Fungus gnats (Diptera: Sciaroidea) associated with dead wood and wood growing fungi: new rearing data from Finland and Russian Karelia and general analysis of known larval microhabitats in Europe

2011·0.25 MB·

by  Jevgeni Jakovlev

©EntomologicaFennica.20December2011
Fungus gnats (Diptera: Sciaroidea) associated with dead wood
and wood growing fungi: new rearing data from Finland
and Russian Karelia and general analysis of known larval
microhabitats in Europe
JevgeniJakovlev
Jakovlev,J.2011:Fungusgnats(Diptera:Sciaroidea)associatedwithdeadwood
andwoodgrowingfungi:newrearingdatafromFinlandandRussianKareliaand
generalanalysisofknownlarvalmicrohabitatsinEurope.—Entomol.Fennica
22:157–189.
Inthiscontributionnewrearingrecordsoffungusgnatsfrompoorlystudiedlar-
val microhabitats are presented. From 61 species of wood growing Basidio-
mycetefungi,6speciesofAscomycetefungiandslimemouldsmostofwhich
hadnotpreviouslybeenthesubjectofrearingstudies,andfromdeadwoodsamp-
leswithfungalmyceliamadeoveraperiodof1994–2009inFinlandandRussian
Karelia,110speciesoffungusgnatswereobtained,98ofthemfromidentified
fungi.Ofthesefor12speciesfungalhostswereformerlyunknownandfor30
specieslarvalmicrohabitatshavebeendiscoveredforthefirsttime.Numbersof
fungusgnatspecieswithknownlarvalmicrohabitats(atotalof498speciesthat
comprises 45.4% of the European fauna) and numbers of known fungal hosts
(some650speciesofmacrofungi)arecalculatedandcategorizedbasedonthis
studyandpreviousrecordsfromEuropeandEastPalaearctic.
J.Jakovlev,FinnishForestResearchInstitute,VantaaResearchUnit,P.O.Box
18,FI-01301Vantaa,Finland;E-mail:[email protected]
Received14July2010,accepted5November2010
1.Introduction cernDiptera.Thisisdueinparttothetechnical
difficulties of obtaining adult flies from larvae
Dipteraareoneofthemajorgroupsofsaproxylic livinginfungalfruitingbodies,andespeciallyin
insects, but the ecology of most species is still decaying wood, which are most difficult to be
poorlyknown.Manyspeciesarerareandthreat- cultured.Notleastistheproblemofidentifying
enedwithextinctionduetolossofwoodlandsand host fungi, especially if they are present in the
impoverishment of what remains. However, the samplesonlyassteriletissues(mycelia)thatare
conservation of them is hindered by lack of oftenusedbyDipteralarvae.
knowledge, particularly poor understanding of Fungus gnats in the broad sense (Diptera:
the larval habitat requirements (Rotheray et al. SciaroideawithoutSciaridae),includingthefam-
2001).Duringrecentyearsmanystudiesofsub- iliesBolitophilidae,Ditomyiidae,Diadocidiidae,
strateassociationsamonginsectsinhabitingdead Keroplatidae, and Mycetophilidae with more
wood and fungi growing on wood have been than1,100speciesthatoccurinEurope(Chandler
made,butwithafewexceptionstheydonotcon- 2004andsubsequentcontributionsbyvariousau-
158 Jakovlev (cid:127) ENTOMOL.FENNICAVol.22
thors)comprisethelargestgroupofDipteraasso- studyingthecollectedmaterialneedconfirmation
ciated with fungi. All known rearing records of (Falk&Chandler2005).Inmorerecentyearsad-
fungus gnats with a few exceptions are from ditionalrearingrecordswereobtainedfromEu-
fungi,eitherfromfruitingbodiesorfromrotten rope by Chandler (1993a), Rimšaite (2000),
wood or soil litter impregnated with fungal Zaitzev (2003), Šev(cid:1)ik (2006) and from Japan
mycelia. Based on this fungus gnat larvae are (Sasakawa&Ishitaki1999),whichraisedtheto-
generally viewed as mycetophagous although it talnumberoffungusgnatspecieswithknownlar-
is uncertain how many species are true fungal valmicrohabitatsto some400 speciesin Palae-
feeders and how many are predatory or sapro- arcticandsome380speciesinEurope.
phagous. Afurther increase in sources of information
Studies focused on discovering fungal host on fungus gnat species with formerly unknown
speciesofthelarvaeoffungusgnatshasalong larval microhabitats appeared when researchers
historyinEurope.Traditionallygoodknowledge started to use emergence traps set up on dead
of the larval microhabitats have been in Great wood(overfallentrunks,branchesandstumps),
BritainsinceaclassicstudybyEdwards(1925) soil,litterandmosscarpets(Jakovlevetal.1994,
and subsequent investigations by Madwar Irmleretal.1996,Økland1999,Šev(cid:1)ík&Rohá-
(1937), Buxton (1960), Trifourkis (1977) and (cid:1)ek2008;Jakovlev,J.,Penttinen,J.,Polevoi,A.,
Chandler(1978,1993a)coveringabouttwohun- Salmela,J.&Ståhls-Mäkelä,G.,inprep.).This
dredfungusgnatspecies.Comprehensivestudies method does not alter the substrate or micro-
on Diptera living as larvae in fungi were con- climaticconditionsandcertainlyproducesacon-
ducted also in certain areas of Central Europe, siderable variety of fungus gnat species. How-
e.g.GermanandAustrianAlps(Eisfelder1955, ever, records obtained by the emergence traps
Plassmann 1971), Hungary (Dely-Draskovits (withafewexceptions,e.g.Cardew&Carrières
1974),Czech&Slovakrepublics(Šev(cid:1)ik2006) (2001)wheretheauthorssampledonlyonefun-
and in Northern Europe, including Finland galspecies)donotalwaysprovidetheexactin-
(Hackman&Meinander1979,Väisänen1981), formation on the fungal hosts because two or
RussianKarelia(Jakovlev1980,1986,1995)and more species of fungi might inhabit the sample
Estonia(Kurina1991,1994,1998). coveredwiththetrap.Thisdoesnotallowcorre-
Rearingrecordsoffungusgnatswerealsoob- lating hatched species of insects with their re-
tained from France (Falcoz 1921, 1923, 1926, spectivefungalhosts,andtherecordsarenotof
Bonnamour 1926, Matile 1962, 1963, 1964, the same value as those obtained with rearing
1990),theNetherlands(Barendrecht1938),Italy fromthesampleofonlyonehostfunguswhichis
(Canzanelli1941),Bulgaria(Bechev1989),Por- correctlyidentified.
