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INTRODUCTION

Potamogeton genus is represented by hydrophytes occurring in a wide range of diverse habitats from standing to fast fl owing waters. They could be found in natural oligo-mesotrophic conditions as well as on hydromorphologically disturbed places with high water eutrophy level (e.g. K and T ,

, K , S and K - , K , J et . , K

, C and B , C and S - , Z -G , Z -G

et . ). They show a high variability in the pheno-type relative to the environmental conditions (P -

a, b, K , Z -G et . ). The majority of these macrophytes are aquatic

perennial and they produce turions and numerous seeds (J et . , Z -G ). In Poland

Potamogeton species and six hybrids occur among - species in total noted worldwide (Z -G -

, Z -G et . ). Most of Po-tamogeton species is freshwater, only a few of them can occur in marine waters e.g. P. fi liformis, P. pectinatus, P. perfoliatus and P. pusillus (Z -G ).

Aquatic vegetation communities with Potamogeton genus are concerned as signifi cant component of bio-logical structure of riverine habitats. In Polish lowland

rivers distinguished by fast fl ow of water diff erent spe-cies could be found of Potamogeton genus. However, the most frequent are P. nodosus, P. berchtoldii, P. crispus, P. pectinatus and P. ×fl uitans, in less range P. polygonyfolius, P. alpinus and P. pusillus could be found (e.g. T -

and K , K , Z --G , S et . , K

and W , Z -G et . ). In fl uvial ecosystem, the investigated P. crispus, P. nodo-sus and P. pectinatus are concerned as characteristic for impoverished type of Ranunculion fl uitantis rivers and represent the habitat Nature – “Water courses of plain to montane levels with the Ranunculion fl uitantis and Callitricho-Batrachion vegetation” (code ) (C -

D / /EEC, A I, II: I - M – EUR ). The mentioned above vascular

plants are also used in quality assessments of rivers in many European countries, according to the Water Frame-work Directive (D / /EC).

The environmental requirements of Potamogeton communities occurring in water course have been still poorly investigated compared to standing waters. Pre-vious studies of parameters determining the distribu-tion of Potamogeton were just related to one species of this genus e.g. P. crispus (S and M , F , J et . ), P. nodosus (P et .

, K and W ) and P. pectinatus

DISTRIBUTION PATTERNS OF THREE POTAMOGETON SPECIES P. CRISPUS, P. NODOSUS AND P. PECTINATUS ALONG VELOCITY AND BASE RICHNESS

GR ADIENTS FROM A LOWLAND RIVER

E J , M G

E. Jakubas, M. Gąbka, Department of Hydrobiology, Institute of Environmental Biology, Faculty of Biology, Adam Mickiewicz University, Umultowska , - Poznań, Poland, e-mail: [email protected]

(Received: April , . Accepted: June , )

ABSTR ACT. This study presents distribution and abundance of three Potamogeton species, namely Pota-mogeton crispus, P. nodosus and P. pectinatus along environmental gradients in the lowland river Wełna (NW Poland). The relationships between environmental factors and the pattern aquatic vegetation distribution along river were investigated. Among ecological factors rarely undertaken in aquatic ecology the light climate was concerned. It is postulated that the Potamogeton communities in the investigated river are strongly connected with water velocity, substrate of bottom and light conditions, in particular dissolved organic matter (DOM). Elodeo-Potametum crispi and algae communities with dominant spe-cies Hildenbrandia rivularis were well developed in the places shading by trees, with high velocity and fairly clean water, mostly with stony bottom. Potametum nodosi was noted in mean values of velocity and medium water quality with high content of organic matter in the bottom substrate. The last investigated community Sparganio-Potametum interrupti was found in poor water quality with the highest values of electric conductivity. The obtained results give a new approach of the ecology and abiotic typology of rivers with macrophytes including abundance of Potamogeton species (Nature habitat, code – “Water courses of plain to montane levels with the Ranunculion fl uitantis and Callitricho-Batrachion vegetation”).

