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University of South Bohemia in České Budějovice

Faculty of Biology

Bachelor thesis

Comparative study of meadow sedges No them ost of IC

Jan Košnar

2003

Supervisor: prof. RNDr. Jan Lepš CSc.

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Košnar, J. (2003): Comparative study of meadow sedge. [Comparative study of meadow sedges.

Bc. Thesis]. - 37 p., Faculty of Biological Sciences, University of South

Bohemia, Ceske Budejovice, Czech Republic.

Annotation:

Responsible of seven Carex species, coexisting in wet meadow, to soil nutrient level

and interspecific competition were studied in two greenhouse experiments and the results

They were compared with data from the field. Differences found in responses suggest niche

Differentiation in the species.

The work was financed from the grant FRVŠ 1284. It is part of the project "Mechanisms

coexistence of species in semi-natural meadow communities ”, the researcher is prof. RNDr.

Jan Lepš CSc.

I declare that I have written this Bachelor Thesis myself, only with use

cited literature.

In České Budějovice on 12.5.2003

Jan Košnar

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Thanks

Here I would like to thank everyone who has helped me somehow during my work. Mine

the supervisor, Jan Lepš - Šusp, for his good ideas and valuable advice to me willingly

whenever it was needed. Jirka Košnar - brother - and Vojta Lant for great

assistance in the field and in the greenhouse. To my faculty friends who have encouraged me many times,

when something went wrong. And to my whole family for their understanding and self-sacrificing support

I've always received from her.

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CONTENT

1. Introduction

2. Methodology

2.1. Study species and their habitat

2.2. Experiments

2.2.1. Study of soil nutrient response

2.2.2. Study of reaction to interspecific competition

2.3. Field observation

2.4. Statistical processing

2.5. Nomenclature

3. Results

3.1. Experiments

3.1.1. Reaction to the amount of soil nutrients

3.1.2. Responses to interspecific competition

3.2. Field observation

4. Discussion

4.1. Experiments

4.1.1. Reaction to the amount of soil nutrients

4.1.2. Responses to interspecific competition

4.2. Field observation

5. Conclusion

27 Mar:

6. Literature

7. Attachments

7.1. Localization of photographed areas

7.2. Phytosociological images

7.3. Schematics of rhizome systems of studied species

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1. INTRODUCTION

Plants are modular organisms (Begon et al. 1997); that their body is composed

a plurality of similar parts (modules); loss of one or more

from these parts does not necessarily lead to the extinction of the whole organism, as the modules may be

replaced by the ability of meristems to retain division activity throughout their lives

(Klimesova & Klimes 1997). This allows for clonal growth, which can be defined as

vegetative formation of modules potentially capable of self-existence (Herben et al. 1994).

Such modules are called arms, sets of arms of the same genotype (ie clones formed)

from one parent organism) then genets. In plant ecology with clonal plants

means those which consist of a system to a different extent morphologically and physiologically linked

ramet; not individuals of the same genotype but isolated from each other as they do

in a broader genetic perspective (Herben et al. 1994).

In some groups of vascular plants, clonal growth is a condition for survival, for others it is not

indispensable but nevertheless advantageous (Mogie & Hutchings 1990). Thanks to clonal growth

on the one hand, it reduces the risk of the disappearance of a particular genotype because, as mentioned above,

As a result of mortality lost shoulders can be replaced with new ones and in addition facilitates

acquisition of resources in spatially and temporally heterogeneous environment, their transport to parts

currently under unfavorable conditions and storage in it

adapted bodies such as tubers and rhizomes (van Groenendael & de Kroon 1990,

de Kroon et al. 1998).

It is a known fact that some plants forming the lateral processes (which is most common

reported, but not, as warned by Klimešová & Klimeš 1997, the most common type of clonal

in a heterogeneous environment may exhibit behavior for which it was adopted from ecology

known term "foraging". It is conditioned by morphology

plasticity (ie environment controlled variability in morphological characteristics, eg length)

processes or their number; de Kroon et al. 1994) and allows the plant to place the arms in the

surfaces ( patches ) with favorable conditions, and conversely to avoid adverse pads. Except

the frequently mentioned foraging strategies, de Kroon & Schieving (1990) described two others,

referred to as "consolidation strategy" and "conservative strategy"

('Maintenance'), the first being characteristic of the species acting as vegetative

dominated, while the latter are exploited by resource-poor habitat plants.

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However, the morphological variability of clonal plants is not necessarily plastic

(de Kroon et al. 1994). According to Krahulec (1994), there are essentially two points of view

variability: the first perceives it as a consequence of selection pressures from the community

and habitats (ie as a consequence of the current, relatively short - term biotic effects)

and abiotic factors), while the latter considers the key long term, especially

phylogenetic processes, which implies that variability of characters cannot be explained only by knowledge

the environment in which the species is present.

As Goldberg (1990) states, plants usually interact with one another

mediator (which may be eg abiotic sources, pollinators, herbivores, etc.),

on the one hand, they influence the amount of the mediator and, on the other hand, the quantity itself

react. In this respect, competition is an example of an interaction where the mediator is abiotic

sources whose plants have a negative impact (they deplete them) and at the same time

they react positively (they use them for growth and reproduction). They are then resistant to competition

plants that can either draw resources faster than others or endure low availability

resources.

Plants compete for light, water and mineral elements (generally nitrogen,

phosphorus and potassium). Competition for light tends to be asymmetric, ie. larger individuals gain much

a larger proportion of the total available resource than the size ratio

competitor (higher plants overshadow lower ones). Competition for soil resources (water and mineral

nutrients), on the other hand, is considered symmetrical (the resource is divided among the individual competitors)

more or less proportional to their size, eg because of the larger ones

Root systems of larger plants often overlap zones, from which individual

roots can acquire resources - Schwinning & Weiner 1998). Fitter & Hay (1989) distinguished

in plants, in relation to the availability of mineral elements, two sets of properties.

They are better applied in rich soils, which are more spatially and temporally variable in the amount of nutrients

species with flexible, plastic growth, rapid formation of above-ground organs and the resulting

greater ability to successfully compete for light; on the other hand, poor soil types are less

morphologically and physiologically flexible, they do not more or less respond to changes in nutrient concentration,

photosynthesis products invest primarily in the formation of roots and grow slowly - these are Grim's

S-strategist.

Sedge ( Carex L.) are typical clonal plants; their shoulders, represented

branches of horizontally growing underground stem (rhizome), bear roots, leaves

as well as flower-bearing stems, so they can provide all vegetative functions and sexual functions

reproduction. According to the growth method of the rhizome system, they were distinguished by Jermy & Tutin (1982)

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four growth forms, differing in the type of branching (monopodial or sympodial), frequency

branching, length of internodes and their number. Most of our sedges (which are in the Czech Republic)

80 species) inhabits wetland habitats as littoral zones of stagnant and flowing waters,

swamp alder, fens, peat bogs and wet meadows (Kubát et al. 2002).

