Biogeographical and ecological description of the Odonata of eastern Vasyugan Plain, West Siberia, Russia

2010·2.3 MB·English

by  Bernard, R

Odonatologica39(I):1-28 March I.2010
Biogeographicalandecological description
of theOdonataof easternVasyuganPlain,
West Siberia, Russia
R. Bernard¹andO.E.Kosterin²
1DepartmentofGeneral Zoology,Adam MickiewiczUniversity, Umultowska89,
PO-61-614 Poznan,Poland; - [email protected]
2Institute ofCytologyand Genetics,Siberian Division, RussianAcademyofSciences,
Academician LavrentevAvenue 10,RUS-630090 Novosibirsk, Russia; — [email protected]
Received January2,2008/ReviewedandAcceptedAugust11,2008
Resultsofthestudies ofodonate fauna,carried outin July2006in theodonato-
logicallyalmostunexploredVasyuganPlain,arepresented.The studiesconcentrat-
edinthenorthern and northeastern parts ofthe largestbogin theworld,theVasy-
ugan Bog, and itssurroundings.Large primeval complexesofSphagnumbogsand
fensandotheraccompanyinghabitats,man-made aswell,werestudied.25localities
arebrieflydescribed and theoccurrenceof35recorded spp. iscommented. Due to
almost total absence oftypicallyEast Palaearcticspp. (onlyShaogomphuspostocu-
larisfound),theaspect oftheodon. faunainthestudiedareaissimilar toacertain
degreetothatknown fromcentral and eastern Europe,but with somedifferences
in the species composition,abundance ofmany spp. and their habitat preferences.
InCoenagrionpuella,C.pulchellum,Enallagmarisi, Nehalennia speciosa,Gomphus
vulgatissimus, Shaogomphuspostocularis,Somatochloraflavomaculata,Leucorrhinia
albifrons, and L.pectoralis, thenorthern limit oftheir distribution appears further
N than it waspreviously known. This suggeststhat theactual northern rangelimit
ofsomeofthesespeciesinWSiberia doesnot descendassharplytotheSasitwas
expected.The firstknown site ofS.postocularisWofthe Ob’riverandonaperfect
plainisalsoworthnoticing.Amongthemostremarkable discoverieswastheregular,
area-wide occurrenceofseveralpreviously poorlyknown inSiberia spp.,such asN.
speciosa, Aeshnasubarctica, G. vulgatissimus and S.flavomaculata. Taxonomically
interestingisthecoexistence,inthestudiedareabut notatthesamelocalities,oftwo
taxa considered assubspeciesorseparatespecies,E. c.cyathigerumand E.(c.?) risi.
The factofclearspatialseparationand atmost aminimum degreeofintergrading
(ifany)ofthese2taxasuggeststheir fullspeciesstatuswhichwould agreewith mor-
phologicalandrecent moleculardata. Theregularand notrarepresence of2 andro-
chrome Calopteryxsplendens 9formsand d6 with the wingscoloured tothetips,
aswellasthe occurrenceofbrownish wing‘smoking’ofmany 9 S. flavomaculata
andS.arcticaarepeculiarfeaturesofthe Vasyuganodon. aspect.The 6 segregation
2 R.Bernard& O.E. Kosterin
inthe ‘triangle’ofpeatbogaeshnids,Aeshnacrenata,A. junceaandA.subarctica,IS
describedand discussed.BetweenA.junceaandA.subarctica itwasvery advanced,
partiallyspatialandpartiallyweather/temporal,betweenA. crenataandA.subarctica
almostcomplete, spatial,and between A.crenataand A.junceaadvanced, weather/
temporal.These observationsconfirm thedominant positionofA.crenataoverthe
watertable,andalso suggestthe lackofcrenata-dominance offthewatertable.The
reliabilityofadult diagnosticfeatures,more andless commonlyused to distinguish
between A.subarcticaand A.juncea,isdiscussed.
INTRODUCTION
Theknowledge ofecology, biology,geneticsandthecurrent stateofNehalennia
speciosa inEurope isalreadyfairlyadvanced(BERNARD &WILDERMUTH,
2005a,2005b; BERNARD & SCHMITT,2007). On the contrary, still very lit-
tleisknownonN.speciosa inWesternSiberiawhereitsrecordswereremarkably
scarce (BELYSHEV, 1973; ZAIKA, 1974; KOSTERIN, 1987; KOSTERIN et
al., 2001; BERNARD & WILDERMUTH,2005a) though theoretically, con-
sidering anincomparably weaker humanimpact, theconservation statusofthe
species thereshould be much more favourable than in Europe. To collect sam-
ples of N.speciosa forDNA studiesand to complement the knowledge of the
species with West Siberian data, an expedition to the odonatologically almost
unexplored Vasyugan Plainwas organized. Thechoiceofthe areawas dictated
bythefactthattheVasyugan Plain, extremely abounding inboggy habitatswith
Carexlimosa,seemedtobe very appropriate for thespecies.