KEY WORDS: ecology, distribution pattern, Potamogeton, gradients, river, environment, Natura

E. Jakubas, M. Gąbka

(S , D and J , L M et . ) or were concerned general studies of aquatic vegetation (O , P , B --P et . ). This is the fi rst study to attempt to determine the eff ect of environmental factors on the distribution pattern of three Potamogeton species and co-occurring plants (mosses, macroscopic algae, vascu-lar plants) in one lowland river. The present publication answers the following questions:

– What is the main ecological factor responsible for the distribution and abundance of Potamogeton species?

– What is the species composition of vegetation patches?

METHODS

Study areaThe study area was the Wełna river, being right tribu-

tary of the Warta river (NW Poland). The fi eld study was performed along km distance of the Wełna river.

This lowland, naturally meandering river fl ows through the feeding system of eight eutrophic lakes. The total length of the river is . km. The study was conducted during one vegetation period (June-September ) (Fig. ).

Field sampling and laboratory analysesIn all research vegetation plots (each m ) con-

taining particular of P. crispus, P. nodosus and P. pecti-natus or each altogether species at the same vegetation plot the physical and chemical parameters of the water as well as hydrological factors were analysed (Phot. - ). The following parameters were measured in situ: temperature (Temp), pH, electric conductivity (EC), dissolved oxygen (DO). In each plot, water velocity was measured at the surface layer ( . m) among the plants using a hydromet-ric universal current meter. The depth at the sites of Pota-mogeton species was measured in centimetres. The width of the channel (Width) in each of the study plots was measured using a laser rangefi nder (type Hilti PD ). The percentage shading by trees (Tree) and the stones percent-age coverage by mosses and algae (Stones) was also noted.

F . . Location of the Wełna river with the distribution of the Potamo-geton species stands

Distribution patterns of three Potamogeton species (P. crispus, P. nodosus and P. pectinatus) along velocity ...

P . - . Potamogeton species and their communities in the Wełna river: - – Potamogeton nodosus, - – Potamo-geton crispus, - – Potamogeton pectinatus (Phot. – M. Gąbka, Phot. - – E. Jakubas)


Standard measurements for the following character-istics were made at each site: forms of dissolved organic matter (DOM nm) were measured spectrophoto-metrically using a Cadas UV-VIS (Dr Lange) spec-trophotomether, with cm quartz cell on fi ltrate (after fi ltration through . μm membrane fi lter, converted into the m- ).

In each of the vegetation plots, the bottom samples were collected using a core cutting sampler, which was divided into layers ( cm). The content of organic matter (OM) was measured with weight method after sample combustion at °C and hydration (H) was analysed after drying of sample in °C to a content weight.

Physical and chemical analyses were performed ac-cording to the standard methods for hydrochemical analyses (APHA ). Water samples for spectropho-tometrical analysis (inherent colour and organic mat-ter forms) and dissolved organic carbon were fi ltered through . μm membrane fi lters.

Numerical analysesStatistical analyses were performed by using CANO-

CO software ( B and Š ). The CCA (canonical correspondence analysis) to explore the rela-tions between the species distribution and environmen-tal variables was used ( B and Š ).

Furthermore in Monte Carlo permutation test (with permutations made to reduce the number of vari-

ables) velocity of water, substrate and light conditions refl ected by dissolved organic matter were statistically signifi cant (P < . ).

The assessment of environmental data in bioindi-cation was made by using software C (J ) including model weighted averaging (WA). This ena-bled to determine optima (weighted-average) and toler-ance range (SD) of species in relation to environmental parameters.

Nomenclature Names of syntax are according to treatise of M

et . ( ), mosses of O et . ( ) and algae of Algaebase.org. Names of vegetation compositions are according to R et . ( ) including study of P ( a, b, ).

