Effects of water quality and hydrologic driverson periphyton colonization on Sparganium erectumin two Turkish lakes with different mixing regimes

Meriç Albay & Reyhan Akçaalan

Received: 27 July 2007 /Accepted: 6 November 2007 / Published online: 10 January 2008# Springer Science + Business Media B.V. 2007

Abstract The aim of this paper is to determine andcompare the environmental factors controlling verticalcolonisation of periphyton on Sparganium erectum ina shallow eutrophic turbid lake, Manyas Lake, and anoligo-mesotrophic deep lake, Sapanca Lake, Turkeyduring the July 1997–November 1998. To investigatethe effect of the environmental factors on periphytoncolonization on S. erectum, the stem was cut abovethe rhizomes and subdivided into three equal sections.Multivariate statistical analyses have been applied toclarify relationships between environmental vari-ables and periphyton colonization on S. erectum.Results indicated that physical disturbance andtrophic level of the lakes influenced the colonizationof the periphyton. Among the measured parameters,low light intensity, total suspended solids, tempera-ture and water level fluctuation were observed asdriving factors in Manyas Lake whereas nutrientdeficiency was found as key factor in Sapanca Lake.The zonation of the periphyton, density, compositionand dominant/subdominant taxa were significantlydifferent in these lakes. However, Oedogonium sp.,Mougeotia sp., Cylindrocapsa sp., Cladophora glom-erata (Linn.), Aulacoseira italica (Ehr.) Simonsen,

Melosira varians C. Agardh, Navicula tripunctata(O.F. Müller) Bory and Fragilaria ulna (Nitzsch)Lange-Bertalot were found as dominant species at allsections of S. erectum both in Manyas Lake andSapanca Lake. It can be thought that these specieshave a broad range of tolerance to several physical,chemical and hydrologic disturbances. This is thefirst study to introduce how much the water qualityand hydrologic drivers have affected vertical coloni-zation of periphyton on S. erectum in two lakes withdifferent mixing regimes.

Keywords Periphyton colonization . Biomonitoring .

Manyas lake . Sapanca lake . Sparganium erectum .

Water quality

Introduction

The littoral zone is regarded very complex area bothof deep and shallow lakes. Studies show that thelittoral region provides almost entire production of thelakes (Meulemans 1998; Kiss et al. 2003), as animportant food source for invertebrates and fish. Theperiphyton is the most important component withmultiple interactions among the organisms in thissystem. It provides high level of primary productivityand a major regulator of nutrients fluxes. Besides laketrophic status, the quantity of the periphyton produc-tion depends on the multiple biotic and abiotic factors

Environ Monit Assess (2008) 146:171–181DOI 10.1007/s10661-007-0069-5

M. Albay (*) : R. AkçaalanFaculty of Fisheries, Istanbul University,Ordu Cad. No: 200 Laleli,34470 Istanbul, Turkeye-mail: merbay@istanbul.edu.tr

including light intensity, temperature, water levelfluctuation, type of the substratum, wave actions,biotic interaction, intra- and inter-specific competi-tion, grazing, host plant architecture etc. Quantity ofthe periphyton and species richness may closelyreflect the trophic state of lakes (Scheffer 1998;Meulemans 1998; Hill et al. 1995).

Light intensity is usually considered to be the mostderiving abiotic factor regulating the periphytoncolonization in lakes, particularly in shallow eutrophiclakes. Irradiance level can affect the epiphytic algalproductivity (Hill et al. 1995). The influence of thelight intensity depends upon the solar energy thatpenetrates the water column in the littoral zone aswell the self-shading within the periphyton (Müller1996). It is worthwhile to point out that most part ofthe light can be either absorbed or scattered into adifferent direction in shallow lakes. Abundance ofmacrophytes and periphyton decreases with turbidityin the shallow areas under the very turbid conditions(Scheffer 1998).

