Distribution of ectomycorrhizal fungi in periodically inundated plant communities on the Pilica...

Beata Sumorok1, Krzysztof Kosi ski2, Dorota Michalska-Hejduk3, Edyta Kiedrzy ska4

AbstractStudies on ectomycorrhizal symbionts in plant communities were developed in the experimental floodplain located in the middle reach of the Pilica River valley near Sulejów (central Poland). This area is a natural floodplain where habitats are perma-nently or periodically flooded. Its vegetation is highly diversified and includes hay-growing meadows (Molinio-Arrhenateretea), rush communities (Phragmitetea, Caricetea), willow shrubs (Salicetea) and pine monoculture. Samples of trees’ and bushes’ roots were collected using a metal probe, sporocarps of ectomycorrhizal fungi (ECMF) were collected together. Researches indicate that wet soil conditions and peri-odic flooding have a significant influence on mycorrhiza presence. The vast majority of ectomycorrhizal fungi were present in trees growing in dry habitats – willow bushes and pine monoculture (Salix species, Betula pendula and Pinus sylvestris). The species composition were determined - in total noted c.a. 40 species of ectomycorrhizal fungi and 15 morphological types. The number and diversity of ectomycorrhizal (ECM) morphotypes increased significantly from flooded to dry habitats. Despite the fact that extremely humid and dry soil conditions make a negative impact on mycorrhiza, sym-biotic parthnership enable floodplain plant communities to tolerate water level fluctua-tions, eliminate water stress and provide plants tissues with e.g. phosphorous. Key words: dry-to-wet gradient, waterlogging, plant cover, mycorrhizae.

1. Introduction

One fundamental assumption of the ecohydro-logical approach is using the evolutionarily estab-lished properties of catchments and providing a

framework for their management and restoration that can enhance the capacity of ecosystems to retain nutrients (Zalewski 2000; Zalewski 2002; Kiedrzy ska et al. 2008). Therefore, the floodplain habitats which appear at the aquatic/terrestrial inter-

Vol. 8No 2-4, 401-4102008

Distribution of ectomycorrhizal fungi in periodically inundated plant communities

on the Pilica River floodplain

Ecohydrological Processes and Sustainable Floodplain Management

1Research Institute of Pomology and Floriculture, Rizosphere Laboratory, 18 Pomologiczna str., 96-100 Skierniewice, Poland, [email protected]

2University of ód , Department of Applied Ecology, 12/16 Banacha str., 90-237 ód , Poland;

3University of ód , Department of Geobotany and Plant Ecology, Banacha 12/16 str., 90-237 ód , Poland;

4International Institute of the Polish Academy of Sciences European Regional Centre for Ecohydrology under the auspices of UNESCO,

3 Tylna str., 90-364 ód , Poland.

DOI:10.2478/v10104-009-0032-x

B. Sumorok et al. 402

face, due the high spatial and temporal dynamics of the hydrological and biological processes there, should be considered not only as the key element for river ecosystem restoration, but also as the start-ing point for implementation of the new way of thinking about sustainable water and ecosystem services (Zalewski 2006). The key to use “ecosys-tem processes as a management tool” in water man-agement (Zalewski 2000) is understanding the hydrology–biocoenosis interplay (Zalewski et al. 1997; Zalewski 2006; Sumorok, Kiedrzy ska 2007; Kiedrzy ska et al. 2008). The floodplain river systems, which play a significant role in nutrient retention (Kiedrzy ska et al. 2004; Kiedrzy ska 2006), sedimentation (Magnuszewski et al. 2005; Kiedrzy ska 2006; Altinakar et al. 2006) and nutrient uptake by plants (Kiedrzy ska et al. 2008), are well developed in Poland. Their buffering capacity depends on the vegetation cover, especially on plants effectiveness in nutrient accumulation (Wagner- otkowska, Kiedrzy ska, Sumorok 2004; Kie-drzy ska et al. 2008). Plant growth conditions are influenced by symbiotic organisms, such as bacteria and fungi present in the soil around roots. The microbial activity of the rhizosphere crucially affects the condition of plants, constitutes natural resistance to pathogens and promotes nutrient absorption of