Ananalysis ofthe literature(BELYSHEV, 1973) suggested thatdragonflies of
boggy landscapes of WestSiberian Lowland had beenstudied only to a small
extent sofar.Therefore, thesecond, parallel aimofthisexpedition was to study,
inallpossible aspects, the ©donatefaunaoftheareapredominated by primeval
largecomplexes ofSphagnum bogs and fens. Thearticlepresents resultsofthe
studiescarriedoutin thelargest bog intheworldandotheraccompanying habi-
tats intheVasyugan Plain.
THEREGIONSTUDIED
TheWest Siberian Lowland and theAmazon Lowland arethe largest lowlands in the world. A
greatpartoftheformer, which iswithinthetaigazone,issubstantiallyboggy,and thetotalareaoc-
cupiedbybogshasincreased ataspeedof0.8km-ayear.Thelargestbogintheworld,theVasyugan
Bog(53000 km2),occupiesapartoftheWestSiberian Lowland - theveryslightlyelevatedand flat
VasyuganPlain situated betweentheriversofIrtysh andOb’.Itextendsfor 500km fromtheWNW
(approx58°N,75°E)totheSSE(approx55°30’N, 83°E)asastrip 75-175km wide. Itembracesabout
800000lakes accumulatingin total400km-’ ofwaterand isdrained bytheriversoftheTara,Om’,
Vasyugan,Parabel’,Chayaandtheir tributaries.Administratively,thisareaisdividedalongitsmain
axis betweenthe Tomsk(N) and Novosibirsk (S)provinces, with thewesternpartbelongingtothe
Omskprovince. Thismostlypeat-mossbogisnotcontinuous:it hasafairlynarrowaxis ofsublatitu-
dinalorientation andside branches whichlike spursprotrudenorthandsouth fromtheaxis. During
Odonate faunaofeasternVasyuganPlain, Russia 3
ourexpedition,weexploredthe twonortheasternmost ‘spurs’ ofthebog,called Bakcharskoe and
IksinskoeBogs, andthe SambusskoeBogincluded inanorthern spur.
The elevations ofthe VasyuganPlain descendfrom 160to 100m a.s. 1. from S toN and NE
(MESHCHERYAKOV, 1962).The surfaceoftheinterfluvesisflat,with numerousshallow depres-
sions ofthermocarstand suffosionorigin.Thereliefisveryshallow andonlyneartherivers(suchas
theShegarka,Iksa, Bakchar)erosionvalleysofsmallwaterflowsareexpressed.Thesoils areformed
fromsubaerial loess-like sedimentswhich cover, with alayer5-40mthick,theclaysand clayeysedi-
ments oflakeorigin.Hydromorphicsoilsplay agreatrolein thearea,withavariableexpressionofa
relichumushorizon formedinmeadow carbonatesoilsduringtheclimaticoptimumoftheHolocene
(LAPSHINAetal.,2000).Thepeatbogscoveringalargepart oftheVasyuganPlain arenotcon-
nectedwith groundwater and areformed duetotheprevalenceofprecipitation overevaporation
inconditions ofanegligiblewateroutflux onalmostperfectlyflatland surfaces (SOROKINetal.,
1999).Thegrowthofthepeatlayer ledtotheformation ofaninvertedrelief,wheretheformerridges
ofmineralground,bearingforestislands,appearedsurroundedby risingpeat-mossbogs.Atlastthey
becamerelief depressionswithopenSphagnumbogsandfenslocallycalled‘galya’.Inthesebogs,the
waterusuallyflowsthroughthe vegetationandpeat withquiteaconsiderable speed(upto20cm/h).
Thesemiresformsourcesofbrooks providing outfluxofthebogwatersinto rivers(LAPSHINAet
al.,2000).
