Table Of Content©EntomologicaFennica. 25 February2011
Parasitoids (Hymenoptera) of leaf-spinning moths
(Lepidoptera) feeding on Vaccinium uliginosum L.
along an ecological gradient in central European peat bogs
AurelI.Lozan,KarelSpitzer,JosefJaros,AndreyKhalaim,MariaConcettaRizzo,
Emilio Guerrieri&AlesBezdék
Lozan, A. 1., Spitzer, K., Jaros, J Khalaim, A., Rizzo, M. C., Guerrieri, E. &
.,
Bezdek, A. 2010: Parasitoids (Hymenoptera) ofleaf-spinningmoths (Lepidop-
tera)feedingon VacciniumuliginosumL. alonganecologicalgradientincentral
Europeanpeatbogs.—Entomol. Fennica21: 243—253.
Parasitoidsofleaf-spinningLepidopteraassociatedwithtwoisolatedcentralEu-
ropean peat bogs were investigated. Five families ofparasitoid Hymenoptera
(Braconidae, lchneumonidae, Eulophidae, Pteromalidae and Encyrtidae) were
recorded. Threecategorieswererecognised: (1)primaryparasitoids,(2)faculta—
tivehyperparasitoidsand(3)obligatoryhyperparasitoids. TenspeciesofBraco-
nidae,fivespeciesandsevenmarkedmorphospeciesamonglchneumonidae,and
three species ofChalcidoidea were identified. Despite ofsome niche-specific
(butlesshost-specific)parasitoids, allthesehymenopteransarelikelytobegen-
eralists andnone ofthemwere confirmedto be habitat and/or host specialists.
Unliketheireurytopic(opportunistictyrphoneutral)parasitoids,theLepidoptera
hostsassociatedwithpeatbogsarepartiallyhighlystenotopic(tyrphobiontsand
tyrphophiles). The occurrence ofparasitoids compared to their potential hosts
was structuredalonganecological(mesoclimatic) gradient, so mostparasitoids
wererecordedfrommarginswhilestenotopic(narrowhabitatadaptation)moths
weremostlydistributednearthecentreoftheboghabitat.
A.I.Lozan,K.Spitzer,J.Jaros&A.Bezdek,BiologyCentre,InstituteofEntomo—
logy, Czech Academy ofSciences, Branisovska 31, 37005 Ceske' Budejovice,
CzechRepublic;E—mails.‘[email protected],[email protected],jaros@entu.
cas.cz, [email protected]
A. Khalaim, ZoologicalInstitute, Russian Academy ofSciences, Universitets—
kaya nab. I, St. PetersburgI99034, Russia;E—mail.‘ [email protected]
M. C. Rizzo, SENFIMIZO Department, Entomology, Acarology and Zoology
Section, Faculty ofAgriculture, University ofPalermo, viale delle Scienze I3,
Ed. 5, Palermo, Italy;E—mail.‘ [email protected]
E. Guerrieri,InstituteforPlantProtection, theNationalResearch CouncilofIt—
aly, Via UniversitaI33,80055Portici(NA),Italy;E—mail.‘[email protected]
Received26January2009, accepted2June2010
244 Lozan etal. ° ENTOMOL. FENNICAVol. 21
1. Introduction nopteran parasitoids (Braconidae), collected by
lighttraps(Lozan2002),werenotrestrictedtothe
Parasitoids ofanecologicalguildofLepidoptera bog as reported for stenotopic Coleoptera and
feedingonrelictboreo-montaneplant Vaccinium Lepidoptera(cf. Bezdeketal. 2006).
uliginosum L., a deciduous circumboreal shrub
characteristic ofpeaty soils ofcoldlands, in two
isolatedcentralEuropeanpeatbogs(Spitzeretal. 2. Material and methods
2003) were studied. The moths rearedfromleaf
spinnings belong to four families (Tortricidae, 2.1. Sitesofinvestigations
Gelechiidae, Chimabachidae and Geometridae)
and include both stenotopic (tyrphobiontic and Leaf spinnings on Vaccinium uligz'nosum were
tyrphophiloustaxa), closelyassociatedwithpeat collected during June 1998—2001 from two
bogs, and opportunistic (tyrphoneutral) species montaneisolatedpeatbogsofthecorezoneofthe
notrelatedtothebogs. gumavaNationalPark, SWBohemia,CzechRe-
PeatbogsofcentralEurope,whichdeveloped public; details aregivenby Spitzeretal. (2003):
under isolation, are true habitat islands for a
unique insectdiversity ofhighly stenotopic taxa a) Mrtvy luhnearVolary (740 ma.s.l., 310 ha)
(”relicts”)amongLepidopteraandparasitoidHy- representsamontaneoligotrophicvalleypeat
menoptera (Spitzer & Banks 2006, Lozan etal. bog more or less closed by the forest. Open
2010). The Lepidoptera community of central treeless areas are covered by V. uligz'nosum
Europeanpeatbogs hasbeenwell studiedunder (andother Vaccinium spp.), Eriophorum va—
the long-term monitoring programmes, but the ginatum andCalluna vulgariswithagradual
parasitoidshavenotbeensubjectofsuchinvesti- transitiontodwarfforestofPinusmugoS. lat.
