within the city. Fer- tonia.WefoundthemamonggarbageinNarva;in
mentingbait.Leg.A.Saura. Tartu they came to fermenting baits. They are
C.amoenaisacommensalofhumanculture easytoculture.
i.e.“domestic”inEurope.Larvaeliveinnutsin
NorthAmerica,butinEuropethespecieshasex- D.histrioMeigen,1830
periencedanicheshiftinthatitnowbreedsinun- Thisspeciesbelongstothehistriospeciesgroup.
ripe fruits (Band et al. 1999). This is by far the Awidespread species that is not common any-
northernmostrecordofthisspecies.Wehaveear- where. It comes to fermenting baits. It can evi-
lierfoundthespeciesinLithuania(Escheretal. dentlynotbecultivatedonstandardmedia.The
2004). larvae live in mushrooms. Kunberg (1981) re-
portsitfromTartu.
C.caudatulaOldenberg,1914
Adevere,13.VI.2000;logsinadeciduousforest. D.immigransSturtevant,1921
Suction.Leg.A.Saura. Valga,21.VIII.2000;deciduousforest.Ferment-
Like C. costata (see below), a very wide- ingbait.Leg.A.Saura.
spreadspecies. BotharefoundfromNorway to D.immigransbelongstotheimmigransspe-
Japan(Bächli&RochaPité1981). ciesgroup.Awidespread,domesticspeciesthat
breedsincompostheaps,etc.Itiseasytocultivate
C.costata(Zetterstedt,1838) andcomestofermentingbaits.
Adevere, 13.VI.2000;deciduousforest. Suction
overlogs.Leg.A.Saura. 4.3.3.SubgenusDrosophila,
Thisisoneofthemostcommondrosophilids quinariaspeciessubgroup
innorthernEurope(Hackmanetal.1971)thatex-
hibitsaclineinthereactiondeterminingthelarval This is a uniform group that breeds predomi-
diapause(Riihimaa1996,Riihimaaetal.1996). nantlyonmushrooms.Courtneyetal.(1999)de-
scribedthegeneralbiologyasamushroom-feed-
C.fuscimana(Zetterstedt,1838) ingdrosophilidguild.Theyareeasytocollecton
Adevere,13June2000;deciduousforest.Suction theirbreedingsitesandcomereadilytoferment-
onlogs.Leg.A.Saura. ingbaits;likewise,they areeasy tocultivateon
standardmedium.
16 Escheretal. (cid:127) ENTOMOL.FENNICAVol.17
D.kuntzeiDuda,1924 D.repletaWollaston,1858
Loobu, Rakvere, 20.VIII.2000; mixed forest. Narva,20.VIII.2000;city.Nettingovergarbage
Suction.Leg.A.Saura. closetoanurinal.Leg.A.Saura.
Thisisthenorthernmostrecordforanother- Awidespread,domesticspecies,thoughless
wise central European species. We have earlier commonthanD. hydei. InnorthernEurope, the
foundthespeciesalsoinLithuania(Escheretal. two species are human commensals: D. hydei
2004). breeds in all kinds of garbage and D. repleta in
urinewells offarms, etc. (Lakovaara&Itämies
D.limbatav.Roser,1840 1994).
Aratheruncommonspeciesthatseemstobeasso-
ciatedwithdecayingplantmaterialandblackcur- D.testaceav.Roser,1840
rant(Ribesnigrum)(A.Sauraunpubl.).Kunberg Loobu, Rakvere, 20.VIII.2000; mixed forest.
(1981)foundthespeciesfromtheLooriver. SuctionoverBoletus.Leg.A.Saura.
Udna,Narva,20.VIII.2000;youngmixedfor-
D.phalerataMeigen,1830. est and lakeshore. Fermenting bait. Leg. A.
GeyspitsandSimonenko(1970)andMuonaand Saura.
Lumme(1981)havedescribedthediapauseofD. Valga, 21.VIII.2000; deciduous forest. Fer-
phalerata and D. transversa (see below). D. mentingbait.Leg.A.Saura.
phaleratagoesupinthenorthtoabout65°Nand Padise, 26.VIII.2000; young, wet deciduous
to North Africa in the south. We found themin forest.SuctiononBoletus.Leg.A.Saura.
Loobu, Rakvere, Saare, Valga, Lihula and Thespeciesbelongstothetestaceasubgroup.