Accordingtoreconstructions reviewed inLAPSHINAetal.(2000),thebogsofthatareaappeared
firstattheendoftheAtlanticperiodandthebeginningoftheSubboreal time:atplaceswitha3.6-3.8
mthickpeatlayer, 5200-5500 yearsago;inmostareas,witha2.5-3mthickpeatlayer, 3000-3500 years
ago.Atfirsttheywererepresentedby swampswithEquisetum,whichoccupiedthe depressions.Very
soontheyextendedbeyonddepressions,mergedandoccupiedinterfluves,leavingburiedrelicmead-
owsoils. Atthattimethebogschangedtoeutrophicswampbiocenoses with domination ofCarex,
which existedforalongtime,leaving 1-1.8mofpeat.Theimpoverishment ofmineral feedingand
anincreasein precipitationtransformedthem intomesotrophicScheuchzeria-Carex,Scheuchzeria-
Carex-Sphagnumand Carex-Eriophorum-Sphagnumassociations. Atthecrucial momentofdevel-
opmentthebogsenlargedsothatwateroutfluxalongtheslopesofthewatershedsbegan.This ledto
the mesotrophicboggingofdark-coniferous andmixedforestonthegentleslopes oftheVasyugan
Plain,whilethe watershedbogspassed intotheoligotrophicstage ofdevelopment(whichhas per-
sisted untilnow).This occurred attheborder-timebetween the Subboreal and Subatlanticperiods,
between 2000 and 2200yearsago.Atthat time overmoistedSphagnumand Eriophorum-Sphagnum
bogsand pine-S'phagnumassociationsappeared.1500-1000 years agothefruticulose-pinepeat-moss
complexes(locallycalled‘ryam’)spreadand becamedominating.InRussian literaturethese Sphag-
numand pineformationsarecalledridge-openbog/fen(‘gryadovo-topyanye’),ridge-lake-openbog/
fen(‘gryadovo-ozerkovo-topyanye’)and ridge-openquakingbog(‘gryadovo-mochazhinnye’)com-
plexes.IntheseRussiantermsthe“ridges”correspondtopatches(withlowpine)onlyslightlyelevated
abovepeat-mossopenbog.
Out ofthestudied bogs,precisestructural andclimaticdatahave been availableonlyforthe Bak-
charskoeBog;size 10-25 x 150km, peatdepth2.6-3 m,waterpH 3.5-5(LAPSHINAetal.,2000),
meanannual precipitation 420 mm (7mm in June), meansnowcover90mm, average humidityin
thewarmseason30-40% inthedaytimeand 100%atnight(SOROKINetal., 1999).In 1986,some
irrigationwasstartedwith theaim ofpeat digging,but itwasnotsuccessfulandleftonlyamarginal
systemofditches.
There arealso fairlynumeroussecondaryaquatichabitatsin theVasyuganPlain. Theyarerep-
resentedby various,mostlysmall man-made waterbodies (pools andpondsonmineral substrate),
situatedinrare villagesand alongroads andtracks, andbyfire-fightingpools,irrigation ditchesand
canalsin peatyareas.
4 R.Bernard &O.E.Kosterin
LOCALITIES
Thelocalities togetherwith datesofvisits and briefdescriptionsofhabitatsarepresentedbelow;
(1) SambusskoeBog, 1 km SoftheroadKyonga- Kedrovyi,56km EofKedrovyi, 57°36’42”N,
80°32’21” E;20July;asmall lake surrounded by theopen Sphagnumbog(few dozens meters
broad)andfurtherby thevastpeatbogovergrownwith densestandsoflow pines Pinussylves-
tris;thewide transition mire zonearoundtheopenwatertableveryflexible andalmostimpen-
etrable(inaccessibleplacesthedepthofwater 15-40cm),with thevegetationtypicalofoligo-
trophicconditions: Sphagnumsp., abundant Carexlimosa and Scheuchzeriapalustriswith the
admixture ofRhynchosporaalba,andverynumerousDroseraanglica;nymphaeidsrepresented
by fairlyabundant Nupharpumila.
(2) Next tothebridgeacrossthe Emelich Riverontheroad Kyonga- Kedrovyi, 11.5km W of
Kyonga, 57°29’06”N, 80°51’41”E;20 July.
(3) TheroadfromVysokii YartoParbig, 15kmNEofParbig,57°17’48” N, 81°34’46” E;20July.
(4) Bakchar town, theGalka River (bridgeon theroad to Bogatyrevkaupto 1.3km S ofit),
57°0T44”-02’25” N,82°03’20”-04’04” E; 19July;thesmallriver(12-17m broad)withthe very
slow current(fasteronly locallyincorridors between vegetation)bringingbrown turbid warm
waters,depthupto2m,muddydeposits,vegetationlocallyabundant:predominatingSparganium
emersum(partlywithfloatingleaves), alsoButomusumbellatus,Carexacuta, Phalarisarundina-
cea,Glyceria sp„ Sagittariasagittifolia, Callitriche verna,manydeadtree trunksand branches
in thewater;surrounded partlybya sprucetaiga,with aspensontheedge and willows onthe
riverbanks, andpartlyby openareas(withherbaceous vegetationand bushes)stretchingfrom
thetown.
Fig. 1.Situation ofstudiedlocalities: (A) intheinvestigatedarea;- (B)intheenvironsofPlotnikovo.
Numbers correspondtothelocalitynumbers giveninthe text.
OdonatefaunaofeasternVasyuganPlain,Russia 5
(5) Bakchar,nearthebridgeontheGalka River, 57°02’22”N, 82°04’04” E;19 July;aman-made
smallpoolsituatedinopenareas,bounded withhighherbaceousvegetationand Carexrostrata,
veryabundantsubmergedandfloatingvegetation:Lemnatrisulca,Spirodelapolyrrhiza,Hydro-
charismorsus-ranae.