gations. Thispaperisbasedonprevious investi- aroundthebog.
gations ofLepidoptera (Spitzer etal. 2003) and b) Chalupskaslat’bognearBorovaLada(900m
aims at assessing parasitoid diversity along an a.s.l., 116ha)isamontaneoligotrophicraised
ecological(mesoclimatic) gradientintwo differ- bog with a central bog-lake and mountain
entpeatbogsystems. Thismesoclimaticgradient pine forest around, with large patches of V.
ischaracterisedbyextremetemperaturesandeX- uligz'nosum (including other Vaccinium spp.)
pressed by local vegetation of “forest-tundra” andBetulapubescens (the gradient between
formationsbetweenbogcentresandbogmargins the centre and margins is not gradual like in
(including“lagg”, seeMaterialandmethods,and theMrtvyluhbog).
Bezdeketal. 2006). Thesetwo centralEuropean
bogs are isolated ancienthabitats ofdistinct cli-
matic andedaphic conditions closeto the forest- 2.2. Sampling
tundrabiomewithahighproportionoflocalcold-
adaptedspecies. For collecting the hosts, leaf-spinnings were
There is an identified distinctmicro-climatic sampledalonganecologicalgradientbetweenthe
gradientfromlagg (the ecotone ofouterparts of outermargins, lagg (innermargins) andthecen-
thebog) to margins and centres ofpeatbogs, in- tresofeachinvestigatedbogsystem.Thecoverof
cluding intermediate zones, where we collected V. uligz'nosum was highernearthe treeless areas
ourfieldsamplesforecologicalanalysis.Thebog of the bogs and the stenotopic host taxa (i.e.
habitat affinity (association or relation to bog tyrphobionts+tyrphophiles) were mostabundant
habitat) ofboth moths and parasitoids from our in the centres (edaphic “tundra-like” formation)
sampleswascorrelatedwithavailablereliablere- when compared with the pine forested margins
cords oflevels ofhost specialisation in order to (including the lagg ecotone, which is an integral
revealthedegreeoftheirstenotopyandeurytopy. partofthebog)ofthebogs.Itwasnotpossibleto
Previouspreliminaryinvestigationsinoneofthe identify exactly which parasitoid species
studiedpeatbogsunderthe sameecologicalgra- emerged from the particular host spinning. The
dient(centreversusmargins) showedthathyme- attemptforhostlarvaeidentificationbyheadcap-
ENTOMOL. FENNICAVol. 21 ° Parasitoidsofleafspinningmoths 245
suleswasnotsuccessfulandnotsufficientlyreli- Table 1. No. ofrearedspecimens/speciesfromthe
able. fourfamiliesofleaf-spinningLepidoptera(Chima-
bachidae, Gelechiidae*,Tortricidae*, Geometridae*;
includespecialist/sten0t0pictaxa:tyrphobionts+
tyrphophiles)intwopeatbogs.
2.3.Lepidoptera
Parasitoid family Specimens Species
The complete listofmoth specieswithquantita- reared reared
tive data (species, reared specimens and leaf
spinnings per site) is given by Spitzer et al. Ichneumonidae 26 12
(2003), here we supply only the list ofspecies Braconidae 56 1O
Eulophidae 4 1
wherestenotopictaxaaremarkedwithanasterisk
Pteromalidae 4 1
(tyrphobionts andtyrphophiles), as follow: Encyrtidae 14 1
Diurnea lipsiella (Denis et Schiffermiiller) Unidentified (Chalcidoidea) 3 ?
andDasystomasalicella (Hiibner)(Chimabachi-
dae);Athrz'pspruinosella(Lienig&Zeller)>’<(Ge- Total 107 25
lechiidae); Aclerz's laterana (Fabricius), Aclerz's
maccana (Treitschke)*, Aclerz's lipsiana (Denis
& Schiffermiiller)*, Cnephasia stephensiana All Braconidae and Chalcidoidea specimens
(Doubleday), C. asseclana (Denis & Schiffer- were identified at the species level except for
miiller), Argyrotaenia [jungiana (Thunberg), three damagedones (presumably Chalcidoidea).
Pandemiscinnamomeana (Treitschke),P. hepa— MostofthecollectedIchneumonidaewereidenti-
rana (Denis & Schiffermiiller), Clepsissenecio— fied atthe species level, some only at the genus
nana (Hubner),Adoxophyes orana (Fischervon level.