The latter type of habitat is located near České Budějovice

Enclosures, economically extensively used (and environmentally friendly in the last nine years)

intensively studied - see ia Špačková et al. 1998, Lepš 1999 ad.) Species-rich meadow,

where twelve species of sedges coexist (Coal 1998).

It is generally believed that coexistence of organisms (even close relatives) is possible if they are

these organisms are at least to some extent ecologically different (Begon et al. 1997)

and thus respond differently to environmental factors, or in other words, their ecological niches

they do not overlap too much and therefore there is no competition exclusion.

Based on this assumption, the objectives of the work were set as follows:

Try to find differences in greenhouse manipulation experiments

selected ecological characteristics of seven species of meadow sedge, namely differences

in replies to:

(a) the amount of nutrients in the substrate

(b) the action of the competing species

2) compare the results obtained with field data obtained from:

a) prof. J. Lepse during a long-term field experiment at the locality Ohrazení

(for details see Better 1999)

b) author of this work during phytosociological imaging of wet meadows with occurrence

sedge.

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2. METHODOLOGY

2.1. Study species and their habitat

The following seven were chosen for experimental study of ecological properties

kinds:

Carex demissa Hornem. - sloping sedge,

Carex hartmanii Cajander - Hartman's Sedge,

Carex pallescens L. - Pale Sedge,

Carex panicea L. - Millet Sedge,

Carex pilulifera L. - Pill Sedge,

Carex pulicaris L. - flea sedge,

Carex umbrosa Host - Shady Sedge.

They are more subtle representatives of the genus Carex (compared to species forming communities

high sedges of the order Magnocaricetalia ). Their rhizomes branch out sympodially, especially

Carex hartmanii is able to produce very long protuberances, somewhat shorter than C. panicea.

C. demissa , C. pallescens and C. umbrosa have a tiny habitus with minimal frequency

and the length of the projections. C. pilulifera with C. pulicaris then represent a "transient type" with protuberances

quite common, though quite short. Schematics of rhizome systems created by

in natural conditions, they are part of the appendix.)

All studied species come from the locality Ohrazení, located 10 km southeast

from České Budějovice (48 ° 57´ N, 14 ° 36´ E, altitude 510 m). Average annual

temperature is 7.8 ° C, average annual rainfall 620 mm (weather station data)

in České Budějovice). It is traditionally extensively managed (once to

twice a year mowed) oligotrophic wet meadow, with species composition corresponding to

most of the Molinion , and to a lesser extent Violion caninae . None of the above

those sedges cannot be regarded as a dominant feature of the enclosure, but Carex hartmanii

and C. panicea in some parts of the meadows achieve greater coverage.

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2.2. Experiments

In two consecutive vegetation seasons (2001 and 2002) were carried out

experiments in the greenhouse, the purpose of which was to identify the reactions of individual species to

manipulated environmental factors - amount of soil nutrients and intensity of interspecific competition.

The starting material in both cases was sterile arms taken initially

vegetation season (2.4.2001, or 4.4.2002) in the locality Ohrazení.

2.2.1. Study of the reaction to the p of func nutrients

The plants were grown for 96 days in pots containing the base substrate (peat

with 1: 2 sand) with various additions of commercial NPK fertilizer (19% N, 6% P,

12% K): one third of all pots were left without fertilizer, to the other one 1 g was added

and up to the remaining 4 g. This created three types of substrate - low, medium and high

mineral nutrients. Five plants were placed in each substrate type (repetition)

from each kind; however, some mortality was lost due to mortality during the experiment

of plants, the actual number of repetitions used in the statistical data evaluation is therefore given

in Table 2-1.

Table 2-1. Repeat counts for statistical evaluation of the manipulated quantity experiment

soil nutrients; inequalities are caused by mortality during the experiment.

amount of nutrients

low

medium

high

C. demissa

C. hartmanii

C. pallescens

C. panicea

C. pilulifera

C. pulicaris

C. umbrosa

All plants were weighed for their fresh biomass (for

this and all other weighings were used with PRECISA laboratory balances with an accuracy of 0.01 g); after

at the end of the experiment, all plants were then harvested, the arms were counted in the clones

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and underground projections, the weights of fresh biomass of the plants were determined and (after drying in progress)

24 hours at 80 ° C) as well as the weight of above-ground dry biomass (leaves, flowering stems)

and underground (roots, rhizomes) parts. Relative values were determined from obtained values

growth rates (RGR) and root / shoot ratio (R / S) according to

of the following formulas:

m 1 - mass of dry biomass at the end of the experiment,

m 0 - mass of dry biomass at the beginning of the experiment, derived from the regression of dry biomass to

the end of the experiment on fresh biomass at the end of the experiment calculated for each species

(the same procedure was used by Sagittarius 2001),

m k - mass of dry root biomass,

m n - mass of dry above-ground biomass.

2.2.2. Study of reaction to interspecific competition

Plants were grown for 98 days, again in peat + sand substrate 1: 2. One third of the pots

it contained only one sedge arm at the beginning of the experiment. In addition, another third was sown

15 grains of Holcus lanatus grass and for the remaining third 45 grains of H. lanatus . After germination she was

H. lanatus density adjusted to one-third of pots containing 5 shoots of this species

and another third of 25 shoots (and these numbers were newly removed during the experiment

accretion shoots of H. lanatus maintained). This created three experimental environments,

with zero, moderate and high intensity interspecific competition. Every environment was

placed, as in the previous case, five repetitions of each type of sedge;

even this time, some losses due to mortality were recorded, the final numbers of repetitions reported

Table 2-2.

Prior to the experiment, the weight of fresh biomass was determined for all plants.

In the course of the experiment, the number of arms was repeatedly determined by 28, 52, 78 and 98

days of the experiment. After the plants were harvested, the subterranean

RGR

SR =

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weights of fresh and dry biomass of aboveground and underground parts were determined and calculated

RGR and R / S.

Table 2-2. Repeat counts for statistical evaluation of the manipulated intensity experiment

interspecies competitions; inequalities are caused by mortality during the experiment.

intensity of competition

none

moderate

strong

C. demissa

C. hartmanii

C. pallescens

C. panicea

C. pilulifera

C. pulicaris

C. umbrosa

2.3. Field observation

Phytosociological imaging was performed during the 2001 and 2002 vegetation seasons

wet meadows with sedges. A total of 9 localities were visited in the south and east

Bohemia and 18 records were made (see appendices). The size of the scanned areas was 5 x 5 m.

