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Taxonomy, habitat choice and distribution of Kimunpsocus flavonimbatus (Rostock, 1879) comb. n. (Psocodea: ‘Psocoptera’: Psocidae) PDF

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©EntomologicaFennica.16September2011 Taxonomy, habitat choice and distribution of Kimunpsocus flavonimbatus (Rostock, 1879) comb. n. (Psocodea: ‘Psocoptera’: Psocidae) GergelyVárkonyi&CharlesLienhard Várkonyi,G.&Lienhard,C.2011:Taxonomy,habitatchoiceanddistributionof Kimunpsocusflavonimbatus(Rostock,1879)comb.n.(Psocodea:‘Psocoptera’: Psocidae).—Entomol.Fennica22:97–105. ThebarklicePsocusflavonimbatusRostock,1879andPtyctachubsugulensis Günther, 1982 are assigned to the genus Kimunpsocus Yoshizawa, 2009. The maleterminaliaofK.flavonimbatusaredescribedforthefirsttimeandcompared tothecorrespondingstructuresoftheotherknownspeciesofthegenus.Previ- ously only known from the type locality in Estonia and the municipality of KuhmoinFinland,K.flavonimbatusisnowreportedfrom13newsitesandasa newspeciesforfourmunicipalitiesineasternCentralFinland.Analysesofanex- tensivematerialstronglysuggestthatK.flavonimbatusisahabitat-specialistspe- cies,confiningitsoccurrencetopristineandsemi-naturalspruce-dominatedfo- rests.Nonetheless,itsoccurrenceseemstobesporadicevenintheold-growthfo- rests.Foreststructureofasubsetofoccupiedandunoccupiedsitesisdescribed andtheconservationbiologyofK.flavonimbatusdiscussed. G.Várkonyi,FinnishEnvironmentInstitute,FriendshipParkResearchCentre, Lentiirantie 342B, 88900 Kuhmo, Finland; E-mail: gergely.varkonyi @ymparisto.fi C.Lienhard,Muséumd’histoirenaturelle,C.P.6434,CH-1211Genève6,Swit- zerland;E-mail:[email protected] Received6April2011,accepted17May2011 1.Taxonomy 1.1.Introduction Kimunpsocus flavonimbatus (Rostock, 1879) Thisinterestingspecieshasbeenredescribedby comb.n.(Fig.1) Lienhard&Kanervo(2002),basedonnewlydis- covered material from Kuhmo, eastern Central PsocusflavonimbatusRostock,1879:129. Finland.Themaleterminaliacouldnotbeexam- Amphigerontia flavonimbatas [sic!] ined for that redescription, because all males (Rostock). Mühlen 1884: 331 (incorrect spell- available were damaged, lacking the abdomen. ing). Thereforeitwasnotpossibletoassignthespecies Psocidusflavonimbatus(Rostock).Smithers to one of the modern genera of the family 1967:108;Lienhard&Smithers2002:444. Psocidae.Thus,intheWorldcataloguepublished Psocus flavonimbatus Rostock. Lienhard & byLienhard&Smithers(2002),itwasassigned Kanervo2002:59(redescription). tothegenusPsocidusPearman(s.l.),aheteroge- 98 Várkonyi&Lienhard (cid:127) ENTOMOL.FENNICAVol.22 Fig.1.AfemaleKimun- psocusflavonimbatus fromTeerisuo-Lososuo MireReserve,Kuhmo, easternCentralFin- land.Photo:G.Várko- nyi neous group within Psocidae containing many comb. n. for Ptycta chubsugulensis Günther, species,whichcannotbeallocatedtooneofthe 1982. genera defined in this family after the redefini- Wehadnowthechancetoexamineaperfectly tion, in a narrow sense, of Psocus Latreille by preserved male from Finland (Suomussalmi, Pearman(1932). Murhisalo, beating, 71765:36448, 31.vii.2008, Lienhard & Kanervo (2002) suggested that leg.P.Kähkönen),whichwascollectedtogether