Roslerstamm), Apotomis sauciana (Frolich)*,
Celypha lacunana (Denis & Schiffermiiller),
2.5.Data analysis
Phiaris bl'punctana (Fabricius)*, Rhopobota
naevana (Hiibner) and Pammene luedersiana We used the same community data analysis
(Sorhagen)>’< (Tortricidae);Rhinoprora(Pasiphi— schemeasinSpitzeretal. (2003). TheCanonical
la)debiliata (Hubner) (Geometridae). CorrespondenceAnalysis(CCA;CANOCOver-
Therearingofatotalof19species(18species sion3.12software)byTerBraak(1987)wasused
oftypical leaf-spinning microlepidopteran spe- todeterminethehabitatpreferencesofparasitoids
cies and one macrolepidopteran leaf-spinning inrelations to the centers andthemargins ofthe
species) was conductedunder laboratory condi- two investigatedbogs.
tions, where larvae were kept and fed on V.
uligz'nosum inglassboxes.
3. Results
Atotalof107 specimensofparasitoids(Table 1)
2.4.Hymenoptera wererearedunderlaboratoryconditions from 19
species ofleaf-spinning Lepidoptera feeding on
All reared hymenopterans were initially pre- typical bog plant V. uliginosum in two central-
servedin80%ethanol,thendriedupwiththehelp European peat bogs ofthe 8umava Mountains.
offilterpaperandmountedintocollections. Tax- Allidentifiedparasitoidspeciescanbeclassified
onomic analysiswas donebyusing themostre- astyrphoneutraltaxa,beinggeneralistsandmore
cent keys (e.g., Graham 1959, Askew 1968, orlesspolyphagousparasitoidsofLepidopteraor
Huddleston1980, 1984,Boucek&Rasplus 1991, oligophagous species associated with smaller
Belokobylskij &Tobias 1998,2000,Guerrieri& numbers of biologically similar Lepidoptera
Noyes 2005), and by comparing the specimens (Table 2). None ofthem was confirmed to be a
with authoritatively identified specimens and habitatand/orhostspecialist,despitethatsomeof
withothermaterialdepositedininstitutionalcol- their leaf-spinning Lepidoptera hosts were bog
lections. specialists.
246 Lozan etal. ° ENTOMOL. FENNICAVol. 21
Table2. Knowngeographicrangeand hostrangeforparasitoid Hymenoptera(lchneumonoidea, Chalcicoidea)
rearedfromleaf-spinning Lepidopterafeedingon VacciniumuliginosumL. intwocentral European peatbogs
(Sumava Mts.).
NumberofLepidoptera recorded
in literature per:
Taxa Geographicrange Families Species
lchneumonoidea: Braconidae
Hormiusmoniliatus Holarctic, Oriental 5 at least 17
Hormiussimilis Transpalaearctic, Oriental ProbablyasH. moniliatus
Oncophanesminutus Transpalaearctic 10 >50
Bracon erraticus Transpalearctic 4 5
Charmonextensor almostCosmopolitan 13 >75
Meteorusictericus Palaearctic, Australian 13 >90
Meteoruscolon Transpalaearctic 1O 25
Earinusgloriatorius West Palaearctic 6 12
Orgiluspimpinellae Transpalaearctic 9 25
Ascogasterbidentula Transpalaearctic 2 2
lchneumonoidea: lcnheumonidae
ltoplectistunetana* Transpalearctic 6 13
Glypta ceratites Europe, Mongolia 6 183
Tranosema carbonellum Holarctic Firstrecord on Lepidoptera
Chorinaeusfunebris Holarctic 7 17
Exochustardigradus Palaearctic Unknown Unknown
Chalcidoidea: Eulophidae
Elachertusartaeus Palaearctic, Nearctic 5 11
Chalcidoidea: Pteromalidae
Termolampapinicola Transpalaearctic 1 2
Chalcidoidea: Encyrtidae
Copidosomafilicorne Holarctic 2 17
*Facultativehyperparasitoid.
According to their known foraging activity, bogwasalmostequalinbothparts, i.e. 15 speci-
they are either primary parasitoids (Braconidae mens and seven species in centre and 16 speci-
and Chalcidoidea, partially Ichneurnonidae) of mens andsiXspecies inmargins (Table4).
Lepidopteraandrarelyofotherinsectgroups, or
obligatory and facultative hyperparasitoids Table3. No. ofrearedspecimensandspeciesper
(amongIchneurnonidae) (see Table3). parasitoidcategoryfromthefourfamiliesofleaf-spin-
Being widely distributed with a wide host ning Lepidoptera(Chimabachidae, Gelechiidae*,
range (Ichneumonoidea) orpartially with a nar- Tortricidae*, Geometridae; *includespecialist/steno-
rower host spectrum among Lepidoptera only topictaxa:tyrphobionts+tyrphophiles)intwopeat
(Chalcidoidea), they have comparatively low bogs.
host specificity and a wide coverage ofvarious
Parasitoid category Specimens Species
habitats,beingnotrestrictedtotheboghabitatat
reared reared
all.