The seven-member Braun-Blanquet scale was used to assess species coverage

abundance and dominance (Dust 2001). Two squares of side were measured in each frame

50 cm and the total above - ground biomass was taken and dried (24 hours)

at 80 ° C) and weighed; the obtained dry weight value served as a measure of productivity

of the site. In addition, it was recorded whether the captured site was mowed or not.

2.4. Statistical processing

Data from both experiments were evaluated using two-way analysis of variance

(or two-way analysis of variance for repeated measurements if the value was variable

collected during the experiment multiple times) performed in Statistica for Windows

5.5 (StatSoft, Inc. 1999).

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Data obtained by site imaging were evaluated by major component analysis (PCA)

and Redundancy Analysis (RDA) in Canoco for Windows 4.5 (Ter Braak & Šmilauer

2002). The use of linear methods was evidenced by the very nature of the data (basically collected

in one type of habitat, ie in relatively homogeneous vegetation), so "control" the gradient length after

computation of detrended correspondence analysis (DCA; according to Lepš & Šmilauer 2000, Herben

& Münzberg 2002).

All the species captured in the images were included in the analysis of the major components,

the phytosociological linkages of studied species were investigated.

To redundancy analysis included only species Carex and was thus ascertained

dependence of composition of meadow sedge communities on environmental characteristics.

The species cover values were converted from ordinal to Braun-Blanquet scale

a seven-member scale (according to Lepš & Šmilauer 2000). Then the data was no longer in any way

transformed and due to the mentioned nature of the data, no standardizations were performed

(according to Herben & Münzberg 2002). Statistical significance of canonical axes and variables

Monte Carlo permutation tests evaluated. The ordination diagrams were constructed

in CanoDraw for Windows 4.0 (Ter Braak & Smiluer 2002).

2.5. Nomenclature

All plant species names have been united by Kubat et al. (2002).

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3. RESULTS

3.1. Experiments

3.1.1. Responding to the amount P of func nutrients

(a) total above-ground biomass production

Influence of both factors, ie species and quantity of nutrients used in

two-way analysis of variance was statistically significant. Factor interaction then

although it was not conclusive at the 5% level of significance, it was found

the probability of error of the first kind was rather low (see last line

Tables 3-1); Obviously, rather than the validity of the null hypothesis was on

five percent significance level inconclusive result due to less

the strength of the test, which resulted from fewer repetitions and disproportions

in the number of repetitions between groups (see page 5 and Table 2-1). When used

one-way analysis of covariance with factor "type" and "amount of nutrients" (resp.

square root values of fertilizer dose weights) as

the linear quantitative predictor (covariate) was the factor interaction

with conclusive covariate (Table 3-2). The obvious interaction shows that:

the amount of nutrients did not have the same effect in all species. Four of them ( Carex

demissa , C. hartmanii , C. pallescens and C. panicea ) at increased availability

nutrients have produced significantly higher above-ground biomass than at low availability,

while the remaining three ( C. pilulifera , C. pulicaris and C. umbrosa ) for changes

they did not react in substrate potency (Fig. 1).

Table 3-1. Results of a two-way analysis of variance for the "total

above-ground biomass production ”. The left column shows the tested factors

and their interactions (denoted by " ×") , in the middle value of F-statistics given

number of degrees of freedom (in parentheses), in the right level of significance.

species

F (6; 65) = 15.781

P <0.001

nutrients

F (2.65) = 5.599

P & lt; 0.01

type × nutrients

F (12.65) = 1.766

P = 0.071

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Table 3-2. Results of a one-way covariance analysis for the “total

above-ground biomass production '; the factor was “kind”, covariate “amount of nutrients”

(Root Transformed Weights of Fertilizer Batches). In the left column are

the tested predictors and their interactions (marked " ×") are shown in the middle

values of F-statistics at given number of degrees of freedom (in parentheses), in the right

achieved significance level.

species

F (6; 78) = 13.336

P <0.001

type × amount of nutrients

F (6; 72) = 3.511

P & lt; 0.01

and

ilu

and

that

í b

and

with

and

with

)

low mn. zivin

Medium Qty. zivin

high pl zivin

Fig. 1. Influence of soil nutrients on total above-ground production

biomass of seven species of meadow sedge. The column corresponds to the average

a positive line for the standard deviation value. CxDemi - Carex demissa ,

CxHart - C. Hartmanii , CxPall - C. Pallescens, CxPani - C. Panicea, CxPilu

- C. pilulifera , CxPuli - C. Pulicaris , CxUmbr - C. umbrosa .

b) root / shoot ratio

The influence of the above factors was significant, but their interaction did not

(Table 3-3): Influence of nutrient levels on root biomass to biomass ratio

above ground was the same for all species, and this ratio was decreasing

with increasing substrate yield (Fig. 2).

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Table 3-3. Results of a two-way variance analysis for the root / shoot variable

ratio ". The left column shows the tested factors and their interactions (labeled

" ×") , in the middle value of F-statistics at given number of degrees of freedom

(in parentheses), at the right level of significance.

species

F (6; 65) = 4.239

P & lt; 0.01

nutrients

F (2.65) = 40.956

P <0.001

type × nutrients

F (12.65) = 0.616

P> 0.8

and

ilu

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

t / s

t ra

tio

low mn. zivin

Medium Qty. zivin

high pl zivin

Fig. 2. Effect of soil nutrients on root biomass ratio

to root / shoot ratio in seven species of meadow sedge. Column

is the mean, the abscissa is the positive standard deviation.

CxDemi - Carex demissa , CxHart - C. hartmanii , CxPall - C. pallescens ,

CxPani - C. panicea , CxPilu - C. pilulifera , CxPuli - C. pulicaris , CxUmbr

- C. umbrosa .

The remaining characteristics to be measured, ie the number of underground projections,

the biomass of the underground parts, the number of arms, and the relative growth rates were not

the amount of nutrients demonstrably affected.

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3.1.2. Responses to interspecific competition

(a) the number of underground projections

C. demissa was excluded from the analysis prior to its implementation ,

C. pallescens and C. umbrosa , which produced none during the experiment

a salient (which is due to their naturally clumsy growth habit, see also

4 and Annexes); leaving them in the analysis would increase the strength of the test,

however, any interpretation of the results would be trivial (the number of projections would be

Logically - it varied primarily between the species that make up them and those that do not).

The values of the variable were demonstrably dependent on species. On

the 5% significance level was the effect of the intensity of interspecific competition,

strictly speaking, inconclusive; however this time the result was caused

much less powerful test (for the reasons already stated on page 9), moreover

Again, the probability of error of the first species was sufficiently low (Table

3-4). Highly conclusive interaction between factors indicated that the effect of competition did

varied among species. There was no count for C. hartmanii , C. pilulifera and C. pulicaris

the processes are demonstrably influenced by the intensity of competition; when

C. panicea, however, the total production of underground processes in the present

competitor significantly decreased (Fig. 3).

