TurkJZool 28(2004)245-265 ©TÜB‹TAK

FaunisticandMorphologicalStudieson(,Ciliophora) fromaSmallPond,withResponsesofPopulationsto ChangingEnvironmentalConditions

NaciyeGülk›zfiENLER*,‹smailYILDIZ** DepartmentofBiology,YüzüncüY›lUniversity,Van-TURKEY *E-mail:[email protected] **E-mail:[email protected]

Received:12.06.2003

Abstract: Sixty-nineciliatewereidentifiedin19samplesfromasmallpondcoveredby Lemnaminor inVan,Turkey. Statisticalanalysiswasemployedtoexaminetherelationshipsbetweenciliatesandthephysicalandbiologicalparameters.Totalciliate abundanceappearstoberelatedmainlywithphysicalandbiologicalfactorssuchasthedissolvedoxygeninthebottomlayerandthe volumeof Lemna andtemperatureinthesurfacelayer.Somespecieswereinvestigatedindetailusingliveobservation,silver impregnation,andmorphometry.

KeyWords: CiliatedProtozoa,Pond,Lemna,PhysicalParameters,CiliateCommunities,Infraciliature,Morphology.

KüçükBirGöletteBulunanSiliyatlar(Protozoa,Ciliophora)Hakk›ndaFaunistikveMorfolojik Çal›flmalarileSiliyatPopulasyonlar›n›nDe¤iflenÇevreselKoflullaraYan›tlar›

Özet: Buçal›flmadaVan’dabulunanLemnaminor ileörtülübirgöletten19kezörneklemeyap›lm›flve69siliyattürütan›mlanm›flt›r. Fizikselvebiyolojikparametrelerilesiliyattürleriaras›ndakiiliflkileriincelemekamac›ylaistatistikselanalizyap›lm›flt›r.Toplamsiliyat bollu¤uesasolarakdipteçözünmüfloksijen;yüzeytabakada Lemna hacmives›cakl›kgibifizikselvebiyolojikfaktörlerileiliflkili görünmektedir.Baz›türlercanl›gözlem,gümüflempregnasyonvemorfometriileayr›nt›l›olarakincelenmifltir.

AnahtarSözcükler: SiliyatProtozoa,Gölet,Lemna,FizikselParametreler,SiliyatKomuniteleri,‹nfrasiliyatür,Morfoloji.

Introduction investigators(Porteretal.,1985;Finlayetal.,1988; Inaquaticenvironments,ciliatedprotozoatogether BeaverandCrisman,1989;Madoni,1991a,1991b; withflagellates,,andsmallmetazoansforma SalvadoandGracia,1991;Cabré,1993;Sommarugaand microbialcommunity.Amongthemembersofthis Psenner,1993;Madoni,1996;Biyu,2000;Holen, communitycloserelationshipssuchaspredationand 2000).However,thereislittlepublishedinformationon competitionoccur(Madoni,1996).Ciliatedprotozoa theciliatedprotozoaoccurringinaquaticecosystems constituteasignificantportionofthemicrobialfoodweb fromTurkey. andplay2mainrolesinaquaticecosystems(Porteretal., Identificationofciliatesisdifficultduetohighspecies 1985;Finlayetal.,1988;BeaverandCrisman,1989; diversityinaquaticecosystems,variabilityofcellsizes, SalvadoandGracia,1991;SommarugaandPsenner, damagecausedbyfixativestocellsandcultivation 1993;Holen,2000):theylinkphytoplanktonand problems.Thedescribedspeciesmayshowmorphological bacteriainthefoodwebtohigherorganisms.Inaddition, variationaswell.Thereforetaxonomicerrorsmayoccur theseciliatesmaybeimportantinnutrient inmanylistsofciliatefauna(FoissnerandO’Donoghue, remineralization. 1990).Althoughthespeciescompositionwasgenerally Freelivingciliatesfromaquaticecosystemsinsome similar,somegeographicalvariationsexistedinthe countrieshavebeencarefullysurveyedbymany occurrenceofsomespecies.Specifically,manynew

245 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions speciesandalsoendemicspeciesweredetectedincertain populationsweremadebydirectobservationoflive areasbyusingbiometryandcytologicalstaining sampleswithin5hofcollection.Taxaweresubjectto techniques(FoissnerandO’Donoghue,1990;Foissner, taxonomic(cytological)proceduresandidentifiedto 1997).SimilarresultscouldbeexpectedinTurkey. specieslevelusingthekeysofCurds(1982),Foissneret Indeed,theciliatecompositioninfreshwaterhabitats al.(1991,1992,1994,1995),FoissnerandBerger ofTurkeyhasneverbeenstudied.Weexaminedsome (1996)andspecificliteraturecitedinthespecies freshwaterciliatesinsomeriversandwatertreatment descriptions.Theciliarypatternandothercytological systems(fienleretal.,1998;fienlerandY›ld›z,1998; detailswererevealedbyvarioussilverimpregnation 1999a,1999b;fienleretal.,1999).Itwouldbeofgreat techniques,allofwhichweredescribedbyFoissner significancetodeterminetheciliatefaunaoftheaquatic (1991)andFernandez-Galiano(1976,1994).The ecosystemsinTurkeyandthenmakecomparisonswith ciliateswerealsosubjecttosupravitalstainingwith faunisticstudiesaboutthefreshwaterfaunaofdifferent methylgreen,methylgreen-salina-formaline(MSF)and geographicalregions. Feulgen’snuclealreactiontodeterminecytological features.Allofthecellmeasurements,giveninthestudy Thepurposeofthisstudywastodeterminetheciliate inmicrometers,wereobtainedwiththeaidofacalibrated speciesinhabitingbothonthesurfaceandinthebottom ocularmicrometer.Morphologicalcharacterizationwas ofaproductivepondcoveredbymacrophytesinVan doneonrandomlyselectednon-dividingcellsafter Castle,Van,Turkey,andtogivemorphologicalvariations impregnation.Permanentpreparationsbelongingto withinthespecies.Statisticalanalysiswasappliedto hypotricheswerenotachievedbecauseoftheirfixation determinetherelationshipsbetweenciliatespecieswith problems.Thereforehypotricheswiththeexceptionof physicalparameters. thespeciesdescribedinvivoweregivenastotal hypotriches. MaterialsandMethods Estimatesofpopulationdensityinthemacrophyte Studyarea: Thepond,situatedatanaltitudeof1700 layerandinthebottomweremadeaccordingtothe minVanCastleintheprovinceofVan,hasanareaof48 techniquesdescribedbyMadoni(1991a,1991b).For m2 andmaximumdepthof0.7m.Thisshallowpondis ciliatecounts,watersampleswereconcentratedtoa 21x7.5mandisfedbyrainfallandlittlewaterbearing smallvolumebya23µmmeshnetandcountedby stratumemergence.Sincethewaterishighlyeutrophic, HydroBios-Kiell(0.5)countingcameraaftersubsampling. Lemnaminor isabundantinthepond.Thepondis Themostconvenientdropsize(0.05ml)andnumberof exposedtotheanthropogeniceffectsofgrazingand replicatecounts(3replicates)wereselectedaccordingto picnics. Madoni(1984).Theabundanceofeachtaxonwas expressedasnumberofindividualspermilliliterof Samplingofciliatedprotozoa: Thestudywas surfaceandbottomarea. conductedatintervals,sometimesmonthlyand sometimesoncein2weeks,fromMay1999toMay Physicalparameters: Temperature(°C),pH, -1 -1 2000.Thepondwassampled19times.Sampleswere dissolvedoxygen(mgl ),conductivity(µmhoscm )and collectedfromboththesurfacelayerandthebottom. salinity(‰)weremeasuredinthestudyareawithaDO Samplingofciliatedprotozoawasperformedusing meter(Jenway,9070)andasalinity-conductivity- Madoni’stechniques(1991a,1991b).Atthesampling temperature(SCT)meter(YSI33). point,thewaterlayerfromthesurfacedowntoabout2 Dataanalysis: Thenormalityassumptionwas cmwascollected;asurfaceareaof500cm 2 was checkedforallmeasurementsofphysicalandbiological explored.Ciliatesfromthebottomofthepondwere variables’distribution,andappropriatetransformations collectedwitha2cmdiameterpipebysiphoning. weremade[Variable=ln(Variable+1)].Partial Identificationandenumerationofciliated correlationanalyseswereperformedtorelatenumbersof microfauna: Sampleswereinvestigatedonthedayof speciesinsamplestothephysicalparametersmeasured. sampling,usingabrightfield,invertedandphasecontrast ThestatisticswereanalyzedusingMINITABandSPSS software.Intheaccompanyingtables,thefollowing microscope.Ineachsample,estimatesofciliate – abbreviationsareused:X,arithmeticmean;SD,standard