Quantitatively, most parasitoids were re- Primary 95 22
corded from margins in the Mrtvy Luh bog, i.e. Facultative hyperparasitoids 1 1
55 specimensbelongingtotwelvespeciesversus Obligatory hyperparasitoids 8 2
Unidentified (Chalcidoidea) 3 ?
only 21 specimens and nine species from the
centre+intermediary parts. The number ofpara- Total 107 25
sitoids reared from hosts ofthe Chalupska slat’
ENTOMOL. FENNICAVol. 21 ° Parasitoidsofleafspinningmoths 247
Table4. Numberofreared parasitoids(Hymenoptera: Ichneumonoidea, Chalcicoidea)fromleaf-spinning moths
(Lepidoptera)on VacciniumuliginosumL. pereach investigated bogsite(éumava Mts.).
Position along the gradient#
Mrtvy luh Chalupska slat’
Taxa Cntr Inter Marg Lagg Cntr Marg+Lagg
Ichneumonoidea: Braconidae
Hormiusmoniliatus
Hormiussimilis
Oncophanesminutus 0—\00000—\—\OO 0000000000 (00000000000 _\00l\)—\—\O‘ll\)00 00040400000 0000000000
Bracon erraticus
Charmonextensor
Meteorusictericus
Meteoruscolon
Earinusgloriatorius
Orgiluspimpinellae
Ascogasterbidentula A
Ichneumonoidea: lcnheumonidae
ltoplectistunetana*
Glypta ceratites
Glypta sp. 000000000000 00—\—\00000000 0000000004404 400000N0000—\00 0—\00—\0000000 000000N—\00000
Campoplexsp. 1
Campoplexsp. 2
Campoplexsp. 3
Campoletissp.
Tranosema carbonellum
Chorinaeusfunebris
Exochustardigradus
Mesochorussp. 1**
Mesochorussp. 2**
Chalcidoidea: Eulophidae
Elachertusartaeus 3 O
Chalcidoidea: Pteromalidae
Termolampapinicola O O
Chalcidoidea: Encyrtidae
Copidosomafilicorne 7 O
OtherChalcidoidea (damaged) 3 0
Total 19 2 23 32 15 16
*Facultativehyperparasitoid.
**Obligatoryhyperparasitoid.
#Abbreviations:Cntr—centre, Marg—margins, Inter—intermediatezone, Marg+Lagg—marginsandIaggecotone.
The stenotopic tyrphobiontic species ofleaf microclimatic gradient. On the contrary, the
spinning Lepidoptera were mostly associated stenotopicspecies spectraofLepidopteraofboth
withshrubsofV.uliginosumnearthecentreofthe bog centreswerevery similar, butvery different
investigatedbogs,whiletyrphoneutraltaxawere inbogmargins (Fig. lb). Thegradientis charac-
distributedinthemarginsoftheboghabitat. Spe- terised by the near ground temperatures which
ciesspectraofparasitoidsrecordedfrombogcen- were extremely low near the bog centres and
tres andbog margins were irregularly dispersed gradually increasingto thebog margins, e.g. the
(Fig. la),sothatonlyfewspecieswerepresentin minimumtemperatures inJuly—Augustreachof-
two or more investigated habitats along the ten —4°C near the centres (margins —2°C at the
248 Loganerof. a ENTOMOL.
MLcentre
CS centre
MLintermediate
ML margin
MLlagg
CS margin + lagg
1.8 1.8 2.0 2.2 2.4 2.8 2.8
MLcentre .
Fig. ’l.Similarities(pres-
enceuabsencedata,sin-
CScentre . glelinkage, Euclideandis"
tance)ofdifferentparts,
Ml. intermediate . fromcentrestomargins
(includinginnermar-
Ml... margin ginsxlagg),ofMrtvyluh
(Ml...)andChalupskaslat’
(CS)peatbogs.ma. Para"
CS margin + lagg
sitoids(numberofspecies
and morphospecies)of
Ml. lagg leaf-spinningmoths.mb.
Leafspinning mothcom
munities;datafrom
1.6 1.8 2.0 2.2 2.4 2.8 2.8 Spltzeretai. (2003).
sametimeperiod) (forbasictemperature dataof Teophoridae, Tortricidae. Pyrahdae) (Belokobyi-
thebogsites see Spitzereral. 2003). ski]? &Tobias 1998).
