Morphological and molecular taxonomy of Dendrolimus sibiricus Chetverikov stat.rev. and allied lappet moths (Lepidoptera: Lasiocampidae), with description of a new species

2008·2.4 MB·

by  Kauri Mikkola

©EntomologicaFennica.13June2008
Morphological and molecular taxonomy
of Dendrolimus sibiricus Chetverikov stat.rev.
and allied lappet moths (Lepidoptera: Lasiocampidae),
with description of a new species
KauriMikkola&GunillaStåhls
Mikkola,K.&Ståhls,G.2008:MorphologicalandmoleculartaxonomyofDen-
drolimus sibiricus Chetverikov stat.rev. and allied lappet moths (Lepidoptera:
Lasiocampidae),withdescriptionofanewspecies.—Entomol.Fennica19:65–
85.
The populations of the well-known forest pest, Dendrolimus sibiricus Chet-
verikov,1908stat.rev.,weresampledintheEuropeanfoothillsoftheUralMoun-
tains,Russia.D.sibiricusisaspeciesdistinctfromtheJapanesetaxonD.supe-
rans(Butler,1877).AnothertaxonfromtheSouthernUrals,taxonomicallyclose
toD.pini(Linnaeus),isdescribedhereasD.kilmezsp.n.Thesyntheticfemale
pheromonespreparedforD.piniandD.sibiricusattractedequallywellallthree
taxapresent,andthuscannotbeusedtoidentifythesespecies.TheUralpopula-
tionsofD.sibiricusshowdifferencesinexternalappearance,andasalreadyin
the1840sEversmannindicatedthatthespecieshadcausedlocalforestdamage,
D. sibiricusmust bealong-established species in theUral area. Thus, natural
spreadingwestwardofthepestisnottobeexpected.ThefiveDendrolimusspe-
ciesofthenorthernPalaearcticandthemalegenitaliaareillustrated,andthedis-
tinguishingcharactersarelisted.TwoMatsumuralectotypesaredesignated.
K.Mikkola,FinnishMuseumofNaturalHistory,P.O.Box17,FI-00014Univer-
sityofHelsinki,Finland;E-mail:[email protected]
G.Ståhls,FinnishMuseumofNaturalHistory,P.O.Box17,FI-00014University
ofHelsinki,Finland;E-mail:[email protected]
Received8January2008,accepted26April2008
1.Introduction years,speciesofDendrolimushavebeenparticu-
larly serious pine forest pests also in China
InRussia,in1977–1998,theannualforestdam- (Zhangetal.2004).
age caused by the caterpillars of the Siberian Chetverikov (1903) and Petersen (1905) re-
pine-tree lappet or Siberian silk moth, Dendro- ported damage caused by the caterpillars of D.
limus sibiricus Chetverikov, 1908, averaged sibiricus in the southeastern parts of the Ural
150,000hectares(variation5,600to574,800ha; Mountains.They notedthatthespecieshas,ac-
Gninenko2000;cf.Kharuketal.2004).Thefo- cordingtotheforesters,invadedtheUralsfrom
restlossesthatoccurredintheearly1900’sonthe theeast. Petersen forecasted it to spread “in the
islandofSakhalinweredescribedasspectacular nearestfuture”intoEurope.Thisviewofrecent
(Matsumura 1925, Kuznetsov 2006). In recent spreadingfromtheeastandexpectedcontinued
66 Mikkola&Ståhls (cid:127) ENTOMOL.FENNICAVol.19
spreading westward is maintained by several lustratethespecies. Usefulmorphologicalchar-
Russianauthors(e.g.Gninenko2000,Orlinskiy actersdistinguishingtheEuropeanpine-treelap-
2000, Konefal & Karnkowski 2003). However, petD.piniandtheSiberiantaxonhave,asfaras
Eversmann (1844) indicated that the pest was weknow,neverbeenpublished.