Table 3-4. Results of a two-way analysis of variance for the variable "calc

underground projections'. The left column shows the tested factors and their

interaction (denoted by " ×") , in the middle value of F-statistics at given numbers

degrees of freedom (in parentheses), at the right level of significance.

species

F (3.48) = 22.281

P <0.001

competition

F (2.48) = 3.171

P = 0.051

type × competition

F (6; 48) = 3.325

P & lt; 0.01

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CxHart

CxPani

CxPilu

CxPuli

č e

that

íc

ě žk

without competition

moderate competition

strong competition

Fig. 3. Influence of interspecific competition on the number of created underground

outcrops in four species of meadow sedge. The column corresponds to the average

a positive line for the standard deviation value. Carex pulicaris in the environment

strong competition did not create any protuberance. CxHart - Carex hartmanii ,

CxPani - C. panicea , CxPilu - C. pilulifera , CxPuli - C. pulicaris .

(b) total above-ground biomass production

The effect of both factors was significant, unlike their interaction

(Table 3-5), therefore the effect of competition did not differ between species - in all cases

there was a reduction in above-ground biomass production in the presence of the competitor

(Fig. 4).

Table 3-5. Results of a two-way analysis of variance for the "total

above-ground biomass production ”. The left column shows the tested factors

and their interactions (denoted by " ×") , in the middle value of F-statistics given

number of degrees of freedom (in parentheses), in the right level of significance.

species

F (6; 77) = 12.013

P <0.001

competition

F (2.77) = 10.166

P <0.001

type × competition

F (12.77) = 0.838

P> 0.6

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and

ilu

0.5

1.5

2.5

3.5

and

that

í b

and

with

and

with

)

without competition

moderate competition

strong competition

Fig. 4. Influence of interspecific competition on total above-ground production

biomass in seven species of meadow sedge. The column corresponds to the average

a positive line for the standard deviation value. CxDemi - Carex demissa ,

CxHart - C. Hartmanii , CxPall - C. Pallescens, CxPani - C. Panicea, CxPilu

- C. pilulifera , CxPuli - C. Pulicaris , CxUmbr - C. umbrosa .

The results (Table 3-6) were very similar to the previous case.

Again an inconclusive interaction between factors points to the same (and negative)

the effect of competition on the production of underground biomass in all species (Fig. 5).

Table 3-6. Results of a two-way analysis of variance for the "total

underground biomass production '. The left column shows the tested factors and

their interaction (denoted by "×"), in the middle value of F-statistics given

number of degrees of freedom (in parentheses), in the right level of significance.

species

F (6; 77) = 11.406

P <0.001

competition

F (2.77) = 7.883

P <0.001

type × competition

F (12.77) = 0.898

P> 0.5

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15 Dec

and

ilu

0.5

1.5

2.5

that

í b

and

with

and

with

)

without competition

moderate competition

strong competition

Fig. 5. Influence of interspecific competition on total underground production

biomass in seven species of meadow sedge. The column corresponds to the average

a positive line for the standard deviation value. CxDemi - Carex demissa ,

CxHart - C. Hartmanii , CxPall - C. Pallescens, CxPani - C. Panicea, CxPilu

- C. pilulifera , CxPuli - C. Pulicaris , CxUmbr - C. umbrosa .

d) relative growth rate

It was shown a significant effect of both factors while their interactions took place

appeared to be insignificant (Table 3-7) - RGR of all species with increasing

intensity of competition decreased (Fig. 6).

Table 3-7. Results of a two-way analysis of variance for the variable “relative

growth rate ”. The left column shows the tested factors and their interactions

(denoted by "×"), in the middle F-statistics at given number of degrees

freedom (in parentheses), at the right level of significance.

species

F (6; 77) = 18.215

P <0.001

competition

F (2.77) = 21.821

P <0.001

type × competition

F (12.77) = 0.805

P> 0.8

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and

ilu

0.005

0.01

0.015

0.02

0,025

0.03

0,035

tiv

í rù

with

and

with

t (d

n -1

)

without competition

moderate competition

strong competition

Fig. 6. Influence of interspecific competition on relative growth rates

seven species of meadow sedge. The column corresponds to the average

a positive line for the standard deviation value. CxDemi - Carex

demissa , CxHart - C hartmanii , CxPall - C. pallescens , CxPani -

C. panicea , CxPilu - C. Pilulifera, CxPuli - C. Pulicaris , CxUmbr -

C. umbrosa .

e) clone growth (rate of formation of new arms)

Statistically significant interactions (Table 3-8) between "species" and "time" factors,

respectively. “Competition” and “time” show that the speed of the creation of new arms has been

dependent on species, respectively. on the number of ramets competing plants

(with increasing competitor abundance the growth of the observed clone slowed).

Highly evident triple interaction between "species", "competition" and "time"

means that the effect of interspecific competition on the rate of ram formation is between

differed species of sedge studied; Carex demissa , C. pallescens ,

C. panicea and C. pilulifera were more strongly affected than C. hartmanii ,

C. pulicaris and C. umbrosa (Fig. 7).

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Table 3-8. Results of two-way analysis of variance for repeatedly measured

variable "number of arms". The left column shows the tested factors and their

interaction (denoted by "×"), in the middle value of F-statistics at given numbers

degrees of freedom (in parentheses), at the right level of significance.

species

F (6; 83) = 14.994

P <0.001

competition

F (2.83) = 12.858

P <0.001

time

F (3.249) = 42.104

P <0.001

type × competition

F (12.83) = 0.969

P> 0.4

type × time

F (18.249) = 2.690

P <0.001

competition × time

F (6; 249) = 24.046

P <0.001

type × competition × time

F (36.249) = 2.030

P <0.001

Fig. 7. Influence of interspecific competition on clone growth (expressed as velocity

creation of new ramets) of seven species of meadow sedge. Points match

mean values from individual measurements (28, 52, 78 and 98 days of experiment);

rings indicate a non-competitive environment, squares with moderate competition

and triangles with strong competition. CexDemi - Carex demissa , CxHart -

C. hartmanii , CxPall - C. Pallescens, CxPani - C. Panicea, CxPilu -

C. pilulifera , CxPuli - C. pulicaris , CxUmbr - C. umbrosa .

č e

t ra

CxUmbr

CxPilu

CxDemi

CxPani

CxHart

CxPall

CxPuli

Page

Root / shoot ratio was then the only variable for which the intensity of competition did not

demonstrable influence.