HormiusSimflis Széphgefi, 1896
Tdiobiont ectoparasitoid, very close to previous
ThedatabelowhavebeensummarizedinTable2, species sharing, probably, same habitatrequire-
accompanied by the geographic ranges of the ments and host range (Belokobyiskij & Tobias
parasitoidspecies. 1998)
Oncophanesminums(Wesmael. 1838)
4.1.. Pfil aryparasi‘mids
idiobiontectoparasitoidofLepidoptera. Thepan-
rasitism on Coieoptera (Cerambycidae, Bostri—
41.]. [Chneumonoidea
chidae)and.Hymenoptera,Symphyta(Tenthredim
Braconidae nidae) is not counted and require continuation
(Belokobyiskij&Tobias1998,,Diaconu&Lozan
Hormiusmonflmms (Nees, 1811) 2000)
TdiobiontectoparasitoidofLepidoptera(cg. Com
ENTOMOL. FENNICAVol. 21 ° Parasitoidsofleafspinningmoths 249
Bracon erraticus(Wesmael, 1838) a wide spectrum ofonly Lepidoptera (Aeschli-
Idiobiont ectoparasitoid covering a wide spec- mann 1975).
trumofhosts,mainlyamongLepidoptera(Tortri-
cidae, Geometridae, Gelechiidae etc.) (Tobias Glypta ceratites Gravenhorst, 1829
1986). The species is both morphologically and Solitary koinobiont endoparasitoid of a wide
ecologically very variable (or there are several spectrumofLepidoptera(Aubert 1978).
species in fact), so parasitism on Coleoptera
(Chrysomelidae), Diptera(Tephritidae) andHy- Glypta sp.
menoptera (phytophagous taxa ofEurytomidae) Members ofthegenus Glypta arekoinobionten-
isnotexcluded. doparasitoidscoveringawidespectrumofmicro-
lepidopteranhostslivinginconcealment(Tortri-
Charmon extensor(Linnaeus, 1758) cidae, Gelechiidae, Geometridae etc.) or other
Larvalkoinobiontendoparasitoidcovering awi- Lepidoptera(Lasiocampidae, Lymantriidae, Ly-
despectrumofLepidopterahosts(typicallyfrom caenidae etc.). There are records oftheirparasi-
families Gelechiidae, Geometridae, Tortricidae, tism on other insect orders as Hymenoptera-
Yponomeutidae etc.) (Billups 1897, Achterberg Symphyta (Cephidae, Diprionidae) andColeop-
1979, Belokobylskij &Tobias 1998). Parasitism tera (Cerambycidae, Scarabaeidae, Curculioni-
on Coleoptera (Cerambycidae, Bostrichidae) dae)(Dasch 1988),howevertheserequireconfir-
and/or Hymenoptera, Symphyta (Tenthredini- mation.
dae)requireconfirmation.
Exochustardigradus Gravenhorst, 1829
Meteorusictericus(Nees, 1811) Hostunknown. Membersofthemetopiinegenus
Solitary koinobiont larval endoparasitoid cove- Exochusareprimarykoinobiontlarval-pupalen-
ringawidespectrumofLepidoptera(Huddleston doparasitoids ofawiderangeofmicrolepidopte—
1980,Tobias 1986).Arecordonadiprionidsaw- rous larvaeinleafrollsorsimilarsituations (Ge-
flyrequires confirmation. lechiidae, Tortricidae,Geometridae,Yponomeu-
tidaeetc.)(Gauld2002,Tolkanitz2007).Records
Meteoruscolon (Haliday, 1835) on Coleoptera and Hymenoptera—Symphyta
Solitary koinobiont larval endoparasitoid ofLe- (Tenthredinidae) areprobablyerroneous.
pidoptera(Huddleston 1980, Tobias 1986).
Tranosema carbonellum (Thomson, 1887).
Earinusgloriatorius(Panzer, 1809) Mentionedaskoinobiontendoparasitoidofsaw-
Solitary koinobiont endoparasitoid ofLepidop- fly Arge pagana Panzer (Argidae) by Hedwig
tera(Nixon 1986). (1939).Thisisthefirstrecordofthisspeciesfrom
Lepidoptera.Membersofthegenusaregenerally
OrgiluspimpinellaeNiezabitowski, 1910 knownas endoparasitoids ofvarious smallLepi-
Solitarykoinobiontlarval endoparasitoidofma- doptera, especiallythefamilyTortricidae.
ny Lepidoptera (Taeger 1988, Belokobylskij &
Tobias 1998). Campoletis sp. and Campoplex spp. (three un-
identifiedspecies).