present in the Ural area around 60 years before ThedistributionofD.sibiricusontheEuro-
the notes of Chetverikov and Petersen (see be- peansideoftheUralMts.isparticularlypoorly
low). Gninenko & Orlinskiy (2002) report the documented. The scientific collections of the
westernmost occurrences as being in western MGU(MoscowStateUniversity)andZIN(Zoo-
Udmurtia, where forest damage occurred in the logicalInstituteofRussianAcademyofScience,
late1950s.Morerecentdefoliationhasbeenob- StPetersburg)toourknowledgedonotholdany
servedinPerm(A.N.Omysheva,oralcomm.). materialsofDendrolimusfromtheUralarea. A
ThecloselyrelatedJapanesespeciesD.supe- field guide to the European “Bombyces” (Rou-
rans (Butler, 1877) is also a pest on coniferous geot&Viette1980)andahandbooktreating,e.g.,
trees (Maeto 1991). The damage caused by the the Western Palaearctic Lasiocampidae (de
European pine-tree lappet D. pini (Linnaeus, Freina&Witt1987)donotmentionthetaxonat
1758)islessdramatic(e.g.Möller2005).Ebert all. A handbook of forest pests (Wellenstein
(1994) gave only historical data about defolia- 1978) expressly notes that the species does not
tions by this species in SW Germany. Saalas occurinEurope.
(1949) mentioned forest damage in Norway in Gninenko & Orlinskiy (2002) claim wide-
1812–1816and1902,andSwedenin1938–1940, spreadoccurrenceofD.sibiricusintheRepublic
but not at all in Finland. Wellenstein (1978) re- ofMariEl,betweenMoscowandtheUralMts.
ported no forest damage in Europe since 1947– Thiscouldnotbeconfirmedbythefieldcollec-
1948. tionsmadeinMariElbytheauthorKMin1997
Themostrecenttaxonomicrevisionofthege- norbytheFinnishmonitoringscheme“Noctur-
nusDendrolimusisbyLajonquière(1973),who na”(cf.Nieminen1996)in1998and1999.The
recognized12species,allfromthePalaearcticre- claimabouttheoccurrenceoftheSiberiantaxon
gion.Mosttaxonomistshaveconsidered,follow- inMariElandintheMoscowandTverregions
ingLajonquière,theSiberiantaxonasasubspe- (Gninenko&Orlinskiy2002)maybebasedonan
ciesoftheJapanesetaxonD.superans,andused erroneous concept of species specificity of the
thecombinationDendrolimussuperanssibiricus synthetic female pheromones used to attract
(e.g. Dubatolov & Zolotuhin 1992, Zolotuhin males(cf.Pletnievetal.1999).
1995, Chistyakov 1999). This combination was To facilitate determination of the quarantine
also adopted by a Russian handbook on forest statusofD.sibiricusbytheEuropeanandMedi-
pestdynamics(Isayevetal.2001:290),bysev- terraneanPlantProtectionOrganization(EPPO),
eral Russian researchers (Gninenko 2000, Kiri- anexpeditiontothelowfoothillsoftheUralMts.,
chenko&Baranchikov2004)andby theCana- toPermandUdmurtia,wasfundedbytheFinnish
dianFoodInspectionAgency(2005).However, Plant Protection Department (KTTK). The trip
theCABI(1996)listandtheEuropeanPlantPro- wasmadeincooperationwiththeRussianquar-
tection Organization (EPPO 2002, 2005) have antineauthoritiesoftheseregions.
followed“themaininternationalopinion”(!)and Thefieldcollectionsalsoaimedatobtaining
use the name D. sibiricus Chetverikov, as do specimens of Dendrolimus for molecular work.