host plants (Azcón-Aguilar, Barea 1992; Smith, Read 1997; Linderman 2000; Miller, Jastrow 2000; Sumorok, Kiedrzy ska 2007). Fungi colonise over 90% plant species in natural ecosystems (Read 1991), and mycorrhizae is widespread in all envi-ronment types (e.g. Allen 1991). Ectomycorrhiza (ECM) predominates in the temperate zone forest areas while arbuscular mycorrhiza (AM) has more importance in herbaceous plant communities also in the tropical zone (Read 1991). In the ectomycorrhizal symbiosis, the mantle is connected to highly branched hyphae that pene-trates the root and grows between the cells. This network of hyphae (hartig net) is the nutrient exchange area. Arbuscular mycorrhizal fungi pro-duce a highly branched, intracellular hyphal struc-ture called arbuscule, that is the site of nutrient exchange. However, symbiotic fungi occurring in periodically flooded vegetation are greatly under-explored (e.g. Clayton, Bagyaraj 1984; Wetzel, van der Valk 1996; Miller 2000; Baar et al. 2002). The distribution of mycorrhizal fungi in plant com-munities depends on moisture level, flooding, tem-perature, pH, and nutrient uptake (e.g. Cox, Tinker 1976; Entry et al. 2002; Tibbet, Sanders 2002). Ectomycorrhizaes are important in the functioning of forest ecosystems and play a crucial role in plant water uptake and nutrient acquisition.

402

Fig. 1. The inundation model of the Pilica River floodplain represents sequences of floodplain overflowing: low flood (L), medium high flood (M), high flood (H), flow conditions (FC) (Kiedrzy ska et al. 2004, Kiedrzy ska et al. 2008, modified).

Ectomycorrhizal fungi in periodically inundated floodplain plant communities 403

Ectomycorrhiza also called sheathing mycor-rhiza, which is one of the most common mycor-rhizal association (Jackson, Mason 1984) occurs in relatively small number, probably 3% of phanero-gams (seed plants). However, this type of mycor-rhiza is significant because of the global impor-tance of arborescent plants (trees, shrubs) covering the huge terrestial land surface as the main produc-ers of timber and their nutrients uptake more than herbaceous plants. Ecto-mycorrhizal associations are able to increase the growth rate and biomass production of their host plant as well as the prolif-eration of the root system. As a result the fungal component may contribute 25%, or more, to the dry weight of an infected root (Isaac, 1992). Because of the importance of symbiotic fungi for plant growth and development, there is a great need to investigate the mycorrhizal status of plants in periodically flooded areas. In this context, the aim of the study was to evaluate mycorrhizal status of arborescent plants in Pilica river floodplain plant communities using mycorrhizal morphotypes and sporocarps of fungi. The hypothesis is tested, that humid soil condition fluctuations and periodic flooding have a significant influence on plant com-munities and ecomycorrhizal fungi distribution. The occurence of ectomycorrhizae is limited in perma-nently humid habitats.

2. Materials and methods

Study area

The research was conducted in growing sea-sons 2003-2005 on the 26.6 hectares of the experi-mental Pilica river floodplain (51o 18’58.46”N and 19o54’10.54”E), upstream of the Sulejów Reservoir. The Pilica River, one of the major tribu-taries of the Vistula, is an important river in Central Poland. Its length is 342 km, and its catchment area comprises 9258 km2. The experimental floodplain is covered by various seminatural plant communities, ranging from wet conditions to dry sites (Fig. 1), there-fore it is a model object for mycorrhizal studies in dry- to-wet gradient. The floodplain is mostly covered by fresh meadows of the order Arrhenetheretalia, mown and occasionally grazed. They occupy the elevated part of the research area, flooded during high water lev-els only. Meadows and sedge rushes are the other group of plant communities in that area (Fig. 2). Grass rushes with dominant Phragmites australis and Phalaris arundinacea, growing along the old river bed, cover local hollows in which water stag-nates throughout the year. Sedge rushes with Carex gracilis as the dominant species cover the areas with a slightly lower ground water level. Wet meadows are formed as an ecotone between rush communi-

ties and fresh meadows, usually the Epilobio-Juncetum effussi association in the grazed parts. Because of different types of floodplain manage-ment in a recent past, various succession stages can be observed, mainly towards the alder forest (Alno-Ulmion) with Alnus glutinosa in the vicinity of the river, and the bog (Ribeso nigri-Alnetum) in local hollows and pine plantation with Pinus sylvestris, Betula pendula and Quercus robur. Willow shrubs of the Salicetum pentandro-cinereae plant associa-tion grow in places with stagnant water, dry places on the river and area elevations (Sumorok, Michalska-Hejduk 2005). The following trees species occur in the flood-plain: Alnus glutinosa, Betula pendula, Pinus syl-vestris, Quercus robur and willows: Salix alba, S. aurita, S. fragilis, S. cinerea, S. pentandra, S. pur-purea, S. triandra, S. viminalis (Sumorok, Kiedrzy ska 2007). Water logged soils (wet) dominate on the floodplain area, mainly ground-gley soils and allu-vial soils. In the part of the floodplain with pine monoculture forest soils from eolian sands occur (Kosi ski 2006 unpubl.).