246 N.G.fiENLER,‹.YILDIZ

deviation;CV,coefficientofvariation;min,minimum; Biyu,2000)andricefields(Madoni,1996).Generally, max,maximum;N,numberofindividualsexamined. ciliatesareconsideredquitecosmopolitanintheir distribution(Madoni,1990;Biyu,2000),whichmay revealwhynoendemicspecieswerefoundduringthis ResultsandDiscussion study. Thevaluesofeachphysicalparametersmeasuredin Averagetotalciliateabundanceordensityatthe thepondaresummarizedinTable1. surfaceandthebottomwas211.90cellsml-1 and385.60 Duringthestudyperiod,69speciesofciliated cellsml-1,respectively.Thepresentstudyhasshownthat protozoawereidentified(Table2).Thetotalnumberof thepond,whichiscoveredby Lemnaminor,hasgreater taxacollectedissimilartothosereportedforpondswith ciliateabundance.Conversely,inthemacrophyte-rich macrophytes;Madoni(1991a,1996)found40and60 lake,theaverageciliatedensitywas13.5cellsml-1 andin taxainapond,collectingsamples17timesovera1-year themacrophyte-poorlaketheaverageciliatedensitywas periodandinaricefield,collectingsamples43timesover 35.5cellsml -1 (Biyu,2000).Moreproductivelakes a4-yearperiod,respectively.Ourresultsarealsosimilar exhibitgreaterabundancethanoligotrophiclakes, tothetotalnumberofciliatespeciesrecordedbymany throughastrongrelationshipbetweentheabundanceof authors(SalvadoandGracia,1991;Cabré,1993). planktonicciliatesandtrophicstate,andphysicalfactors Moreover,Finlayetal.(1988)reportedasignificantly togetherwithbiologicalrequirementssuchasfood, highernumberoftaxainasmallpondwithmacrophytes protectionandreproduction(BarbieriandOrlandi,1989; similartothoseoftheinvestigatedpondwithina48h BeaverandCrisman,1989;Biyu,2000). period.Thepondsupportedahighdiversityofciliated Inthepresentstudy,23specieswereobservedonly protozoainthepresentstudy.Biyu(2000)showedthat atthebottom,and21specieswerefoundonlyinthe themacrophyte-richlakehadgreaterciliatespecies surfacelayercoveredby Lemnaminor ;25ofthese numbers,butlowerciliateabundancethanthe speciesoccurredinbothhabitats.Themajorityofthe macrophyte-poorlake.Theresearchernotedthat observedspecieswerebactivorousciliates.Predator macrophytesmaysustainmorediverseciliate formswereobservedonlyinthe Lemna layerwithafew communities,probablythroughthemodificationofciliate exceptions: Acineriauncinata and Amphileptus foodresourcesandspacesheterogeneity. pleurosigma werefoundbothinthesurfacelayerandat Ourstudyindicatesthattheciliatedprotozoa thebottom.Sessileciliatesfeedingonbacteria communitycontainsrelativelysimilarciliatespecies,at representedarelativelymoreimportantcomponentin leastineutrophicponds.Themajorityofciliatedprotozoa the Lemna layer(seeTable2).Thesessileperitrich determinedinthepondhavebeenrecordedpreviouslyas Epistylisdigitalis wasobservedasepizoiconlyattachedto commontofreshwatersystemssuchaslakes(Madoni, copepods.Thealgivorousspecies bursaria 1990;Biyu,2000),ponds(Prattetal.,1986;Finlayet and Trithigmostomasteini occurredonlyinthesurface al.,1988;Madoni,1991a;SalvadoandGracia,1991; layer.Amongthesespecies,vorticellidciliatesand

Table1.Physicalcharacteristicsfromthepond(N=19).

Parameter Mean Range

Temperature(°C) 11.62 7.1-20.0 PH Surfacelayer 7.49 7.00-7.97 Bottom 7.33 6.83-7.78 DissolvedOxygen(mgl-1) Surfacelayer 4.63 1.2-7.00 Bottom 2.91 0.9-6.5 Conductivity(mmhoscm-1) 736.25 680-820 Salinity(‰) 0.46 0.30-0.70

247 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Table2. Feedinghabit,frequencyandabundance(individualnumberml -1)ofciliatespeciesobservedalongthestudyperiodinthepond.P, predator;Ba,bacteria;SB,sulfurbacteria;Al,algae;O,omnivorous;Di,diatoms;Fl,heterotrophicflagellates;Cy,cyanobac teria.

Surfacelayer Bottom – – Species Nutrition Frequency X (min-max) Frequency X (min-max)

Acineriauncinata P 21.05 0.34(0.00-2.00) 31.58 3.85(0.00-26.60) Amphileptuspleurosigma P 31.58 0.42(0.00-2.48) 5.26 0.35(0.00-6.60) Apsiktrata sp. ? 68.42 12.87(0.00-86.60) Aspidiscacicada Ba 52.63 2.57(0.00-15.34) 89.47 16.74(0.00-40.00) Aspidiscalynceus Ba 21.05 0.24(0.00-2.00) 26.32 2.44(0.00-20.00) Bothrostomaundulans SB 10.53 0.70(0.00-6.60) Brachonellaspiralis SB,Al,Fl,Ba 10.53 0.70(0.00-6.60) Caenomorphauniserialis Ba,SB 5.26 0.05(0.00-1.00) Caenomorpha sp. Ba,SB 5.26 0.35(0.00-6.60) Carchesiumpolypinum Ba 31.58 12.9(0.00-200.00) Chilodonellauncinata Ba 47.37 2.11(0.00-22.00) 5.26 0.71(0.00-13.40) Cinetochiliummargaritaceum Ba,Al 63.16 5.56(0.00-44.00) 73.68 22.08(0.00-106.60) Cirrantermobilis ? 21.05 2.19(0.00-15.00) hirtus O 26.32 1.95(0.00-23.00) 47.37 9.47(0.00-33.40) Cothurniaannulata Ba 5.26 0.02(0.00-0.33) Cyclidiumglaucoma Ba 47.37 1.88(0.00-10.80) 63.16 27.00(0.00-100.00) Cyclidiumheptatrichum Ba 47.37 15.77(0.00-46.60) Dexiotrichagranulosa Ba 73.68 26.05(0.00-80.00) Dexiotrichidescentralis Ba 5.26 0.35(0.00-6.60) nasutum P 5.26 0.05(0.00-1.00) sp. P 5.26 0.11(0.00-2.00) Epalxella sp. SB 15.79 8.42(0.00-100.00) Epistylisdigitalis Ba 47.37 13.90(0.00-201.70) aediculatus O 63.16 3.08(0.00-15.34) angusta O 10.53 0.29(0.00-4.46) 63.16 13.58(0.00-46.60) Glaucomascintillans Ba 15.79 0.16(0.00-2.30) Halteriagrandinella Ba,Al 26.32 0.24(0.00-2.00) 15.79 3.42(0.00-53.40) Holophryadiscolor O 78.95 22.98(0.00-113.40) Holophrya sp. ? 26.32 5.25(0.00-46.60) Holostichapullaster Ba,Di,Al 26.32 0.87(0.00-7.14) 26.32 2.45(0.00-13.40) Homalozoon sp. O 21.05 2.11(0.00-13.40) Lacymariaolor P 5.26 0.05(0.00-1.00) Lagynuselegans O 57.89 7.41(0.00-26.60) Litonotuscygnus P 15.79 0.26(0.00-3.00) Litonotus sp. P 26.32 0.37(0.00-3.00) striatus Al,Di,Cy 43.37 8.76(0.00-60.00) es Ba,Fl,Al 15.79 1.04(0.00-6.60) Metopusstriatus Ba,Fl,Al 5.26 0.35(0.00-6.60) Myelostomaanatinum ? 10.53 0.05(0.00-0.46) 84.21 18.68(0.00-60.00) Monodiniumbalbiani P 5.26 0.02(0.00-0.44) Opisthonecta sp.? 15.79 0.73(0.00-7.84) Parameciumbursaria Ba,Al,Di 68.42 3.09(0.00-14.00) Parameciumcaudatum Ba,Al 15.79 0.12(0.00-1.00) 15.79 1.40(0.00-13.40) Plagiopylanasuta Ba,Al,Fl,SB 5.26 0.02(0.00-0.46) 47.37 4.37(0.00-20.00) Platycoladecumbens Ba,Al,Di 5.26 0.05(0.00-1.00) Pseudoconhilembuspusillus Ba 57.89 19.29(0.00-93.40) Satrophilus sp. ? 15.79 1.40(0.00-13.40) ambiguum Ba,Fl,Al 5.26 0.02(0.00-0.46) 15.79 2.19(0.00-20.00) Spirostomumminus Ba 15.79 2.37(0.00-20.00) Spirostomumteres Ba,Al,Di,SB 5.26 0.44(0.00-8.40) 31.58 7.71(0.00-80.00) coeruleus O 21.05 0.15(0.00-1.00) 15.79 1.04(0.00-6.60) Stentormuelleri Ba,Al,Di 57.89 1.97(0.00-11.00) 5.26 0.35(0.00-6.60) Strobilidiumcaudatum Di,Al,Ba 10.53 5.59(0.00-80.80) sp. ? 31.58 3.11(0.00-29.00) Tachysomapellionellum Ba,Cy,Al,Di 42.11 2.66(0.00-12.64) Thigmogasteroppositovacuolatus Ba 26.32 0.32(0.00-2.30) 5.26 0.35(0.00-6.60) Trithigmostomacucullulus Di,Al,Cy,Ba 31.58 7.36(0.00-60.00) Trithigmostomasteini Di 26.32 1.32(0.00-13.60) Trochiliaminuta Ba 21.05 1.03(0.00-9.50) 26.32 8.77(0.00-80.00) Urocentrumturbo Ba,Di 21.05 11.60(0.00-208.90) 52.63 18.98(0.00-125.00) Uronemanigricans Ba,Fl 10.53 0.16(0.00-2.00) 42.11 8.77(0.00-53.40) Urotrichaglobosa Ba,Al 94.74 29.55(0.00-80.00) Urotrichasp. ? 47.37 8.25(0.00-33.40) Vaginicolatincta Ba 5.26 0.04(0.00-0.66) campanula Ba,Al 84.21 39.70(0.00-207.40) 26.32 3.52(0.00-20.00) Vorticellaconvallaria Ba 100 71.90(2.00-298.30) 21.05 1.75(0.00-13.40) Vorticellamicrostoma Ba,Al 36.84 2.87(0.00-11.00) Vorticellaoctava Ba 89.47 9.07(0.00-26.00) 10.53 2.11(0.00-33.40) Vorticellapicta Ba,Al 21.05 0.90(0.00-12.00) OtherHypotriches 78.95 7.81(0.00-53.34) 73.68 19.99(0.00-106.60)