some Russian scientists (e.g. Gninenko & PreviousDNAstudiesonDendrolimusspp.con-
Orlinskiy 2002) and some other forest pest re- cernmicrosatellitevariationofeightChinesespe-
searchers(e.g.Øklandetal.2004).Thus,thereis ciesandsubspecies(Zhangetal.2004),andare
“significantdisagreementaboutthetrueidentity notinformativeforthepresentstudy.MtDNAis
ofthespeciesanditsproperscientificname”(Da- particularly usefulfor species-levelanalyses, as
visetal.2005). demonstratedinmanystudiesofinsectevolution-
The‘UralMts.’isoneofthetypelocalitiesof ary relationships. We employed mitochondrial
D.sibiricus(Chetverikov1908),assoonnotedby COI(cytochromecoxidasesubunitI)sequences
Grünberg(1911).Gaede(1932)wasthefirsttoil- ofthe3’and5’regions(fromourlaboratoryand
ENTOMOL.FENNICAVol.19 (cid:127) TaxonomyofD.sibiricusandalliedmoths 67
fromtheBarcodingofLifelaboratoryinCanada,
respectively) in conjunction with DNA charac-
tersfromthenuclearITS2(secondinternaltran-
scribedspacer)generegion.
Theaimofthepresentstudyisto1)stabilize
the nomenclature of the Siberian taxon, 2) de-
scribe distinguishing morphological characters
forallfiveDendrolimusspeciesthatoccurinthe
northernlatitudesofthePalaearcticregion,and3)
useDNAsequencedatatoaugmentthemorpho-
logicaltaxonomy.Thus,weemploytheconcept
ofintegrativetaxonomybyusingtraditionalmor-
phologicalcharactersinadditiontomoderntech-
niques of eversion of internal genitalia (cf.
Mikkola 2007, 2008), and by combining DNA
datafrommultiplesourceswiththemorphologi-
calmaterialandinterpretingtheproduceddatain
lightofavailableinformationaboutspeciesdistri-
butions.
2.Materialandmethods
Fig.1.SampledlocalitiesinRussiaandFinland.
2.1.Collections
On1.–15.VII.2002,samplesofD.sibiricuswere mainlyforsequencing.Wealsosequencedacat-
collectedfromtwoconiferousforests,inPerm(at erpillarofD.sibiricusdonatedbyB.I.Kovalev.
60°30’N 56°20’E, ca. 50 km NW of Cherdyn) Comparativesamplesofmothswereobtained
andabout500kmSWofCherdyn,inUdmurtia from Tammisaari, Finland (KM, D. pini), from
(at57°00’N51°05’E,outskirtsofKilmez,cf.Fig. Mari El, Mariy Chodra (KM, D. pini and sp.),
1),bothsitesinthelowland-likewesternfoothills from Miass, Chelyabinsk region, southeastern
of the Ural Mountains. In both forests, Abies Uralarea(K.NupponenandM.Ahola,D.sibiri-
sibiricagrewtogetherwithPiceaabiesandPinus cus,piniandsp.),fromPrimorye,theRussianFar
silvestris. East(V.S.Kononenko,D.sibiricus)andfromJa-
Thefollowingcollectingmethodswereused: pan(severalsites;M.Owadaandassociates,D.
(1)twoportablelighttrapsequippedwith125W superans and spectabilis) as well as from mu-
mercuryvapourbulbs,runbya1kWHondagen- seum materials collected earlier in Siberia
erator,and(2)femalepheromonesforD.piniand [mainly Altai Mts. and Buryatia; Finnish Mu-
D.sibiricus,preparedintheNetherlands(Phero- seum of Natural History (MZH), D. sibiricus],
bank)andinRussia(cf.Pletnievetal.1999,Klun Allspecimensintheillustrationsare,ifnotother-
etal.2000),respectively.Theformerwereused wiseindicated,fromtheMZH.
incommercialgluetraps,thelatterinFinnish-de-
signedbaittraps.
By running light traps, 10 and 21 males of 2.2.Morphologicalstudies
Dendrolimus were collected at Cherdyn (D.
sibiricus) and Kilmez(D. sibiricusand sp.), re- Themorphologicaldistinctionofthetaxaisbased
spectively.Inaddition,thepheromonetrapscol- onmalegenitalia;onlysinglefemalesofD.pini
lectedseveraldozenmales.Becauseoftheglue and D. sibiricus were dissected. Microscopic
and erratic flying in the bait traps, these speci- genitalia slides were prepared by conventional
mens were mostly in weak condition and used methods, but the male vesicae and the female
68 Mikkola&Ståhls (cid:127) ENTOMOL.FENNICAVol.19
Fig2.ThemalegenitaliaoffivespeciesofDendrolimus,totheleft,theopenedexternalgenitaliaand,tothe
right,theevertedvesica,photographedinvitro.–a.D.pini,showingvesicaalsoinalateralview(lowerfigure).