RESULTS

Species richnessAquatic vegetation surveys including performance

of species composition and their cover in the channel showed fl oristic communities among the study sites ( vegetation plots). The macrophyte composition of

vegetation plots contains: three species of algae, two mosses and vascular plants including P. crispus, P. nodosus and P. pectinatus. Abundance according to V M scale ( ) and frequency of occur-rence under study are presented in Table . The analysed plots were quite rich in species. Among algae, the most often Hildenbrandia rivularis occurred, among mosses Fontinalis antypyretica and vascular plants were mainly represented by Sagittaria sagittifolia (submerged form) and Nuphar lutea (submerged form). In Potamogeton ge-nus P. nodosus ( stands), P. crispus ( stands) and P. pectinatus ( stands) dominated respectively.

The Potamogeton nodosus association (Potametum nodosi Segal ( ) ) composition was distin-guished by vascular plants mostly by S. sagittifolia (sub-merged and emergent form) and N. lutea (submerged form). In this case pleustophytes occurred in smaller range and no species of grasses were found. The vegeta-tion compositions of Potamogeton crispus (Elodeo-Pota-metum crispi (Pignatti ) Pass. ) was dominated by macrophytes preferring fast fl ow of water and stones as substrate of bottom e.g. algae such as Hildenbrandia

T . Frequency and mean and max value of cover van der Maarel scale of Potamogeton species and other species in vegetation samples (n = )

Species Symbol I II III

Algae Hildenbrandia rivularis Hilriv . ( - ) . ( . - ) . ( . - )

Heribaudiella fl uviatilis Herfl u . ( . - ) . ( . - ) . ( . - )

Rhizoclonium sp. Rhizsp . ( . - ) . ( . - ) –

Mosess Fontinalis antypyretica Fonant . ( . - ) . ( . - ) . ( . - )

Leptodictum riparium Leprip . ( . - ) . ( . - ) . ( . - )

Vascular plants

Potamogeton nodosus Potnod ( . - ) . ( , - ) . ( . - )

Potamogeton crispus Potcri . ( . - ) ( . - ) . ( . - )

Potamogeton pectinatus Potpec . ( . - ) . ( . - ) ( . - )

Sagittaria sagittifolia Sagsag . ( . - ) . ( . - ) . ( . - )

Nuphar lutea Nuplut . ( . - ) . ( . - ) . ( . - )

Sparganium emersum Spaeme . ( . - ) . ( . - ) . ( . - )


T – cont.

Lemna trisulca Lemtri . ( . - ) . ( . - ) –

Schoenoplectus lacustris Scilac . ( . - ) . ( . - ) . ( . - )

Sparganium erectum Spaere . ( . - ) . ( . - ) . ( . - )

Myriophyllum spicatum Myrspi . ( . - ) . ( . - ) . ( . - )

Spirodella polyrhiza Spipol . ( . - ) . ( . - ) . ( . - )

Rorippa amphibia Roramp – . ( . - ) . ( . - )

Butumus umbellatus Butumb . ( . - ) . ( . - ) . ( . - )

Veronica anagalis-aquatica Verana . ( . - ) . ( . - ) –

Lemna gibba Lemgib . ( . - ) . ( . - ) . ( . - )

Berula erecta Berer . ( . - ) . ( . - ) . ( . - )

Hydrocharis morsus-ranae Hydmor – . ( . - ) . ( . - )

Elodea canadensis Elocan – . ( . - ) –

Ceratophyllum demersum Cerdem – . ( . - ) –

Lemna minor Lemmin . ( . - ) – . ( . - )

Alisma plantago-aquatica Alipla . ( . - ) – –

Phalaris arundinacea Phaaru – – . ( . - )

Phragmites australis Phraus – – . ( . - )

Abbreviation: I – P. nodosus ( stands), II – P. crispus ( stands) and III – P. pectinatus ( stands).