AscogasterbidentulaWesmael, 1835 Members of these Campopleginae genera are
Solitarykoinobiontegg-larval endoparasitoidof known as primary koinobiont larval (rarely lar-
only Lepidoptera (typically families Geometri- val-pupal)parasitoidsofsmallandmedium-sized
dae and Tortricidae) (Huddleston 1984, Tobias Lepidoptera ofvarious families (Townes 1969,
1986) Kasparyan 1981, Gauld 1991, Shaw &Aeschli-
mann 1994, Wagener et al. 2006). Records of
Ichneumonidae theirparasitism on other insect groups (Coleop-
tera) and/or their hyperparasitism on other Ich-
Chorinaeusfunebris(Gravenhorst, 1829) neumonidae require confirmation (Townes
Larval-pupalkoinobiontendoparasitoidcovering 1945)
250 Lozan etal. ° ENTOMOL. FENNICAVol. 21
4.1.2. Chalcidoidea 4.3. Obligatoryhyperparasitoids
Eulophidae Ichneumonoidea: lchneumonidae
Mesochorus spp. (two unidentified species).
Elachertusartaeus(Walker, 1839) Membersofthislargegenusareendoparasitichy-
GregariousectoparasitoidofvariousLepidoptera perparasitoids ofeitherHymenoptera (e.g. para-
(Gelechiidae, Gracillariidae, Noctuidae, Noto- sitoids ofthe families Braconidae, lchneumoni-
dontidae, Tortricidae), attacking larvae offree- dae)ortachinidparasitoids(Diptera:Tachinidae)
movingleafspinning/rollers,orlarvaeofleaf-mi- (Wahl 1993, Schwenke 1999).
ners which spin cocoons intheir last stage often
byrollingtheedgeoftheleaf(Boucek&Askew
1968,Noyes2003). 5. Discussion
Pteromalidae 5.1.Parasitoiddiversity
TermolampapinicolaBoucek, 1961 Parasitoids in our samples can be classified as
KnownasaprimaryparasitoidofRetiniaresinel— shown in Table 3. The members ofthe family
la (L.) (Lepidoptera, Tortricidae) onPinusthun— Braconidae were the most numerous and they
bergz'z'(Parl.)andPinussp.(Noyes2003).Ourre- were only primary parasitoids. The lchneumo-
sults show this species is probably associated nidaewere twice less numerous thanthe Braco-
with a tortricid or gelechiidmoth other than the nidaeandweremorediversified,e.g.primaryand
onlyknownhost,whichcanbeconsideredanew secondary (obligatory and facultative) parasi-
hostrecord.Thespeciesisthereforelikelytobeat toids. Identified species ofChalcidoidea (Ptero-
leastoligophagous. malidae,EncyrtidaeandEulophidae)wereallre-
portedasprimaryparasitoids. Forthelistofspe-
Encyrtidae cies seeTable2.
Primary parasitoids of families Braconidae
Copidosomafilicome(Dalman, 1820) and lchneumonidae can be classified as poly-
Egg-larvalendoparasitoid, oneofthemostcom- phagous species associated with various Lepi-
monspeciesofthegenusinEuropewitharelati- doptera (except probably Bracon erraticus,
velynarrowhostspectrum,limitedtothefamilies whichinliteratureisalsorecordedfromotherin-
Gelechiidae and Tortricidae (Lepidoptera) (Gu- sect orders (see below)). Primary parasitoids of
errieri &Noyes2005). family Chalcidoidea are often more niche-spe-
cific than taxon-specific, with a moderate host
rangeofbiologically similarhosttaxabelonging
4.2.Facultativehyperparasitoids toamoreorlesslargetaxonomicrange(Askew&
Shaw 1986, Lawton 1986).
Ichneumonoidea: lchneumonidae Theonlyonefacultativehyperparasitoidspe-
cies (Itoplectis tunetana, lchneumonidae) is not
Itoplectistunetana (Schmiedeknecht, 1914) veryhabitatselectiveandgenerallypolyphagous,
Primary parasitoids of various Lepidoptera with wide geographic range. The obligatory
(Acrolepiidae, Choreutidae, Gelechiidae, Plutel- ichneumonid hyperparasitoids ofthe genusMe—
lidae,Tortricidae,Yponomeutidae)andhyperpa- sochorus attack probably a series of other pri-
rasitoids ofother parasitoid Hymenoptera (Bra- mary parasitoids, e.g. among parasitoid Hyme-
conidae:Aleiodesrossz'cus) (Kasparyan 1981). noptera and Diptera (Wahl 1993). Although
hyperparasitic intraguild strategy frequently oc-
curs in parasitoid Hymenoptera, it is difficultto
reveal and detect such connections in the field
samples. The multitrophic parasitism in our
samples is undoubtedly only a small part ofthe
ENTOMOL. FENNICAVol. 21 ° Parasitoidsofleafspinningmoths 251
intrinsic and almost unknown parasitoid food (Taeger1988),consistentlywithoursamples.All
web ofpeatbogs. these species ofLepidoptera and Hymenoptera
Mostparasitoidspeciesrecordedinthisstudy (Braconidae) were represented in our samples
areconsideredtobewidelydistributedinthePa- and were also encountered during our previous
laearctic (except Earz'nus gloriatorius, recorded studies in the investigated bog sites (e.g. Lozan
recently only from Europe) and several species 2002, Spitzeretal. 2003, Bezdeketal. 2006).