Totalciliates 211.90(6.30-569.10) 385.60(53.00-799.80)

248 N.G.fiENLER,‹.YILDIZ

Epistylis havebeenreportedasrelativelycommoninthe Onlyspecieswithasignificantfrequency(>10%) surfacelayerofeutrophicponds(Madoni,1991a;Cabré, wereselectedforpartialcorrelationanalysis.Table3 1993). representstherelationshipbetweentheciliatesandsome physicalparameters. Ingeneral,atthebottom,theciliatecommunity prevalentlyconsistedofmicroaerophylicandanaerobic Althoughciliateabundancedidnotcorrelatewith species. Bothrostomaundulans , Brachonellaspiralis , physical-chemicalvariables,probablybyusingsimple Caenomorpha spp., Cirrantermobilis , Epalxella sp., correlation(Biyu,2000),thepresentstudyhasshown Lagynuselegans , Loxodesstriatus,Metopus spp., thatthereisanobviouscorrelationbetweensomeciliate Myelostomaanatinum and Plagiopylanasuta represent abundanceordensityandphysicalvariablessuchas thecharacteristiccomponentsoftheciliatefaunaof temperature,dissolvedoxygenand Lemna volume(see sulfureta(Curdsetal.,1983;Madoni,1990;1991a; Table3)byusingpartialcorrelation.Itisapparentthat PattersonandHedley,1992;Cabré,1993;Foissnerand temperatureisthemostimportantparameterinthe Berger,1996).Theirdensitiesaregenerallyratherlow surfacelayer.Thehighestvaluesforthecorrelation coefficientwerefoundbetween Lemna andthe (seeTable2).Theseareanaerobicorganismscapableof temperatureofthesurface.Macrophytes,infact,were feedingondifferenttypesofsulfhurbacteria.Other associatedwithhightemperatures.Thesamecorrelations ciliateslike Apsiktrata sp., Cyclidiumheptatrichium , wereobservedfortheciliate Parameciumbursaria and Dexiotrichagranulosa,Holophryadiscolor, taxasuchasperitrichs Vorticella spp.wereassociated Pseudocohnilembuspusillus and Urotricha spp. withhighLemna volumeandtemperatureofthesurface. representedarelativelymoreimportantcomponentof Conversely,taxasuchas Chilodonellauncinata , thespeciesassociatedwiththebottom.Theirfeedingis Tachysomapellionellum and Trochiliaminuta were verydiverse,i.e.bacteriatoProtozoa(Foissnerand associatedwithlowmacrophyteandtemperaturelevels. Berger,1996). Urotrichaglobosa wasthemost commonlyobservedspeciesatthebottom.Barbieriand Thepositiverelationshipbetween Urocentrumturbo Orlandi(1989)found Urotricha spp.inbothhabitatsin andtemperatureofthebottomindicatesthathigh temperaturesfavorthedevelopmentof Urocentrum aneutrophicreservoir,butwithgreaterdensityinsurface turbo.Incontrast, Dexiotrichagranulosa,Holosticha samples.However,inafreshwaterreservoir, Urotricha pullaster andLoxodesstriatus hadanegativerelationship sp.arefrequentlyfoundonthesurfacewithorwithout withthetemperatureofthebottom. Lemna gibba (SalvadoandGracia,1991).Foissnerand Berger(1996)reportedthattheciliatecommunityof Inconclusion,whentotalciliatesareconsidered,the pelagialandsmall,stagnantwaterbodiesalsocontained correlationanalysissuggestedwhich Lemna volumeand Urotricha spp.thatareabletoliveinmucuoustubes temperatureofthesurfaceanddissolvedoxygenofthe attachedtodebrisonthebottom,andfeedonbacteria. bottomhavethegreatestinfluenceontotalciliate abundance.Similarresultsfor Spirostomumteres were Amongtheciliatesobservedbothinthesurfacelayer observedbyMadoni(1991b).Itisapparentthatfood andatthebottom,onlyafewspecies, Aspidiscacicada, availabilityismajorfactorcontrollingtheabundanceof Cinetochiliummargaritaceum,Cyclidiumglaucoma and ciliatedProtozoa(Madoni,1990).Physicalconditions Urocentrumturbo,occurredwithhighfrequency. suchasthepresenceorabsenceofoxygenand Thesurfacecontainedsomesporadicandaccidental temperaturehavealsobeendescribedashavinganeffect speciesfromthebottomlike Myelostomaanatinum and ontheabundanceanddistributionoftheseorganisms. Plagiopylanasuta,whichwererepresentedbyonlyalow Finlay(1982)reportedthatoxygenavailabilityisan abundanceandfrequency.Thebottomalsocontained importantfactorcontrollingthedensity,biomassand sometypicalsurfaceciliateslikeVorticella spp.withlower communitystructureofbenthicciliatedProtozoa.The frequency.Theseunexpectedresultsincommunity presentstudyalsoconfirmedthisview. structureareduetoavarietyoffactorsincludingthe shallownessofthepondandeffectivemixingbywinds, Thedescriptionofspecies anthropogeniceffectsandrun-offfromthesurrounding picnicareas. Wecarriedoutathroughinvestigationofthespecies retrievedfromthesamples,butnonewspecieswere

249 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Table3.Partialcorrelationcoefficientsbetweenciliatedspeciesandsomephysicalvariables.*P<0.05;**P<0.01;***P <0.001

Species Lemnaminor Temperature(°C) DissolvedOxygen(mgl -1)