T . Summary of water chemistry collected within stands of Potamogeton species

Property Abbreviation Mean Min Max

Physicochemical indicators of water and substrate

Temperature (°C) Temp . . .

pH pH . . .

Conductivity (μS·cm- ) EC . . .

O dissolved (mg·O l- ) DO . . .

Dissolved organic matter (m- ) DOM . . .

Organic matter (%) OM . . .

Hydration (%) H . . .

Stones as bottom substrate covered by mosses and algae (%)

Stones . . .

Shaded by tree (%) Tree . . .

Hydrological indicators of water

Surface velocity of water (m·s- ) H_surf . . .

Velocity of water in plants (m·s- ) H_veg . . .

Hydrological indicators of river

Depth of water (m) Deep . . .

Width (m) Width . . .


F . . Biplots of canonical analysis (CCA) for species. Hildenbrandiarivularis – Hil-riv, Heribaudiellafl uviatilis – Herfl u, Rhizoclonium sp. – Rhizsp, Fontinalisantypyretica – Fonant, Leptodictumriparium – Leprip, Sagittariasagittifolia – Sagsag, Nupharlutea – Nuplut, Potamogetonnodosus – Potnod, Potamogetoncrispus – Potcri, Potamogeton-pectinatus – Potpec, Sparganiumemersum – Spaeme, Sparganiumerectum – Spaere, Scirpuslacustris – Scilac, Spirodellapolyrhiza – Spipol, Veronica anagalis – aquatica – Verana, Lemnatrisulca – Lemtri, Lemnagibba – Lemgib, Myriophyllumspicatum – Myrspi, Berulaerecta – Berer, Butumusumbellatus – Butumb, Rorippaamphibia – Roramp, Lemna minor – Lemmin, Hydrocharismorsus – ranae – Hydmor, Lemnatrisulca – Lemtri, Ali-smaplantago – aquatica – Alipla, Elodea canadensis – Elocan, Ceratophyllumdemersum – Cerdem, Phalarisarundinacea – Phaaru, Phragmites australis – Phraus.

rivularis and mosses like Fontinalis antypyretica. For this association any species of grasses were not also noted. The fl oristic community of Potamogeton pectinatus (Sparganio-Potametum interrupti (Hilbig ) Weber

) consisted mostly of N. lutea (submerged form) and Fontinalis antypyretica.

Only in three of investigated vegetation plots above-mentioned Potamogeton species occurred alto-gether in following system Potamogeton crispus-no-dosus-pectinatus, whereas in seven stands occurred two among three species. They created two diff erent systems: ) Potamogeton nodosus-pectinatus and ) Potamogeton crispus-pectinatus. Nevertheless P. no-dosus and P. crispus have never been presented in one individual composition. Potamogeton nodosus oc-curred alone or in the whole group of Potamogeton species.

Environmental factors The Potamogeton species were found in the river

section where the width of the river channel was . -- . m, at . - . m depth. The surface velocity values of water ranged - . m·s- and water velocity among plants was always smaller - . m·s- . Potamo-geton species preferred warm . - . °C and alka-line water in the range . - . pH. These vascular plants occurred in quite high values of light conditions e.g. dissolved organic matter was . - . m. In the

vegetation plots substrate of bottom was mostly with low content of organic matter. These data are presented in Table .

Potamogeton species along environmental gradientsThe used biplots of canonical analysis (CCA; Fig. )

showed distribution pattern of aquatic vegetation in relation to environmental gradients. The occurrence of Potamogeton nodosus which was the most common species among all Potamogeton genus in the Wełna river was strongly connected with light parameters such as dissolved organic matter (DOM), Potamogeton crispus with strong water velocity in shallow, stony places and the abundance of Potamogeton pectinatus was associ-ated with higher electric conductivity.