Nutrient availability is the most important factorcontrolling periphyton; several studies demonstratedthat when aquatic ecosystems are subjected toeutrophication, a subsequent loss of diversity can bedetermined (Cederwall and Elmgren 1990; Wormet al. 1999). However, there are controversial studiesthat nutrient enrichment can either increase ordecrease species diversity (Pringle 1990; Miller etal. 1992). Several workers reported that some domi-nant species can give a brief idea about the waterquality. For instance, Marker and Collett (1997) statedthat Cocconeis placentula Ehr. grows well under awide range of conditions and Niztschia amphibia,Niztschia palea and Gomphonema parvulum aregenerally recognised to be indicators of pollutedwaters. They also concluded that Achnanthes minu-tissima, Navicula minisculus, Nitzschia dissipata andFragilaria ulna are generally found in less pollutedbut eutrophic waters.

Periphyton is widely used as an indicator of waterquality studies since it is sensitive to both hydrologicand water quality changes, and responds rapidly tothese changes (Gaiser et al. 2004) and responses ofthis community to pollutants can be measured at avariety of time scales determining the physiological ofthe community-level changes (Strand and Weisner1996; Buczkó and Ács 1998; Ács et al. 2000).Therefore, structural and functional characteristics of

the periphyton have been used for several decades asindicators of water quality (Biggs 1989; Albay andAkçaalan 2003). Vis et al. (1998) stated that thecharacteristics of periphyton that respond to changesin water quality are not always the same. Theyemphasized that certain studies showed the correlationbetween indices of diversity or species autoecolgy andwater quality whereas other studies revealed strongerresponses of biomass indices (Biggs 1989). Accord-ingly, Albay and Akçaalan (2003) reported that somespecies (such as Aulacoseira italica, Melosira var-ians, and Planktolyngbya limnetica) have a broadrange of tolerance to physical disturbance includingwater level fluctuation, large amounts of suspendedsolids, and low Secchi disc depth.

This paper aimed to compare the environmentalfactors and hydrologic conditions controlling verticalcolonisation of periphyton on Sparganium erectum ina shallow eutrophic turbid lake (Manyas Lake) and anoligo-mesotrophic deep lake (Sapanca Lake). Thespecific goal was to clarify the possibility of the usageof the periphyton as bioindicators of anthropogeniccontamination in two lakes with different mixingregimes.

Materials and methods

Study sites

Descriptions of the studied lakes and complete data-sets are reported in Albay et al. (2003) and Albay andAkçaalan (2003). The main morphological andhydrological characteristics of the lakes are summa-rized in Table 1, Manyas Lake is a shallow-turbid lakewith a surface area of 159 km2. The investigated areaof the lake is covered with mostly emerged macro-phytes particularly, Phragmites australis, Typha sp. S.erectum and has a large littoral region where there islittle water movement because of dense macrophytestand, and is surrounded by agricultural land. ManyasLake is one of the most important world wide naturalreserve (Ramsar site) of Turkey because of itsecological values. The lake lies on a migratory waterbirds route and is rich in food for birds. The lakesupports 23 fish species, crayfish, otter, turtles, watersnakes etc. (Balik 1989; Erkakan 1997; Çelik 2006).

Sapanca is a moderately deep lake, located in thenortheast of the Marmara region of Turkey, has a

172 Environ Monit Assess (2008) 146:171–181

55 m maximum depth, 46.8 km2 surface area and ca.250 km2 catchment area. The lake is fed by severalcreeks and significant amount of ground water(Tuğrul and Morkoç 1991). The catchment area wasused for agricultural activities. The unexpecteddevelopment of tourism and recreation in the begin-ning of the 1990s has given rise to excessive nutrientinput to the lake. Then, first fish kill was observedafter the luxuriant growth of Planktothrix rubescensin 1997 (Albay et al. 2003). At present, the lakeprovides drinking water for the district of Sakarya. Itis also used for agriculture and abstracted for industrialuse.