Psocusflavonimbatuscouldbeclosetooreven withtypicalfemales.AsindicatedbyLienhard& identical with Ptycta chubsugulensis Günther, Kanervo (2002), the specific identification of described from Mongolia (Günther 1982). Un- bothsexesiseasyduetotheircharacteristicfore- fortunately,asonlythemaleholotypeisknownof wing pattern. The general morphology of both thisspecies,andnointactmaleswereavailableof sexes and the female terminalia have been de- Psocusflavonimbatus,itwasimpossibletomake scribedindetailbytheseauthors.Themaletermi- anydetailedcomparisonofthesespecies. naliaareheredescribedforthefirsttime,basedon ThemorphologyofmaleterminaliaofPtycta the above mentioned specimen. The following chubsugulensisindicatesthecloserelationshipof measurementsofthismalecompletethedataal- thisspecieswithKimunpsocustakumaiYoshiza- ready given in the redescription of the species wa, recently described from northern Japan (Lienhard&Kanervo2002):bodylength(BL)= (HokkaidoIsland)andconsideredbyYoshizawa 3.2mm;forewinglength(FW)=5.0mm;length (2009)asthetypespeciesofanewmonotypicge- ofhindfemur(F)=0.95mm;lengthofhindtibia nus,KimunpsocusYoshizawa.Thereforewepro- (T)=1.85mm;lengthofhindtarsomere1(t1)= pose here the following new combination: Ki- 600 µm; length of hind tarsomere 2 (t2) = 175 munpsocus chubsugulensis (Günther, 1982) µm;minimumdistancebetweencompoundeyes ENTOMOL.FENNICAVol.22 (cid:127) ThebarklouseKimunpsocusflavonimbatus 99 Fig.2.Kimunpsocus flavonimbatus(Ro- stock),male.–a.Ter- minalia(lateralview). –b.Hypandrium,apical partofmedianstrap (anteriorview). –c.Hypandrium(pos- teriorview). dividedbytheantero-posteriordiameterofcom- slightly clubbed seta near its anterior base (see poundeye,indorsalview(IO/D)=1.2.Thechar- detail of Fig. 3a), trichobothrial field oval, not actersofmaleterminaliaconfirmthecloserela- subdivided. Hypandrium (Fig. 2a–c) with a tionshipofthisspecieswithKimunpsocuschub- slightly asymmetrical, strongly sclerotized me- sugulensiscomb.n.butjustifytheseparationof dianstrapandwithapartlymembranous,partly the European and the Mongolian forms as two sclerotizedroundedlaterallobeoneachside.The distinctspecies.Thus,thisEuropeanspecieshas apical part of the median strap antero-ventrally alsotobeassignedtothegenusKimunpsocus. curved and asymmetrically trilobate, one of its terminallobesroundedanddenticulate(Fig.2b– c). Phallosome (Fig. 3c) distally slightly asym- 1.2.Descriptionofmaleterminalia metrical, with a toe-shaped smooth submedian (Figs.2a–c,3a–d) process,oneachsideindistalhalfwithasclero- tizedventro-laterallongitudinalband. Cluniumcharacteristicallyshaped,anteriormar- gin with a deep medio-dorsal notch (Fig. 3d), postero-lateralregionslightly concaveinlateral 1.3.Discussion view (Fig. 2a). Epiproct chair-shaped in lateral view (Fig. 2a), its basal lobe broad and rugose, Kimunpsocusflavonimbatus(Rostock)isclosely slightlyprominentmedially(Fig.3b).Paraproct related to the two other members of the genus (Fig. 3a) lacking laterally projecting basal lobe, KimunpsocusYoshizawa,theMongolianspecies withapointedapicalhookandawell-differenti- K.chubsugulensis(Günther)comb.n.