Linnamäe, either on mushrooms or attracted by These small flies breed in mushrooms; they are
fermentingbait,inAugust2000.Kunberg(1981) easy to collect over themand using baits. They
reports the species from Saaremaa, Samliku are also easy to cultivate. Handling them, how-
(Pärnu area), Luua, Tartu, Sangaste, Rõuge, ever, is difficult, as they are very sensitive to
VapramäeandObinitse. ether. Overwintered flies are generally black,
whilesummerfliesarepaleyellow.Thedistribu-
D.transversaFallén,1823 tionareacoversthePalaearctic.Kunberg(1981)
SpreadfromNorwayacrossAsia;anorthernspe- earlierreportedthespeciesfromKaaveandLuu.
cies(upto71°N)comparedwiththeformer.We
caughtthemeitherfrommushroomsorusingfer- 4.3.5.SubgenusDrosophila,
menting bait in Kiviõli, Kohtla-Järve, Iisaku, virilisspeciesgroup
Saare,Valga,Lihula,Padise;allinAugust2000.
Kunberg(1981) reportsitfromSaaremaa, Saka Four North-European species belong to this
(Kohtla-Järve area), Kaave, Luua, Samliku group: D. ezoana Takada and Okada, 1957, D.
(Pärnu area), Tartu, Tähtvere, Vapramäe, littoralis Meigen, 1830, D. lummei Hackman,
PikasillaandRõuge. 1972 and D. montana Patterson & Wheeler,
1942.Theyallinhabitlittoralzonesoflakesand
4.3.4.SubgenusDrosophila, rivers;onlyD.littoralisisrestrictedtoEurope.D.
repletaspeciesgroup ezoana and D. lummei evidently occur across
AsiaallthewaytoJapan,whileD.montanagoes
D.hydeiSturtevant,1921 even further as it inhabits also much of North
Narva,20.VIII.2000;city.Nettingovergarbage. America. There is extensive literature on the
Leg.A.Saura. diapause(e.g.Lumme&Lakovaara1983),court-
Tallinn,26.VIII.2000;city.Nettingovergar- ship behavior (Hoikkala & Lumme 1987,
bage.Leg.A.Saura. Hoikkala & Aspi 1993) and eclosion rhythm
Awidespread,domesticspecies. (Lankinen1986)onnorthernEuropeanD.virilis
groupspecies.
Theseflies comereadily to fermenting baits
andareeasytocultivate.Maleshaverathervari-
ENTOMOL.FENNICAVol.17 (cid:127) DrosophilidaeofEstonia 17
ablegenitalmorphologybutthefemalesaremor- 4.5.1.SubgenusSophophoraSturtevant,1939,
phologically indistinguishable. We made a spe- melanogasterspeciesgroup/melanogaster
cialefforttocollectthese.Theresultwasasingle subgroup
femalefromthecitymoatofPärnu,caughtusing
a fermenting bait on 16 June 2000. Kunberg D.melanogasterMeigen,1830
(1981) reports “D. littoralis” from Saaremaa, Thebestknowninsect.Awidespread,domestic
Viljandi,Luua(Jõgevaarea)andTartu.Accord- speciesthatcomestofermentingbaits;canalso
ingly,wecanonlynote“D.virilis”group,inde- becollectedovergarbageetc.Kunberg(1981)re-
terminate,fromEstonia. portsthatitiscommonalloverEstonia.Wefound
it, often in greatnumbers, in aboutalllocalities
influencedbyhumansandtheirgarbage.
4.4.GenusLordiphosaBasden,1961 D.simulansSturtevant,1919isanothercos-
mopolitananddomesticspeciesthatveryproba-
Flies of this group do not come to fermenting blyoccursinEstonia.InnorthernEuropeitisfar
baitsandtheycanevidentlynotbecultivatedon less common than D. melanogaster. Only the
standardmediaeither. males can be distinguished fromthe ones of D.
melanogaster.Wecheckedroutinelyallmalesof
L.fenestrarum(Fallén,1823) melanogaster-groupfliesbutdidnotfindasingle
This is a common species in northern Europe. D.simulans.Thebestwaytofinditwouldbeto
Kunberg (1981) reported the species from sweepoverfruitshelvesofstoresinSeptember–
SaaremaaandSolbi(Võruarea). November and look at the male genitalia under
microscope.
L.nigricolor(Strobl,1898)
Aratherrarespecies,foundinFinlandandtheSt. 4.5.2.SubgenusSophophoraSturtevant,1939,
Petersburg area as well as in central Europe. obscuraspeciesgroup
Kunberg(1981)founditfromSaaremaa,Viljandi,
Puhtu, Kaave (Jõgeva area), Musti (Valga area) Flies belonging to this group are dark brown to
andObinitsa(Võruarea). shinyblack.Theventralsideofmaleabdomenis
red, that of females, white. All come readily to
fermenting baits and are tolerably easy to culti-
4.5.GenusScaptodrosophilaDuda,1924 vate.Mostareindifferenttohumanculture.The
breeding sites of these often exceedingly com-
S.deflexa(Duda,1924) monfliesremainlargelyunknown.