Periphyton and chlorophyll-a

The samples of the periphyton were collected fromthe stems of the S. erectum from the Sapanca andManyas Lake between July 1997 and November 1998at monthly intervals. In a few cases, samples were notcollected for practical reasons during the studiedperiod. Each of the S. erectum stem was cut abovethe rhizome and subdivided into three equal sections(25 cm each) called “upper,” “middle” and “lower”ones. Periphyton matt was removed manually andcollected in 200 ml distilled water. A one hundredmillilitres of periphyton-containing water quantita-tively subsample for algal counting according toUtermöhl (1958). For each course, 600–800 cellswere counted. To determine the diatoms, watersamples were treated with H2O2. The remainingslurry (100 ml) filtered from the glass-fibre filters(Whatman GF/C) and chlorophyll-a measured accord-ing to Nusch (1980). The water level fluctuation

(WLF) data was kindly provided by the GeneralDirectorate of Electrical Power Resources Survey andDevelopment Administration. The methods for deter-minations of the epiphytic algal biomass, algaltaxonomy and water quality parameters have beendescribed by Albay and Akçaalan (2003).

Statistical analysis

The relationships between epiphytic algal groups, chl-a,and water quality parameters were explored byCanonical Correspondence Analysis (CCA) usingCANOCO 4.5 software (ter Braak and Smilauer2002). Relationship between environmental parametersand dominant species were evaluated using Spearmanrank correlation test. Changes in periphyton dynamicswere examined using Shannon’s diversity index, H′,calculated from the biovolume data as;

H' ¼Xs

i¼1

pi log2 pi

Where pi is the relative biovolume of species and i ands are number of species (Sommer 1993). Equitability(E) was calculated as H/Hmax.

Results

Manyas lake

The results presented in Table 2 shows the physicaland chemical conditions of Manyas Lake. The lakewater has showed eutrophic characters with lowphotic depth (zeu), high concentration of nutrients,conductivity and TSS because of intense sediment–water contact during the day. The CCA ordinationdemonstrated the importance of the different environ-mental factors in determining dominant epiphyticalgal species. The position of the dominant speciese.g. Rhoicosphaenia curvata (Kütz.) Grun. ex Rabh.,Nitzschia linearis (Agardh) W. Smith, Stigeocloniumsp., Cylindrocapsa sp., Limnothrix sp. were placedclosed to temperature and pH vectors; Melosiravarians C. Agardh and Nitzschia palea (Kütz.) W.Smith were closed to NO2+NO3, Secchi depth, o-PO4

and Chl-a vectors; Pediastrum boryanum (Turpin)Meneghini and Aulacoseira italica (Ehr.) Simonsenwere placed to near the TSS vector in the ordination

Table 1 Morphological characteristics and locations ofManyas and Sapanca lakes

Parameters Unit Manyas Lake Sapanca Lake

Coordinates 40°12′N–28°00′E 30°15′E–40°42′NAltitude M 18 30Maximum depth M 3.60 55Volume m3 265.3×106 1.2×109

Catchment area km2 3,352 250Surface area km2 159 45Retention time Year 0.71 UnknownMixing type Polymictic Warm-monomicticSpecial status National parka No special status

a Ramsar side

Environ Monit Assess (2008) 146:171–181 173

diagram of CCA (Fig. 1). Only 21 dominant algalspecies are shown in Fig. 1. The eigenvalues of CCAaxis 1 (0.35) and axis 2 (0.25), constrained to theeight variables account for 50% of the cumulativevariance in the species data. The species–environmentcorrelations of CCA axis 1 and 2 are high and the firsttwo axis account for 59.3% of the variance in theepiphytic algae–environment relationships (Table 3).

The seasonal variation of chlorophyll-a (chl-a)concentration showed considerable difference amongthe sections (p<0.05; Fig. 2). Mean chl-a concentra-tion was 21.4, 37.5 and 31.2 µg cm−2 on lower,middle and upper sections, respectively. It showedsimilar seasonal trend with algal biovolume at allsections (r=0.68, 0.72, 0.58; p<0.01 on lower,middle and upper sections). In the samples obtainedduring the 1997, maximum chl-a was recorded in Julyon the middle section as 59.4 µg cm−2 and minimumchl-a was measured in November on lower section as9 µg cm−2.