withalmostcosmopolitanrange(sometaxaintro- Many ofthe obtainedparasitoids are known
duced).Allsuchspeciesseemtobecharacteristic also as true bioregulators operating in orchards
invariousbiotopes,includingpeatlands,buthav- and forests (Oncophanes minutus, Charmon
ingno specificrequirementstoboghabitats. extensor,genusMeteorusetc.),andsomeoftheir
Ahugeamountofhostrecordshasthoroughly potential tyrphoneutral hosts were abundant in
beenanalysed(forexample,Taxapad)inorderto theinvestigatedpeatbogs.
beableto assesstherealhostrangeanddistribu-
tion of each parasitoid species we managed to
identify in our samples. However, many pub- 6. Conclusions
lishedrecords ofparasitismon a large spectrum
oflepidopteran families and/or on other insects In the two isolated central European peat bogs
than Lepidoptera (such as Hymenoptera-Sym- studied here, the diversity ofparasitoids (super-
phyta (Tenthredinidae, Cephidae, Diprionidae) families Ichneumonoidea and Chalcidoidea) is
and Coleoptera (Cerambycidae, Scarabaeidae, slightlyhighercomparedwiththeirlepidopteran
Curculionidaeetc.))mightbewrongand,atmini- hosts. The overall “tyrpho-association” ofboth
mum, all thesepublished connections wouldre- leaf-spinning Lepidoptera and their potential
quireconfirmation.Ultimately,onlyanumberof parasitoids (Ichneumonoidea + Chalcidoidea)
morelimitedandmorereliablesourceswerecon- represents:
sidered, byjudging the host ranges as a context
that has reasonable boundaries (e.g. ‘generalist’ a) 19 species of Lepidoptera, of which seven
versus‘specialist’parasitoid,seeShaw1994)and stenotopic taxa (tyrphobionts+tyrphophiles)
byusingdatafromtaxonomiccollectionsthatwe and 12tyrphoneutrals(cf. Spitzeretal. 2003,
managed to examine elsewhere (see also Ac- Spitzer&Danks2006);
knowledgements). b) 25speciesofmultitrophicparasitoids;noneof
the identified species was a bog stenotopic
taxon, therefore, all of them are tyrpho-
5.2.AssociationwithbogLepidoptera neutrals (eurytopic components of various
ecosystems).
It was really difficult to reveal the precise rela-
tionships of emerged parasitoids to the reared Associatedwithpeatbogs,thereareafewknown
host(s) in our leafspinning samples. The braco- stenotopic taxa among Braconidae (Lozan &
nidendoparasitoid Charmon extensor, whichat- Tobias 2002, Lozan etal. in press) and Ichneu-
tacks a large spectrum ofinsects throughoutthe monidae(Tereshkin 1996)sofar.However,none
world, is known from a strictly tyrphophilous ofthem matched our data ofrearings from leaf
tortricid moth Aclerl's maccana (Billups 1897). spinnings. Investigations of parasitoid Hyme-
The same host species is recorded for a poly- nopteraassociatedonlywithpeatbogsweregen-
phagous endoparasitoid braconid, Meteorus erallyneglectedandnotmany dataare available
ictericus, which is a widely distributed species (cf. Krogerus 1960, Spitzer&Danks2006).
notonlyinthePalaearcticregion.Anotherbraco- The ability ofparasitoids (including multi-
nidendoparasitoid,Orgiluspimpinellae,whichis trophic levels) topopulate awide range ofhabi-
Lepidopterahostorientedandwidelydistributed tatsandhosts(i.e.eurytopy)contrastswiththatof
in the Palaearctic, has been recorded from the stenotopic leaf-spinning Lepidoptera species.
tyrphobionticgelechiidmothAthripspruinosella Thehistoricalpatternsofevolutionofparasitoids
associated with the host plant V. ulz'ginosum are more important determinants than current
252 Lozan etal. ° ENTOMOL. FENNICAVol. 21
ecologicalcharacteristicsinan ‘isolatedhabitat’, Askew,R.R. &Shaw,M. 1986:Parasitoidcommunities:
whichreflects acertaindegree ofenvironmental theirsize,structureanddevelopment.7In:Waage,J.
& Greathead, D. (eds), InsectParasitoids: 2257264.
tolerance (cf Hawkins 1994, Hawkins & Mills
AcademicPress, SanDiego. 389pp.