–b.D.kilmezsp.n.–c.D.sibiricus.–d.D.superans.–e.D.spectabilis.
bursae were first stained using chlorazol black 2.3.Molecularmethods
and then everted using isopropyl alcohol injec-
tionswithasyringe.Anumberofvesicaethatare Wefirstproducednucleotidesequencesofthe3’
highlythree-dimensionalwere,beforemounting fragment of mitochondrial COI and the nuclear
them on slides, photographed in vitro (Fig. 2). ITS2generegionsfromspecimensthat,basedon
When molecular and morphological differences externalmorphology,wereidentifiedasD.pini,
betweenD.sibiricusandD.superansaswellas D.sibiricusandD.superans,andfromspecimens
between D. pini and D. sp. had been detected, ofD.spectabilisandLasiocampatrifolii(Denis
moremalesofeachtaxonweredissectedandse- &Schiffermüller)asoutgroups.Forcomparison,
quenced;theobserveddifferencesremainedcon- P.Hebert(Guelph,Canada)kindlyprovidedse-
stant. To construct a Hennigian argumentation quencesoftheCOI–5’fragmentgeneratedfrom
scheme,thekeysmustbeconsulted. fiveofoursequencedspecimens, andfromfive
additionalspecimensprovidedbyus.
DNAwasextractedusuallyfrom2–3legsof
ENTOMOL.FENNICAVol.19 (cid:127) TaxonomyofD.sibiricusandalliedmoths 69
specimenspreservedin70–95%alcoholorfrom kilmezsp.n.,andforfourspecimensofD.pini.As
relativelyfreshpinnedmuseumspecimens(Table wehadonlyfivespecimensfromwhichbothCOI
1).DNAwasextractedusingtheNucleospinTis- sequence fragments were achieved, we had no
sue Kit (Machery-Nagel, Düren, Germany) ac- reasontocombinethedatasets.
cordingtomanufacturer’sprotocols. Themoths Our pruned data matrix of the ITS2 region
werepreservedasDNAvoucherspecimensinthe comprised 31 ingroup specimens and four
ZoologicalMuseum(MZH)oftheFinnishMu- outgroupspecimenswithsequencelengthvaria-
seumofNaturalHistory(FMNH),Helsinki,Fin- tion of 473–478 nt among ingroup taxa. The
land. DNAvoucher specimens carry labels, in- alignedITS2sequencematrixcomprised484nt
cludinglabcodesandlocalityinformation,asin- characters.
dicatedinTable1.
PCR and sequencing procedures followed
those described in Mengual et al. (2006). The 2.4.Parsimonyanalyses
primersusedforamplifyingandsequencingfor
the COI were C1-J-2183 (alias Jerry, 5’- Analyses of the ITS2, COI-3’ and COI-5’ se-
CAACATTTATTTTGATTTTTTGG-3’) and quencematriceswereconductedusingtheparsi-
tl2-n-3014(aliasPat,5’-TCCAATGCACTAAT- mony program NONA v. 2.0 (Goloboff 1998)
CTGCCATATTA-3’)(Simonetal.1994)andfor spawn from Winclada (Nixon 1999), with the
theITS2fragmentITS2A5’-TGTGAACTGCA- command line “hold 100000; hold/50, mult*
GGACACAT-3’andITS2B5’-TATGCTTAAA- 1000; max*”. All characters were equally
TTCAGGGGGT-3’(Beebe & Saul 1995). The weighted,andgapsweretreatedasfifthcharacter
sequenceswereeditedandassembledusingSe- statesintheITS2analysis.Toillustrateandde-
quenceNavigator™(version1.01). scribe COI haplotype diversity among included
InapreliminaryparsimonyanalysisofITS2 specimens, sequences were manually compared
sequences, we used Dendrolimus punctatus and variable nucleotide positions were docu-
(Walker,1855)(China;datafromGenBank)asan mented.