Surface Bottom Surface Bottom

Acineriauncinata 0.3073 Amphileptuspleurosigma 0.4346 0.6844** 0.5202 Bothrostomaundulans -0.3281 Brachonellaspiralis 0.3240 Chilodonellauncinata -0.4737* -0.5273 Cirrantermobilis -0.3024 -0.4282 Cyclidiumheptatrichum -0.3175 -0.5098* Dexiotrichagranulosa -0.5021* -0.4430* Dexiotrichidescentralis -0.3166 Epistylisdigitalis -0.3165 Glaucomascintillans -0.3490 Holophryadiscolor -0.4494* Holophrya sp. -0.3090 Holosticapullaster -0.4597* -0.4399 Litonotus sp. 0.3209 Homalozoon sp. 0.6227** Loxodesstriatus -0.4977* -0.4301 Monodiniumbalbiani -0.3269 Myelostomaanatinum 0.3669 0.3068 0.3249 Parameciumbursaria 0.5628** 0.5074* Parameciumcaudatum -0.6706** Plagiopylanasuta -0.4276 Satrophilus sp. 0.5104* Strobilidiumcaudatum 0.3964 Stylonychia sp. 0.3442 0.4903* Tachysomapellionellum -0.4379* Trochiliaminuta -0.5865** -0.5171* -0.3585 Urocentrumturbo 0.5517** -0.4609* Urotrichaglobosa -0.3311 Urotricha sp. 0.4098 Vaginicolatincta 0.4627* Vorticellacampanula 0.3014 0.4002 Vorticellaconvollaria 0.6735*** 0.6954*** 0.5494** 0.3729 Vorticellamicrostoma 0.5128 Vorticellaoctava 0.6082** 0.6185** Vorticellapicta 0.3025 Totalciliates 0.6031** 0.4672* -0.4691* Temperature 0.8123***

found.However,sometaxawerenotdetermined,mainly Halteriagrandinella (MUELLER,1773)DUJARDIN, becausefewspecimenswerefound,whichmadethe 1841 determinationtoodifficult.Verylikely,someofthese (Figure1;Table4) unidentifiedtaxawerenewspeciestoo. ThisspeciesbelongstotheorderOligotrichida,family Halteriidae.Invivoitmeasures22.50-37.50x15.00-

250 N.G.fiENLER,‹.YILDIZ

Figure1. Halteriagrandinella ,invivo.AZM,adoralzoneof membranelles;Arrow,jumpingbristles;Bar:8µm.

Table4.Morphometriccharacterizationof Halteriagrandinella.

– Character X SD CV Min Max N Figure2. Euplotesaediculatus,silverimpregnation.AZM,adoralzone Body,length 36.80 4.50 11.66 30.00 45.00 15 ofmembranelles;Ma,;SLS,silverlinesystem; Body,width 37.13 4.70 12.66 30.00 45.00 15 arrow(inb),;arrow(ina),kinety;Bar:20 Length/Width (L/W) 1.04 0.04 4.24 1.00 1.11 15 µm. Macronucleus, length 17.87 3.36 18.79 12.00 23.00 15 Macronucleus, width11.40 1.84 16.18 8.00 15.00 15 Table5.Morphometriccharacterizationof Euplotesaediculatus. Micronucleus, diameter 3.87 0.52 13.34 3.00 5.00 15 – Character X SD CV Min Max N

– – Body,length 98.85 7.71 7.80 70.00 110.00 40 32.50µm(X =26.56,SD=4.99,CV=18.79,N=8;X Body,width 58.45 7.90 13.52 39.00 74.00 40 Length/Width =22.81,SD=5.08,CV=22.27,N=8)withtheL/W (L/W) 1.71 0.19 11.09 1.35 2.20 40 ratio1.19(1.00-1.67),almostsphericalwithan Dorsalkineties, number 10.14 0.58 5.73 8.00 11.00 29 equatorialgirdleofstiffcirri.Anadoralzoneof Adoralzoneof membranelles(AZM)surroundstheanteriorendofthe membranelles, length 68.56 3.63 5.30 62.00 77.00 16 cell.Macronucleusovoidwithasphericalmicronucleus, Adoralzoneof anditsdiameterisonaverage3.87µm. membranelles, number 51.78 4.74 9.15 45.00 58.00 9

Euplotesaediculatus PIERSON,1943 x110.00µm.Bodyovoidtoexpandedellipsoid;ventral (Figures2a-c;Table5) sideofthebodyisflat,dorsalsideconvexwith10.14 ThisspeciesbelongstotheorderHypotrichida,family (8.00-11.00)longitudinalkineties.Dorsalsilverline Euplotidae.Itssize,afterfixation,isapproximately70.00 systemisdouble-eurystomustype.Dorsalkineties

251 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions numberisdifferentinthosereportedbyFoissneretal. Table6.Morphometriccharacterizationof Spirostomumminus. (1991).Researchersfound8,rarely9kineties.The – adoralzoneofmembranelles,withlength68.56µm Character X SD CV Min Max N (62.00-77.00),contains45.00-58.00membranelles. Body,length MacronucleusC-shapedwithadjacentmicronucleus. (invivo) 808.80 126.10 15.59 490.00 950.00 13 Body,width (invivo) 38.08 6.30 16.54 30.00 50.00 13 AZM,length Spirostomumminus ROUX,1901 (invivo) 273.10 54.40 19.92 150.00 350.00 13 AZM/Bodylength (Figures3a,b;Table6) (invivo) 0.34 0.06 17.33 0.25 0.48 13 Macronucleus, Spirostomum spp.belongtotheorderHeterotrichida, number 18.31 7.80 42.60 4.00 38.00 39 familySpirostomidae.Bodyelongate,worm-likeshape, Macronucleus, highlycontractile.Sizeinvivo490.00-950.00x30.00- length 25.19 9.78 38.82 14.00 47.50 24 Macronucleus, 50.00µm,withtheL/Wratioonaverage21.52(16.33- width 8.77 4.89 55.77 4.00 20.00 24 28.33).Thelengthofadoralzoneofmembranelles (AZM)isonaverage273.10µm,extendsto34%ofthe Spirostomumteres CLAPARÈDE&LACHMANN, celllength.Macronucleusmoniliform,about4.00-38.00 1858 individualmacronuclearsegments,eachellipsoidal. Generalorganizationofthespeciesisconsistentwiththe (Figure4;Table7) previousdescriptions(AugustinandFoissner,1992; Ourspecimensmeasureinvivo300.00-490.00x Foissneretal.,1992). 25.00-80.00µm,withtheL/Wratio10.46(3.94-14.00) andnearlycorrespondtothosereportedbyFoissneretal. (1992)andAugustinandFoissner(1992).Macronucleus islocatedapproximatelyinmiddleofbody,andis ellipsoidal.Contractilevacuoleislocatedattheposterior endandacanalrunstowardstheanteriorofcell.AZMon

Figure3. Spirostomumminus (a,silverimpregnation;b,nuclear reaction).Ki.Kinety;Ma.Macronuclearsegments;Arrow. mouth;Bar:45µm. Figure4. Spirostomumteres,silverimpregnation.Ma,Macronucleus; Arrow,adoralzoneofmembranelles;Bar:30µm.

252 N.G.fiENLER,‹.YILDIZ

Table7.Morphometriccharacterizationof Spirostomumteres. Table8.Morphometriccharacterizationof Stentorcoeruleus.

– – Character X SD CV Min Max N Character X SD CV Min Max N

Body,length Body,length (invivo) 377.92 50.66 13.40 300.00 490.00 36 (invivo) 410.70 120.40 29.32 240.00 735.00 15 Body,width Body,width (invivo) 37.36 8.90 23.82 25 80.00 36 (invivo) 127.00 14.49 11.41 105.00 155.00 15 AZM,length Macronucleus, (invivo) 140.14 20.62 14.71 100.00 215.00 36 length 12.88 3.04 23.60 10.00 18.00 8 AZM/Bodylength Macronucleus, (invivo) 0.37 0.08 12.78 0.26 0.46 30 width 8.63 2.20 25.51 6.00 12.00 8 Macronucleus, Frontalfield, length 30.44 6.61 21.71 22.50 55.00 36 numberofkineties 24.20 4.27 17.64 20.00 30.00 5 Macronucleus, width 12.04 2.73 22.69 7.50 20.00 36 theleftbodyedgeisrelativelylong,extendsabout37% averageis15.21x10.4µm.Thereare24.20(20.00- ofcelllength,anditslengthonaverage140.14µm. 30.00)kinetiesinthefrontalfield(peristomialfield), enclosedbyadoralzoneofmembranelles.The morphologicalcharactersobservedonthespecieslargely Stentorcoeruleus (PALLAS,1766)EHRENBERG, correspondtothoseofFoissneretal.(1992),butour 1831 specimensarerathersmaller. (Figures5a,b;Table8)

ThisspeciesbelongstotheorderHeterotrichida, Parameciumbursaria (EHRENBERG,1831) familyStentoridae.Ourspecimensmeasureinvivo FOCKE,1836 240.00-735.00x105.00-155.00µm,withhighly contractileturquoisebody.Macronucleusmoniliform,the (Figures6a,b;Table9) numberofmacronuclearsegmentsisabout9,sizeon ThisspeciesbelongstotheorderHymenostomatida, family.Ourspecimensmeasureinvivo

Figure5. Stentorcoeruleus (a.invivo;b.silverimpregnation).Ma,macronuclearsegments;Cv,contractilevacuole;AZM,adoralzoneof membranelles;UM,undulatingmembrane;Triplearrows,frontalfield;Arrow,oralopening;Bar:35µm.