(=Ptycta ated subapical prominence, the latter bearing a chubsugulensisGünther,seeSection1.1.)andthe 100 Várkonyi&Lienhard (cid:127) ENTOMOL.FENNICAVol.22 Fig.3.Kimunpsocus flavonimbatus(Ro- stock),male.–a.Left paraproct.–b.Epiproct (posteriorview). –c.Phallosome(ven- tralview).–d.Clunium (dorsalview,spread out). JapaneseK.takumaiYoshizawa.Maleterminalia tinctly trilobate epiproct lobe (see Yoshizawa areverysimilarinallthreespecies,inparticular 2009:Fig.2B).Themedianstrapofthehypan- their characteristically shaped phallosomes are driumofK.takumaiisprobablyverysimilartoK. almostidentical(hereshowninventralview,see flavonimbatus(accordingtothesomewhatsche- Fig.3c;sameviewinFig.2EbyYoshizawa2009; maticFig.2DinYoshizawa2009). depictedindorsalviewbyGünther1982:Fig.5). Theforewingpatternisessentiallythesamein However,themaleofK.chubsugulensiscanbe allthreespecies,butofvaryingintensity(proba- distinguishedfromthemaleofK.flavonimbatus blyalsowithconsiderableintraspecificvariabil- bytheshapeoftheantero-ventrallycurvedapical ity): highly contrasted pattern as depicted by partofthemedianstrapofthehypandrium:itis Lienhard & Kanervo (2002: Fig. 1a, b) for K. simpleinK.chubsugulensis(seeGünther1982: flavonimbatus; medium contrast in Yoshizawa Fig.4)andtrilobateinK.flavonimbatus(Fig.2b– (2009:Fig.1)forK.takumai;weaklycontrasted c).Thisstrikingdifferencecouldbeconfirmedby pattern in Günther (1982: Fig. 2) for K. chub- theexaminationoftheholotypeofPtyctachub- sugulensis.Thefemaleofthelatterspeciesisnot sugulensis (mounted on three microscopical known,inthetwootherspeciesthemorphology slidesbyK.K.Günther,depositedintheZoologi- offemaleterminaliaisverysimilar(Lienhard& calInstituteoftheRussianAcademyofSciences, Kanervo2002:Fig.1g,i;Yoshizawa2009:Fig. St.Petersburg),whiletheremainingterminaliaof 3A, B). However, Lienhard & Kanervo (2002: this specimen (i.e. clunium, epiproct, paraproct, Fig.1h)illustrateaweaklysclerotized,asymmet- phallosome)arethesameasdescribedabovefor rically structured spermapore region (internal K. flavonimbatus. The JapaneseK. takumai can plate)forK.flavonimbatus;inK.takumaithein- be distinguished from these species by its dis- ternalplateis“withoutobvioussclerotizationnor ENTOMOL.FENNICAVol.22 (cid:127) ThebarklouseKimunpsocusflavonimbatus 101 Fig.4.DistributionofKimunpsocusflavonimbatusineasternCentralFinland.Thesmallsquaresymbolsdenote the1-km2squareswherethespecieshasbeenrecordedduring1997–2008.Thinblacklinesdelineatemunici- palities,ofwhichKuhmoisshowninitsentirety.Thetwomostcentralsquares,isolatedfromlargesemi-natural forestareas,denoteremnantpopulationsofK.flavonimbatusthathavegoneextinctduringthepastdecadedue toforestrymeasures. pigmentation” (Yoshizawa 2009: 153). In this East of Ussuriysk, Kamenushka village, contextitmaybeofinteresttomentionasingle 19.vi.1992, leg. S. Kurbatov), deposited in the female from Eastern Russia (Primorski Kray, collection of the Geneva Natural History Mu- 102 Várkonyi&Lienhard (cid:127) ENTOMOL.FENNICAVol.22 seum and tentatively assigned to K. 27.vii.–11.viii.2006 and 18.vii.