tugal(Ribeiro1990),Lithuania(Rimšaite2000) Currentknowledgeofthehostfungioffungus
and some areas of Central Russia (Sakharova gnat species relies mostly upon rearing records
1977, Zaitzev 1984c, Khalidov 1984, Krivo- from soft fruiting bodies of agarics, russulas,
sheinaetal. 1986), SouthernRussiaandTrans- boletiandsomesoftpolypores,whereasassocia-
caucasian republics (Krivosheina & Mamajev tionswithwood-encrustingfungiandwithfungal
1968,Zaitzev1994,2003),SiberiaandRussian mycelia in dead wood, soil and litter are still
Far East (Ostroverkhova 1979, Zaitzev 1982, poorly investigated. What little is known of
1994,2003)andJapan(Okada1939,Sasakawa& saproxylic mycetophilids has been summarized
Ishizaki1999). fortheBritishIslesbyAlexander(2002),chiefly
Accordingtomycompilationclassifyingthe on the basis of the accounts of Edwards (1925)
presentknowledgeofrearingrecordsofPalaearc- and Chandler (1978, 1993a). For other areas of
ticDipterafromfungalfruitingbodiesandfrom thePalaearcticregionrearingrecordsfromwood-
othermediacontainingfungalhyphae(Jakovlev encrustingfungiandslimemouldsarescattered
1994), identified fungal hosts were known for inafewpublicationsofwhichthemostcompre-
some300speciesinthePalaearcticRegionand hensive are Krivosheina et al. (1986), Jakovlev
some 280 species in Europe, of these some old (1994),Zaitzev(1994,2003)andŠev(cid:1)ik(2006).
rearing records which cannot be checked by Itisevidentthatarequirementisastudyofthe
ENTOMOL.FENNICAVol.22 (cid:127) Fungusgnatsrearedfromwood 159
Table1.Listofstudysiteswithinformationontheirlocationandforesttype.Thelocalitynameistheclosest
nameforeachsiteona1:20000map.Thefollowingabbreviationsareusedforthesitetypes:ogf=oldgrowth
forest,omf=oldmanagedforest,city=urbanforestpatchesincities;bcc=clear-cuttreatedwithprescribed
burning.Coordinates:Finland=E27grid;RussianKarelia=E33grid.
Site Pro- Municipality Locality Coordinates Sitetype Years
No vince
1 Ab Karjalohja KarkaliSNR 66851:33221 ogf,lime 2004–2008
2 N Espoo NuuksioSNR 66939:33621 ogf,spruce 2005
3 Ta Lammi KotinenSNR 67944:33964 ogf,aspen 2004–2008
4 Ta Lammi KotinenSNR 67946:33965 ogf,spruce 2003
5 Ta Padasjoki VesijakoSNR 68061:33988 ogf,spruce 2005,2008
6 Pp Tervola PisavaaraSNR 73536:34154 ogf,spruce 2003
7 Kon Kondopoga KivachSNR 69088:35503 ogf,spruce 1994,1998,2000
8a Ok Kuhmo UlvinsaloSNR 70998:36638 ogf,spruce 2004,2005
8b Ok Kuhmo PaljakkaSNR 71834:35502 ogf,spruce 2004,2005
8c Kpoc Kostomuksha KostamusSNR 71707:36590 ogf,spruce 2005
9 N Sipoo Rörstrånd 67068:34008 ogf,spruce 2005,2009
10 N Sipoo Sipoonkorpi 66915:33983 ogf,spruce 2008,2009
11 N Kirkkonummi Fagerö 66694:33661 omf,alder 2007
12 Ta Lammi Pappilanlehto 67731:33946 omf,aspen 2004,2007,2008
13 Ta Luopioinen Kuohijoki 68017:33813 omf,aspen 2007,2008
14 N Helsinki Haltiala 66859:33855 city,spruce 2007,2008
15 N Helsinki Herttoniemi 66801:33909 city,spruce 2007
16 N Vantaa Kuusijärvi 66909:33962 city,spruce 2001,2003,2004
17 N Tuusula, Ruotsinkylä 66971:33898 city,spruce 2005
18 Kon Petrozavodsk Lososinka 68519:35571 city,spruce 1996
19 Li Kilpisjärvi Saanamountain 76751:32533 mountainbirch 2006
20 Ta Lammi Evo,Leipäsuonaho 67899:33959 bcc 2003,2005,2008
21 Ta Lammi Evo,Saarijärvi 67927:33959 bcc 2003,2004,2005
fungusgnatspecieswhichmightberearedfroma 2.Studyareas,materialandmethods
widerangeofwood-growingfungithathavenot
been sufficiently examined by entomologists. 2.1.Samplingsites
Manyspecieswithunknownlarvalmicrohabitats
couldberestrictedtospeciesortogroupsoffungi Thedatapresentedinthispaperresultfrommate-
thathavebeenlittlestudied.Theotheroptionis rials collected in Russian Karelia during 1994–
thattheydonotcolonizefruitingbodiesbutuse 2000 and in Finland during 2001–2009. The
decayingwoodasashelterandfungalmyceliaas study areas were located both in hemiboreal,
nutritionthatmakethemdifficulttofindandrear. southern-,mid-andnorthernborealzonesandac-
ForthepastfifteenyearsIhaveworkedtofill cording to Heikinheimo & Raatikainen (1971),
thisgapinknowledgeandthispaperpresentsre- include biogeographical provinces of Regio
sultsofmyrearingexperimentsfromdeadwood aboensis(Ab),Nylandia(N),Tavastiaaustralis,
and wood-encrusting fungi in Finland and Rus- (Ta),Ostrobottniakajanensis(Ok),Ostrobottnia
sian Karelia that have not been published yet. borealis (Ob), Lapponia enontekiensis (Le) in
Most of the fungi involved had not previously Finland, Karelia onegensis (Kon) and Karelia
beenthesubjectofrearingstudies.Ialsotriedto pomoricaoccidentalis(Kpoc)inRussianKarelia.