Sampling

Based on the Digital Terrain Model four sites representing various heights and inundation levels were established in the floodplain: low flood (L), medium high flood (M), high flood (H), flow conditions (FC), with arborescent plant species in different moist conditions (Fig. 1).

Soil Soil analyses of granulometric composition were provided by Bouyoucosa-Casagrande areo-metric method in Proszynski’s modification. Physical and chemical analyses of soil as well as N, P, K content were measured according to the standard analytical methodology used at the Laboratory of Regional Chemical-Agricultural Station in ód . The measured parameters are presented in Table I.

Plants Field studies of meadows and sedge com-munities were conducted at the end of July 2003 and the beginning of August 2005. The plant communities were investigated using the com-monly applied Braun-Blanquet method, including modifications by Matuszkiewicz (2001). Twenty six phytosociological releves were executed in randomly selected sites. The syntaxonomic sys-tem and the syntaxonomic affiliation of species were adopted after Matuszkiewicz (2001); Kucharski, Michalska-Hejduk (1994), and Balátova-Tulá ková (1978). Nomenclature of species was based on the Checklist of vascular plants of Poland (Mirek et al. 1995).


Mycorrhizal samples Plant roots were collected in spring, sum-mer and autumn in 2003, 2004 and 2005 sea-sons. Ectomycorrhizal samples were collected from under the trees and bushes species. The roots were thoroughly rinsed in tap water and analysed quantitatively and qualitatively under a light and stereoscopic microscope examination. Ectomycorrhizal fungi sporocarps and mycor-rhizal types were identified using conventional procedures with keys (e.g. Moser 1978; Moser et al. 1985-1987; Nespiak 1975, 1981, 1990; Galli 1996; Skirgie o 1960, 1991, 1998; Breitenbach, Kränzlin 1984-1995) under a light microscope Nicon Eclipse 400 and under a ster-eoscopic microscope. Different mycorrhizal morphotypes were described and their viability were determined by Agerer methods (1986,

1987-1997, 1991). The length of mycorrhizal roots was assessed using the gridline intersec-tion method and the frequency F of ECMF myc-orrhizae were determined using the gridline intersection method (Tennant 1975; Vogth et al. 1983). For ECM roots, it was calculated using the formula (Kosi ski 2006 unpubl.; Sumorok, Kiedrzy ska 2007):

F [%] = (number of mycorrhizal roots) · 100 · (number of all roots)-1

Similarity index

Similarity of species composition according to Sörensen index between ectomycorrhizal fungi communities (S0-S3 sites) was calculated using the formula:

404

Fig. 2. Plant cover map of the experimental Pilica River floodplain (Sumorok, Kiedrzy ska 2007 modified).

Parameters Site

pH (in KCl)

Total N (%)

P2O5 (mg 100g-1)

K2O (mg 100g-1)

Mg (mg 100g-1)

FC (flow condition) 5.23 <0.01 4.3 1.5 0.5

L (low flood) 3.65 0.29 1.9 3.5 1.3

M (medium high flood) 4.32 0.24 1.1 1.5 1.8

H (high flood) 5.17 0.21 3.1 1.5 0.7

Table I. Chemical properties of soil samples from sites on the Pilica River floodplain.


P [%] = 2c 100· (a + b)-1

where: P – similarity index; c – number of species or morphotypes common for a and b; a – number of species or morphotypes in first site; b – number of species or morphotypes in second site.

3. Results

The majority of ectomycorrhizal fungi were present in trees in dry habitats – willow bushes and pine monoculture (Salix species, Betula pendula and Pinus sylvestris). The species com-position were was determined - in total c.a. 40 species of ectomycorrhizal fungi was detected

and 15 morphological types (Table. II). The diversity of ECMF morphotypes increased sig-nificantly from flooded to dry habitats. For H sites the largest number of ectomyc-orrhizal fungi and morphotypes were observed – 31 species, for M sites 23 species, whereas the least for L and FC sites, accordingly 13 and 6 species and morphotypes. Ectomycorrhizal fungi sporocarps connected with Pinus sylvestris and Betula pendula were very common species from genera Thelephora (Fig. 3), Russula, Lactarius, Amanita and Cortinarius especially for M and H sites. From species connected with Alnus glutino-sa very often noted Russula decolorans and Naucoria sp. Four willow species (Salix alba, S.