253 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Figure6. Parameciumbursaria,(a.invivo;b.silverimpregnation).Arrows,symbioticalgae;Ki,kinety;Ma,macronucleus;OA,oralapparatus;Bars: 25µm.

Table9.Morphometriccharacterizationof Parameciumbursaria. Frontoniaangusta KAHL,1931 (Figures7a-c;Table10) – Character X SD CV Min Max N ThisspeciesbelongstotheorderHymenostomatida, Body,length 143.19 15.13 10.57 120.00 172.50 21 familyFrontonidae.Ourspecimensmeasureinvivo – Body,width 88.33 14.84 16.71 70.00 115.00 21 125.00-205.00x40.00-142.50µm(X =156.10,SD= – Length/Width 19.97,CV=12.79,N=25;X =63.00,SD=14.03,CV (L/W) 1.63 0.18 11.19 1.33 2.07 21 Macronucleus, =22.37,N=25),withtheL/Wratio2.55(1.77-3.28), length 38.12 4.36 11.45 30.00 47.50 21 andarelargelydifferentfromthosefoundbyFoissneret Macronucleus, al.(1994).Bodyiscigar-shapedorscutiform,witha width 19.95 3.07 15.39 15.00 25.00 21 slighttaperingposteriorly.Nuclearapparatusis Micronucleus, representedbyelongatedmacronucleuswithone length 14.76 2.49 16.85 10.00 22.50 21 Micronucleus, micronucleus.Simple,vesicularcontractilevacuoleis width 7.20 0.80 11.10 5.00 7.50 21 locatedinthemiddleofthebody.Endoplasmbright Somatickineties, brown,withfoodvacuolescontainingdiatom,algae.Oral number 117.62 11.29 9.60 102.00 136.00 8 apparatustypicalfor(Foissneretal.,1994; AlekperovandAsadullayeva,1999).Foissneretal. – 95.00-137.50x45.00-70.00µm(X =118.68,SD= (1994)reported4vestibularkinetiesontherightofthe – 12.40,CD=10.45,n=19;X =57.63,SD=7,09,CD oralaperturebutthepresentresultsconflictedwiththeir =12.30,N=19),withtheL/Wratio2.08(1.73-2.60). view.Inourspecimensvestibularciliatureiscomposedof Comparedtothepopulationmorphometrically 3kineties.Somatickinetiesuniformwithpre-oraland characterizedbyFoissneretal.(1994),therewasno post-oralsutures,terminatingatposteriorpole.Somatic differenceinbodysize,butourspecimenshavemany kinetiesnumber78.00-113.00andarecomposedof somatickineties. pairedkinetosomes,with6-7postoralkineties.

254 N.G.fiENLER,‹.YILDIZ

Figure7. Frontoniaangusta ,silverimpregnation.Ki,kinety;VKi,vestibularkineties;UM, undulatingmembrane;M1-3,adoralemembranelles;Arrows,postoralkineties;c, silverlinesystem;Bar:20µm.

Table10.Morphometriccharacterizationof Frontoniaangusta.

– Character X SD CV Min Max N

Body,length 145.67 31.34 21.51 92.50 250.00 45 Body,width 75.00 21.81 29.08 40.00 142.50 45 Length/Width (L/W) 1.99 0.27 13.44 1.38 2.42 45 Macronucleus, length 35.56 8.75 24.61 22.50 55.00 27 Macronucleus, width 22.13 6.07 27.43 10.00 35.00 27 Somatickineties, number 97.92 9.46 9.66 78.00 113.00 26

Urocentrumturbo (MUELLER,1786)NITZSCH, 1827 (Figures8a-d;Table11) ThisspeciesbelongstotheorderHymenostomatida, familyUrocentridae.Cellsinvivo57.50-72.50x42.50- – – 55.00µm(X =64.00,SD=5.03,CD=7.86,N=19;X Figure8. Urocentrumturbo,silverimpregnation.Ma,macronucleus; =49.50,SD=4.05,CV=8.18,N=19)withtheL/W Mi,micronucleus;FP,frontalplate;AKi,kinetiesofanterior ratio1.30(1.21-1.39),ovaltospherical.Ourspecimens ciliaryband;PKi,kinetiesofposteriorciliaryband;Sc, arerelativelysmallerthanthoseofFoissneretal.(1994) scopula;OA,oralapparatus;Arrows,kinetiesofequatorial ciliaryband,c,tuft;d,scopula;Bar:20µm. andMartín-Gonzálezetal.(1986).Table11illustrates thebiometricalcharacterizationofourpopulation.The theposteriorpartofthecell.Infraciliaturewasdescribed nuclearapparatusconsistsofahorseshoe-shaped indetailbyFoissneretal.(1994)andMartín-Gonzáleset macronucleusandasphericalmicronucleus,situatedin al.(1986).Thesomaticinfraciliaturecomprisesanterior

255 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Table11.Morphometriccharacterizationof Urocentrumturbo.

– Character X SD CV Min Max N

Body,length 89.87 7.50 8.35 77.50 102.50 20 Body,width 84.00 9.01 10.73 67.50 100.00 20 Length/Width (L/W) 1.07 0.07 6.17 1.00 1.23 20 Macronucleus, length 34.37 5.61 16.32 22.50 42.50 20 Macronucleus, width 13.00 2.51 19.33 7.50 17.50 20 Macronucleus, length 33.12 5.25 15.85 20.00 40.00 20 Macronucleus, width 13.12 2.28 17.34 7.70 15.00 20 Micronucleus, length 3.33 0.62 18.51 3.00 5.00 15 Micronucleus, width 3.07 0.26 8.42 3.00 4.00 15 Tuft,length Figure9. Uronemanigricans,silverimpregnation.Ma,macronucleus; (invivo) 20.25 3.62 17.88 15.00 25.00 10 OA,oralapparatus;Ki,kinety;Bar:10µm. Scopula,kineties number 9.83 0.72 7.30 9.00 11.00 12 Table12.Morphometriccharacterizationof Uronemanigricans. Anteriorciliary band,kineties – number 148.50 3.89 2.62 142.00 155.00 13 Character X SD CV Min Max N

Body,length 43.29 6.16 14.23 34.00 51.00 7 ciliaryband,equatorialciliarygirdleandposteriorciliary Body,width 18.86 4.78 25.34 12.00 27.00 7 band.Distanceoftheunciliatedfrontallamtoanterior Length/Width ciliarybandisabout34.5µm.Theanteriorbandconsists (L/W) 2.40 0.64 26.67 1.89 3.75 7 Macronucleus, of148.50(142.00-155.00)kinetieswith19 length 8.57 1.27 14.84 7.00 11.00 7 kinetosomes,thefirst2ofwhichlieveryclosetoeach Macronucleus, other.Equatorialgirdlehasalmostthesamenumberof width 7.14 1.35 18.83 5.00 9.00 7 kinetiesastheanteriorone,andeachkinetyconsistsof5 Micronucleus, kinetosomes.Thekinetiesoftheposteriorciliaryband length 2.00 0.63 31.6 1.00 2.00 6 aremoreirregularinappearancethanthoseinthe Micronucleus, width 1.83 0.41 22.26 1.00 2.00 6 anteriorciliaryband.Intheposteriorciliaryband,there Unciliatedfrontal areveryargentophilickineties(ciliaryfieldorscopula) plate(bulge), consistingof9.83(9.00-11.00)kinetiesthatandin length 2.40 0.55 22.83 2.00 3.00 5 caudalcirrus(tuft).Thelengthofthetuft(invivo)is Somatickineties, 20.25µm(15.00-25.00). number 15.33 1.51 9.82 14.00 17.00 6

descriptions.Measurementsafterfixationaregivenbelow Uronemanigricans (MUELLER,1786)FLORENTIN, withbiometriccharacterization.Onaverage43.29-18.86 1901 µminsize;ovoid,withflattenedanteriorbulge(frontal (Figure9;Table12) plate),inanteriorthirdwithsmallindentationindicating ThisspeciesbelongstotheorderScuticociliatida, oralopeningwithasmalloralapparatus.Somaticciliature familyUronematidae.Wehavenodataonliving isonaveragecomposedof15.33longitudinalkineties. specimens.Thegeneralorganizationofthespecies,body Kinetiesbeginafterunciliatedfrontalplate,itslengthis shape,movement,nuclearapparatus,caudalcilium,and 2.40µm(2.00-3.00),terminatingaroundthecaudal contractilevacuolewereconsistentwiththeoriginal ciliumintheposteriorpole.