outgroup,butsinceD.spectabilis(Japan;EMBL
accessionnumberAJ577260)asanoutgrouppro-
duced an identical topology, only the latter is 2.5.Pheromonestudies
shown.WecouldnotamplifytheITS2fragment
forLasiocampatrifolii(SWFinland)ortheCOI- Experimentsusingfemalepheromonesmanufac-
3’fragment for D. spectabilis. Hence, we were tured for D. pini and D. sibiricus (The Nether-
forcedtoconstructthecladogramsusingdifferent lands,Pherobank,andKrasnoyarsk,seeKlunet
outgroups,andwedidnotcombineITS2andCOI al. 2000, respectively) were conducted in the
sequences.TheCOI-5’fragmentcouldbeampli- fieldinPermandUdmurtia,Russia,andatTam-
fied for D. spectabilis apparently due to suffi- misaari,Finland.
ciently universally conservative primers, as op-
posedtotheprimersitesoftheCOI-3’fragment
(Hebert & Lafontaine, pers. comm.). Thus, D. 3.Results
spectabiliswasusedasoutgroupintheITS2ma-
trixandintheCOI-5’dataset,andL.trifoliiinthe Our results confirmthe occurrence of the Sibe-
COI-3’dataset. riantaxonD.sibiricusontheEuropeansideofthe
TheCOI-3’fragmentcomprised729nucleo- UralMts.,andwere-examineitshistorythereas
tides for 29 ingroup specimens (see Table 1 for wellasseparateittaxonomicallyfromtheJapa-
GenBank accession numbers), and we obtained neseD.superans,anddescribeanewsisterspe-
the COI-5’sequences of 658 nt for 10 ingroup ciesofD.pinifromthesouthernpartsoftheUral
specimens.Thelatterwereobtainedfortwospec- Mts.
imens each of D. sibiricus, D. superans and D.
70 Mikkola&Ståhls (cid:127) ENTOMOL.FENNICAVol.19
Table1.ListofDendrolimusandLasiocampatrifoliispecimensusedformolecularstudy,includingGenBank
(GB)accessionnumbers.
Labcode Taxon Locality GBacc.COI
Date,Collector GBacc.ITS2
MZH_KM1 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn, AM946694
5.–6.VII.2002,K.Mikkolaleg. AM946744
MZH_KM2 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn AM946695
5.–6.VII.2002,K.Mikkolaleg. AM946745
MZH_KM3 D.sibiricusChetverikov,1908 Russia,Perm,Solikamsk, AM946696
Kovalevleg.,2002 AM946746
MZH_KM4 D.kilmezsp.n. Russia,Udmurtia,Kilmez, AM946707
8.VII.2002,K.Mikkolaleg. AM946727
MZH_KM5 D.sibiricusChetverikov,1908 Russia,Udmurtia,Kilmez, AM946697
8.VII.2002,K.Mikkolaleg. AM946756
MZH_KM6 D.sibiricusChetverikov,1908 Russia,Udmurtia,Kilmez, AM946698
8.VII.2002,K.Mikkolaleg. ---
MZH_KM7 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn, ---
4.VII.2002,K.Mikkolaleg. AM946747
MZH_KM8 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn, ---
4.VII.2002,K.Mikkolaleg. AM946748
MZH_KM9 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn, AM946699
4.VII.2002,K.Mikkolaleg. ---
MZH_KM12 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn, ---
6.VII.2002,K.Mikkolaleg. AM946749
MZH_KM13 D.sibiricusChetverikov,1908 Russia,Perm,Cherdyn, ---
6.VII.2002,K.Mikkolaleg. AM946750
MZH_KM19 D.sibiricusChetverikov,1908 Russia,Udmurtia,Kilmez, AM946701
9.VII.2002,K.Mikkolaleg. AM946751
MZH_KM20 D.sibiricusChetverikov,1908 Russia,Udmurtia,Kilmez, AM946702
9.VII.2002,K.Mikkolaleg. AM946752
MZH_KM21 D.sibiricusChetverikov,1908 Russia,Udmurtia,Kilmez, ---
9.VII.2002,K.Mikkolaleg. AM946753
MZH_KM23 D.kilmezsp.n. Russia,Udmurtia,Kilmez, AM946705
9.VII.2002,K.Mikkolaleg. AM946728
MZH_KM24 D.kilmezsp.n. Russia,Udmurtia,Kilmez, AM946706
9.VII.2002,K.Mikkolaleg. AM946729
MZH_KM25 D.pini(Linnaeus,1758) Finland,N:Tammisaari, ---
Gullö,15.–17.VII.2002, AM946723
K.Mikkolaleg.