summarizeallrearingrecordsfromtheliterature The sites from which fungi were sampled
to categorise substrate associations of fungus (Table 1) were selected to obtain a diversity of
gnatspeciesoccurringinEurope. wood-growingfungiandwere,therefore,gener-
allysituatedinprotectedareaswherefallenwood
islefttodecayontheground.Altogethertensites
160 Jakovlev (cid:127) ENTOMOL.FENNICAVol.22
consistedofold-growthforest(ogf)instrictna- inKuhmo,EasternFinlandandKostamusStrict
turereserves(SNR)inFinland(sites1–6,8a,b) NatureReserve,RussianKareliawerepassedto
andinRussianKarelia(sites7,8c).Anotherten mebyGergelyVárkonyiin2004–2005andthose
sites (10–18) were in small reserves for old- reared from fruiting bodies of polypores in
growth (sites 9–10), old-managed forest (omf, Pisavaara,northernFinlandthatIobtainedfrom
sites11–13)andurbanforestpatches(city)pre- DmitrySchigelin2004arealsoincludedinthis
servedwithinthecitiesofHelsinki(sites14–15), paper.
Vantaa,Tuusula(sites16–17)andPetrozavodsk
(site18).Allthesesitesrepresentseminaturalco-
niferous forest, whether spruce-dominated or 2.3.Rearingtechniques
mixedherb-richforestonthelimestoneandfer-
tile soils with a large proportion of deciduous In1994–2007forrearingadultfliesfromfungal
trees,chieflyaspen,birch,alder,andinsomesites hostsIusedthetraditionalmethodbyplacingthe
alsolimeandhazel. sampleoffungalfruitingbodiesorpiecesofdead
Additional rearing records were obtained wood with mycetophilid larvae into rearing
from non-forested sites like oroarctic mountain chambersonalayerofsterilizedpeatsoil(turf).A
birchmeadowsbehindthetimberlineinFinnish peat (substrate) was covered with damp moss
Lapland (site 19) and, to allow fire-dependent (Sphagnum, etc.) on which the samples were
fungitobesampled,fromclearcutstreatedwith placed. To minimize disturbance of the larval
prescribedburning(bcc)in1997–2001andwith websItriedtosamplerelativelybigpiecesofde-
some trees retained, in southern Finland (sites cayingwoodwithfungiusing1.5–2lplasticcon-
20–21). tainers(sizesvaryfrom114×110mmto180×
180mm)astherearingchambers.Thecontainers
werekeptoutdoorsuntilthefirstfrostinOctober
2.2.Samplingmethods –November,andthenwereremovedtothelabo-
ratorywheretherearingprocesswascontinuedat
Adult fungus gnats were reared from fruiting roomtemperature.Thecontainerswereregularly
bodiesgrowingondeadwoodorfrompiecesof checked to collect emerging adults, keeping
deadwoodimpregnatedwithfungalmycelium.I moistureandremovingmoulds,predaceousbee-
aimedtocoverawiderangeofwood-decayfungi tlesandmites.
andthereforetriedtocheckdifferenttreespecies In2008–2009toavoidhighmortalityoflar-
in different stages of decay (including strongly vae the rearing techniques were slightly modi-
decayedlogsovergrownbysurroundingground fied:apartoftherearingchambersweremoved
vegetation)andindifferentconditions.Thestudy intothelaboratory,whiletheotherpartwasoper-
areas were walked through and dead wood ap- ateddirectlyintheforestsitestoprovidethelar-
pearingtobesuitablehabitatforfungusgnatlar- vaewithamorenaturalpupationhabitat.Samples
vae (fallen logs, logging residues and naturally weresortedintwopartsbythekindsofmaterial:
brokenbranches,stumpsandstandingdeadtrees) (1)larvaeinsidefungalfruitingbodiesor(2)out-
were carefully examined. Mosses, lichens and side fruiting bodies, or in/on wood, under bark,
liverwortsgrowingondecayingwoodwerealso etc.ForthelattergroupofsamplesIremovedthe
examined for the presence of Diptera larvae. bottomsfromtheplasticcontainersandembed-
Samplescontaininglarvaeweretakenforfurther dedthemflushwiththesoildirectlyintheforest
rearing. The common species of fungi (e.g. site, next to the log or stump where the larvae
Fomes fomentarius, Fomitopsis pinicola, etc.) were found. Then I put the sample with larvae
were identified directly in the field, but of the within the container on a layer of damp moss.
more difficult species I collected specimens of Containerswerecoveredwithaperforatedtrans-
fruitingbodiesforfurthermicroscopicalidentifi- parentcoverandprotectedfromrainwithakind
cationbytheexperts(seeAcknowledgements). ofhip-roofmadefromlaminatedpaper.Ichecked
AfewspecimensofMycetophilidaecollected thecontainersone-twotimesamonthduringthe
duringaninvestigationintothesaproxylicinsects season, opening the cover and placing inside a
ENTOMOL.FENNICAVol.22 (cid:127) Fungusgnatsrearedfromwood 161
smallpieceofcottonwithetherforashorttime. 3.1.Surveyofspecies
ThenIcarefullysearchedemergingadultswitha
pocketlamp.Adultfungusgnatsthatemergedin The list of fungus gnat species obtained during
therearingchamberswereidentifiedtothespe- this study is constructed in the following order:
cieslevel(allmalesandalsothefemalesinsome species name, number of reared individuals
genera where it was possible) and preserved in (males, females), locality according to Table 1,
70%alcoholusing2mlplastictubeswithscrew speciesoffungus(iffungusfruitingbodypresent
caps. Male terminalia for detailed observation in the sample), tree species and details (trunk,
wereseparatedfromtheabdomenandheatedina stump,branch),collectingdate,emergencedate.