Otepää,21.VIII.2000;beachanddeciduousfor- Inadditiontothebelow-listedspecies,weex-
est.Fermentingbait.Leg.A.Saura. pect that D. alpina Burla, 1948 and D.
Valga, 21.VIII.2000; deciduous forest. Fer- subsilvestris Hardy & Kaneshiro, 1968 will be
mentingbait.Leg.A.Saura. found in Estonia. They are locally abundant on
This species comes to fermenting baits and suitable biotopes in southern Finland. We also
canbecultivated;thelarvaegotodiapause,the have found D. subsilvestris Fallen, 1823 from
detailsofwhicharenotknown. Latvia(Escheretal.2002).
S.rufifrons(Loew,1873) D.ambiguaPomini,1940
Rather uncommon, found in the St. Petersburg Lihula,25.VIII.2000;deciduousforest.Ferment-
areabutnotinFinland(Hackman1957).Itmay ingbait.Leg.A.Saura.
have adult diapause. Reported earlier from Tallinn,26.VIII.2000;city.Suctionovergar-
Tähtvere(Tartuarea)byKunberg(1981). bage.Leg.A.Saura.
ThisisadomesticspeciesinnorthernEurope.
18 Escheretal. (cid:127) ENTOMOL.FENNICAVol.17
D.bifasciataPomini,1940 Thesefliescometofermentingbaits.Theyare
Pärnu,15.VI.2000andValga,21.VIII.2000;de- spreadovernorthernEuropebutareseldomfre-
ciduousforest.Fermentingbait.Leg.A.Saura. quentatalocality.Thelarvaeliveinmushrooms.
SpreadfromNorwaytoJapan;mostcommon
in northern Scandinavia (Saura 1974). Kunberg
(1981) reported the species from Viljandi, 4.7.GenusScaptomyzaHardy,1849
Tommuski (Viljandi area), Tartu, Vapramäe
(Tartuarea)andElva.Theserecordsmay,inpart, Hackman(1959)haswrittenamonographonthis
beincorrect,asthekeyofShtakelberg(1970),the genus.Davis(2000)hasshownthatthegenushas
one that Kunberg used, does not include D. originatedfromDrosophilaontheHawaiianIs-
subobscura that may have been more common lands and from there colonized the rest of the
thanD.bifasciata. world.WiththeexceptionofS.pallida,thelarvae
are leaf miners and some are agricultural pests.
D.obscuraFallén,1823 Theadultsmaycometofermentingbaits.
Otepää,21.VIII.2000;beachanddeciduousfor-
est.Fermentingbait.Leg.A.Saura. 4.7.1.SubgenusScaptomyzaHardy,1849
Valga, 21.VIII.2000; deciduous forest. Fer-
mentingbait.Leg.A.Saura. S.flava(Fallén,1823)
Pärnu, 16.VI.2000; city moat (hundreds of Thisisanagriculturalpestinmany partsofthe
flies).Fermentingbait.Leg.A.Saura. world.Kunberg(1981)reportedthespeciesfrom
Tartu, 13.VI.2000; park. Hundreds of flies Puhtuon25July,1951andon14July,1969.
overfermentingbait.Leg.A.Saura.
CommoninnorthernEurope(Lakovaaraand S.graminum(Fallén,1823)
Saura 1971). Kunberg (1981) reported the spe- Acommonspecies.Kunberg(1981)recordedthe
ciesfromViljandi. speciesfromSaaremaa,Viidumäe,Ruila(Harju
area), Treppoja, Puhtu, Luua, Tartu, Uderna
D.subobscuraCollin,1936 (Tartu area), Tähtvere, Sulbi, Pikasilla
Tartu, 13.VI.2000, city park. Fermenting bait. Rannaküla,Sangaste.
Leg.A.Saura.
Narva, 20.VIII.2000; city. Suction over rot- S.griseola(Zetterstedt,1847)
tingapple.Leg.A.Saura. Kunberg (1981) reported this species from
Otepää, 21.VIII.2000; beach and deciduous Viidumägion8July,1967.
forest.Fermentingbait.Leg.A.Saura.