(6) Bakchar,in the centreofthe town, 57°0r22” N, 82°03’52” E; 19 July; aman-made largeeu-
trophicpondwithaclearwater, withoutshorevegetationbutwitha5-15mbroadzoneofvery
abundant submergedand floatingvegetation; mostlyCeratophyllumdemersumand Spirodela
polyrrhiza,additionallyLemna trisulcaand Hydrocharismorsus-ranae.
(7) Bakcharskoe Bog, 14kmW-WSWofPlotnikovo,4.5km Softheroad toBakchar, 56°5ri7”
N, 82°51’Ol”E; 13July;areasaround theecologists’shut- asub-marginalzoneofthe large
complexofSphagnumbogs andfens; ‘dwarfforests’oflowpines Pinussylvestris, dense shrub-
bery ofChamaedaphnecalyculata, Betula nana, locallyLedumpalustre, Andromedapolifolia,
Rubuschamaemorus;old drainageditches;avastopenpoorfen(mainlyCarexlasiocarpawith
anadmixture ofEquisetumlimosum)sosuffusedwith waterthatimpenetrable,inthesummer
thewateronlylocally emergesabovemosses, but inthespringitcertainlyoccursinlargequan-
titiesonthesurface;afire-fightingpoolwithdark waterand locallysedgesonthebanks.
(8) BakcharskoeBog, 14kmSWofPlotnikovo,9kmSoftheroad toBakchar.56°48’44”N,82°51’18”
E; 13July;oligotrophicpools(inthegiantcomplexofpeat bogsand fens),surrounded byan
open Sphagnumbogmostlywith Rhynchosporetumalbae (Rhynchosporaalba,Carexlimosa.
Droseraanglica,locallyChamaedaphnecalyculata)andridgesofverylow pinesPinussylvestris;
observations mainlyin azoneofapartlyfloodedtransition mirevegetation(‘Schwingmoor’)
alongtheshores (Sphagnum,abundantScheuchzeriapalustris,C. limosa,Menyanthestrifoliata,
also R.alba, Utriculariasp.;NupharpumilaandrareNymphaeatetragonafloatingonthewater
table),marginallyinadjacentsmall depressionsin Sphagnummats,with waterand sparse C.
limosaand S.palustris.
(9) Plotnikovo village,0.2-0.3km Wofthebridge onthe Iksa River,next theroad toBakchar,
56051’31”N,83°03’48”E; 17July;forestswampswith Callapalustris, Carexsp.,abundantLem-
naminor,numerousdeadspruce andbirch trees, abeaverlodge.
(10) Plotnikovo,0.1km WSWofthebridgeontheIksaRiver,neartheroad toBakchar, 56°51’30”
N, 83°03’52”E; 14,21 and22 July;smallwaterbodies mostprobablypost-excavation remains,
situatedindepressionsin the very warmgladebounded by willows and aspruce-birchtaiga;
astaticandpartlytemporarypools(onelargeranddeeper,andagroup ofsmalland shallow)
overgrownwith abundant Ceratophyllumsubmersum and Lemnatrisulca,withSpirodelapolyr-
rhiza andfloweringHydrocharismorsus-ranaecoveringalargepart ofthewatertables;locally
Carexpseudocyperus,Alismaplantago-aquatica,Typhaangustifolia,and Callapalustris.
(11) Plotnikovo,the IksaRiver and itsclosesurroundingsS andN ofthebridge,56°5r20-41” N,
83°03’58”-04’21” E;mostly 12and 14July,marginally21 and 22July;the 15-25wideriverwith
brownturbid water,thecurrentveryslow but fasterlocallybetween patchesofvegetation,bot-
tomdepositsmuddyandlocallysandy-gravely,vegetationlocallypoorer, mostlyonthebanks,
locallyabundantovergrowingalsoalargepart oftheriverbed: Carexacuta, Butomusumbella-
tus,Sparganiumemersum,stripsofNupharlutea;close surroundingspartly with adarktaiga,
but mostlyopenandsunny with richherbaceous vegetationandnumerouswillows.
(12) Plotnikovo,a former oxbow ofthe Iksa River Wof it (0.6km Wof Spart ofthe village),
56°50’49” N, 83°04’20” E; 16(mostly),22(marginally)July;abow-shapedopenareacovered
with abog-fencomplexpredominatedby Sphagnumand veryabundant Oxycoccuspalustris,
with Carexrostrata, C.lasiocarpa, C.diandra,Drosera rotundifolia,Scheuchzeriapalustris,Eq-
uiselum limosum;in themiddle,astripofComarumpalustrewith sparseMenyanthestrifioliata,
Typhalatifolia;and Thelypterispalustrisandwith waterlocallyappearingonthesurface; addi-
tionallya‘window’ ofopenwateramongthepeat-moss,50x20cminsize andquitedeep.