solution of KOH for maceration, then washed Larvae and cocoons, if not specified otherwise,
withaceticacidanddistilledwaterforneutraliza- werefoundin/onthefruitingbody.Fruitingbod-
tionandinsertedintoglycerine. After examina- iescollectedfromthesametreeareconsideredas
tion,terminaliawerestoredintheglycerineme- onesample
diuminspecialplasticmicrovials.Thematerialis Specieswithformerly unknownnamedfun-
depositedintheauthor’scollection. galhostsaremarkedwithoneasterisk(*)infront
ofthespeciesname,thosewithtotallyunknown
larvalmicrohabitats(thefirstrearingrecords)are
3.Results marked with two asterisks (**). Specimens ob-
tainedfromD.SchigelandG.Várkonyiarede-
The pooled material obtained during this study notedwiththecollector’snamewithinbrackets,
consistsofmorethan400rearedindividualsand otherwiseallmaterialwascollectedbytheauthor.
110 species of mycetophilids, 98 of them from Higher taxonomy of mycetophilids follows
identified fungi, the others from rotten wood Bechev(2000),hierarchyandnumbersofspecies
withoutfungalfruitingbodies.Theywerereared occurring in Europe are given, if not specified
from61speciesofBasidiomycetefungi,includ- otherwise,accordingtotheFaunaEuropaeaon-
ingagarics,polyporesandnon-polyporespecies, linedatabase(Chandler2004).Thenomenclature
5speciesofAscomycetefungiandonespeciesof offungigenerallyfollowsCABI,Bioscienceda-
slimemoulds(seeAppendix). tabases (2008) and Kotiranta et al. (2009) for
Among110fungusgnatspecieslistedinthis polyporesandotheraphyllophoroidfungi.
paper 68 species have previously been reared Data on known larval microhabitats is pre-
fromfungi.Fortheother12species,viz.:Diado- sented (if the original record is not otherwise
cidiaspinosula,Orfelianigricornis,O.unicolor, specified)accordingtoJakovlev(1994)andfur-
Macrocera fasciata, M. pilosa, Mycomya bico- ther rearing records from fungal fruiting bodies
lour,Boletinanigricans,Ectrepesthoneuracoly- by Chandler (1993a), Kurina (1998), Rimšaite
eri,Dynatosomareciprocum,Trichontahamata, (2000), Zaitzev (2003) and Šev(cid:1)ík (2006). Re-
T. subfusca, T. subterminalis fungal hosts were cordsobtainedwithemergencetrapsoverdecay-
unknown and for 30 species, including Orfelia ingwood,soilandleaflitter(Jakovlevetal.1994,
nemoralis,Mycomyanitida,M.forestaria,M.ru- Irmler et al. 1996, Økland 1999, Šev(cid:1)ík &
ficollis, Neoempheria pictipennis, Boletina ed- Rohá(cid:1)ek 2008; Jakovlev, J., Penttinen, J., Pole-
wardsi, B. populina, Acnemia falcata, Phthinia voi,A.,Salmela,J.&Ståhls-Mäkelä,G.,inprep.)
congenita,P.mira,Sciophilafenestella,S.jakuti- areindicatedseparately.
ca,S.setosa,Syntemnadaisetsuzana,S.penicil-
la,S.stylatoides,Leiapicta,Epicyptafumigata,
Mycetophilaabiecta,M.bohemica,M.dziedzick- 3.1.1.FamilyBolitophilidae
ii,M.lubomirskii,M.xanthopyga,Phroniaunica,
Zygomyiapictipennis,Z.vara,Z.zaitzevi,Brevi- GenusBolitophilaMeigen
cornu serenum, Exechiopsis pulchella and Bolitophila is a genus comprising 36 species In
Pseudobrachypezahelveticalarvalmicrohabitats Europe, of these there are rearing records for
werediscoveredforthefirsttime. twentyonespecies,allfromfungalfruitingbodi-
es. Most species are associated with Agaricales
162 Jakovlev (cid:127) ENTOMOL.FENNICAVol.22
(both lignicolous and terrestrial) and Boletales. nen 2001), Postia and Leptoporus (Chandler
Ofthesetherearesomepolyphagousspecies(e.g. 1978,Šev(cid:1)ík2006,Schigeletal.2006).
B. cinerea Meigen, 1818, B. tenella Winnertz,
1863, B. hybrida Meigen, 1804, B. pseudo- 3.1.2.FamilyDiadocidiidae
hybrida,Landrock1912)thatwererearedfroma
number of fungal hosts and, vice versa, species GenusDiadocidiaRuthe
thatareprobablyconfinedtoparticulargroupof Five species in Europe; collecting records ob-
fungal hosts, e.g. B. basicornis Mayer, 1871 – tainedusingemergencetrapsoverrottingwood
fungi of the family Cortinariaceae, B. aperta – exist for the three most common ones: D.
Strophariaceae,B.bimaculataZetterstedt,1838, ferruginosaMeigen,1830,D.validaMik,1874,
B. glabrata Loew, 1869 and B. maculipennis D. spinosula Tollet, 1948 (Irmler et al. 1996,
Walker,1836–Tricholomataceae,B.nigrolinea- Økland1999)andfortherecentlydescribedspe-
taLandrock,1912–Paxillaceae,B.rossicaLan- cies D. trispinosa Polevoi, 1995 (Jakovlev, J.,
drock, 1912 – Boletaceae. Among wood-gro- Penttinen, J., Polevoi, A., Salmela, J. & Ståhls-
wing fungi Bolitophila species usually colonize Mäkelä,G.,inprep.).Preimaginalstagesarede-
agaricslikeArmillaria,Flammulina,Hypholoma scribedonlyforD.ferruginosathatliveaslarvae
andPholiota.Onlythreespecies:B.occlusaEd- in long dry silken tubes under bark or in rotten
wards,1913,B.obscuriorStackelberg,1969and wood(Edwards1925)andprobablyfeedonfun-
B. rectangulata Lundström, 1913 are chiefly or gal mycelia (Zaitzev 1994) or spores (Matile
exclusively associated with soft polypores, of 1997).