A total 123 species of periphyton were identified.Periphyton assemblages on S. erectum were identicalmostly in the eutrophic lakes. The predominantspecies at all sections were mostly filamentousmulticellular green algae, Oedogonium sp. and Sti-geoclonium sp., Cylindrocapsa sp., Spirogyra sp.,Mougeotia sp. In addition to these species, filamen-tous diatom, Aulacoseira italica, A. muzzanensis(Meist.) Krammer, filamentous blue-green algae, P.limnetica Lemm., loosely attached taxa F. ulna, theadnate species Navicula tripunctata, Surirella linearisW. Smith, and Chlorococcal green algae Pediastrum

Table 2 The physical, chemical and biological characters of Sapanca and Manyas lake between July 1997 and November 1998

Manyas Lake Sapanca Lake

Parameters Unit Mean (max–min) Mean (max–min)

Temperature °C 18,6 (7.1–27.1) 19.9 (8–27)pH 8.1 (7.4 –8.9) 8 (7.8–8.4)Conductivity µS cm−1 2,830 (980 –8,860) 205 (157–253)TSS mg l−1 94.4 (61–134) 3.89 (0.6 –6.8)SRP µg l−1 168 (40.4–324) 5.2 (2–16)TP µg l−1 628 (164 –922) 12.4 (4.5–38)NO2+NO3 µg l−1 614 (188–1,144) 27.1 (2–92)Secchi depth cm 21.6 (16–34) 276 (100 –380)Zeu cm 60.5 (44.8–95.2) 773 (280 –1,064)Total biomass mm3 cm−2 4.21 (0.17–11.35) 14.79 (2.29–31.86)Shannon diversity H′ 3.01 (2–3.7) 3.34 (2.4 –4.4)Number of species n 22 (13– 40) 27 (15–38)

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pedsim

fraul

moug

plali

pedbor

stigeTemp.

pH

TSS

o-PO4

TP

NO2+NO3

WLF

Sec dep.

Fig. 1 Biplot of first and second CCA axis for periphyton inManyas Lake. Environmental variables: achlin, Achnantheslinearis; aulit, Aulacoseira italica; cyli, Cylindrocapsa; cymlan,Cymbella lanceolata; fraul, Fragilaria ulna; limno, Limnothrix;melvar, Melosira varians; natri, Navicula tripunctata; nacry,Navicula cryptocephala; nitlin, Nitzschia linearis; oed1, Oedo-gonium1; moug, Mougeotia; pedbor, Pediastrum boryanum;pedsim, Pediastrum simplex; plali, Planktolyngbya limnetica;rhcur, Rhoicosphaenia curvata; sceco, Scenedesmus ecornis;scqu, Scenedesmus quadricauda; stige, Stigeoclonium; syber,Synedra berolinensis; Temp, Temperature; Sec dep, SecchiDepth

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simplex Meyen and P. boryanum grew at all sections(Table 4). The stalked diatom species, Cymbellalanceolata (Ehr.) Kirchne, filamentous and chroococ-cal Limnothrix sp., Microcystis aeruginosa (Kütz.)Kütz., filamentous chlorophytes as Cladophora sp.reached high biomass on the middle sections, whereasfilamentous and chroococcal cyanobacteria, Merismo-pedia glauca (Ehr.) Kütz., and Anabaena sp. andloosely attached species as N. linearis (Agardh) W.Smith grew considerably both on upper and middlesections. Scenedesmus quadricauda (Turpin) Breb.,M. varians and Navicula cryptocephala Kütz. werefound in high biomass both on middle and lowersections during the study period. When M. aerugi-

nosa reached high numbers in phytoplankton insummer, it became biovolume dominant in periphytonon upper section. The prostrate - motile species,Gyrosigma sp., adnate species, Surirella robusta Ehr.and filamentous green algae Ulothrix sp. reached highbiomass only on the lower section. Although Achnan-thidium minutissimum (Kütz.) Czarnecki and N. paleawere found in high numbers at all sections, they neverbecame biovolume dominant nor subdominant inperiphyton because of their small size.