256 N.G.fiENLER,‹.YILDIZ

Dexiotrichagranulosa (KENT,1888)FOISSNER, approximately40.00x18.33µm,egg-shaped,about BERGER&KOHMANN,1994 twiceaslongasbroad,2.00x2.43.Oralapparatus (Figures10a,b;Table13) subapical,lightlyindented.Distanceanteriortomouthis 7.00µm(5.00-9.00).Macronucleus,sphericaltoovoid, ThisspeciesbelongstotheorderHymenostomatida, issituatednearlyinmid-body,withaspherical familyLoxocephalidae.Ourspecimensmeasureinvivo micronucleus.Thedistancebetweentheanteriorendand themacronucleusis28.89µm(19.00-37.00).Thereisa contractilevacuoleaboutinthemiddleofthecell.Somatic andoralinfraciliatureweregivenindetailbyAugustin andFoissner(1992)andFoissneretal.(1994). Longitudinalkinetiesextendfromunciliatedanteriorfield (bulge),itslengthisapproximately2.32µm,toposterior pole.Cytoplasmcontainsheavilyringshaped,refractile granules.Ourspecimensinmanycharacteristics correspondtothosefoundbyFoissneretal.(1994),but withfewerkineties.Somaticciliatureconsistsof20 (18.00-26.00)kineties.Inthisrespectourspecimens correspondto Dexiotrichatranquilla (Augustinand Foissner,1992).

Cyclidiumglaucoma MUELLER,1773 (Fiures11a,b;Table14) ThisspeciesbelongstotheorderScuticociliatida, Figure10.Dexiotrichagranulosa ,silverimpregnation.FP,frontal familyPleuronematidae.Sizeinvivo22.50-30.00x – plate;Ki,kinety,Arrows,ciliaryband;Bar:10µm. 10.00-15.00µm(X =25.16,SD=1.70,CV=6.76,N – =16;X =12.34,SD=1.70,CV=13.77,N=16),with Table13.Morphometriccharacterizationof Dexiotrichagranulosa. theL/Wratio2.07(1.67-2.50).Nuclearapparatus – Character X SD CV Min Max N consistsofasinglemacronucleusandaspherical micronucleus.Contractilevacuoleisterminallysituatedin Body,length 46.95 3.51 6.17 40.00 51.00 21 posteriorofcell.Whenwecompareourfindingswith Body,width 23.95 5.77 24.09 16.00 37.00 21 Length/Width thoseofFoissneretal.(1994),ourspecimensareslightly (L/W) 2.05 0.39 19.98 1.30 2.77 21 largerandshowawiderrangeofsomatickineties Macronucleus, number,13.20(11.00-15.00).Foissneretal.(1994) length 11.15 2.21 19.79 7.00 15.00 20 observed10-11longitudinalkineties.Unciliatedfrontal Macronucleus, field,orbulge,isconspicuouswith3.84µm(3.00-5.00) width 8.40 2.14 25.44 5.00 13.00 20 Micronucleus, length.Somatickinetiesarearrangedlongitudinally, length 2.50 0.76 30.44 2.00 4.00 20 whichareusuallydikinetidsintheanteriorofeachrow. Micronucleus, Allkineties,withtheirlength20.31µm(13.00-31.00), width 2.20 0.62 28.00 1.00 4.00 20 havemoreorlessshortenedinposteriorportionmaking Distanceanterior alargeunciliatedposteriorfieldthelengthofwhichis endtooralopening 7.00 1.53 21.83 5.00 9.00 7 Distanceanterior 5.69µm(3.00-8.00).Oralapparatusistypicalforthe 1 endtomacronucleus 28.89 4.65 16.10 19.00 37.00 18 genus Cyclidium,extendingtoabout /2 ofcelllength, Unciliatedfrontal withprominentundulatingmembraneonitsright field(Bulge),length 2.32 0.67 28.97 2.00 4.00 19 margin.Table14illustratesthebiometrical Bulgelength/body characterizationofourspecimens.Song(2000) length 0.05 0.02 33.88 0.04 0.10 19 Somatickineties, examined Cyclidium spp.bycomparingthepopulations number 20.00 2.57 11.28 18.00 26.00 12 morphometrically.

257 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Urotrichaglobosa SCHEWIAKOFF,1892 (Figures12a-c:Table15) ThisspeciesbelongstotheorderProstomatida,family Plagiocampidae.Ourspecimensmeasureinvivo12.50- – 25.00x10.00-22.50µm(X =19.67,SD=3.76,CV= – 19.14,N=15;X =17.17,SD=4.3,CV=25.16,N= 15)withtheratio1.77(1.00-1.50)andarealmost spherical.Themorphologicalcharactersdeterminedby liveobservationareconsistentwiththosereportedby

Figure11.Cyclidiumglaucoma,Silverimpregnation.FP,frontalplate; Ki,kinety;UM,undulatingmembrane;Arrows,adoral membranelles(M1-3);Ma,macronucleus;Mi,micronucleus; Cc,caudalcilium;b,binaryfission;Bar:10µm.

Table14.Morphometriccharacterizationof Cyclidiumglaucoma. Figure12.Urotrichaglobosa,silverimpregnation.OA,oralapparatus; – Ki,kinety;Ma,macronucleus;Mi,micronucleus;Cc,caudal Character X SD CV Min Max N cilium;Arrows,brosse(adoralorganelle);Bar:10µm. Body,length 28.33 3.36 11.84 21.00 35.00 36 Body,width 21.53 3.33 15.45 11.00 27.00 36 Length/Width(L/W) 1.34 0.22 16.17 1.08 2.36 36 Table15.Morphometriccharacterizationof Urotrichaglobosa. Macronucleus,length 9.70 1.88 19.38 6.00 13.00 33 – Macronucleus,width 7.58 1.69 22.15 5.00 12.00 33 Character X SD CV Min Max N Micronucleus,diameter 2.46 0.52 21.08 2.00 3.00 13 Unciliatedfrontalplate Body,length 24.79 4.58 18.48 17.00 34.00 28 (Bulge),length 3.84 0.60 15.67 3.00 5.00 19 Body,width 23.14 4.69 20.26 15.00 32.00 28 Distanceanteriorend Length/Width(L/W) 1.08 0.11 10.47 1.00 1.56 28 tooralapparatus 7.89 3.52 44.61 4.00 12.00 9 Distanceanterior Macronucleus,length 6.29 1.61 25.56 4.00 11.00 28 endto 21.12 3.00 14.20 15.00 25.00 8 Macronucleus,width 5.29 1.08 20.51 4.00 7.00 28 Oralapparatus,length 13.62 4.31 31.64 8.00 20.00 8 Micronucleus,length 2.32 0.67 28.87 1.00 4.00 28 Kinetosomesinparoral Micronucleus,width 2.18 0.77 35.43 1.00 4.00 28 membranella,number 37.56 9.82 26.14 16.00 49.00 9 Caudalcilium,length 11.31 1.65 14.62 9.00 13.00 13 Somatickineties, Somatickineties, number 13.20 0.99 7.53 11.00 15.00 35 number 22.15 2.77 10.23 18.00 26.00 26 Somatickinety,length 20.31 4.27 21.02 13.00 31.00 13 Somatickinety,length 14.80 1.42 9.62 13.00 18.00 15 Somatickinety,number ofkinetosomes 13.13 1.45 11.08 12.00 16.00 24 Somatickinety,number Unciliatedfieldin ofkinetosomes 16.00 2.92 18.22 11.00 20.00 9 posterior,length 5.69 1.25 21.98 3.00 8.00 16 Bodylength/somatic Caudalcilium,length 13.89 1.36 9.82 11.00 15.00 9 kinetylength 1.60 0.13 8.13 1.33 1.80 15