3.2. Field observation

Results of analysis of main components (PCA) of species composition variability

imaging wet meadows communities are summarized in Table 3-9 and shown in Figure 3-9

Fig. 8. On the main gradient found (corresponds to the first ordination axis) are on one

At the end, the species of intensively managed or neglected meadows were placed

( Alopecurus pratensis , Carex hirta , Festuca pratensis , Hypericum perforatum , Lolium

perenne , Scirpus sylvaticus ), while at the end opposite species tied to meadows

extensively farmed ( Achillea ptarmica , Galium boreale , Gentiana

pneumonanthe , Molinia coerulea , Succisa pratensis ), with which they positively correlate

studied sedge species.

Table 3-9. Results of analysis of main components (PCA) of species composition of communities

wet meadows.

ordinal axis number

eigenvalue axis (ratio of explained variability)

0.186

0.152

0,123

0,102

cumulative percentage of explained variability

18.6

33.8

46.1

56.3

Page

19 Dec

Fig. 8. Results of analysis of main components of species composition of meadow communities

sedge. The horizontal axis corresponds to the first, the vertical axis to the second ordination axis. The species studied are

highlighted in bold. Abbreviations are explained in Table 3-13.

To determine the relationship between the composition of sedge communities shot wet

The ordination model was created with the recorded environmental characteristics

two limited (canonical) axes of variability. Monte Carlo permutation test of both

canonical axes showed statistically significant dependence of species composition on these

axes (199 permutations; F = 2.262; P <0.005), both of which together explained approximately

23% of the total variability in species data (Table 3-10).

Page

20 May

Table 3-10. Results of redundancy analysis (RDA) with two canonical axes,

created by a linear combination of two real environmental characteristics, the "presence of mowing"

(two-valued nominal variable) and "habitat productivity" (continuous variable).

Results of Monte Carlo permutation tests of canonical axes and real variables are presented

in the text.

ordinal axis number

eigenvalue axis (ratio of explained variability)

0.18

0.052

0.244

0.169

the proportion of variability explained by canonical axes

0.232

The first axis was strongly negatively correlated with the variable "presence of mowing" while

the second weaker and positively with “habitat productivity” (Table 3-11, Figure 9).

Table 3-11. Correlation of RDA canonical ordination axes with real characteristics

environment (the numbers in the second and third lines are correlation coefficients).

canonical axis number

habitat productivity

0,634

0.7734

the presence of mowing

-0.9678

0.2516

The contribution of individual environmental characteristics was investigated by the gradual method

selection ( forward selection ) and subsequent permutation tests. In the case of “productivity

habitat ”there was no detectable effect (199 permutations; F = 1.18; P> 0.3);

on the contrary, the presence of mowing had a significant influence on the composition of the communities (199 permutations;

F = 3.31; P <0.005). The coverage of most sedges (including studied species) is missing

data for Carex pulicaris and C. umbrosa , which due to its relatively rare

were not captured in any of the randomly placed frames) with this variable

positively correlated (Fig. 9), as shown by species scores on the canonical axis of the partial RDA

with a single explanatory variable - the presence of mowing (Table 3-12).

Page

Table 3-12. Score of studied species on the canonical axis of partial redundancy analysis

with a single explanatory variable (environmental characteristic).

canonical axis (environmental characteristics)

the presence of mowing

C. demissa

0.3693

C. hartmanii

0.5959

C. pallescens

0.2357

C. panicea

0.4140

C. pilulifera

0.4573

Fig. 9. Results of redundancy analysis of species composition of meadow sedge communities.

The horizontal axis corresponds to the first, the vertical axis to the second canonical axis. The species studied are

highlighted in bold. Abbreviations are explained in Table 3-13.

Page

22nd

Table 3-13. Explanations of abbreviations of the species names used in ordination diagrams (Figs. 8 and 9).

AchiMill Achillea Millefolium CentJace Centaurea jacea

MentArve Mentha arvensis

AchiPtar Achillea ptarmica

CeraHolo Cerastium holosteoides MoliCoer Molinia coerulea

AgroCapi Agrostis capillaris

CirsPalu

Cirsium palustre

MyosPalu Myosotis palustris

AjugRept Ajuga reptans

DescCesp Deschampsia planes PlanLanc Plantago lanceolata

AlchemSp Alchemilla sp .

EquiArve Equisetum arvense

PoaTriv

Poa trivialis

AlnuGlut Alnus glutinosa

EquiSylv Equisetum sylvaticum

PoteErec Potentilla erecta

Alopecrat pratensis FestPrat

Festuca pratensis

RanuAcri Ranunculus acris

AngeSylv Angelica sylvestris

FestRubr Festuca rubra

RanuAuri Ranunculus auricomus

AvenPube Avenulla pubescens GaliAlbu Galium album

RanuFlam Ranunculus flammula

BrizMedi Briza media

GaliBore Galium boreale

RanuRepe Ranunculus repens

BetoOffi Betonica officinalis Galli uliginosum

RumeAcet Rumex acetosa

CareEchi Carex echinata

GentPneu Gentiana pneumonanthe SangOffi Sanguisorba officinalis

CarDemi Carex demissa

HolcLana Holcus lanatus

ScirSylv

Scirpus sylvaticus

CareFlac Carex flacca

HypePerf Hypericum perforatum SeliCarv Selinum carvifolium

CareHart Carex hartmanii

JuncArti

Juncus articulatus

StelGram Stellaria graminea

CareHirt Carex hirta

JuncCong Juncus conglomeratus

SuccPrat Succisa pratensis

CareMuri Carex muricata

JuncEffu Juncus effusus

TrifPrat

Trifolium pratense

CareNigr Carex nigra

LathPrat

Lathyrus pratensis

TrifRepe Trifolium repens

CareOval Carex ovalis

LoliPere

Lolium perenne

VeroCham Veronica chamaedrys

CarePall Carex pallescens

LuzuCamp Luzula campestris

Vicici cracca

CarePani Carex panicea

LychFlos Lychnis flos-cuculi

ViolPalu Viola palustris

CarePil Carex pilulifera

LysiVulg Lysimachia vulgaris

Page

4. DISCUSSION

4.1. Experiments

4.1.1. Responding to the amount P of func nutrients

Greater substrate potency (in terms of greater nitrogen, phosphorus and potassium concentration) has led to

increase in total above-ground biomass production in only four of the studied species - Carex

demissa (the most significant change here), C. hartmanii , C. pallescens and C. panicea - while

the remaining three - C. pilulifera , C. pulicaris and C. umbrosa - for the amount of nutrients in no way

did not respond. This is more or less the data of Dostál (1989) reported by C. demissa ,

C. hartmanii and C. panicea as "breeding" soils (in the case of C. panicea even

fertilized, which is in contrast to the results of the Better 1999 experiments and observations

Miner 2001). C. pilulifera often and successfully inhabits acidophilic habitats where it is possible

expect a low availability of mineral nutrients (Fitter & Hay 1989), similar to that of

peat substrates most frequently hosted by C. pulicaris (Kubát et al. 2002).