MZH_KM27 D.pini(Linnaeus,1758) Finland,N:Tammisaari, AM946708
Gullö,15.-17.VII.2002, AM946724
K.Mikkolaleg.
MZH_KM28 D.pini(Linnaeus,1758) Finland,N:Tammisaari, AM946709
Gullö,15.-17.VII.2002, AM946725
K.Mikkolaleg.
MZH_KM29 D.pini(Linnaeus,1758) Finland,N:Tammisaari, AM946710
Gullö,15.-17.VII.2002, ---
K.Mikkolaleg.
MZH_KM30 D.pini(Linnaeus,1758) Finland,N:Tammisaari, AM946711
Gullö,15.-17.VII.2002, AM946726
K.Mikkolaleg.
MZH_KM31 D.sibiricusChetverikov,1908 Russia,Burjatia,Baikal AM946703
AM946754
MZH_KM32 D.sibiricusChetverikov,1908 Russia,Burjatia,Baikal AM946704
AM946755
MZH_KM41 D.kilmezsp.n. Russia,Udmurtia, AM946693
2002,K.Mikkolaleg. AM946730
ENTOMOL.FENNICAVol.19 (cid:127) TaxonomyofD.sibiricusandalliedmoths 71
Table1continued.
Labcode Taxon Locality GBacc.COI
Date,Collector GBacc.ITS2
MZH_KM42 D.pini(Linnaeus,1758) Russia,Novosibirsk, ---
1.VII.1984 ---
MZH_KM43 D.pini(Linnaeus,1758) Russia,Mari, ---
12.VI.1997 ---
MZH_KM_J1 D.superans(Butler,1877) Japan,Honshu,Nagano, AM946712
Minami-Azumi,Tazawa, AM946735
2.VIII.2003,M.Owadaleg.
MZH_KM_J2 D.superans(Butler,1877) Japan,Honshu,Nagano, AM946713
Minami-Azumi,Tazawa, AM946736
2.VIII.2003,M.Owadaleg.
MZH_KM_J5 D.superans(Butler,1877) Japan,Honshu,Nagano, AM946714
Minami-Azumi,Tazawa, AM946734
2.VIII.2003,M.Owadaleg.
MZH_KM_J6 D.superans(Butler,1877) Japan,Honshu,Nagano, AM946715
Minami-Azumi,Tazawa, AM946731
2.VIII.2003,M.Owadaleg.
MZH_KM_J7 D.superans(Butler,1877) Japan,Honshu,Nagano, AM946716
Minami-Azumi,Tazawa, AM946732
2.VIII.2003,M.Owadaleg.
MZH_KM_J12 D.superans(Butler,1877) Japan,Idenemaizawa,Kawabe, AM946717
AkitaPref., AM946737
1.VIII.2003,A.Sasakileg.
MZH_KM_J13 D.superans(Butler,1877) Japan,Moriyashizawa,Moriyoshi, AM946718
AkitaPref., AM946738
29.VII.2003,M.Tanakaleg.
MZH_KM_J14 D.superans(Butler,1877) Japan,Moriyashizawa,Moriyoshi, AM946719
AkitaPref., AM946739
29.VII.2003,M.Tanakaleg.
MZH_KM_J15 D.superans(Butler,1877) Japan,Moriyashizawa,Moriyoshi, AM946720
AkitaPref., AM946740
29.VII.2003,M.Tanakaleg.