these B. rectangulata is restricted to the only Diadocidia ferruginosa Meigen, 1830. 1 #,
fungalhosts,Laetiporussulphureus. Site 18, ex. Peniophora laurentii on fallen pine
Bolitophila (Bolitophila) tenella Winnertz, branch, 12.–29.VI.1996. Chandler (1993a) re-
1863.4##,Site5,ex.Armillariamellea-group portedtherearingofthisspeciesfromthefungus
on aspen, 18.VIII.–4.IX.2003. Formerly reared Peniophorasp.
from many species of agarics, mostly wood- * Diadocidia spinosula Tollet, 1948. 3 ##,
growingArmillaria,HypholomaandPholiota. Site 18. Reared from decaying wood of burnt
Bolitophila (Cliopisa) aperta Lundström, spruce stump bearing Antrodia xantha, 12.–
1915.7##,3$$,Site12,ex.Pholiotasquarro- 29.VI.1996.
saonbirch,24.IX.–1.XI.2007;2##,Site15,ex.
Hypholoma capnoides on spruce stump, 2.IX.– 3.1.3.FamilyKeroplatidae
13.X.2008.FormerlyhasbeenrearedfromHyp-
holoma capnoides (Hackman & Meinander SubfamilyKeroplatinae
1979), H. fasciculare, Cortinarius trivialis and
Tricholoma focale (Jakovlev 1995), all rearing GenusKeroplatusBosc
recordswereobtainedfromFinlandandRussian AllfivespeciesoccuringinEuropearewebspin-
Karelia. nersonbracketfungiandprobablysporefeeders
Bolitophila (Cliopisa) occlusa Edwards, (Matile 1990). Larvae of the most widespread
1913. 5 ##, 4 $$, Site 10, ex. Postia alni on species,K.testaceusDalman,1818andK.tipu-
aspen stump on clear-cut, 4.–18.IX. and 22.IX. loidesBosc,1792havebeenrepeatedlyrecorded
2008;2##,1$,Site5,exPostiacaesiaonspru- living in webs, which they construct under the
celog,24.VIII.–25.IX.2008;5##,4$$,Site9, bracketsandonadjacentbark(Chandler1993b,
ex. Postia stiptica on spruce log, 19.X.–5.XI. Jakovlev1994).K.reaumuriiDufour,1839was
2009; 5 ##, 3 $$, Site 6, ex. Postia alni on rearedbyChandler(1993b)fromlarvaefoundon
aspenlog(D.Schigelleg);2##,sameplaceex. undetermined encrusting fungi on fallen bran-
Leptoporusmollisonsprucelog(D.Schigelleg.). ches. Šev(cid:1)ík (2006) reared K. tuvensis Zaitzev,
FormerlyhasbeenrearedfromHypholomafasci- 1991 fromPolyporus varius. Accociations with
culare (Bogatyreva 1979), Pleurotus (Zaitzev ThelephoraterrestrisforK.disparDufour,1839
1984c)andfromsmallsoftpolyporesasPolypo- citedinJakovlev(1994)actuallyrelatedtoK.tes-
rus (Ostroverkhova1979), Amylocystis (Komo- taceus.
ENTOMOL.FENNICAVol.22 (cid:127) Fungusgnatsrearedfromwood 163
KeroplatustestaceusDalman,1818.1#,Site natedforest.Larvaeinwebsontheundersideof
17, ex. Trametes hirsuta on birch log, 10.VI.– fallendampaspenbranchesbearingfungalmyce-
5.VII.2002; 1 #, 2 $$, Site 2, ex Bjerkandera lium and resupinate fruiting bodies of
adusta on dead standing grey alder, 10.VIII.– Byssomeruliuscorium,29.V.–18.VII.2008;1#,
9.X.2005;2##,1$,Site2,exFomitopsispini- Site 8a, reared from a piece of decaying aspen
cola on fallen grey alder log, 16.VI.– log,9.–16.VII.2004(G.Várkonyietal.leg).For-
24.VII.2005;1#,Site10,ex.Pycnoporuscinna- merlylarvaewerefoundinrottingwood(Chan-
barinus on birch log, 14.V.–12.VII.2008. This dler1978),underbarkofdeadtrunkofJuglans
speciesdevelopsinwebsontheundersideoflogs regia (Zaitzev 1994) and imago obtained with
bearingencrustingfungiorbeneaththebrackets emergence traps over beech logs (Irmler et al.
ofpolypores.TherearerecordsfromFomes,Fo- 1996).
mitopsis, Hapalopilus, Phellinus, Polyporus, Orfeliaunicolor(Staeger,1840).2##,Site
Pycnoporus,Stereum,SerpulaandTrametesspe- 1, reared from fallen moist and moss-covered
cies(Falk&Chandler2005). trunkofwillow,Salixcaprea.Larvaeinwebson
KeroplatustipuloidesBosc,1792.1#,2$$, lowerside,onthesurfaceofrottenwoodwithout
Site 4, ex. Fomes fomentarius on birch, 9.VI.– fungal fruiting bodies. 26.V.–30.VII.2006. For-
12.VII.2003. Formerly was reared repeatedly merlyrearedfromfallensprucetrunk,larvaeun-
fromF.fomentariusandwas,therefore,classified der bark, on the surface of dead wood covered
as monophagous on this fungal host (Jakovlev withfungalmycelia(Zaitzev1994)andobtained
1994,Cardew&Carrières2001). in emergence traps over beech and alder logs
(Irmleretal.1996).