The epiphytic algal diversity fluctuated slightlybetween the sections. The Shannon index (H′) rangedfrom 2.3 to 3.6 on lower section, 1.8 4 on middlesection, 1.7 3.9 on upper section, respectively. In

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Fig. 2 Seasonal composi-tion of chlorophyll-a andepiphytic algae on lower,middle and upper sections ofSparganium erectum inSapanca (a and b) andManyas Lakes (c and d)

Table 3 Summary statistics for the first four axes of CCA performed on the Manyas and Sapanca lakes

Ordination axes

Manyas Lake Sapanca Lake

1 2 3 4 1 2 3 4

Eigenvalues: 0.348 0.248 0.158 0.116 0.239 0.137 0.101 0.086Species-environment corr.: 0.956 0.952 0.932 0.977 0.948 0.888 0.891 0.969Cumulative percentage varianceof species data: 29.2 50.0 63.2 73.0 22.8 35.8 45.4 53.7of species-environment relation: 34.7 59.3 75.1 86.7 32.6 51.2 65.0 76.8

Environ Monit Assess (2008) 146:171–181 175

general, high values are found during mid spring tomid autumn, and low values were found during lateautumn to early spring months and maximum valueswere recoded in summer months (Fig. 3). Positivecorrelations were recorded between Shannon diversityand number of species on all sections; a modestpositive correlation was recorded on lower and uppersections (r=0.46 and r=0.62; p<0.01, respectively)and a strong positive correlation was found on middlesection (r=0.69; p<0.01). Temperature and Suspended

Solids were the factors which affect the ShannonDiversity profoundly during the study period.

Sapanca lake

Table (2) summarizes the measured physical andchemical characters of Sapanca Lake for the periodJuly 1997 November 1998. The seasonal dynamics ofnutrient concentrations and other water qualityparameters in Sapanca Lake differed notably from

Table 4 Dominance of most abundant species according to biovolume units on Sparganium erectum in Sapanca and Manyas Lake

Sapanca Lake Manyas Lake

Low. S. Mid. S. Upp. S. Low. S. Mid. S. Upp. S.

Aulacoseira italica * * * * * *Aulacoseira muzzanensis * * * * *Cocconeis placentula * * *Cymbella lanceolata * * * *Encyonema minuta * * *Fragilaria construens *Fragilaria crotonensis * * *Fragilaria ulna * * * * * *Gyrosigma sp. * * * *Melosira varians * * * * *Navicula cryptocephala * * * * *Navicula tripunctata * * * * * *Nitzschia linearis * *Nitzschia palea * *Rhoicosphaenia curvata *Surirella linearis * * * *Cladophora sp. * * * *Coleochaete sp. *Cosmarium formulosum * * *Cylindrocapsa sp. * * * * * *Draparnaldia sp. *Mougeotia sp. * * * * * *Oedogonium sp. (1) * * * * * *Oedogonium sp. (2) * * *Pediastrum boryanum * * *Pediastrum simplex * * *Scenedesmus quadricauda * *Stigeoclonium sp. * * *Ulothrix sp. * * *Merismopedia glauca * *Oscillatoria limnetica *Planktolyngbya limnetica * * *Pseudoanabaena sp. * * *