(13) Plotnikovo,meadows N ofthevillage,56°53’10”N,83°04’14”E;ISJuly.
6 R.Bernard& O.E. Kosterin
(14) Plotnikovo,field stationoftheInstitute ofSoil Scienceand Agrochemistry,the Siberian Divi-
sion oftheRussianAcademy ofSciences, 56°5141’ N, 83°04’28” E; 12July; meadowpatches
with treestandsdescendingtothehighbank oftheIksa River.
(15) Plotnikovo,next tothe tracktoBorodinsk (1.3km N of theroad toTomsk), 56°52T0” N,
83°04'37”E;mainly 16,additionally15 and 21 July;aman-madeeutrophicpond(usedby cat-
tle)with apastureononesideand anaspengroveontheother,thewatercloudyandtheveg-
etationratherpoor(especiallyemergent):onlyrare Alismaplantago-aquatica,Glyceriasp. and
Carex sp„ locally,a 1m wide shorebelt of Ceratophyllumsubmersum coveredwith Spirodela
polyrrhiza.
(16) Plotnikovo,theYarushkaStreamandthetrackjusttotheS ofthevillage,56°50’31”N,83°05'14”
E;21 July;a2-4.5 m wide forest stream,mostlyshaded, bringingbrownish waterwitha slow
current between Callitriche mats.
(17) Plotnikovo,nexttothetrack Softhevillage(2.2-3kmSoftheroadtoTomsk),56°49’42”-50’09”
N,83°05'11-17”E;21July;twosmallman-made poolswithveryabundant Ceratophyllumsub-
mersum,coveredwith Spirodelapolyrrhiza and Lemnaminor,andbounded byabruptlyending
walls ofCarexsp. and Glyceria sp., andlocallyby Typhalatifolia,Eleocharispalustris•and Al-
ismaplantago-aquatica.
(18) Plotnikovo,0.5 km Eofthe village,next totheroadtoTomsk, 56°5IT7” N. 83°05’51” E; 12
and21July;asmall (0.15ha)roadside pool(surroundedbymeadows)withclearshallowwater
onaloess-like substrate;abundant emergent vegetation:widespreadEleocharispalustris inter-
spersedwith scattered Alisma plantago-aquatica,somepatches ofEquisetumlimosum,Carex
acuta,Phragmitesaustralisand Typhalatifolia;onthewatersurfacesomefloatingleavesofPota-
mogetonalpinus;submergedvegetation:abundantmossesandLemna trisulca.
(19) Iksinskoe Bog.S part, 2.4km S oftheroad Plotnikovo-Tomsk, 56°50’05” N, 83°16’35” E; 18
July;twosmalllakessituatedinthemarginalzoneofthegiantpeat bog,inthewidestripofopen
Sphagnumbogwith Scheuchzeriapalustris,Rhynchosporaalba,Drosera(anglicamoreabundant
thanrotundifolia), Oxycoccuspalustris, Eriophorumvaginatumand side pineridges(‘gryady’);
thelakesseparatedandsurrounded bymesotrophicfenswithdense Carexrostrata andbymore
oligotrophicbogs;thewatertablebounded withatransitionmire (‘Schwingmoor’verysuffused
with water,partlyflooded)with Sphagnum,abundant Carex limosa and C.rostrata, and some
admixtureofMenyanthestrifoliataand Scheuchzeriapalustris;nymphaeids:Nupharpumilaand
rarer Nymphaeatetragona.
(20) IksinskoeBog,Spart. 1 kmSoftheroad Plotnikovo-Tomsk,56°50'55”N.83°17’33”E; 18 July;
afire-fightingpool withdark but clearwaterandpeatyshores, old drainageditches and peat
bogsovergrown with pine.
(21) Iksinskoe Bog,N part,4.0-4.4kmNoftheroad Plotnikovo-Tomsk,56°54'00-18”N,83°17’05-
19”E;23 July;three small lakessituated in thevast Sphagnumbog,initsopenstrip;onemes-
otrophic, rimmedwith Carex rostrata,onepoorer in nutrients with <C.rostrata and C.limosa,
and oneoligotrophicwithmuchrarerC.rostrata and abeltalongtheshores formedmostlyby
C.limosa, Menyanthestrifoliataand Scheuchzeriapalustris;nymphaeids:Nupharpumilaand
Nymphaeatetragona;inthe surroundings,Sphagnummatsverysuffused withwaterovergrown
withRhynchosporaalba, C. limosa,Drosera anglica andD.rotundifolia.