The optimum and tolerance of particular Potamoge-ton species compared to selected environmental factors are presented in Table . In particular, Elodeo-Potame-tum crispi and algae communities with Hildenbrandia rivularis were developed in high velocity and clean water, in shady places and stones as substrate of bot-tom. Potametum nodosi was noted in mean values of water velocity with high content of organic matter in substrate. The last investigated community Potametum pectinati was found in water of poor quality with the highest values of electric conductivity.

It seems that Potamogeton nodosus showed the most fl exible compared to shading by trees while P. pectinatus


the lowest. As for factors such as width of channel and depth of water the highest tolerance had also P. nodo-sus. Potamogeton crispus preferred places the hardest overgrown by mosses and algae contraire to P. nodosus. However, P. crispus had the narrowest tolerance of con-ductivity and pH gradient and this species could be very sensitive to change of these parameters (Table ).

DISCUSSION

Species compositionOur study in the lowland river ecosystem revealed

that investigated Potamogeton species such as P. nodo-sus, P. crispus, and P. pectinatus belong to the species inhabiting the river current, where mean velocity of water was . m·s- . These vascular plants, mainly P. nodosus are believed as characteristic for running waters (e.g. G et . , P , Z -G -

, K and W ). Nevertheless, it should be noted that all of them commonly occur also in lakes and ponds (K , , S -

and K , J et . , C and B , Z -G ). The vegetation analysis showed that P. nodosus and P. pectinatus were closer together and occurred among vascular plants such as Schoenoplectus lacustris, Myriophylllum spicatum and Nuphar lutea, while P. crispus was located further, among macrophytes preferring high water velocity such as Hildenbrandia rivularis and mosses such as Fontinalis antypyretica and Leptodictyum riparium. Previous lit-erature on vegetation communities in watercourse indi-cated that same of Potamogeton species like P. nodosus

is a characteristic of patches of Potametum nodosi or Ra-nuculetum fl uitantis habitat with plant communities of pleustophytes e.g. Lemnetum gibbae Miy. et J. Tx. (K and W ). This association of Po-tametum nodosi has been categorised in Poland as very rare and endangered (R et . ). However, in the Wełna river Lemna trisulca occurred in fast fl ow water as submerged form and it was strongly adhered to stones, which substrate of bottom was overgrown in this case by P. crispus. These species of Potamogeton are typi-cal of Lemnetum communities and were noted previously (O , G et . ).

Furthermore, the species structure analysis together with CCA explains that P. nodosus and P. crispus have never created common composition, therefore they are isolated from each other and prefer diff erent environ-mental conditions. That P. crispus could be able to form communities corresponding to Elodeo-Potametum crispi has been well known from many lowland rivers in Eu-rope (e.g. P , F , J et . , C and S ).

According to earlier studies of aquatic vegetation the fl oristic lists of Potamogeton communities including P. gramineus, P. lucens, P. perfoliatus, P. natans, P. pu-sillus and other vascular plants such as Myriophyllum spicatum, M. verticillatum, Ceratophyllum demersum and Elodea canadensis (O , G et

. , P ). These mentioned, co-occuring vascular plants could be considered as a constant for Potamogeton communities. Some of them e.g. Myrio-phyllum spicatum, Ceratophyllum demersum and Elo-dea canadensis were also rarely noted in the Wełna river.

T . Optimum tolerance of particular Potamogeton species compared to stud-ied environmental parameters

Parameter I II III

Shaded by tree (%) . ( . ) . ( . ) . ( . )

Stones as bottom substrate covered by mosses and algae (%)

. ( . ) . ( . ) . ( . )

Conductivity (μS·cm- ) ( . ) ( . ) ( . )

Dissolved organic matter (m- ) . ( . ) . ( . ) . ( . )

Surface water velocity (m·s- ) . ( . ) . ( . ) . ( . )

Velocity of water in plants (m·s- ) . ( . ) . ( . ) . ( . )

Depth of water (m) . ( . ) . ( . ) . ( . )

Width (m) . ( . ) . ( . ) . ( . )

pH . ( . ) . ( . ) . ( . )

Temperature (°C) . ( . ) . ( . ) . ( . )

O dissolved (mg·O l- ) . ( . ) . ( . ) . ( . )

Hydration (%) . ( . ) . ( . ) . ( . )

Organic matter (%) . ( . ) . ( . ) . ( . )

Abbreviation: I – P. nodosus ( stands), II – P. crispus ( stands) and III – P. pectinatus ( stands).