GenusOrfeliaCosta
Fifteen species in Europe, all are web spinners GenusNeoplatyuraMalloch
chieflyassociatedwithdeadwoodbut,according FourspeciesinEurope,rearingrecordsexistfor
toHutsonetal.(1980)andSmith(1989)couldbe onlyonespecies,N.flava,larvalhabitsaresimi-
alsofoundinturf,grasstussocks,underlogsand lartothoseofOrfelia–thelarvaeliveinwebsun-
boulders, in worm tunnels, among mosses and derbark,underfallenwoodandinsoil.
liverworts. Rearing records do not indicate Neoplatyura flava (Macquart, 1826). 1 #,
namedfungalhostswiththeexceptionofChand- Site10,rearedfromdampmoistbirchbranches
ler(1993a)whohasreportedOrfeliaunicolorfor stored in heaps at the edge of clear-cut area.
thefirsttimefromthepupasuspendedinthreads 29.V.–18.VI.2007.Thelarvainwebsontheun-
onTrametesversicolor. dersideofbranchesbearingdeadfruitingbodies
*Orfeliafasciata(Meigen,1804).1#,Site of Fomes fomentarius. Formerly was obtained
13, rearedfromdampaspenstump;thelarvain with emergence traps over decaying pine wood
webs under the loose bark bearing dead sporo- (Jakovlevetal.1994)onsoil,groundvegetation
phores of Trametes ochracea, 29.V.–18.VII. and moss carpets (Økland 1999) and has been
2008. Formerly was reared by Edwards (1925, rearedinWestSiberiafromDaldiniaconcentrica
p.530)from“larvaefoundfeedingonmouldsun- andChalciporuspiperatus(Bogatyreva1979).
derloosebutwetbark(poplar)”andobtainedin
emergencetrapsoverbeechstumps,logsandon SubfamilyMacrocerinae
litter(Irmleretal.1996).
** Orfelia nemoralis (Meigen, 1818). 1 #, GenusMacroceraMeigen.
Site13,rearedfromslashresiduesinaspen-domi- Thisgenuscontains46speciesinEurope.Incon-
natedforest.Larvaeinwebsontheundersideof trastwithKeroplatinae,larvaeofMacroceraare
fallendampaspenbranchesbearingfungalmyce- quitedifficulttofindandthatseemstobetherea-
lium and resupinate fruiting bodies of Bysso- sonofthetotalabsenceofrearingrecordsfrom
meruliuscorium,29.V.–18.VII.2008.Noformer namedfungalhosts.Edwards(1925)mentioned
rearingrecords. thattheearlystagesofMacroceraarepractically
*Orfelia nigricornis Fabricius 1805. 2 ##, unknown,inspiteofthefactthatmanyofthespe-
Site13,rearedfromslashresiduesinaspen-domi- cies are quite common. According to Falk &
164 Jakovlev (cid:127) ENTOMOL.FENNICAVol.22
Chandler (2005) larvae of this genus have been develop also in soil and litter, e.g. M. annulata
reared from a range of situations including (Meigen,1818),M.britteniKidd,1955,M.levis
clumpsofturf,rottingwoodandcavewallsand Dziedzicki,1885,M.marginata(Meigen,1818)
areconsideredpredaceous. and M. shermani Garrett, 1924, that have been
Theoldestrearingrecordsexistfortwocom- reared from soil and ground vegetation using
monspecies:M.fasciataMeigen,1804fromlar- emergence traps (Jakovlev et al. 1994, Økland
vaefeedingonfungusgrowinginacellar(Enslin 1999, Šev(cid:1)ík & Rohá(cid:1)ek 2008) and M. nitida
1906) and M. stigma Curtis, 1837 from a de- (Zetterstedt, 1852) found in burrows of rodents
caying trunk of Carpinus betulus (Winnertz (Hackman 1963). Although larval habits of
1863).Furthercollectionsusingemergencetraps Mycomya are similar to those of Orfelia and
(Jakovlevetal.1994,Irmleretal.1996,Økland Macroceratheirlarvaldietisunknown.Accord-
1999, Šev(cid:1)ík & Rohá(cid:1)ek 2008; Jakovlev, J., ing to Laštovka (1972) Mycomya larvae are
Penttinen, J., Polevoi, A., Salmela, J. & Ståhls- viewedaszoophagous,althoughthestudyofgut
Mäkelä,G.,inprep.)hasrevealedM.anglicaEd- contents of M. marginata has verified their
wards,1925,M.angulataMeigen,1818,M.ater- mycophagy(Parmenter1953).
rima Stackelberg, 1945, M. centralis Meigen, KnownfungalhostshavebeenlistedbyVäi-
1818,M.parvaLundström,1914,M.pilosaLan- sänen(1984)andJakovlev(1994)butaccording
drock,1917,M.stigmaCurtis,1837andM.stig- toFalk&Chandler(2005)accuracytheoldrear-
moides Edwards, 1925 from rotten wood, M. ingrecords,e.g.forM.griseovittata(Zetterstedt,
crassicornisWinnertz,1863,M.stigma,M.vitta- 1852), M. ornata (Meigen, 1818) and M.
taMeigen,1830–fromsoilandlitterandM.fa- punctata(Meigen,1818)isdoubtful.Additional
scipennis Staeger, 1840 from tussocks of the fungal hosts have been reported by Chandler
grassScirpussylvaticus. (1993a) and by this investigation. The list of
* Macrocera fasciata Meigen, 1804. 2 ##, knownfungalhostsincorporatesmostlybark-en-
Site3,rearedfromdampmoistlogofaspenbear- crusting fungi, bracket polypores, and wood-
ingDatroniamollis,27.VI.–24.VII.2006.Larval growing agarics but also some terrestrial fungi,
webswereundertheloosebarkontheunderside e.g. Väisänen (1981) has reared M. circumdata
ofthelog.Formerlyitwasrearedfromsomefun- (Staeger,1840),M.permixtaVäisänen,1984,M.
gus growing in a cellar (Enslin 1906) and from tenuis (Walker, 1856) and M. trilineata (Zetter-
larvae found in hollows of an aspen tree stedt, 1838) from decaying fruiting bodies of
(Plassmann1971). Leccinum.