*Available

176 Environ Monit Assess (2008) 146:171–181

Manyas Lake. When the seasonal averages of thesevariables were regressed against periphyton, it can beseen that the variables e.g. TSS, SRP, NO2+NO3,WLF and Secchi depth are correlated with periphyton(Fig. 4). Total algal biovolume was negativelycorrelated with TP (r=−0.59; p<0.05, respectively)whereas positively correlated with chl-a (r=0.96; p<0.01) and Secchi depth (r=0.57; p<0.05). The CCArevealed a modest relationship between speciesdistribution and environmental variables. Only 29dominant algal species are shown in Fig. 4. Theeigenvalues of CCA axis 1 (0.24) and axis 2 (0.14),constrained to the eight variables account for 35.8%of the cumulative variance in the species data. Thespecies–environment correlations for CCA axes 1

and 2 are high and together these two axesaccounted for 51.2% of the variation in theperiphyton environment relationship. Monte Carlopermutation tests showed that both axes werestatistically significant (p<0.01).

The chl-a concentrations varied greatly among thesections. Mean chl-a concentration was 41.1, 53.2and 70.4 µg cm−2 upon lower, middle and uppersections, respectively (Fig. 2). The temporal changeof chl-a and epiphytic algal biovolume showedsimilar trends during the study period. A first chl-apeak was recorded on upper section in July 1997,when diatoms dominated 62% of periphyton andlowest values were recorded during the autumn 1997.Second peak was recorded on all sections in March

Fig. 3 Species richness ver-sus Shannon Diversity onupper (a), middle (b) andlower (c) section ofSparganium erectum inSapanca Lake and upper(d), middle (e) and lower (f)section of Sparganiumerectum in Manyas Lake

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Environ Monit Assess (2008) 146:171–181 177

1998, when more than 95% of the periphytoncomprised of diatoms. A last and highest chl-a peakwas measured in June 1998 when more than 86% ofthe biovolume consisted of chlorophytes.

A total 109 species of periphyton were identifiedChlorophytes, diatoms, and cyanobacteria were foundas major algal groups in the periphyton on S. erectumin Sapanca Lake. However, the zonation of theperiphyton, density, composition and dominant/sub-

dominant taxa were significantly different in theselakes; mean algal biomass was considerably high inSapanca Lake whereas the number of the species washigh in Manyas Lake (Table 2). Mean diatom andcyanobacteria biomass showed a modest differenceamong the sections whereas mean chlorophyte bio-mass was found considerably high on upper section(p<0.01; Table 5). A. italica, F. crotonensis, C. lanceo-lata, N. cryptocephala, N. tripunctata, M. varians,Gyrosigma sp., Cylindrocapsa sp., F. ulna, C. placen-tula, Oedogonium spp., Pseudoanabaena sp., Mougeo-tia sp., and Cladophora glomerata (Linn.) Kütz. grewon all sections whereas N. palea, and Ulothrix sp. grewon lower and middle sections, Cosmarium formulosumW.E. Hoffmann ex Nordsted and Oedogonium sp.grew on middle and upper sections. Fragilariaconstruens var. venter (Ehr.) Grun., Draparnaldia sp.and Oscillatoria limosa (Dill.) C. Agardh wererecorded in high biomass only on the lower sectionand Cocconeis scutellum (Ehr), Surirella linearis W.Smith, Epithemia adnata (Kütz.) Breb., Coleochaetesp., R. curvata were found in high biomass only on theupper section (Table 4).

Figure 3 shows the seasonal differences in epi-phytic algal diversity on S. erectum in Sapanca Lake.In general, the diversity values were slightly higherthan Manyas Lake; They ranged from 1.8 to 4.6 onlower section, 2.1 to 4.4 on middle section and 1.9 to4.3 on upper sections, respectively. Contrary toManyas Lake, the highest diversity values were foundin early spring and the lowest values were found insummer months. Shannon diversity index and numberof species showed a very strong positive correlationon the lower section (r=0.87; p<0.01)) whereas amodest positive correlation was recorded on themiddle and upper sections (r=0.49 and r=0.42 p<0.01, respectively).

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Sec dep.