(22) Tazyrachevo(Tyzyrachevo) village,anoxbowoftheShegarkaRiverSWoftheroad toTomsk,
56°44"29” N, 83°33’46” E; 17July;thelargeoxbowsurrounded bya‘cedar-forest(Pinussibiri-
ca)ononeside,andbirch groves andadegradedmeadow ontheotherside;muddygroundand
eutrophicbutclear waterwith a thick rimof highBolboschoenus maritimus and someCicuta
virosa;in theopenwaterabundant Lemnatrisulcaand Ceratophyllumsubmersumcoveredwith
Spirodelapolyrrhiza; patchesofNupharlutea.
(23) Tazyrachevo(spellingonmaps;accordingtotheroadsign- Tyzyrachevo),theShegarkaRiver
anditssurroundings,fromthebridgeto0.25km NEofit,56°44'39-45”N, 83°34’08-20” E; 15
Odonate fauna ofeastern VasyuganPlain,Russia 7
and 17July;theriver atthestudiedsection isslightlydifferent than Iksa andGalka:colderand
faster,withamorerecognizablecurrent,thewaterslightlyturbidbut notbrown,bottomdeposits
muddyandsandy-gravely, locallyargillaceous,less(onlylocally)vegetatedin shorezoneswith
Carexsp.,Butomus umbellatus and Sparganiumemersum;theriver ispartlysheltered byfairly
highriverslopesovergrownwith willows,andsurrounded bothbyforestsand openareas.
(24) ObskoeSwampandadjacentmeadows,neartheroadMel’nikovo - StarayaShegarka,56°32’45”-
33’06”N,84°06’32”-07’02” E;15July;theverywidestripofswampsinthefloodplainoftheOb’
River,hardlypenetrable,overgrownwithlowbirches,willow thicketsandabundantherbaceous
vegetation(mostlyCarexpaniculata,C.pseudocyperus, C.rostrata, Calthapalustris, Comarum
palustre,and Menyanthestrifoliata),alsoinreeds andquakingbogpatches;intheswamps,apool
ofopenwaterwith Typhalatifolia,Acoruscalamusand Carexlasiocarpa;between theswamps
anda side western branch ofthe Ob’ River,anovergrazed,rather ruderal meadow,bordered
withtall Salixalba.
(25) SideWbranch oftheOb’Riverneartheroad Mel’nikovo- StarayaShegarkaandthe Obskoe
Swamp,56°32’59” N, 84°07’06” E; 15 July;theriver with turbid water and afairly fastcur-
rent.
ANNOTATED LISTOFSPECIES
Intheannotatedlist ofspecies thenumbers oflocalities correspondtothenumbers givenabove.
Thefollowingabbreviationsareused: VA:veryabundant,- A: abundant,—FA:fairlyabundant,
— M:inmoderatenumbers, — FR: fairlyrare, — R:rare, —VR:veryrare, —S;single; —Col.:col-
lected,— ten.:presence ofteneral individual(s),- ex.: exuviae.The collected specimens arein the
collections of;i:R.Bernard, —ii:FacultyofBiology,Adam MickiewiczUniversity,Poznan,Poland,
- iii:O.Kosterin,—iv:Siberian ZoologicalMuseum,InstituteofAnimalSystematicsand Ecology,
Novosibirsk, Russia, and - v:HessischesLandesmuseum,Darmstadt,Germany.
Calopteryxsplendens (Harris, 1782)
4; VA,alsoandrochrorae females,Col. 27c5 (21typical, 6 coloured tothe tips ofthewings), 192
(6typical,6androchrome coloured tothetipsofthewings,7androchrome f.faivrei);-11:FA,also
androchrome females.Col. 16,69(3typical,2 androchromecoloured tothetips, 1 androchrome f.
faivrei)',- 23:M,only 1androchrome female,Col.26, I?.
Seemstobe widespreadandmostly abundant, especially invegetation-rich sec-
tionsof therivers. Atloc. 4, visual counting andcollecting showedafairly con-
siderable proportion ofmales withthe wings colouredto the tips (6/27 count-
ed, 22%) anda slight advantage ofandrochromefemales(19/34 counted, 56%)
overtypical ones(15/34,44%). Morethanonethirdoftheandrochromefemales
(37%) represented f.faivreiwith gradually clearing tips offore wings while al-
mosttwothirds(63%) hadthewingscoloureduptothetips inclusive.Generally,
theformerconstitutedca21%of allcountedfemales(7/34) and thelatter 35%
(12/34). Theinnerendofthecolorationvaried. Inthemostextensively coloured
females itreached 50-60%ofthe length between thenodusand the wingbase.