1996,Tschamtkeetal.2002).Itmayalsosuggest Aubert,J.F. 1978:LesIchneumonidesouest—palearctiques
thatany singleperturbationinthepeatbogenvi- etleurshotes2.BanchinaeetSuppl.auxPimplinae.7
ronment might shift the host-parasitoid evolu- Laboratoired’EvolutiondesEtresOrganises,Paris&
tionaryrelationshipsnotinfavourofLepidoptera EDIFAT-OPIDA,Echauffour. 318pp.
Belokobylskij,S.A.&Tobias,V.I.1998:FamilyBraconi—
hosts which seem to be much more habitat de- dae.7In:Lehr,P.A.(ed.),KeyoftheInsectofRussi—
pendent (often cold—adapted “relict” taxa) than anFarEast(Neuropteroidea,Mecoptera,Hymenopte—
their opportunistic parasitoids (Spitzer & Danks ra) 4(3). Dal’nauka, Vladivostok. 708 pp. [In Rus—
2006). All parasitoids are therefore only eury- sian.]
topic taxa when compared with the steno- Belokobylskij S.A.&TobiasV.I.2000:FamilyBraconi—
topic+eurytopic combination of leaf-spinning dae.7In:LehrP.A.(ed.),KeyoftheInsectofRussi—
anFarEast(Neuropteroidea,Mecoptera,Hymenopte—
Lepidoptera hosts. From the conservation point ra) 4(4). Dal’nauka, Vladivostok. 651 pp. [In Rus—
ofview,thecomplexoftheisolatedpeatboghab- sian.]
itats, hydrological conditions and the very spe- Bezdék,A.,Jaros,J. &Spitzer,K. 2006: Spatialdistribu—
cificlocalmicro-andmesoclimatemightprevent tion ofground beetles (Coleoptera: Carabidae) and
moths (Lepidoptera) in the Mrtvy luh bog, Sumava
the potential impact ofthe population outbreak
Mts(CentralEurope): atestofhabitatislandcommu-
or/and invasion of common opportunistic nity.7BiodiversityandConservation 15: 3957109.
parasitoids unless human and climatic impacts Billups, T. R. 1897: Additionalnotes onthehymenopte-
areexcluded. rousanddipterousparasites,bredbymembersofthe
SouthLondonEntomologicalandNaturalHistorySo-
Acknowledgements. Fieldandlaboratorystudies(rearing, cietyduringtheyears 1891 and1892.7Proceedings
oftheSouthLondonEntomologicalandNaturalHis-
identification, habitat comparative analysis) were sup—
torySociety 1896: 80787.
ported by the Czech Academy of Sciences (Grant
Boucek,Z. &Askew,R.R. 1968:PalaearcticEulophidae
1QS500070505)andpartiallybytheEUSynthesys(grant
sineTetrastichinae.IndexofEntomophagousInsects.
GB-TAF-4159) and European Science Foundation
(BEPARGrantNr.1667).WethankD.R.Kasparyan,V.I. 7In:Delucchi,V.&Remaudicre,G.(eds),Indexof
TobiasandS.A.Belokobylskij(ZoologicalInstitute,Saint EntomophagousInsects3.LeFrancois,Paris.254pp.
Boucek,Z.&RasplusJ.Y. 1991:IllustratedKeytoWest—
Petersburg)foridentificationofsomeIchneumonidaeand
Palaearctic Genera of Pteromalidae, Hymenoptera
allowance ofBraconidae collection; G. Broad (Natural
HistoryMuseum,London)andC.vanAchterberg(Natio— Chalcidoidea.7INRAEditions,Paris. 140pp.
Dasch, C. E. 1988: Ichneumon—flies ofAmericanorthof
naalNatuurhistorischMuseum, Leiden) forthe accessto
the Braconidae collections; our technical assistant H. Mexico: 10. SubfamilyBanchinae, tribeGlyptini.7
Zikmundova (Biology Centre, Ceske’ Budéjovice) who MemoirsoftheAmericanEntomologicalInstitute43:
17644.
helpeduswithlaboratoryrearingofbothLepidopteraand
Diaconu,A. &Lozan,A. 2000: Ectoparasitoidbraconids
Hymenoptera.WearethankfultoMarkShawforhishelp-
(Hym.: Braconidae) ofcertain species ofleafrollers
fill comments andvaluable suggestions on early draft of
theMS.Lastbutnotleastwearegratefultothetwoanony- (Lep.: Tortricidae) offruittrees. 7Mitteilungender
Schweizerischen Entomologischen Gesellschaft 73:
mousreviewersfortheirusefulcomments.
5711.
Gauld,I.D.2002:TheIchneumonidaeofCostaRica,4.7
MemoirsoftheAmericanEntomologicalInstitute66:
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