FC (flow conditions)

L (low flood)

M (medium high flood)

H (high flood)

Hebeloma sp. Inocybe sp. Phialocephala sp.

Morphotype Salix 1 Morphotype Salix 2 Morphotype Salix 3

.

Amanita muscaria (L.) Boletus edulis Bull. Cantharellus cibarius Fr. Cenococcum geophilum Fr. Laccaria laccata (Scop.) Cooke Lactarius torminosus (Schaeff.) Gray Lactarius turpis (Bull.) Pers Lycoperdon perlatum Pers. Lycoperdon umbrinum Pers. Paxillus involutus (Batsch) Fr. Russula decolorans (Fr.) Fr.

Russula sp. 1 Russula sp. 2

Scleroderma citrinum Pers. Suillus variegatus (Sw.) Kuntze Tricholoma flavovirens (Pers.) S. Lundell Xerocomus badius Fr. (Kühner)

Morphotype Boletus Morphotype Chroogomphus Morphotype Cortinarius Morphotype Rhizopogon Morphotype Russula Morphotype Tuber Morphotype Xerocomus

Amanita citrina (Schaeff.) Pers. Amanita fulva (Schaeff.) Fr. Amanita muscaria (L.) Cenococcum geophilum Fr. Cortinarius flexipes (Pers.) Fr.

Cortinarius sp. 1 Cortinarius sp. 2

Hebeloma pusillum J.E. Lange Inocybe lacera (Fr.) P. Kumm. L. camphoratus (Bull.) Fr Lactarius rufus (Scop.) Fr. Lactarius sp. Lactarius torminosus (Schaeff. Gray Lactarius vietus (Fr.) Fr. Leccinum scabrum (Bull.) Gray Lycoperdon perlatum Pers. Paxillus involutus (Batsch) Fr. Phialocephala sp. Rhodocollybia butyracea (Fr.) Russula chamaleontina (Lasch) Fr. Russula aeruginea Fr. Russula emetica (Schaeff.) Pers. Russula fragilis Fr.

Russula sp. 2 Russula sp. 3

Scleroderma citrinum Pers. Thelephora terrestris Ehrh. Xerocomus chrysentheron (Bull.)

Quél. Morphotype Russula Morphotype Suillus Morphotype Salix 1 Morphotype Tuber

Amanita citrina (Schaeff.) Pers. Cenococcum geophilum Fr. Hebeloma sp. Inocybe sp. Phialocephala sp. Russula paludosa Britz. Thelephora terrestris Ehrh. Xerocomus chrysentheron (Bull.) Quél.

Morphotype Inocybe Morphotype Cortinarius Morphotype Salix 2 Morphotype Salix 3 Morphotype Tomentella

Number of species and morphotypes

6


23


31


13

Table II. Occurrence of ectomycorrhizal species and symbiont morphotypes in the arborescent plant roots in the experimental Pilica River floodplain (in years 2003-2005).


fragilis, S. pentandra and S. triandra) forming ectomyc-orrhizae were recorded. Other willow species (Salix aurita, S. cinerea, and S. viminalis) formed either both arbuscular mycorrhizae (AM) and ectomycorhizae (ECM) or AM only, e.g . Salix purpurea (Sumorok, Kiedrzy ska 2007). Under the the bushes Salix alba, S. cinerea and S. fragilis noted sporocarps from genera Inocybe and Hebeloma. Taking mycorrhizal sporo-carps into account for M sites a wide variety of sporo-carps was noted, whereas for more dry sites like L a larger diversity of morphotypes (Table II). Ectomycorrhizal morphotypes of birch Betula pendula, scotch pine Pinus sylvestris black alder, Alnus glutinosa and willows Salix sp, col-lected over 3-year vegetative seasons on flood-plain were characterized by morphological fea-tures the total of 15 described. Seven identified types belonging to genera: Russsula, Lactarius, Paxillus, Tuber, Rhizopogon, Chroogomphus and Cortinarius were noted on site M and H. Three types unidentified exhibited hyphal mantle struc-ture very similar to Inocybe, Naucoria and Hebeloma genera. The Cenoccocum geophillum and other black mantle species prevailed in the roots of trees and bushes in periodically flooded habitats and in dry habitats. Other dark mantles, Phialocephala sp. morphotypes, were also the most frequent ectomycorrhizal genera in wet hab-itats (Table II). All of the willow species noted in the Pilica River floodplain can form mycorrhizae - both ectomycorhizal fungi and arbuscular fungi were symbionts of Salix viminalis and Salix cinerea. Ectomycorrhizal fungi were present in trees’ roots in dry habitats there are Salix alba, S. triandra, S. pentandra a n d S . f r a g i l i s ( S u m o r o k , Kiedrzy ska 2007). Three uni-dentified types belong to willow species – morphotype Salix 1-3. After calculation of the simi-larity index of species composi-tion according to Sörensen for-mula the greatest similarity between FC and H sites (accord-ingly 50% and 80%) was found, taking occurence of mycorrhizal