Intheleastcoloured, itoccurred atnodusleveloronly slightlyexceeding it,and
suchcases werenotrare. Theproportions evaluateddonotfitany simple model
ofinheritancewhichwouldexplain all the alternativewingcolour morphs ob-
served in both sexes through asingle genetic factor. Anotherpopulation witha
8 R.Bernard &O.E.Kosterin
high proportionofandrochromefemales(including f.faivrei) andmaleswiththe
wings colouredto thetips was discoveredby O.KosterinintheTodzhaHollow
innortheasternTyva Republic, CentralSiberiain2000(KOSTERIN &ZAIKA,
2003) and2004 (unpublished). Before theserecords, androchromefemaleswere
consideredextremelyrareinSiberia(cf. BELYSHEV, 1973),butneverthelesspro-
vokeda persisting falsenotionof theexistenceofa secondspecies, Calopteryx
johanseni Belyshev, 1955(originally describedin thesubspecies rank) whichis in
factidenticalto f.faivrei. Therangeof spotsin thewings of somemales as well
as the high percentageof androchromefemalespartly resemble some Turkish
populations ofthespecies knownasC. s. intermedia(DUMONT, 2006).
Noteworthy is thecomplete absence ofCalopteryx virgo(L.)whichis stillun-
knownforthelarge area ofWest Siberiabetween theTobol-Ishiminterfluve(a
mappoint in BELYSHEV, 1973)andTomsk (BARTENEF, 1910).TheAltaian
localitiesgiven by BELYSHEV(1973) concern infactCalopteryxjaponica Selys,
1869, consideredby this authoras conspecific to C. virgo(MALIKOVA. 1995).
However, C. virgo was recordedinSiberiaevenfurthereast,fromBunbuy, Kras-
noyarskii Krai(3/6-VI-1915, Valdaevleg.), noteworthy that sympatrically with
C.japonica (MALIKOVA, 1995).
LestesdryasKirby, 1890
10:R,Col. 18,15;- 17:R,Col. 38;-18:A,tandems,Col. 18,15.
Not widespread, recordedonly in secondary habitatsonmineralgrounds.
Lestessponsa(Hansemann, 1823)
1:A,ten.,Col. 28, 1ex.;-5: VR,Col. 18;-8:A,ten.,Col. 18,29;- 10:R.ten.,Col. 12;-11;
- 15:Col,28;- 17:A,tandem,Col. 28, 12;-19:A,ten.,Col. 18,12;-21:A,ten..Col. 38,6ex.;
- 22:FA, tandem,Col. 28;- 23:R,tandem.Col. 18;- 24:R,Col. 28.
Widespread, common andabundant,inhabiting bothprimaryandsecondary
habitats.
Lestes virens (Charpentier, 1825),ssp. vestalis(Rambur, 1842)
8:Col. 1(J ten.;- 18:VA.ten.,Col. 1<J, 15.
Scarce, however,maybe locally veryabundant.Therarity isprobably aresult
ofsituationofthestudiedareaatthenorthernborderofthespecies’ distribution
(cf. BELYSHEV, 1973).Thethoracicpatternis typical ofthessp. vestalis, how-
ever,theindividualsobservedin thefieldseemedtobesmaller incomparison to
thosefromcentralEurope. BELYSHEV(1973)considering theWestSiberianvi-
renswrote aboutits transitionalsizebetweenEastEuropean vestalisandasmall
Asiansubspecies, marikovskii.
Sympecma paedisca (Brauer, 1877)
11;Col. 12 old;- 15: FR,only ten..Col. 12;-22:FA,only ten..Col. 32.
The teneralindividualsof the new (summer) generation were observed only
Odonatefauna ofeasternVasyuganPlain,Russia 9
fourdaysafter collecting thelastrepresentative oftheold(spring) generation in
thesamevillage. Theserecordssuggesta‘time-touching’ ofthesegenerations in
the scaleofasmallarea.Thisphenological phenomenon israthertypical in Si-
beria (e.g. KOSTERINetal., 2001 andunpublished data),but in Europe there
israther ashortbreak(several-week-long, mostly in July)betweengenerations.
Although veryrare records showapossibility ofoccasionaloverlapping ofgen-
erationsalso inEurope, butnobody has confirmeditinthescaleofa smallarea
sofar(cf. JODICKE, 1997). Theoverlapping of generations in Siberiamayre-
sult froma shorterandlatewarm season ofthe year.To adapt the lifecycle to
these conditions, thelarval development is certainly fasterand shorter.Though
eggs maybelaidin Europe already inApril,anemergencestartsinthelastpen-
tadeofJuly(JODICKE, 1997).InPlotnikovo, thereis nopossibility to oviposit
beforeearly or mid-May, but on 16 July the emergenceof the population was
already quite intensive.Theshorteregg/larval development combinedwith the
delayedperiod ofspring activityof imagines, andhencetheirlaterdeath,make
‘themeeting of generations’possible.
Coenagrion hastulatum(Charpentier, 1825)
4:VR;- 5:R,Col. 23, 12;-6;S;- 7 (pool):Col. Id;- B:M,copulation,Col. 2d,32;- 10:R,
tandem.Col. 12;-11: M,tandems,oviposition, Col. 1d;- 15: A,tandems.Col. 3d, 12 andro-
chrome;- 18:A,tandems,Col. Id;- 19:VR,Col. 2d, 12;-20(pool):VR;-22:FA,Col, 3d;-23:
VR.