*MacrocerapilosaLandrock,1917.1#,Site Mycomya (Mycomya) annulata (Meigen,
1, reared from rotten fallen trunk of Corylus 1818). 2 ##, Site 7, reared from strongly de-
avellana. Larval webs were on decaying sap- cayedpinetrunksandbranchesretainedinheaps
woodcoveredwithresupinatefruitingbodiesof lyingonthegroundafterthinninginpineforest.
Antrodiella romellii, 16.V.–25.VI.2007. For- Larvae in webs on the fungal mycelia, 24.V.–
merly obtained in emergence traps over beech 14.VI.1994;2##,Site3,rearedfromdecaying
logsandsprucestumps(Økland1999). pine log partly bearing loose bark and covered
with resupinate fungus Skeletocutis biguttulata,
3.1.4.FamilyMycetophilidae 14.IX.–14.XI.2007. Formerly reared from
Polyporus sp. (Ostroverkhova 1979) and from
SubfamilyMycomyiinae Gyromitragigas(Jakovlev1995).
*Mycomya(Mycomya)bicolor(Dziedzicki,
GenusMycomyaRondani 1885).1#,Site21,ex.Gloeophyllumsepiarium
Among the 89 species of Mycomya recorded in on burned spruce stump, larvae in webs on the
Europerearingrecordsexistforaboutone-third lowersurfaceoffruitingbodies,27.VII.–12.VIII.
of them. The larvae of this genus spin delicate 2003.Formerlyreportedas“larvaeonpolypores”
slimywebsusuallyontheundersurfaceofbark- (Plassmann 1971, Sakharova 1977, Väisänen
growing fungi, or on fungal mycelium under 1984) and “Fagus, under bark” (Plassmann
bark.Thereare,however,somespeciesthatcould 1971).
ENTOMOL.FENNICAVol.22 (cid:127) Fungusgnatsrearedfromwood 165
Mycomya (Mycomya) cinerascens (Mac- stedt,1838).1#,Site4,exLaxitextumbicoloron
quart,1826).1#,Site17,ex.Trameteshirsutaon rottingbirchlog,larvaeinwebsonthesurfaceof
birch log, 13.VIII.–6.X.2002. Formerly reared fruitingbodies,14.–30.VIII.2003;1#,Site5,ex.
fromlarvaefoundonfruitingbodiesofStereum Phlebiatremellosaonbirchtrunkbearingloose
(Edwards 1925, Chandler 1978), Thelephora bark, larvae in webs on fruiting body, 27.VII.–
terrestris (Jakovlev 1995) and Cortinarius sp. 4.IX.2005. Formerly reared from rotting
(Kurina 1994). Besides fruiting bodies the spe- Leccinumscabrum(Väisänen1984).
cieshasbeencollectedwithemergencetrapsover Mycomya (Mycomya) wankowiczii (Dzied-
beechlogsandstumps,alderandsprucestumps zicki,1885).1#,Site8b,rearedfromapieceof
(Irmleretal.1996). decaying silver birch log, 27.VII.–2.VIII.2005
** Mycomya (Mycomya) nitida (Zetterstedt, (G.Várkonyi et al. leg.). Formerly reared from
1852).1#,Site20,ex.Cylindrobasidiumlaeve Stereum on fallen birch branches (Edwards
onsprucestumpunderloosebark,larvaeinwebs, 1925), Phallus impudicus (Plassmann 1971,
29.V.–21.VI.2004. No former rearing records. Hutson et al. 1980) and Hypholoma lateritium
Adults have been found in burrows of rodents (Eisfelder1955).
(Hackman1963).
** Mycomya (Mycomya) forestaria Plass- GenusNeoempheriaOstenSacken
mann,1978.1#,Site18,rearedfromdecaying Larvaearewebspinnersonfungiorrottingwood
spruce stump in moist spruce-dominated forest (Falk & Chandler 2005). Fungal hosts are re-
alongtheriver.Larvaeinwebsunderbark,25.V.– corded for two of eight European species; N.
4.VI.1996.Noformerrearingrecords. proxima(Winnertz,1863)whichhasbeenreared
Mycomya (Mycomya) marginata (Meigen, fromBjerkanderaadusta(Eisfelder1955)andN.
1818). 1 #, 2 $$, Site 5, ex Sparassis crispa, striata(seebelow).
24.IX.–24.X.2008.Formerlyrearedfromvarious **Neoempheriapictipennis(Haliday,1833).
wood-growingfungi,mostly bracketpolypores, 1#,1$,Site1,rearedfromlarvaeinwebson
corticoidandjellyfungi(cf.Jakovlev1994),also moist damp wood of fallen trunk of Corylus
from Pleurotus (Hutson et al. 1980), Naucoria avellana, 23.VII.2007–8.VIII.2007. No former
(Chandler (1993a), Simocybe (Alexander 2002) rearingrecords.
and from fungoid wood (Zaitzev 1994). Col- Neoempheriastriata(Meigen,1818).3##,2
lected by emergence traps over dead wood $$,Site4,ex.Laxitextumbicoloronfallenbirch
(Irmleretal.1996)andontussocksofthegrass trunk, 18.VIII.–2.IX.2003; 2 ##, Site 10, ex.
Calamogrostis epigejos (Šev(cid:1)ík & Rohá(cid:1)ek Trichaptum pargamenun on birch stump, 14.–
2008). 29.VII.2003;1#,Site16,rearedfromlarvaeon
Mycomya (Mycomya) occultans (Winnertz, thesurfaceoffruitingbodiesofTrameteshirsuta
1863).1#,Site4,ex.Laxitextumbicoloronrot- on birch stump, 30.VI.–24.VII.2009. Formerly
ting birch log, 18.VIII.–5.IX.2003. Formerly was reared from larvae on fruiting bodies of
reared from bracket fungi growing on broad- wood-growing fungi: Trametes suaveolens
leaved trees, viz: Daedalea, Piptoporus (Dufour 1842), T. versicolor (Zaitzev 1994),
(Winnertz 1863, Landrock 1927), Lenzites Auricularia auricula-judae (Falcoz 1923),
betulina, Plicaturopsis crispa (Eisfelder 1955), Thelephoraterrestris(Jakovlev1995).Khalidov
Inonotus radiatus (Väisänen 1984, record with (1984)publishedarearingrecordofthisspecies
question mark) and obtained with emergence from Tapinella atrotomentosa which is very
trapsoverbeechlimbs(Schiegg1999). rarely infestedwithinsectlarvae.Matile(1963)
** Mycomya (Mycomya) ruficollis (Zetter- found larvae of N. striata in webs on pine
stedt,1852).1#,Site7,ex.Trichaptumparga- branches lying on the ground and considered
menum on fallen birch trunk, larvaein webs on themtobecarnivorousonnematodeswhichbe-
the lower surface of fruiting bodies, 19.V.– cameimmobilised(probablywithoxalicacid)on
14.VI.2000.Noformerrearingrecords. contactwiththeweb.