258 N.G.fiENLER,‹.YILDIZ

Foissneretal.(1994).Nuclearapparatusconsistsofa Table16.Morphometriccharacterizationof Colepshirtus. – sphericalmacronucleusandamicronucleussituatedinthe Character X SD CV Min Max N mid-body.Contractilevacuoleinposteriorend. Infraciliatureistypicalforthegenus Urotricha.Somatic Body,length 50.47 5.65 11.19 40.00 62.00 36 kinetiesareinalongitudinalarrangementandcomposed Body,width 28.61 6.36 22.23 18.00 48.00 36 of22.15(18.00-26.00)with11.00-16.00kinetosomes, Length/Width(L/W) 1.82 0.32 17.10 0.94 2.50 36 extendingtoaverage14.80µmofthecelllengthfrom Macronucleus,length 11.05 1.62 14.61 8.00 13.00 19 theanteriorend,therestofthebodyisnotciliated.The Macronucleus,width 10.47 1.78 16.95 7.00 13.00 19 Micronucleus,length 2.83 0.75 26.58 2.00 4.00 6 lengthofcaudalciliuminvivo,whichiscentrallylocated Micronucleus,width 2.67 0.82 30.60 2.00 4.00 6 atposteriorpole,isabout11.31µm(9.00-13.00). Somatickineties,number 14.67 0.97 6.59 14.00 16.00 21 Brosse(adoralorganelle)composedof3kineties, anteriorrowwith6kinetids,middlerowwith5to6 kinetids,posteriorrowwith6kinetids. posteriorside–plateseachwith2windows,anteriorand posteriormid-plateswith4windows.Platestructureis thesameasthatof C.amphacanthus (Foissnerand Colepshirtus (MUELLER,1786)NITZSCH,1827 O’Donoghue,1990)butthenumberofwindowsis (Figures13a,b;Table16) differentfromthatofC.amphacanthus.Thewindowsof C.hirtus areverysimilarinshapetothoseoftheother ThisspeciesbelongstotheorderProstomatida,family species, C.amphacanthus and C.spetai (Foissnerand Colepidae.Ourobservationsonthelivematerialusually O’Donoghue,1990;Foissneretal.,1994),whichare coincidewiththoseofFoissneretal.(1994),butour barrel-shaped.InfraciliatureisgivenindetailbyFoissner specimensareslightlysmallerandtheirsizesinvivoare – etal.(1994)andFoissner(1984b);somaticinfraciliature 37.50-55.00x22.50-25.00µm(X =47.03,SD=5.10, – iscomposedof14.67(14.00-16.00)longitudinal CV=10.84,N=16;X =24.22,SD=1.20,CV=4.94, kineties.Sphericalmacro-andmicronucleusareaboutin N=16),withtheL/Wratio1.94(1.50-2.22).Thebody themid-body.Contractilevacuoleisintheposterior. iselongatedandbarrel-shapedtocylindrical.Numberof windowsinpellicularplatescorrespondtothosegivenby Foissneretal.(1994)andFoissner(1984b):anteriorand Lagynuselegans (ENGELMANN,1862) QUENNERSTEDT,1867 (Figures14a-c;Table17) Thisspeciesiswellknownasamemberofthebottom faunabutitssystematicpositionisnotyetclear.The speciesbelongstotheorderGymnostomatida,family Lacrymariidae(Foissneretal.,1995).However,Foissner (1988)recommendedthatthegenus Lagynus shouldbe includedinthefamilyMetacystidae(Prostomatida).Sola etal.(1990)suggestedthatthisgenusbelongstoanew family,Lagynidae,intheorderProrodontida,becauseof theabsenceofloricaandthepresenceofabrosse.The generalorganizationofthespecieswasconsistentwith somedescriptions(Solaetal.,1990;Foissneretal., 1995).Thecellof L.elegans isprolongedamphoraor lamp-chimneyinshape.Anteriorendofthecellis annulated.Thecellconsistsof3parts:anteriorcone,its size6.76x9.62µm,andtheoralopeningislocated there,themainpartofthecellisthewidestpartofthe Figure13.Colepshirtus (a,invivo,b,silverimpregnation).M,mouth; cell,theposteriorpartofthecellisnarrowerand T,thorn;Arrow,windowsinarmorplates;Bar:10µm.

259 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Figure14.Lagynuselegans ,silverimpregnation.Ma,macronucleus; Pe,circumoralinfraciliature;Arrows,nematodesmata,c, silverlinesystem;Bar:15µm.

Table17.Morphometriccharacterizationof Lagynuselegans. – Character X SD CV Min Max N

Body,length 75.15 11.63 15.48 56.00 103.00 27 Body,width 31.04 7.54 24.28 22.00 57.00 27 Length/Width(L/W) 2.49 0.42 16.68 1.79 3.24 27 Anteriorcone,length 6.76 1.20 17.75 5.00 9.00 25 Anteriorcone,width 9.62 2.25 23.36 6.00 15.00 21 Vacuole,length 30.48 7.63 25.16 19.00 45.00 25 Somatickineties, number 29.56 3.43 11.60 26.00 38.00 9 flattened,andalargecontractilevacuoleispresent.Its lengthis30.48µm(19.00-45.00).Macronucleusis reniform,locatedinmid-body,withamicronucleus,its lengthabout17µmand14µm.Somaticinfraciliatureis composedof29.56(26.00-38.00)longitudinalkineties, withsinglekinetosome.

Didinumnasutum (MUELLER,1773)STEIN,1859 (Figures15a,b;Table18) ThisspeciesbelongstotheorderSpathidiida,family Didinidae.Ourspecimensmeasureinvivo80.00-107.50 Figure15.Didinumnasutum ,silverimpregnation.AC,apicalcone; – x45.00-67.50µm(X =92.31,SD=7.32,CV=7.93,N ACb,anteriorciliaryband,PCb,posteriorciliaryband;Ma, – macronucleus,Arrows,extrusomes;Bars:25µm. =13;X =54.23,SD=6.49,CV=11.97,N=13)with theL/Wratio1.73(1.28-2.11).Generalorganizationof aresmaller.With2ciliarybands:1anteriorcomposedof thespeciesisconsistentwiththosereportedbyFoissner about83kinetiesand1equatorialcomposedofabout70 etal.(1995)andFoissner(1984b),butourspecimens kineties.Macronucleusishorseshoeinshapeandlocated

260 N.G.fiENLER,‹.YILDIZ

Table18.Morphometriccharacterizationof Didiniumnasutum. Table19.Morphometriccharacterizationof Monodiniumbalbiani. – Character X SD CV Min Max N – Character X SD CV Min Max N Body,length 104.89 17.89 17.06 82.00 135.00 18 Body,length 83.55 18.63 22.30 49.00 120.00 33 Body,width 76.61 16.33 21.32 52.00 105.00 18 Body,width 67.76 16.75 24.72 35.00 93.00 33 Length/Width(L/W) 1.38 0.11 7.63 1.22 1.58 18 Length/Width(L/W) 1.25 0.10 7.81 1.12 1.57 33 Apicalcone,length 9.94 1.47 14.82 7.00 13.00 18 Apicalcone,length 8.68 1.82 20.92 6.00 13.00 31 Apicalconelength/body Apicalconelength/body length 0.10 0.02 21.45 0.06 0.13 18 length 0.11 0.03 23.92 0.07 0.17 31 Distancebetween2 Somatickineties, ciliarybands 44.15 6.59 14.93 35.00 60.00 13 number 78.83 5.12 6.49 75.00 88.00 6

– cilia.Sizeinvivoabout50.00-87.50X37.50-50µm(X nearthemid-body,itswidthisapproximately20.33µm, – withasphericalmicronucleus(6.00-5.33).Anteriorpart =65.38,SD=10.94,CV=16.73,N=13;X =49.04, oftheciliateisflattenedandhasaprojectingconewith SD=6.58,CV=13.42,N=13)withtheL/Wratio1.34 mouthatthetipofbody,itslengthonaverage9.94µm (1.14-1.75).Somatickinetiesonciliarybandnumber (7.00-13.00). 78.33(75.00-88.00).Macronucleuswidth15.8µm, diameterofthemicronucleus4µm.