Widespread andthecommonest coenagrionid inthestudiedarea. Fairlyeury-
topic. Itdoesnotavoidevencurrent waters,however, theoccurrence ofimagines
islimitedonly to lenticshorezonesof veryslowandbrownrivers.The question
ariseswhethertheseimagines are nativeororiginate fromstanding waters inthe
surroundings. The observed reproductive behaviourat the Iksa River weigh in
favourofthe formeroption.
Coenagrion johanssoni(Wallengren, 1894)
1:FA,Col.3d,32;-8:FA,tandems.Col.3d,22;-10:FA,ten.,ovipositions,Col. 1d,12;-11;
1 d,allochthonous;- 17:S,Col. Id;-18:VA,ovipositions,Col. 2d,22;- 19: FA,ten..Col. 2d,
12;-21:M,Col.6d,22.
Widespread and locally fairly abundant, related to standing waters, inhabit-
ing both primary and secondary habitatsbutwith apreference to the former.
Animportant proximate cue forthespecies is anemergentfairlylow vegetation
(up to50-60cm high) providing important ultimate factors, atleasttheactivity
spaceandwindshield.Thisrequirement, stressed intheliterature(BERNARD
& SAMOL/\G, 2002), underlay the clear opposition observed in the studied
area,betweeninhabitedanduninhabitedlocalities.Theformer(lakes inSphag-
numbogs, aroadside pool)wererich insuch vegetation whileinthelatter(man-
madelarge ponds andsmallfire-fighting pools) itoccurred atmostinvery small
numbers.Twoexceptions onlyconfirmthe rule. Smallpools in Plotnikovo(No.
10 R.Bernard& O.E.Kosterin
10) werepoorin theemergent vegetation butthey were situatedinlanddepres-
sionssurroundedby densebushesandtrees whichcreatedawarmandsufficient-
lycalmmicroclimate. Thesignificance of dense shrubbery for this highly wind
sensitivespecies was communicatedby BELYSHEV(1966). On theotherhand,
anoxbow(No. 22) aboundedinthe emergentvegetationalong theshorebutnot
ofthe preferred structure as being too tallanddense. What is more, water was
probably too eutrophic for C.johanssoni as the species prefers waters poorerin
nutrients.
Atloc. 18, atandemof C.johanssoni was caughtby maleL. dubia.When the
malehadbeen eaten,thefemalemanaged toescape.Thiswas anexcellentexam-
pleofthe significance ofmorecryptic coloursoffemales as theattack of dubia
wasdirected atthe muchmorebrightmale.
Coenagrion lunulatum(Charpentier, 1840)
6:only 1male.Col. 1<J.
Therarityofthespecies was relatedmostprobably totheveryend ofitsflight
season.
Coenagrion puella (Linnaeus, 1758)
5:S,Col. 13;- 6: FA.tandem.Col. 4c?, 1 10:FA,tandems.Col.4c?,29;- 15:FA, tandems.
Col. 1c?, 19;- 17:S,Col. I6;- 22:R,Col. It?;-23: VR,Col. 15.
Theknown distributionofthespecies inSiberiareached55-56°N(BELYSHEV,
1973;KOSTERIN etal., 2001). Therefore, therecords inthestudiedarea, where
thespecies is localbutfairly widespread,expand thespecies’ knowndistribution
limitsignificantly (at least 1°) north, up to 57°02’. Considering the moderate
abundanceofC. puella inthisareaas wellas itsEuropean rangereaching above
60°(DIJKSTRA, 2006a), an occurrence ofthe species even athigher latitudes
inWSiberiaisvery probable. In theVasyugan Plain, C. puella is afairlyregular
inhabitantofsecondary habitats, i.e.villageandroadside ponds andpools, and
veryrarely occurs inprimary habitats,exactly only inanoxbowoftheShegarka
River.Anabundantsubmerged vegetationandahighertrophy aretheircommon
features. A totalabsence of the species in water bodies in Sphagnum bogs and
fens is noteworthy in comparison with the situation in centraland eastern Eu-
ropewhere thespecies inhabitssimilarbiotopes regularly. Thenarrowedhabitat
spectrumin theborderlandsofthedistributioniscertainly an effectofa greater
sensitivenessofthespecies insuch asituationcombinedwith more severe micro-
climaticconditionsin somehabitatsthere. Thisphenomenon isknownalso for
manyotherdragonfly species, e.g.Erythromma lindenii(BERNARD, 2000) and
Aeshnacrenata (BERNARD,2002).
Coenagrion pulchellum(Vander Linden, 1825)
6:M,tandems,Col.4d, 19.
InSiberia, thespecies was known intheSW part upto ca 56°20’N, however.