Even in species whose biomass production is stimulated by greater substrate yield, however

increased availability of nutrients does not induce in complex conditions of the whole community

their greater relative representation. Better (unpublished results) found for all of the above

of the said sedge species a negative correlation with increasing habitat eutrophication (induced by

addition of fertilizer to permanent areas in the locality of the enclosure); in this case it is interesting

a comparison of the extent to which each species has been affected by eutrophication and how this is related

with their resistance to the competitor (see 4.1.2. and Table 4-1).

Due to the fact that the substrate fertility was in the studied sedge species

only the total production of above-ground biomass was increased while the production of underground biomass

(or root) remained more or less constant, with higher levels of nutrients

to decrease the root / shoot ratio. Change this

Klimeš & Klimešová experimentally recorded the ratio under similar conditions

(1994), as a general rule for plants (and whole clones) found in homogeneous plants

her environment is mentioned by Hutchings & Wijesinghe (1997). It can be interpreted as a manifestation of trade-

off between investment in photosynthesizing resp. mineral nutrients of the acquiring organs. When small

the availability of resources in the soil (i.e. their low concentration) is preferable for the plant to maintain

their root system, keeping the ability to draw nutrients from a larger volume of substrate,

Page

even at the cost of reducing the formation of aboveground structures; with sufficient soil nutrient concentrations

then this limitation is ignored (and the development of above-ground photosynthetic organs is important,

since competition for light is now crucial - see Better 1999 and the references inside).

4.1.2. Responses to interspecific competition

Significant decrease in production of both components of total biomass (both aboveground and underground,

therefore, there were no significant changes in the root / shoot ratio) that occurred in all

The species studied indicates that sedges are competitively compared to Holcus lantus

weaker plants. This may be because, for example, their relative growth rates (i.e.

the amount of biomass that a plant can produce from the initial unit

the amount of biomass per unit of time) are (even under conditions where no competitor is present)

and therefore the resources should be sufficient) lower than the RGR of the aforementioned grass species. Sagittarius (2001)

for H. lanatus, the most common RGR value is 0.06, while the RGR of the study sedge

are in the range of about 0.01 ( Carex umbrosa , C. pulicaris ) to 0.03 ( C. demissa ), i.e.

two to six times lower. The weaker competitive abilities of the studied sedges are well worth it

decreases in relative growth rates alone in presence environments

of a competing species.

The formation of underground rhizome protuberances in sedge largely depends on

morphological characteristics of individual species, which predetermine the growth habit of the clone. Their

the reflection is, for example, the well-known distinction between 'sedge' and 'protuberant' sedge

(with possible further clarifications; eg Chater 1980, Jermy & Tutin 1982), the first

of these, they do not create rhizomes, their length and frequency are very small

compared to the second group (which resulted in a limited time

In the experiment, tufted species, ie C. demissa , C. pallescens and C. umbrosa , did not form any

extension). Decrease in the number of processes with increasing intensity of competition is evident in C. panicea

corresponds to the assumption of de Kroon & Schieving (1990) that some clonal plants

(among them eg Carex flacca , ie also sedge with longer rhizomes)

In an unfavorable environment, resources are invested in the arms, favoring their survival over

lateral propagation by the protuberances; the variability of the C. rhizome system rhizome appears to be

plastic in the sense of de Kroon et al. (1994). In the remaining species ( C. hartmanii , C. pilulifera

and C. pulicaris ), however, the influence of the competitor on the production of the processes was not unambiguous,

no signs of morphological plasticity were observed.

Page

Although all species studied are sensitive to the action of the competitor, they do

of the experiment, it seems that in C. demissa , C. pallescens , C. panicea and C. pilulifera is

the negative impact is stronger in the sense that, in addition to reducing biomass production, it is also noticeable

the rate of formation of new arms and thus the growth of the clonal system was also affected. C. demissa se

in nature it often occurs in places that are exposed to disturbances (eg at the edges of roads,

in temporarily flooded terrain depressions; self-observation); vegetation cover is here

due to disturbance quite loose, the plant can thus easily avoid competitors

and - due to its higher growth rate (compared to other meadow sedge) - this

habitat successfully populate. As reported by Jermy & Tutin (1982), it may have a similar mechanism

use C. pilulifera , regenerating and spreading after fires in heathland.

Table 4-1. Comparison of data from field experiment prof. J. Lepse (column “eutrophication”) with results

greenhouse experiments during which the amount of nutrients was manipulated (column "substrate maintenance"

and the intensity of cross-species competition (column " H. lanatus competition "). The symbols +, - and 0 represent

response (positive, negative, or none) of individual species to increasing values of variables,

number of power answer symbol. No data from the field experiment was available for Carex demissa .

eutrophication

substrate fertility

competition H. lanatus

C. demissa

- -

C. hartmanii

C. pallescens

- -

C. panicea

- -

C. pilulifera

- - -

- -

C. pulicaris

C. umbrosa

When comparing experimentally obtained results with the data provided by prof. J. Lepse (see

Table 4-1; it was a species score on the first canonical RDA axis examining species dependence

composition of meadow communities for eutrophication, mowing and removal of dominant species -

unpublished data from a long-term field experiment described by Lepš 1999)

show that C. pilulifera was the most negatively affected by eutrophication , which, according to the above

These experimental findings are among the sedges studied competitively

weaker and (unlike C. pallescens ) fail to benefit from greater substrate yield.

In C. pulicaris , C. umbrosa (species more resistant to competition) and C. panicea (species capable of

Page

to produce more biomass while increasing nutrient availability but at the same time low resistance to nutrients

the negative effect of eutrophication was less and in the case of C. hartmanii (species competitively)

more durable and capable of utilizing a more nutritious substrate).

4.2. Field observation

It can be stated that the unifying element of all results of multivariate field analyzes

The data collected is a confirmation of the well-known fact that most sedges prefer the natural state close

habitats, ie those where the impact of human activity is not very strong (although some species, eg

Carex hirta , can be considered resistant to anthropogenic influences - Schütz 2000 and references

inside). Closer interperetation of phytosociological bonds of studied sedge species on the basis of

however, this data could be due to a small set of frames (and low frequencies)

some species) quite misleading.

Intensive human (agricultural) activity also manifests itself in the case of meadow ecosystems

completely abandoning low-productive and eutrophication of more productive habitats;

the consequence is a decrease in species richness due to the disappearance of competitively weaker species

1999). As the experiments described above have shown, these include the sedge species studied.