–14.viii.2008. flavonimbatusbecauseofthepresenceofahighly Beatingwasperformedinearly morningswhen contrasted wing pattern and of the above men- theambienttemperaturewaslowenoughtopre- tioned asymmetrical structure of the internal ventbarklicefromflyingawayfromthebeating plate.Intheabsenceofinformationonthefemale sheetputunderthesprucebranches.Thoughthe of K. chubsugulensis and on details concerning number ofindividuals sampled was recorded in thestructureoftheinternalplateofK.takumai, eachsite,onlytheinformationonthepresenceor this specific identification remains tentative. absenceofthefocalspecieswasusedintheana- However,thegenusKimunpsocusappearstobe lyses. an eastern Palaearctic element, reaching north- Thestudyareaincludedtheentiremunicipal- easternEuropewithK.flavonimbatus. ity of Kuhmo and adjacent areas as follows: southernpartsofSuomussalmiandHyrynsalmi, eastern parts of Ristijärvi and Sotkamo (region 2.Habitatchoiceanddistribution Kainuu),north-easternValtimo,aswellasnorth- ern parts of Nurmes and Lieksa (region North 2.1.Introduction Karelia)(Fig.4). To address the basic habitat requirements of Kimunpsocusflavonimbatus(Rostock)wasorig- K. flavonimbatus, we first compared its inci- inallydescribedonthebasisofasingleindividual dences in semi-natural vs. managed mature foundnearLakePeipus,easternEstonia,in1878 spruce-dominatedforestsusingallavailabledata, (Rostock1879).Itwasnotuntil1997thatthespe- collected using various methods between 1997 cieswasrediscoveredinKuhmo,easternCentral and2008,attwospatialscales.Atthescaleofin- Finland(seeLienhard&Kanervo2002).During dividualstudysites,beatingsites,siteswithwin- the following decade, a total of 42 specimens dowtraps,asinglesitewithalighttrapandasin- were obtained from semi-natural spruce-domi- glesitewhereK.flavonimbatuswasrearedfroma natedforestsinKuhmo(seeKanervo&Várkonyi sampleofwoodblock,wereconsideredassamp- 2007). Since K. flavonimbatus was never col- ling units (for details see Kanervo & Várkonyi lectedinanyotherhabitatnorinanyotherregion 2007andFig.1within).Atacoarser‘landscape’ in Finland, Kanervo & Várkonyi (2007) sug- scale,a1-km2gridwasputonthemapshowing gested that this species is associated with old- all sampling sites, and sites sharing a grid cell growth forests and restricted to the easternmost were combined as follows: if K. flavonimbatus part of the country. Nonetheless, these authors occurredinanyofthesiteswithinacell,thecell alsoaddressedtheneedforfurtherstudy. was considered as occupied. At both spatial Inthisstudy,ouraimwastoexplorethedistri- scales,ifasitewasrepeatedlysampledindiffer- bution pattern and habitat preferences of K. entyears,asingleoccurrencedetectedinanyyear flavonimbatus by extensive sampling in both rendered the site occupied. The incidence of K. semi-naturalandmanagedmaturespruce-domi- flavonimbatus in semi-natural vs. managed fo- natedhabitats.Thesamplingareawasextended restswasanalysedusingFisher’sexacttest(e.g. toadjacentmunicipalitiesofKuhmointhenorth, Sokal&Rohlf1995).Thistestissuggestedtobe westandsouth.Additionally,weaimedatmea- usedinModelI2×2tablesifthecountnumbers suring and comparing the forest structure of a inanycellarelowerthanfive(Rantaetal.1997). subsetofoccupiedandunoccupiedstudysites. Foreststructurewasmeasuredwithina10-m- radiuscirclearound44studysitesin2008.Dueto the relative rarity of mature managed spruce- 2.2.Materialandmethods dominated forests in