sporocarps and morphotypes into account, where-as for L - dry and M semi-wet sites index for spo-rocarps was 33% and for morphotypes 25% (Table III). Ectomycorrhizal root length was the greatest in medium water level (M) and highly (H) flood-ed habitats, for example (in 100 cm3 of soil core): Alnus glutinosa - 491 mm, Betula pendula – 424 mm, Pinus sylvestris – 333 mm, Salix fragilis – 326 mm. In contrast to Pinus sylvestris (50 mm in 100 cm3 of soil core) in dry habitats. The mycorrhizal frequency depended on the tree species and moisture gradient. It ranged between 65 and 90% on the average, and rarely reached 100%. A decrease in mycorrhizal colonisa-tion occurred during flooding. The highest ECMF frequencies were recorded for Pinus sylvestris and Betula pendula (100%) and Salix cinerea (95%), Salix fragilis (85%) (Sumorok, Kiedrzy ska 2007) in dry habitats. In contrast, in highly flooded habi-tats the value for Pinus sylvestris and Betula pen-dula was 57% and 77% respectively.

406

Fig. 3. Thelephora terrestris morphotype (photo: Piotr Mleczko).

Sporocarps /morphotypes FC L M H

FC (flow conditions) - 0 0 50

L (low flood)

0 - 33 8

M (medium high flood)

20 25 - 24

H (high flood)

80 18 0 -

Table III. Similarity of species composition according to Sörensen index (QS) between ectomycorrhizal fungi (sporocarps and morphotypes) in plant communities various sites located at the floodplain Pilica River floodplain.


4. Discussion

Soil inundation inhibits root formation and branching, and growth of existing roots and myc-orrhizae. Waterlogging induces multiple physio-logical disfunction in plants eg. absorption of macronutrients is decreased in flooded plants because of root mortality, loss of mycorrhizae, and suppression of root metabolism (Kozlowski 1997). Despite the fact that extremely humid or dry soil conditions make a negative impact on mycor-rhiza, symbiotic partnership enable floodplain plant communities to tolerate water level fluctuations and eliminate water stress. On the floodplain study site mycorrhizal herbaceous and arborescent plants actively accumulate, especially in their leaves and roots tissues, large amounts of phosphorous com-pounds. Samples of different plants tissues indicate that higher phosphorous concentration occurs in leafs and roots than in the soil. Due to this mycor-rhizal activity the excess of this compounds is being eliminated from soil and water of Pilica river. The studies on the mycorrhizal status of plants, conducted in the Pilica River floodplain, show that the moisture gradient and water flow dynamics regulate the occurrence of plant and ectomycorrhizal symbionts communities. Fifteen morphotypes of ectomycorrhizal sym-bionts and 40 species were recorded in the flood-plain. This diversity level is high for medium water-logged habitats and dry sites. A higher occur-rence frequency of mycorrhizae was noticed in dry habitats (L) and at medium high water le-vels (M). The similar relationship was noted by Davis and Shearer (1999) The nature of the first formed mycorrhiza and the extent to which either type of colonization per-sists into the adult condition appears to depend upon local soil conditions. Each arborescent plant species grows in part-nership with the typical fungi species or forms a partnership with common fungi species (Molina et al. 1992). Fungi species observed on Pilica flood-plain sites belong to common species, which asso-ciate with different trees as pine, birch and alder. Pine and birch are exclusively ectomycorrhizal but some arborescent plants like willow or alder may also form arbuscular mycorrhizas. Members of Salicaceae in particular have such ability and can be predominantly colonized by arbuscular or ectomyc-orrhizal fungi (Sumorok, Kiedrzy ska 2007). Arbuscular mycorrhizas are typical of Salix or Populus species growing on mineral- or nutrient-rich soils, while ectomycorrhizas predominate in organic soils (Smith, Read 1997). Some of the mycorrhizal fungi species like Laccaria, Hebeloma or Thelephora prefer more humid areas and belong to hydrophilic fungi (Unestam, Sun 1995; Colpaert, Vertuyft 1999), what presented research findings confirm. Hil-