MZH_KM_J27 D.superans(Butler,1877) Japan, AM946721
24.VIII.2003 ---
Outgroups
MZH_KM_J19_KS1 D.spectabilis(Butler,1877) Japan,Kamikawazi,Iida, ---
Nagano,23.VIII.2003, AM946722
K.Shikataleg.
MZH_KM_J21_KS3 D.spectabilis(Butler,1877) Japan,Kamikawazi,Iida, ---
Nagano,23.VIII.2003, AM946743
K.Shikataleg.
MZH_KM_J24 D.spectabilis(Butler,1877) Japan,SagaPref.,Kyushu, ---
Shimotogawa,Ushizu-cho, AM946741
21.VIII.2003,M.Furukawaleg.
MZH_KM_J25 D.spectabilis(Butler,1877) Japan,SagaPref.,Kyushu, ---
Shimotogawa,Ushizu-cho, AM946742
21.VIII.2003,M.Furukawaleg.
MZH_KM38 L.trifolii(Denis& Finland,Ab:Örö, AM946692
Schiffermüller,1775) 31.VII.-10.VIII.2005,lighttrap, ---
J.Kullbergleg.
72 Mikkola&Ståhls (cid:127) ENTOMOL.FENNICAVol.19
Fig.3.MalesofthespeciesofDendrolimusbeingstudied,thecolumnsalsoshowingsomeintra-andinter-
populationvariation.–a–c.DendrolimuspiniFinland:a.Al,Mariehamn;b.Ab,Houtskär;c.N,Tammisaari.
–d–f.D.kilmezsp.n.Russia:d.Udmurtia,Kilmez10.VII.2002(holotype);e–f.Chelyabinskobl.,Miass
28.VI.1997(paratypes).–g–h.D.superans2##Japan,Karuizawa.–i.D.spectabilis1#Japan,Tokyo.
–j–l.D.sibiricusAsiaticRussia:j.SWSiberia,Altai;k.CentralSiberia,Khamar-Daban;l.FarEastRussia,
Primorye.–m–o.D.sibiricus3##Russia,NUrals,Perm,50kmNWCherdyn.–p–r.D.sibiricusRussia,
SUrals:p–q.2##Udmurtia,Kilmez;r.1#Miass.
ENTOMOL.FENNICAVol.19 (cid:127) TaxonomyofD.sibiricusandalliedmoths 73
Fig.4.DarkmalesofDendrolimuspinifromthePonto-Mediterraneanarea:–a.Greece,Arkadia.–b.Ukraine,
Ivano-Frankovskobl.–c.CentralKavkaz,Kabardino-BalkarskN.P.Scaleina=1cm.
3.1.IdentificationofDendrolimusspecies Our work was slowed down by an unexpected
basedonexternalappearanceandon taxonomical observation. What in the field was
morphologyofgenitalia believed to be a unicolorous brownish colour
morph of D. pini, was, based on evidence from
D.spectabilisiseasilyrecognizedonthebasisof DNAsequences of the mitochondrial gene COI
thedisproportionatelyminusculeorlackingdisc- fragments3’and5’,ataxondistinctfromD.pini
alwingspot(Fig.3i).D.superansmaylookmis- (Figs3d–f).Asthisfindingwassupportedbythe
leadinglysimilartosomepolymorphicformsof morphologyofmalegenitalia,whichrevealeda
D.sibiricus,butmostlythemothsofD.superans combination of small morphological characters
are larger and richer in brownish or rusty hues consistentlyseparatingthetaxonfromD.pini,it
(Figs3g–h).D.pini,whichismostlyeasilyrec- isbelowdescribedasanewspecies.