Mycomya (Mycomyopsis) trilineata (Zetter-
166 Jakovlev (cid:127) ENTOMOL.FENNICAVol.22
SubfamilyGnoristinae likely that they are liverwort feeders. Falk &
Chandler(2005)mentionedthatBoletinaandre-
GenusApolephthisaGrzegorzek lated genera develop in or on fungi, decaying
Thisgenusincludesonlyoneknownspecies,A. woodorbryophytes.Ihave,however,neversuc-
subincana,whichlivesaslarvaeinmucilaginous ceededinfindingBoletinalarvaeinliverwortsor
tubes on the bark of deciduous trees encrusting other bryophytes and also failed to rear any
withfungi(Madwar1937). Boletinaspeciesfromfungalfruitingbodies.
Apolephthisasubincana(Curtis,1837).1#, ** Boletina edwardsi Chandler, 1992. 1 #,
Site 5, reared from larvae in webs under loose Site3,rearedfromdecayingpinelog,partlybear-
barkonsoftdecayingbirchlog(whiterot)bear- ingloosebarkandfruitingbodiesofpolypores.
ingresupinatefungusHyphodontiabarba-jovis, The larvae were found within soft damp rotten
24.VIII.–19.IX.2008;1#,Site8b,rearedfroma woodbearingtheresupinatefungusSkeletocutis
pieceofdecayingaspenlog,31.VIII.–6.IX.2005 biguttulata,22.V.–14.VII.2007.Noformerrear-
(G.Várkonyi et al. leg.). Formerly reared from ingrecords.
Hyphodontia paradoxa (Edwards 1925) and BoletinagriphaDziedzicki,1885.4##,Site
Phlebiaradiata(Trifourkis1977). 18,rearedfromdampdecayingwood(brownrot)
ofsprucestump,9.V.–1.VI.1996;2##,1$,Site
GenusBoletinaStaeger 5, reared from huge spruce log bearing loose
This big genus comprises about 90 members in bark; larvae on the surface of decaying wood
thePalaearcticandmorethan70speciesinEu- (white rot) covered with Resinicium bicolor,
rope(Chandler2004,2009,Jakovlev&Penttinen 28.V.–21.VI.2008.Formerlyrearedfromdecay-
2007). Rearing records exist only for seven of ingwoodofpinewherelarvaeliveinlargecolo-
them viz: B. basalis (Meigen, 1818), B. dubia nies (Jakovlev et al. 1994), spruce (Jakovlev
(Meigen, 1804), B. gripha Dziedzicki, 1885, B. 1995), from soil in pine forest (Jakovlev et al.
nigricansDziedzicki,1885,B.nigricoxaStaeger, 1994)andfromfruitingbodiesofSuillusbovinus
1840,B.trivittata(Meigen,1818)andB.trispi- Kurina(1998).
nosa Edwards, 1913. Of these the commonest * Boletina nigricans Dziedzicki, 1885. 1 #,
fourspecies–B.basalis,B.gripha,B.nigricans Site 3, reared from fallen birch trunk, larvae in
andB.trivittatawereobtainedinemergencetraps whitesapwoodunderbarkbearingHyphodontia
in a range of situations including rotting wood barba-jovis, 22.IV.–11.VI.2006. Formerly was
andsoillitter(Jakovlevetal.1994,Irmleretal. obtainedinemergencetrapsonsoil, theground
1996,Økland1999).Rottingwoodisindicatedas vegetation, moss carpets (Jakovlev et al. 1994,
larvalmicrohabitatforB.trispinosaby Schiegg Økland1999)andoverrottenwood(Jakovlev,J.,
(1999) and soil litter for B. nigricoxa by Plass- Penttinen, J., Polevoi, A., Salmela, J. & Ståhls-
mann(1971).Thespeciesofcloselyrelatedgene- Mäkelä,G.,inprep.).
ra, Saigusaia flaviventris (Staeger, 1840) and Boletina nigricoxa Staeger, 1840. 1 #, Site
Aglaomyia ingrica (Stackelberg, 1948) develop 11, reared from larvae found in thick litter of
in rotten wood as well (Chandler 1978, Zaitzev fallenleavesofAlnusglutinosawoodland.Lar-
1994). vaecollectedon27.IX.2007overwintered;adults
TherearetwocasesofrearingrecordsofBo- emerged next spring, 25.IV.2008. Plassmann
letinafromother substrates. B. griphahasbeen (1971)citedtheoldrecordofBeling(1875)who
rearedfromfruitingbodiesofSuillusbovinusby found larvae of this species between decayed
Kurina(1998)andB.dubiahasbeenrearedfrom leavesinsoillitterindeciduousforest.Myfind-
liverwortsbyCheetham(1920).Edwards(1925), ingsuggestsapossibleassociationoflarvaewith
basedonthelatter recordandhisownobserva- rottingplantmaterial.
tionsthattheadultsofmanyofthelargerBoletina **BoletinapopulinaPolevoi1995.1#,Site
speciesaremostfrequentlyfoundalongbanksof 3,rearedfromlarvaecollectedinrottenwoodof
mountainstreams,supposedthattheymightde- dampsprucelogcoveredwithAntrodiaserialis,
velop amongst bryophytes and it seemed quite 22.V.–14.VI.2007.Noformerrearingrecords.