the study area, only five studysitesoutofthe44representedthistypeof Kimunpsocusflavonimbatuswassearchedforon habitat,whiletheremaining39sitesweresemi- lowbranchesofNorwayspruce(Piceaabies(L.) natural spruce-dominated forests. However, the Karsten)bythebeatingmethod(e.g.Sutherland rarityofK.flavonimbatus,foundonlyin12outof 1996)duringthepeakflightperiodofthespecies, 44sites,warrantedustocomparetheforeststruc- ENTOMOL.FENNICAVol.22 (cid:127) ThebarklouseKimunpsocusflavonimbatus 103 Table1.HabitatchoiceofKimunpsocusflavonimbatusineasternCentralFinlandwithsignificanceofFisher’s exacttests(2-sided).Forexplanationsofthespatialscalesemployed,seeSection2.2. Spatialscale Semi-naturalsites Managedsites P present absent present absent Individualsites 29 88 0 16 0.022 1-km2gridcells 22 45 0 12 0.017 Table2.Foreststructurein10-m-radiuscirclesaroundoccupiedandunoccupiedsitesofKimunpsocus flavonimbatusineasternCentralFinland.Allvariablevaluesareexpressedasmean±SD. N Numberoflivingtrees Standarea Stumps Spruce Birch Pine Total (m2/ha) Occupied 12 24.5±7.3 2.9±3.1 2.5±5.0 30.2±9.8 28.1±4.8 0±0 Unoccupied 32 23.8±7.0 1.7±1.7 0.5±0.9 26.2±6.7 28.1±4.6 1.7±4.4 tureofoccupiedandapparentlyunoccupiedsites. associated with the semi-natural spruce-domi- In each study site, werecorded thenumber and natedforestsatbothspatialscales(Table1). diameter (measured at 1.3 m height) of living Owing to (i) the general scarcity of mature trunksforeachtreespeciesoccurringinthestudy managed spruce-dominated forests in the study area, i.e. Norway spruce, Silver birch (Betula area,(ii)theapparentrarityofK.flavonimbatus pendula Roth), Scots pine (Pinus sylvestris L.), eveninsemi-naturalsitesand(iii)theriskoffalse Europeanaspen(PopulustremulaL.),Greyalder zerosintheincidencedata,wedidnotattemptto (Alnus incana (L.) Moench) and Goat willow analyse the differences of forest structure be- (SalixcapreaL.).Wealsorecordedthenumberof tweenoccupiedandunoccupiedsites.Instead,we man-madestumpsfoundinthecircularplotsand rather aimed at describing the basic statistics of estimatedthebasalareaofthetreesintheforest themeasuredvariables(Table2).Thestructureof standsusingarelascope. the forest stands surrounding the sampling sites was, on the average, fairly similar in occupied and unoccupied sites. The only notable differ- 2.3.Results enceswerefoundinthenumbersoflivingpines andofman-madestumps. Thesurveysconductedin2006and2008yielded atotalof23K.flavonimbatusspecimens.Kimun- psocusflavonimbatuswasdiscoveredin13new 2.4.Discussion sites (cf. Kanervo & Várkonyi 2007) and re- cordedasanewspeciestothemunicipalitiesof Oneofthemaingoalsofthepresentstudywasto Suomussalmi, Hyrynsalmi and Sotkamo in clarify the basic habitat requirements of K. Kainuu,andtoValtimoinNorthKarelia(Fig.4). flavonimbatus.OurdatastronglysuggestthatK. Atotalof133sites,ofwhich16and117sites flavonimbatusisahabitatspecialistspecies,con- represented managed mature and semi-natural finingitsoccurrencetopristineandsemi-natural spruceforests,respectively,wereincludedinthe spruce-dominated forests. This hypothesis was habitat-choice analyses. Consistent with earlier originally put forward by Kanervo & Várkonyi results(Kanervo&Várkonyi2007),notasingle (2007), but only our extensive survey in both K.flavonimbatuswasfoundinthemanagedsites. (semi-)naturalandmanagedspruceforestsmade TheoccurrenceofK.flavonimbatuswasclearly itpossibletotestthehypothesis. 