szcza ska (2003, 2005a, 2005b) and Hilszcza ska, Sierota (2006) were finding Thelephora terestris species in laboratory as well as in field researches, stating that this species is widely spread in many terrestrial ecosystems. Genus Alnus is reported to be AM under some circumstances and shows growth responses to inoculation with ECM fungi, what has a connec-tion with soil reaction and water level fluctuation (Smith, Read 1997). Presented research data indi-cate that AM occurs in young specimens, while ECM are present in older ones in acid substrate for L and M sites (pH 3.65 and 4.32). On the Pilica floodplain ectomycorrhizal genera Russula and Naucoria were observed, similar species were noted by Becerra et al. (2005). In flow conditions sites (FC) and high water-logged habitats (H) black mantle morphotypes were recorded as dominant. Cenococcum geo-philum is considered to be widely spread in dry habitats (Piggot 1982), however it was the most common morphotype in often flooded willows and birches, which are easily adapted to survive water-logging conditions (Sumorok, Kiedrzy ska 2007). In Hilszcza ska experimental research (2003, 2005b) Cenoccocum geophilum was found only on Pinus seedlings from dry treatment. However Coleman et al. (1989) pointed out that some strains of Cenococcum geophilum, immune to water-log-ging, exist. In the presented publication, qualitative and quantitative studies are conducted as fine roots are considered to be ecological indicators (Vogt et al. 1983), and the diversity and vitality of mycorrhizae indicate the status of the entire rhizosphere. Flooding typically decreases absorption of major macronutrients, especially N, P, K by flood intolerant plants. Nitrate N is rapidly depleted by denitrification under condition of soil hypoxia. Inhibition of uptake of nitrate also is associated with effects low oxygen tension on root metabo-lism (Kozlowski, Pallardy 1984). Decreasing the O2 concentration around Pinus elliotti roots inhib-ited absorption of P, Ca and Mg (Shoulders, Ralston 1975). Flood-tolerant species often absorb more mineral nutrients in response to soil inundation than unflooded, well-watered plants e.g. Acer rubrum, Populus deltoids, these plants has been attributet to several adaptation e.g. aerenchyma tis-sue and adventitious roots which play role in oxy-gen transport in trees and oxidation of the rhizo-sphere. In well-aerated soils, mycorrhizae increase mineral uptake, largely because of their extensive absorbing surface. The high absorbing capacity of mycorrhizal roots is also associated with reduction in air gaps between the roots and soil particles, low resistance of the fungus to ion transport, increased rate of root growth, and more rapid hydrolysis of certain soil nutrients (Kozlowski 1997).


The present study suggests that mycorrhizae may increase the size and the phosphorus concen-tration of wetland plants under both dry and wet conditions. Consequently, phosphorus content in the soil and ground water was additionally exam-ined. Soil properties and moisture conditions affect ectomycorrhizal fungal communities. Because of the importance of mycorrhizae in nutrient assimi-lation, including phosphorus assimilation, it is highly recommended to explore in depth the influ-ence of flooding and nutrient concentrations on the formation of mycorrhizae in wetlands.

Conclusions

Fifteen morphotypes of ectomycorrhizal symbionts and 40 species were recorded in the floodplain. The studies on the mycorrhizal status of plants, conducted in the Pilica River floodplain, show that the moisture gradient: water flow and stagnation, regulate the occurrence of plant and ectomycorrhizal symbionts communities. Cenococcum geophilum prevailed in the roots of trees and bushes in periodically flooded habitats and in dry habitats. Because of the importance of mycorrhizae in nutrient assimilation, including phosphorus assimilation, it is highly recommended to explore in depth the influence of flooding and nutrient concentrations on the formation of mycorrhizae in wetlands.

Acknowledgments

The research was supported by the Polish Ministry of Education and Science, projects:3 PO4G 020 24, 2 PO4F 053 28 and Demonstration Project under the auspices of UNESCO and UNEP “Application of Ecohydrology and Phytotechnologies for Water Resources Management”. The authors would like to express their thanks to dr Wojciech To oczko for making soil data available from the common researches, dr Marcin Kiedrzy ski for the map of plant cover and Renata Ch ci ska for her technical support, devoted spare time in helping the researchers and effort devoted to the research.

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