Fig. 4 Biplot of first and second CCA axis for periphyton inSapanca Lake. Environmental variables: achlin, Achnantheslinearis; aulit, Aulacoseira italica; clad, Cladophora; cocpla,Cocconeis placentula; cocscu, C. scutellum; cosfor, Cosmariumformulosum; cyli, Cylindrocapsa; cymlan, Cymbella lanceolata;cytum, C. tumida; encmin, Encyonema minutum; epad, Epi-themia adnata; epsor, E. sorex; frcon, Fragilaria construens;frcro, F. crotonensis; fraul, F. ulna; gomol, Gomphonemaolivaceum; gopar, G. parvulum; melvar, Melosira varians;moug, Mougeotia; nacry, Navicula cryptocephala; natri, N.tripunctata; nipal, Nitzschia palea; oed1, Oedogonium1; oslim,Oscillatoria limosa; rhcur, Rhoicosphaenia curvata; surlin,Surirella linearis; Temp, Temperature; Sec dep, Secchi Depth

Table 5 Vertical colonization of major epiphytic algal groups on Sparganium erectum

Sapanca Lake Manyas Lake

Unit Low. S. Mid. S. Upp. S. Low. S. Mid. S. Upp. S.

Chlorophytes % 56.8 58 70.9 43.4 60.3 61.5Diatoms % 38.7 37.4 25.3 48.6 32.8 24.5Cyanobacteria % 4.4 4.4 3.8 8 6.9 14

Low S Lower Section, Mid S Middle Section, Upp S Upper Section

178 Environ Monit Assess (2008) 146:171–181

Discussion

Studies attempting to understand the complex eco-logical interrelationships existing in aquatic environ-ments have frequently overlooked the role of theperiphytic community (Sheldon and Boylen 1975).The factors affecting the colonization of epiphyticalgae on macrophytes and artificial substrates havelong been studied by several workers (Sheldon andBoylen 1975; Ács and Kiss 1991; Vis et al. 1998;Albay and Akçaalan 2003). Temporal and spatialvariability in physico-chemical parameters acts toorganize epiphytic algal assemblages, regulate speciesinteractions and, by providing conditions that supportdifferent taxa under various environmental conditionsmay be crucial for maintaining biodiversity (Vis et al.1998; Wellnitz and Ward 1998; Cardinale et al. 2000).

In the present study, we found distinct patterns ofperiphyton colonization related to water quality andhydrologic drivers. Our results indicate that three majorenvironmental gradients (high amount of TSS, lowlight climate and WLF) control the vertical coloniza-tion of periphyton assemblages for Manyas (Table 2).Mean periphyton biomass was 4.21 mm3 cm−2 inManyas lake whereas it was found considerably high(14.79 mm3 cm−2) in Sapanca lake. Scheffer (1998),stated that light is predominantly absorbed by par-ticulate matter such as phytoplankton cells, detritusand suspended sediment particles in many shallowlakes and shallow, slowly flowing side arms (inSoroksári Danube, Szabó et al. 2001). Accordingly,Müller (1994) stated that the amount of light reachingthe algal cells depends on external factors, such as thePAR and length of the macrophyte stand, and onfactors in the water such as Secchi depth andthickness of the periphytic layer. The high amountof Suspended Solids values (mean: 21.6 cm) limitedthe light penetration in the water in Manyas Lake.Therefore, it is likely to conclude that low lightintensity might have given rise to low amount ofperiphytic biomass on S. erectum in this lakecompared to Sapanca Lake (Table 2).