The communities of P. pectinatus created simple, underwater meadows while P. crispus and P. nodosus occurred in richer species compositions.

Ecological parametersOur results from the Wełna river suggest that dis-

tribution pattern of three Potamogeton species, P. nodo-sus, P. crispus, and P. pectinatus is distinctly connected with water quality and velocity as well as type of bottom (stones) and light conditions. Potamogeton nodosus and P. pectinatus were found in conditions of higher values of dissolved organic matter (DOM). The signifi cance of light availability e.g. on turion germination was con-cerned in previous literature compared to P. crispus (J et . ). Moreover, the experiment of resource allocation for long term survival of P. pectinatus has been made. This allocation model emphasized the im-portance of light conditions in biomass development of P. pectinatus (V D and J ). Nevertheless, another study indicated that P. nodosus prefers moder-ate light conditions (Z et . ).

The CCA analysis (Fig. ) revealed that occurrence of P. crispus is dependent on surface water velocity and water temperature. A number of studies confi rmed that the ecological amplitude of Potamogeton genus is wide and could be associated with above mentioned factors such as light climate, temperature of water and pH gra-dient (T and K , K

, C and S , Z -G , Z -G et . , K and

W ). However, the occurrence of Potamogeton species

can be dependent on completely diff erent factors e.g. L M et . ( ) has studied impact of forag-ing by migrating trumpeter swans on tuber and rhizome density and biomass of P. pectinatus and co-occurring species compositions. Another study has reported that also planting depth infl uences growth of P. pectinatus (S ). A detailed comparison of all three Po-tamogeton species enables to say that P. nodosus prefers the deepest water (optimum . , tolerance . ), P. pec-tinatus can be found both in streams and river in shal-lower places ( . - . m) as well as in intermediate deep habitats in standing waters ( - m; G et . ).

Furthermore, plant communities could be also de-pendent on stream and rivers management. The land-use of catchment and hydromorphological changes infl uence the occurrence of slow-growing species in-cluding Potamogeton species (B -P et . ). In response to environmental conditions they show signifi cant variable of plastic (K , Z -G ). For instance P. nodosus by its fl exibility could be able to reduce the turbulent ki-netic energy of vertical mixing (P et . ). From aquatic ecology point of view the really interesting is signifi cance of communities with three investigated Potamogeton species in relation to the quality of wa-ter. Previous studies e.g. C and S ( ) clearly indicated that Potamogeton genus show diff er-ent requirements of the trophic level from mesotrophy (P. crispus), through eutrophy (P. nodosus) to hypertro-phy (P. pectinatus) in rivers and streams. Our results revealed similar pattern of macrophytes compared to

light conditions and base richness in the Wełna river. Hence, environmental data mostly physico-chemical pa-rameters of water, could be useful in defi ning separate-ness plant communities in the lowland rivers.

Our results emphasize the signifi cance of some Pota-mogeton species in the aquatic vegetation compositions of lowland rivers. The obtained results of the species structure and their habitats provide a new information of ecology of rivers and macrophytes compositions with Potamogeton species communities (Nature habitat, code ).

AcknowledgementsSpecial thanks to the Oborniki Forest District Of-

fi ce, especially, the Forest District Manager Eng. Wło-dzimierz Kowal and Mr. Jarosław Bator for their valuable help during fi eld studies. Thanks to dr Tomasz Joniak for his help in the laboratory analysis. The authors would also like to thank Stanisław Rosadziński for mosses spe-cies identifi cation and valuable information of Potamo-geton species in western Poland.

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