104 Várkonyi&Lienhard (cid:127) ENTOMOL.FENNICAVol.22 Found in one-fourth (24.8%) of the old- LososuoMireReserve,wheremostFinnishindi- growthsites,K.flavonimbatusseemstoberela- vidualshavebeencollectedfrom.Whileitseems tivelyrareeveninthesemi-naturalforests.Some to be clear that K. flavonimbatus is confined to ofitsobservedraritymayaccountforfalsezeros, pristineandnatural-likespruce-dominatedboreal i.e.undetectedoccurrencesinthesampledsites. forests,itislikelythatfurtherpopulationsineast- Suchsamplingerrormayarisefromthesampling ernmostFinlandandinNorthwestRussiawillbe methodemployed.Beatingcanonlybeusedfor discovered.Nonetheless,giventherapidandex- samplingfromthelowerbranchesoftreesand,on tensivelossandfragmentationofold-growthfo- theotherhand,thereisastochasticriskthatarare rests in this region during the past 60 years, it specieswillnotbesampledduetotheshortdura- seemslikely thattheFinnishpopulationwillbe tionofthesamplingevent.Windowtraps,inturn, decliningeveninprotectedareasduetodelayed were operated for the entire flight season of K. population dynamic responses to habitat loss flavonimbatus, which dramatically lowered the (alsoseeFig.4). risk of undetected occurrences. Nonetheless, In the current Red List of Finnish Species even in south-western Kuhmo, where window (Kanervo & Söderman 2010), the conservation traps were used extensively, K. flavonimbatus status of K. flavonimbatus (as Psocidus flavo- wasonlyfoundin9outof35(Teerisuo-Lososuo nimbatus) was assessed as Near Threatened. MireReserve)and2outof21(Jauhovaara)1-ha Kanervo&Söderman(2010)suggestedthatthe areas, each containing four window traps (see causeofthreatfacingthisspeciesis“reductionof Kanervo&Várkonyi2007).Thereforewecon- old-growthforestsandthedecreasingnumberof clude that the observed rarity K. flavonimbatus large trees” (category Mv). While acknowledg- doesmainlyaccountforarealpatternofsporadic ingthis,westresstheimportanceofunderstand- occurrence,ratherthanforasamplingeffect. ing the underlining population dynamic pro- Thoughthesmall-scalehabitatrequirements cessesforthesuccessfulconservationofthisrare ofK.flavonimbatuscouldbespecifiedusingthe psocidspecies. presentdata(Table2),foreststructurecouldnot beusedtoexplaintheoccurrenceofthespecies. Acknowledgements.Thisworkmuchbenefittedfromour Themajority(32outofthe44)ofthesites,where earliercollaborationwithJussiKanervo(Turku,Finland). WethankDr.VictorA.Krivokhatsky(ZoologicalInstitute, forestmeasurementswereperformed,wasunoc- St.Petersburg,Russia)forarrangingtheloanoftheholo- cupied and only 5 out of these situated in man- typeofPtyctachubsugulensis.WeareindebtedtoPauli agedforests.Thus,mostofbothoccupiedandun- Kähkönen (Friendship Park Research Centre, Kuhmo, occupied sites represented semi-natural forests Finland) who conducted much of the field survey of and,obviously,therewasnoreasontoexpectma- Kimunpsocus flavonimbatus. We thank Mikko Viinanen andJuhaKyllönenfortheirhelpduringfieldwork,and jor differences in the