Monodiniumbalbiani FABRE-DOMERGUE,1888 Plagiopylanasuta STEIN,1860 (Figure16;Table19) (Figures17a,b;Table20) M.balbiani isverysimilarto Didiniumnasutum , differingfromthatspeciesmainlywithasinglebandof ThisspeciesbelongstotheorderGymnostomatida, familyPlagiopylidae.Ovoidshape,afterfixationon average91.82x42.42µminsize.Oralopening

Figure16.Monodiniumbalbiani,silverimpregnation.AC,apicalcone; Figure17.Plagiopylanasuta,silverimpregnation.OA,oralapparatus; Cb,ciliaryband;Arrows,extrusomes;Bar:25µm. Ki;kinety;Sb,stripedband;Bar:20µm.

261 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions

Table20.Morphometriccharacterizationof Plagiopylanasuta.

– Character X SD CV Min Max N

Body,length 91.82 17.48 19.04 67.00 135.00 25 Body,width 42.42 15.74 37.11 20.00 80.00 25 Length/Width(L/W) 2.32 0.52 22.39 1.59 3.50 25 Distanceanteriorend tooralopening 17.86 4.26 23.85 11.00 27.50 25 Mouthindentation, length 21.29 6.90 32.40 12.00 38.00 17 Oralopening,length 5.37 0.83 15.48 5.00 8.00 19 Somatickineties, number 52.83 4.22 7.99 48.00 60.00 6 subapical,onaverage5.37µmwidth,atrightanglesto longaxisofthecell.Distancebetweenanteriorendofcell andoralopeningisonaverage17.86µm(11.00-27.50). Thelengthofmouthindentationis21.29µm(12.00- 38.00).Stripedbandcurvesfromvestibuleandextends alongdorsalside.Macronucleusovoid,34x22µm,with asphericalmicronucleus,itsdiameterisapproximately Figure18.Amphileptuspleurosigma (a.silverimpregnation;b,invivo). 2.5µm.Thespeciespossessesonaverage52.83(48.00- E,extrusomes;Ma,macronuclearsegments;Arrows, contractilevacuoles;Bars:40µm. 60.00)kineties.Ourobservationsonthegeneral organizationofthespecieswereconsistentwiththoseof Table21.Morphometriccharacterizationof Amphileptuspleurosigma. Solaetal.(1988)andFoissneretal.(1995). – Character X SD CV Min Max N

Body,length 187.80 40.00 21.30 110.00 250.00 22 Amphileptuspleurosigma (STOKES,1884) Body,width 50.02 15.03 30.05 25.00 75.00 22 FOISSNER,1984 Length/Width(L/W) 3.91 0.74 18.96 2.59 5.71 22 Anteriormacronucleus, (Figures18a,b;Table21) length 19.08 4.73 24.79 12.00 27.50 18 ThisspeciesbelongstotheorderPleurostomatida, Anteriormacronucleus, width 13.03 2.63 20.16 9.00 17.50 18 familyAmphileptidae.Ourspecimensinvivomeasure – Posteriormacronucleus, 150.00-280.00x37.50-62.50µm(X =228.50,SD= – length 18.94 4.36 23.02 12.00 25.00 18 36.63,CV=16.03,N=15;X =53.00,SD=8.30,CV Posteriormacronucleus, =15.66,N=15),withtheL/Wratio4.36(3.18-5.83). width 13.00 2.31 17.75 10.00 17.50 18 Distancebetween Thebodyisspindle-like,taperinganteriorlyand macronuclearsegments 22.10 6.09 27.56 11.00 35.00 15 posteriorly,moderatelycontractile.Macronuclear Somatickineties,number 19.18 4.29 22.37 13.00 27.00 11 segmentslocatedapproximatelyinmiddleofbody, elipsoidal,almostequalinsize(19.08x13.03µm;18.94 Ontherightside,approximately19.18(13.00-27.00) x13.00µm).Distancebetween2macronuclearsegments kinetiesconvergeoneachotherintheanteriorregion. isonaverage22.10µm(11.00-35.00).Single micronucleusisellipsoidal(10x5µm),positioned Loxedesstriatus (ENGELMANN,1862)PENARD, betweenmacronuclearsegments.Manycontractile 1917 vacuolesarepresent,locatedalongdorsalandventral edges.Extrusomesarearrangedalonganteriorend,and (Figure19;Table22) dispersedthroughoutcytoplasm.Infraciliatureistypical ThisspeciesbelongstotheorderKaryorelictida, forthegenus(Foissner,1984a;Foissneretal.,1995). familyLoxodidae.Sizeinvivo150.00-50.00x50.00-

262 N.G.fiENLER,‹.YILDIZ

Table22.Morphometriccharacterizationof Loxedesstriatus. – Character X SD CV Min Max N

Body,length 79.43 13.02 16.39 62.00 120.00 23 Body,width 23.87 3.48 14.58 17.00 34.00 23 Length/Width(L/W) 3.34 3.39 11.61 2.76 4.29 23 Anteriormacronucleus, length 4.65 0.49 10.47 4.00 5.00 23 Anteriormacronucleus, width 4.44 0.51 11.43 4.00 5.00 23 Posteriormacronucleus, length 5.04 0.37 7.27 4.00 6.00 23 Posteriormacronucleus, width 4.61 0.50 10.83 4.00 5.00 23 Anteriormicronucleus, length 1.78 0.42 0.09 1.00 2.00 23 Anteriormicronucleus, width 1.44 0.51 35.33 1.00 2.00 23 Posteriormicronucleus, length 1.87 0.34 18.42 1.00 2.00 23 Posteriormicronucleus, width 1.52 0.51 33.57 1.00 2.00 23 Distancebetween macronuclearsegments 20.26 5.37 26.51 12.00 32.00 23 Oralopening,length 22.54 3.67 16.28 16.00 30.00 13 Cytopharynxlength 14.60 4.33 29.66 9.00 20.00 10 Bodylenght/oralopening length 4.26 0.50 11.75 3.67 5.20 13 Bodylength/cytopharynx Figure19.Loxedesstriatus ,silverimpregnation.OA,oralapparatus; length 7.27 2.15 29.58 5.00 12.22 10 Arrows,Müllervesicles;Bar:20µm. Somatickineties,number 9.34 1.10 11.72 8.00 12.00 32

75.00µm,withtheL/Wratio2.92(2.67-3.08). Flattenedciliateswithacrescent-shapedmouth,itslength onaverage22.54µm(16.00-30.00),situatedanteriorly alongtheventraledge.Oralapparatusendswithan obviouscytopharynx,itslengthonaverage14.60µm (9.00-20.00).Nuclearapparatuscomposedof2diploid macronuclei,almostsphericalandequalinsized,and2 sphericalmicronuclei.Distancebetweenmacronuclear segmentsisonaverage20.26µm(12.00-32.00). Somatickinetiesarebipolar.Theyrunfromtheanterior totheposteriorend,andconsistofpairedkinetosomes. Thereare9.34(8.00-12.00)kinetiesontherightside andonly2ontheleftside,aventralanddorsalone. Cytoplasmgranulated,containingMüllervesicles,a peculiarorganellecharacteristicofloxodidciliates,and amplepigmentgranules,locatedparalleltothekineties. ThestructureandfunctionofMüllervesicleswere examinedindetailbyFenchelandFinlay(1986). Inthisstudy,descriptionsoftheperitrichciliatesare basedononlyliveobservations. Epistylisdigitalis (LINNAEUS,1758)EHRENBERG,1830(Peritrichida, Epistylididae)iscolonialwithanon-contractilestalk,and epibiontoncopepods.Invivoitmeasures60.00-87.50x – 25.00-37.50µm(X =72.88,SD=9.00,CV=12.35,N – Figure20.Carchesiumpolypinum (MSF).M,myonema;Arrows, =13;X =28.46,SD=3.61,CV=12.68,N=13),with discontinuousmyonema;Bar:60µm.

263 FaunisticandMorphologicalStudiesonCiliates(Protozoa,Ciliophora)fromaSmallPond,withResponsesofCiliatePopulations to ChangingEnvironmentalConditions theL/Wratio2.58(2.17-3.30). Carchesiumpolypinum specimensmeasureinvivo70.00-112.50x50.00-72.50 – – (LINNAEUS,1758)EHRENBERG,1830(Peritrichida, µm(X =87.73,SD=16.26,CV=18.53,N=11;X = Vorticellidae)(Figure20)iscolonialwithacontractile 54.09,SD=18.04,CV=33.35,N=11)withtheL/W stalk.Myonemainthestalkisdiscontinuous.Our ratio2.65(1.30-1.54).

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