The positive effect of mowing on their abundance in communities can be ascribed to reality (intuitively

expected and experimentally verified by Sagittarius 2001) that it limits this type of management

excessive growth of above-ground biomass of strong competitors (grasses) and periodically returns habitat

conditions to a state where the intensity of the competition for light is low (or tolerable).

On the other hand, habitat productivity was not a decisive factor for sedges.

However, this may be due to the way productivity was determined, that is, its productivity

by estimating the amount of total biomass (or its dry matter weight) at random

plots, which is a relatively rough method. E.g. determination of nutrients (nitrogen, phosphorus)

and potassium) in soil samples would provide a much more accurate productivity rate and can

to expect that the impact of such an environmental characteristic would be significant as appropriate

the results of Better (1999).

Page

27 Mar:

5. Close Ì R

In relation to the objectives stated in the introduction of this work, the summary of the findings is as follows:

(1) Seven species of sedge, which co-exist in the locality of the enclosure, are among themselves

they differ to some extent in their response to soil conditions (substrate potency) and resistance

against strong competitors. They can be divided into four groups:

(a) Carex demissa , C. pallescens and C. panicea

- species benefiting from a more fertile substrate (producing larger above-ground biomass),

at the same time, they are not resistant to competitors (their clones grow significantly

slower and total biomass production decreasing);

(b) C. hartmanii

Species benefiting from a more fertile substrate, sensitive to the competitor 's effects (production

biomass decreases) but more resistant than others (clone growth is not slowed);

- non - responsive (tolerant to low nutrient - tolerant) substrate species,

not very resistant to competitors;

(d) C. pulicaris and C. umbrosa

- species not responsive to substrate potency, sensitive to competitors, however

relatively more durable.

These differences may contribute to some differentiation of ecological niches

species and allow their coexistence.

(2) Experimentally determined sensitivity of studied sedge species to the competitor

corresponds with the knowledge of the conditions in the field. Occurring in meadow communities

Sedge species are plants that prefer habitats in an extensive manner

management that does not lead to the rapid development of competitively strong species.

Page

6. LITERATURE

Begon, M., Harper, JL, & Townsend, CR (1997): Ecology: individuals, population,

communities. - Publisher of Palacký University, Olomouc.

Dostál, J. (1989): New Flora of Czechoslovakia, vol. 2. - Academia, Prague.

de Kroon, H. & Schieving, F. (1990): Resource partioning in relation to clonal growth

strategy. - In: van Groenendael, J. & de Kroon, H. [eds.]: Clonal growth in plants:

regulation and function. SPB Academic Publishing, The Hague, pp.113-130.

de Kroon, H., Stuefer, JF, Dong, M. & During, HJ (1994): On plastic and non-plastic

variation in clonal plant morphology and its ecological significance. - Folia Geobotanica

et Phytotaxonomica 29: 123-138.

de Kroon, H., van der Zalm, E., van Rheenen, JWA, van Dijk, A. & Kreulen, R. (1998):

The interaction between water and nitrogen translocation in a rhizomatous sedge ( Carex

flacca ). - Oecologia 116: 38–49.

Fitter, AH & amp; Hay, RKM (1989): Environmental physiology of plants. - Academic Press

London.

Goldberg, DE (1990): Components of resource competition in plant communities. - In:

Grace, JB & Tilman, D. [eds.]: Perspectives on Plant Competition. Academic Press, San

Diego, CA 27-49.

Herben, T. & Münzbergová, Z. (2002): Processing of geobotanical data in examples.

Part I. Species composition data. - Study Material, Faculty of Science, University

Charles, Prague.

Herben, T., Hara, T., Marshall, C. & Soukupova, L. (1994): Plant clonality: biology

and diversity. - Folia Geobotanica et Phytotaxonomica 29: 113-122.

Horník, J. (1998): Comparative population biology of sedges. - Ms. [Bachelor thesis, Biological

Faculty of South Bohemia; depon. in: Common Library of Biological Institutes of the CAS

and BF JU, České Budějovice].

Horník, J. (2001): Comparative population ecology of sedges. - Ms. [Diploma thesis, Biological

Faculty of South Bohemia; depon. in: Common Library of Biological Institutes of the CAS

and BF JU, České Budějovice].

Hutchings, MJ & Wijesinghe, DK (1997): Patchy habitats, division of labor and growth

Dividends in clonal plants. - Trends in Ecology and Evolution 12: 390–394.

Page

Chater, AO (1980) Carex . - In: Tutin, TG et al. [eds.]: Flora Europaea, vol

University Press, Cambridge 290-323.

Jermy, AC & Tutin, TG (1982): The Sedges of the British Isles. - The Botanical Society of the

British Isles, London.

Klimes, L. & Klimesova, J. (1994): Biomass allocation in clonal vine: effects of

intraspecific competition and nutrient availability. - Folia Geobotanica et Phytotaxonomica

29: 237-245.

Klimešová, J. & Klimeš, L. (1997): Clonal Plants: Phylogeny, Ecology and Morphology. -

Biological sheets 62: 241-263.

Krahulec, F. (1994): Clonal behavior in closely related plants. - Folia Geobotanica

et Phytotaxonomica 29: 277-289.

Kubat, K., Hrouda, L., Chrtek, J. jun., Kaplan, Z., Kirschner, J. & Stepanek, J. [eds.] (2002):

The key to the flora of the Czech Republic. - Academia, Prague.

Lepš, J. (1999): Nutrient status, disturbance and competition: an exeprimental test

Of relationships in a wet meadow. - Journal of Vegetation Science 10: 219-230.

Lepš, J. & Šmilauer, P. (2000): Multivariate analysis of ecological data. - Study

material, Faculty of Biology, University of South Bohemia, České Budějovice.

Mogie, M. & Hutchings, MJ (1990): Phylogeny, ontogeny and clonal growth in vascular

plants. - In: van Groenendael, J. & de Kroon, H. [eds.]: Clonal growth in plants: regulation

and function. SPB Academic Publishing, The Hague 3 - 22.

Prach, K. (2001): Introduction to vegetation ecology (geobotany). - Script, Faculty of Biology

University of South Bohemia, České Budějovice.

Schütz, W. (2000): Ecology of seed dormancy and germination in sedges ( Carex ). -

Perspectives in Plant Ecology, Evolution and Systematics 3: 67–89.

Schwinning, S. & Weiner, J. (1998): Mechanisms Determining the Degree of Size Asymmetry

in competition among plants. - Oecologia 113: 447–455.

StatSoft, Inc. (1999): Statistica for Windows (Computer program manual). - Tulsa, OK.

Strelec, M. (2001): Comparative ecology of meadow grasses. - Ms. [Bachelor thesis,

Faculty of Biology, University of South Bohemia; depon. in: Common Biological Library

departments of AS CR and BF JU, České Budějovice].