forest structure of those. PirjoAppelgrenforherhelpinpreparingFig.4(allFriend- Nevertheless,wenoticedthatinsomeoftheoc- shipParkResearchCentre).Dr.PetriMartikainen(Univer- cupiedsites,thenumberofpinetreeswasconsid- sityofEasternFinland)kindlysharedwithushisunpub- erablyhigherthanonaverage,andtheirbasalarea lished records of K. flavonimbatus. We also thank two exceeded the basal area of spruce. This is the anonymous referees for their valuable comments on an earlierdraftofthemanuscript. usual case in mesic semi-natural forests, where themaintreeagecohortiscomposedbyspruce andbirch,scatteredwithsomeoldandlargepines References representing an earlier generation of trees. The higheraveragefrequencyofman-madestumpsin Günther,K.K.1982:EinigebemerkenswerteStaubläuse theunoccupiedsitescanbeexplainedbythein- ausderMongolei(Psocoptera:Psocidae).—Deutsche clusionofmanagedsitesinthecomparison. entomologischeZeitschrift29:393–399. Kimunpsocus flavonimbatus was recently Kanervo, J. 2010: Psocidus flavonimbatus (Rostock, discovered in two old-growth forest areas in 1879).—In:TheFinnishExpertGrouponHemiptera (ed.), The Psocoptera of Finland. [www document] Finnish North Karelia (Petri Martikainen, per- URLhttp://www.sci.utu.fi/projects/biologia/elainmu- sonalcommunication;Kanervo2010).Thisarea seo/hemi/psoc/Psocflav.pdf. Updated 15 November issituatedsome100kmsouth-eastofTeerisuo- 2010.(Sitevisitedon11February,2011) ENTOMOL.FENNICAVol.22 (cid:127) ThebarklouseKimunpsocusflavonimbatus 105 Kanervo,J.&Söderman,G.2010:Psocoptera.—In:Ras- Ranta,E.,Rita,H.&Kouki,J.1997:Biometria:tilastotie- si, P., Hyvärinen, E., Juslén, A. & Mannerkoski, I. dettäekologeille.(Biometry:statisticsforecologists.) (eds.), The2010RedListofFinnishSpecies:423– Sixthedition.—Yliopistopaino,Helsinki.569pp.[In 425. Ympäristöministeriö & Suomen ympäristökes- Finnish.] kus,Helsinki.685pp. Rostock, M. 1879: Ueber eine besondere nordrussische Kanervo,J.&Várkonyi,G.2007:OccurrenceofPsocop- Psocus-Art.—EntomologischeNachrichten5:129– tera in boreal old-growth forests. — Entomologica 130. Fennica18:129–137. Smithers,C.N.1967:AcatalogueofthePsocopteraofthe Lienhard,C.&Kanervo,J.2002:RedescriptionofPsocus world.—AustralianZoologist14:1–145. flavonimbatus Rostock (Psocoptera: Psocidae) from Sokal,R.R.&Rohlf,F.J.1995:Biometry:theprinciples Finland.—EntomologicaFennica13:58–62. andpracticeofstatisticsinbiologicalresearch.Third Lienhard,C.&Smithers,C.N.2002:Psocoptera(Insecta): edition.—W.H.FreemanandCompany.887pp. World Catalogue and Bibliography. — Instrumenta Sutherland,W.J.1996:Ecologicalcensustechniques:a Biodiversitatis5:xli+745pp.Muséumd’histoirenatu- handbook.—CambridgeUniversitypress,Cambrid- relle,Genève. ge.336pp. Mühlen, M. vonzur1884:DiePsocidenLiv-, Est-und Yoshizawa,K.2009:DescriptionofKimunpsocustakumai Kurland’s.—SitzungsberichtederNaturforscher-Ge- n. gen. & n. sp. from Hokkaido, Japan (Psocodea: sellschaftbeiderUniversitätDorpat6:329–334. ‘Psocoptera’:Psocidae:Ptyctini).—InsectaMatsu- Pearman,J.V.1932:NotesonthegenusPsocus,withspe- murana,NewSeries65:149–155. cialreferencetotheBritishspecies.—TheEntomolo- gist’sMonthlyMagazine68:193–204.

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