ognizedbyitselegantcontrastbetweenthegrey
background and the nutbrown subterminal field
(Figs.3a–b),sometimesshowsauniformlyash- 3.2.HistoryofDendrolimussibiricus
greybackgroundcolour(cf.Fig.4),andapoly- intheUralarea
morphicrustybrownmorphoccursatalowfre-
quency(Fig.3c).InD.kilmezsp.n.,mostspeci- Thepresentstudydoesnotsupportthepostulated
mensareofthelattermorph(Figs3d–f).Ifsucha recentspreadingofD.sibiricusfromtheeast,and
morphoccursinD.sibiricus(Fig.3l),itmaybe thus,continuedspreadingwestward.Thehabitus
difficulttoseparatefromD.piniandkilmez,but, of the moths from Perm and Udmurtia differ
actually,inD.sibiricusthebrownmorphoccurs clearlyfromeachother,theformerbeingmainly
ataverylowfrequency,andpredominantlyinthe larger(mean±SDwingexpanse62.4±3.4mm,
Eastern Palaearctic (where D. pini does not oc- n=10##,Figs3m–n)andwithverydarkcolor-
cur). ation, the latter mostly small (58.6 ± 3.4 mm,
TheidentificationofspeciesofDendrolimus n=17##:), paler and with a more restlessly
byappearanceiscomplicatedbythewidepoly- irrorated appearance (Figs 3o–p). No molecular
typism and polymorphism of most species. As differencewasdetectedbetweenthepopulations.
mentioned,D.sibiricuslooksdifferentindiffer- The following facts speak against an eastern
ent parts of the Ural Mts. (Figs 3m–p), and the spreadingofthespecies:
UralmothsdifferfrommostmothsfromtheAltai,
Khamar-Daban and Buretian Mountains (Figs (1)Thedifferenthabitusofthepopulationsindi-
3j–k). InD. pini, singlemothsfromtheCentral catesthattheyhavelivedintheseareasovera
EuropeanMountainsaswellasmostmothsfrom longperiod,probablythousandsofyears.
thePonto-Mediterraneanareaarereminiscentof (2) Notwithstanding that Eversmann (1844) did
D.sibiricus(Fig.4). not distinguish D. sibiricus fromD. pini, he
D. sibiricus could be distinguished from D. revealed the presence of D. sibiricus in the
superansbygenitaliacharacters,thatwereillus- Volgo-Uralareaintwoways:
tratedalreadybyLajonquière(1973),butwhich (a) Eversmann(1844)firstpresentedatraditional
he interpreted as effects of climatic differences. descriptionofD.pini,butthen,undervaria-
74 Mikkola&Ståhls (cid:127) ENTOMOL.FENNICAVol.19
Fig.6.ITS2,strictconsensustreeofthreeequallypar-
simonioustreesoffourspeciesofDendrolimusand
Lasiocampatrifoliiasanoutgroup,length41steps,CI
=0.97,RI=0.99.Forexplanations,seeFig.5.
piniisrareorlackinginthoseareasandasit
Fig.5.COI-3’,strictconsensustreeof12equallypar- seldom“destroys forests”, the pest in Spask
simonioustreesoffivespeciesofDendrolimus(one musthavebeenD.sibiricus,asitisstilltoday.
asanoutgroup),length68steps,CI=0.91,RI=0.92.
Eachterminalrepresentsonespecimen.Opencircles
Spreading species do usually not show differ-
indicatenon-uniquechanges,whilefilledcirclesde-
encessuchasthosefoundbetweenpopulationsof
noteuniquechanges.Forthephylogenetictrees,see
D. sibiricus. The recently found note by Evers-
text.
mann(1844)confirmsthatintheSouthUralsthe
tion,hesaid(fromLatin)“allwingsferrugi- specieshascausedforestdamageforatleast60
neous [this fits both D. pini and kilmez; see yearsbeforethenotesofChetverikovandPeter-
above] – or forewings grey, or ashy, with sen.
whitespotandthreedark[transverse]lines”
[intheUralsthisfitsonlyD.sibiricus],and
(b) He wrote that (from Latin) “The larva that 3.3.Pheromones
feeds on Pinus silvestris [probably from the
Latinname]sometimesdestroysforests;soin BothinFinland(onlyD.pini)andinPerm(only
1842 in the Spask [SW Urals] area.” As D. D.superans)aswellasinUdmurtia(D.sibiricus