The nutrients level and other measured waterquality parameters in Manyas Lake are indicative ofthe lake’s present highly eutrophic status (Table 2)whereas Sapanca Lake is still in oligo-mesotrophicconditions according to OECD (1982). The commu-nity structure and vertical colonization of epiphyticalgae on S. erectum showed a distinct difference

between the two lakes. The diatoms were recorded asthe dominant group with 49% in Manyas Lake,whereas green algae accounted for 57% of theperiphyton in Sapanca Lake on the lower section(Table 5). The middle and upper sections weredominated by green algae both in Manyas Lake andSapanca Lake. The highest epiphytic algal biomasswas recorded on the middle section in Manyas Lakewhereas highest biomass values were found on uppersection in Sapanca Lake. The WLF might havebrought about this situation since it fluctuate ±31 cmin Manyas Lake during the study period. It can bethought that WLF has a most pronounced effect onperiphyton colonization in shallow lakes than deep lakesbecause of their morphometric character. Rychkova(1989) stated that the periphytic biomass is largelyeliminated by changes in the WLF and wave action.

Totally eight taxa (Oedogonium sp., Mougeotiasp., Cylindrocapsa sp., Cladophora sp., A. italica, M.varians, N. tripunctata, and F. ulna) were found asdominant species at all sections of S. erectum both inManyas Lake and Sapanca Lake (Table 4). Similarresults were found by Müller (1994) in a small,moderately-deep, hard-water German lake, LakeBelau. It can be thought that these species have abroad range of tolerance to several physical andchemical disturbances including nutrients, TSS, WLFand light intensity (Table 2). Oedogonium sp. was thepioneer species (mean 0.51 and 7.16 mm3 cm−2 inManyas Lake and Sapanca Lake, respectively) duringthe study period. The other filamentous chlorophytes,Stigeoclonium sp. and Spirogyra sp. reached highbiomass values in Manyas Lake, whereas Coleo-chaete sp., Draparnaldia sp., were found in highbiomass values in Sapanca Lake occasionally. Theexcessive growth of some filamentous chlorophytes,such as Oedogonium sp., Spirogyra sp., and Clado-phora glomerata, has long been associated witheutrophication (Cattaneo 1987; Albay and Akçaalan2003). Cattaneo (1987) stated that filamentous greenalgae were positively correlated with TP. Amongthese species Cladophora glomerata has primarilyresponded to eutrophication (Danilov and Ekelund2001) and to a lesser degree Ulothrix, Spirogyra andOedogonium. It can be thought that this species shouldbe used as a key species in lake monitoring studies.

The correlation between diversity (H′) and numberof species was positive however it was not highlysignificant at all sections except lower section of S.

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erectum in Sapanca Lake. Seasonal variation inepiphytic algal diversity and number of speciesshowed highly different pattern between two lakes(Fig. 3 and Table 2). The highest diversity valueswere recorded in summer and lowest values wasrecorded in late autumn in Manyas Lake whereashighest and lowest diversity values were recorded inearly spring and summer months in Sapanca Lake. Itis worthwhile the point out that the highest diversityvalues were recorded when periphyton was dominat-ed by several epiphytic algal species and lowestvalues were recorded when periphyton was dominat-ed by few species in both lakes. Ács et al. (2000)stated that changes of number of species and evennessin periphyton depend on the immigration and repro-duction rate of algae. They concluded that immigra-tion causes an increase in the number of specieswhereas reproduction maintains species richness byreducing the losses deriving from death.

The periphyton community was dominated byspecies which is tolerant to eutrophic conditions inthe Manyas Lake whereas meso-oligotrophic taxadominated in the periphyton of Sapanca Lake. Ourresults showed that certain water quality and physicaldisturbances e.g. high amount of suspended solids,WLF and nutrients and low light intensity have had adistinctive effect on the vertical colonization ofperiphyton, number of species, diversity and compo-sition. The results also revealed that periphyton canbe used as bioindicators of anthropogenic contamina-tion in lakes. More information is needed to examinethe factors governing periphyton colonization in lakeswith different mixing and trophic level. This assess-ment is urgently required, given the need to beginimplementation of the Water Framework Directives(Flynn et al. 2002).

Acknowledgements We are thankful to Eva Acs and JorgenChristiansen for reading and comments on the draft of themanuscript. The present work was supported by the ResearchFund of Istanbul University. Project No: 993/090597 and UDP-156/04062003.

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