Valga, 21.VIII.2000; deciduous forest. Fer- 4.7.2.SubgenusHemiscaptomyzaHackman,
mentingbait.Leg.A.Saura. 1959
Pärnu, 16.VI.2000; city moat. Fermenting
bait.Leg.A.Saura. S.unipunctum(Zetterstedt,1847)
Thisotherwisecommonspeciesexperienced FoundearlierinSika(Võruarea)on10Septem-
apopulationcrashin1994(Sauraetal.1998)in ber,1959(Kunberg1981).
northernEurope,andthepopulationsarestillre-
covering.Wefoundonlysingleflies. 4.7.3.SubgenusParascaptomyzaDuda,1924
S.pallida(Zetterstedt,1847)
4.6.GenusLeucophengaMik,1886 Acommensalofhumanculture,foundingardens,
etc.Comestofermentingbaitsandcanbeculti-
L.quinquemaculataStrobl,1893 vatedonDrosophilamedia.Foundat21localities
Otepää,21.VIII.2000.beachanddeciduousfor- byKunberg(1981).
est.Fermentingbait.Leg.A.Saura.
Tartu,13.VI.2000;citypark.Fermentingbait.
Leg.A.Saura.
ENTOMOL.FENNICAVol.17 (cid:127) DrosophilidaeofEstonia 19
4.8.GenusSteganaMeigen,1830 Escher,A.S.,Ekenstedt,J.,Karpa,A.&Saura,A.2002:
TheDrosophilidae(Diptera)ofLatvia.—LatvianEn-
tomologist39:62–68.
S.furta(Linnaeus,1767)
Escher,A.S.,Ekenstedt,J.,Pakalni(cid:1)kis,S.,Rim(cid:1)aite,J.&
This species belongs to the subgenus Stegana
Saura,A.2004:TheDrosophilidaeofLithuania.—
Meigen, 1830. The species is idespread in Eu- ActaZoologicaLituanica14:48–55.
rope.Theydonotcometofermentingbaits.The Frydenberg,O.1956:TheDanishspeciesofDrosophila
larvae live under bark of trees. Kunberg (1981) (Dipt.).—EntomologiskeMeddelelser27:249–294.
Geyspits,K.F.&Simonenko,N.P.1970:Anexperimental
reported the species from Saaremaa, Puhtu,
analysisofseasonalchangesinthephotoperiodicreac-
Kaarepere(Jõgevaarea),Viljandi,andVapramäe
tion of Drosophila phalerata Meig. (Diptera,
(Tartuarea). Drosophilidae).—EntomologicalReview49:46–54.
Gornostaev,N.G.1993:Newspeciesandnewrecordsof
S.coleoptrata(Scopoli,1763) Drosophilid flies (Diptera, Drosophilidae) from
Kazakhstan.—EntomologicalReview71:79–82.
AwidespreadspeciesofthesubgenusSteganina
Hackman,W.1954:DieDrosophila-ArtenFinnlands.—
Wheeler, 1960. The specimens do not come to
NotulaeEntomologicae34:130–139.
baits.Kunberg(1981)reportedthespeciesfrom Hackman, W. 1957: Beiträge zur Kenntnis der Droso-
Saaremaa,Ogandi(Hiiumaa),Rannaküla(Tartu philidendesLeningradgebietes.—NotulaeEntomo-
area),Rõuge,andSangasteandElva. logicae37:17–22.
Hackman,W.1959:OnthegenusScaptomyzaHardy(Dip-
t., Drosophilidae) with descriptions of new species
fromvariouspartsoftheworld.—ActaZoologica
Acknowledgements.WewishtothankDr.AinaKarpafrom
Fennica97:1–73.
theUniversityofLatvia,andDr.SauliusPakalni(cid:1)kisand
Hackman, W., Lakovaara, S., Saura, A., Sorsa, M. &
hisco-workersfromAkademijos2inVilnius,fortheirhos-
Vepsäläinen,K.1970:Onthebiologyandkaryology
pitality and help during our collecting trip. We are also
ofChymomyzacostataZetterstedt,withreferenceto
gratefultoDr.GerhardBächlioftheZoologicalMuseum
thetaxonomyanddistributionofvariousspeciesof
oftheUniversityofZurich,Switzerland,forsharinghis
Chymomyza (Dipt., Drosophilidae). — Annales
knowledgeandhelpfulsuggestionsregardingthemanu-
EntomologiciFennici36:1–9.
script.
Hoikkala, A. & Aspi, J. 1993: Criteria of female mate
choice in Drosophila littoralis, D. montana and D.
ezoana.—Evolution47:768–777.
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