Špačková, I., Kotorová, I. & Lepš, J. (1998): Sensitivity of seedling recruitment to moss, litter

A dominant removal in an oligotrophic wet meadow. - Folia Geobotanica 33: 17-30.

Page 34

Ter Braak., CJF & Smiluer, P. (2002): CANOCO Reference Manual and CanoDraw for

Windows User's Guide: Canonical Community Ordination Software (version 4.5). -

Microcomputer Power, Ithaca, NY.

van Groenendael, J. & de Kroon, H. (eds.) (1990): Clonal growth in plants: regulation and

function. - SPB Academic Publishing, The Hague.

Page

48 ° 57´12.0´´ N

14 ° 35´35.1´´ E

48 ° 57´27.9´´ N

14 ° 35´24.4´´ E

49 ° 49´25.3´´ N

16 ° 09´17.8´´ E

49 ° 49´32.7´´ N

16 ° 08´57.0´´ E

49 ° 48´55.3´´ N

16 ° 09´20.9´´ E

48 ° 59´52.1´´ N

14 ° 26´27.0´´ E

49 ° 00´18.6´´ N

14 ° 25´54.9´´ E

49 ° 49´06.8´´ N

16 ° 07´13.6´´ E

49 ° 48´13.3´´ N

16 ° 09´30.2´´ E

7. C Ø ANNEXES

meadow in valley "V Kvíčalnici" 1,5 km SW from

municipality Jarošov

450

510

meadow 1 km NE from the village Ohrazení

PP Kaliště

meadow 0,5 km W from village Jarošov

meadow 0,5 km SSW from the settlement Vranice

meadow from V adjacent to the dam of the Černiš pond

S from České Budějovice

meadow 0,5 km SW from the village Bor u Skutče

meadow 1 km W from Budislav village

shot

location

sou of Coordinates

5 - 6

7 - 8

9 - 10

12 - 14

15 - 16

17 - 18

1 - 2

3 - 4

510

500

450

414

Former tankodrome S from České Budějovice

454

67. Bohemian-Moravian Highlands

38. Budějovická pánev

67. Bohemian-Moravian Highlands

38. Budějovická pánev

7.1. Localization of photographed areas.

380

39. Třeboň Basin

69a. Železnohorské foothills

390

fytochorion

the altitude of ská

height (m)

Page

frame number

15 Dec

date

30.7.01 31.7.01 31.7.01 1.8.01 11.8.01 12.8.01 26.8.01 26.8.01 29.8.01 29.8.01 24.6.02 26.6.02 26.6.02 26.6.02 24.7.02 24.7.02 25.7. 02 25.7.02

slope (°)

orientation

WITH

Coverage E 2 (%)

Coverage E 1 (%)

100 ALIGN!

Coverage E 0 (%)

15 Dec

Agrostis capillaris

Achillea millefolium

Achillea ptarmica

Ajuga reptans

Alchemilla sp.

Alnus glutinosa (juv.)

Alopecurus pratensis

Angelica sylvestris

Arrhenatherum elatius

Avenula pubescens

Betonica officinalis

Briza media

Caltha palustris

Campanula rotundifolia

Cardamine pratensis

Carex demissa

Carex echinata

Carex flacca

Carex hartmanii

Carex hirta

Carex muricata

Carex nigra

Carex ovalis

Carex pallescens

Carex panicea

Carex pilulifera

7.2a. Phytosociological images.

Page

frame number

15 Dec

Centarium erythraea

Centaurea jacea

Cerastium holosteoides

Cirsium arvense

Cirsium oleraceum

Cirsium palustre

Colchicum autumnale

Crepis paludosa

Cynosurus cristatus

Dactylis glomerata

Danthonia decumbens

Deschampsia caespitosa

Epilobium montanum

Epilobium sp.

Equisetum arvense

Equisetum palustre

Equisetum sylvaticum

Eriophorum angustifolium

Festuca pratensis

Festuca rubra

Filipendula ulmaria

Fraxinus excelsior (juv.)

Galeopsis pubescens

Galium album

Galium boreale

Galium uliginosum

Gentiana pneumonanthe

Glechoma hederacea

Glyceria fluitans

Gnaphalium sylvaticum

Holcus lanatus

Hypericum perforatum

Juncus articulatus

7.2b. Phytosociological images.

Page 38

frame number

15 Dec

Juncus conglomeratus

Juncus effusus

Juncus tenuis

Knautia arvensis

Lathyrus pratensis

Leucanthemum vulgare

Lolium perenne

Luzula campestris

Lycopus europaeus

Lychnis flos-cuculi

Lysimachia nummularia

Lysimachia vulgaris

Medicago lupulina

Mentha arvensis

Molinia caerulea

Myosotis palustris

Nardus stricta

Persicaria amphibia

Phleum pratense

Picea abies

Pimpinella major

Plantago lanceolata

Plantago major

Poa trivialis

Potentilla anserina

Potentilla erecta

Prunella vulgaris

Quercus robur (juv.)

Ranunculus acris

Ranunculus auricomus

Ranunculus flammula

Ranunculus repens

Rumex acetosa

7.2c. Phytosociological images.

Page

frame number

15 Dec

Rumex crispus

Rumex obtusifolius

Salix aurita (juv.)

Salix cinerea (juv.)

Sanguisorba officinalis

Scirpus sylvaticus

Selinum carvifolium

Stellaria graminea

Succisa pratensis

Taraxacum sp.

Thalictrum lucidum

Trifolium pratense

Trifolium repens

Urtica dioica

Veronica chamaedrys

Veronica officinalis

Vicia cracca

Viola palustris

Viola reichenbachiana

7.2d. Phytosociological images.

Page

7.3a. Schemes corms system of the studied species of .

All the plotted plants come from the locality Ohrazení; rhizome systems were

uncovered at the end of the growing season, in the second half of October 2001 ( Carex hartmanii ), respectively.

2002 (remaining species).

Explanations of tlivky: empty circles - Ramet position with vivid leaves, filled circles - position Ramet

without live leaves, double lines - rhizome processes, M - maternal arm, numeric labels -

the length of the projections in millimeters.

Carex hartmanii

Carex panicea

160

230

300

370

230

100 ALIGN!

350

500

300

270

150

400

230

170

110

210

120

270

120

50 35

WITH

Page

7.3b. Schemes corms system of the studied species of .

Note 1: The projections of the following species are displayed ten times compared to the previous ones

on a larger scale.

Note 2: The projections of C. umbrosa are rather protruding, the distances between the positions

ramet are therefore somewhat distorted (overestimated) on the scheme.

WITH

Carex pilulifera

Carex demissa

Carex pallescens

Carex umbrosa

(fragment of massive clump)

Carex pulicaris

20 May

15 Dec

20 May

15 Dec

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