POLISH JOURNAL OF ECOLOGY
(Pol. J. Ecol.)
54
1
15–27
2006
Regular research paper
Beata MESSYASZ1, Natalia KUCZYŃSKA-KIPPEN2
Department of Hydrobiology, Institute of Environmental Biology,
Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland
2
Department of Water Protection, Institute of Environmental Biology, Adam Mickiewicz University,
Drzymały 24, 60-613 Poznań, Poland, e-mail: [email protected]
1
PERIPHYTIC ALGAL COMMUNITIES:
A COMPARISON OF TYPHA ANGUSTIFOLIA L.
AND CHARA TOMENTOSA L. BEDS IN THREE SHALLOW
LAKES (WEST POLAND)
ABSTRACT: The examination of the composition and dynamics of periphytic communities
took place in three lakes of similar morphometry
– Lake Wielkowiejskie, Budzyńskie and Dębiniec,
situated in the Wielkopolska region, (Western
Poland). These lakes are typical macrophytedominated, shallow and polymictic water bodies.
Periphyton was collected from two macrophytecovered stations, including the rush vegetation
station (Typha angustifolia L.) and the stonewort
stand (Chara tomentosa L.) in each examined
lake. The material was sampled three times in
2003, including the spring (April), summer (July)
and autumn (September) seasons, from the same
parts of the lakes.
The aim of the study was to find out whether
there is an influence of particular macrophyte
species, differing spatially and morphologically
and representing two different ecological types
of aquatic vegetation, on the development of particular periphytic species. There was also an attempt at answering the question of what is more
important in the structuring of the periphytic
communities – the specific architecture of the
macrophyte substratum or the physical-chemical
features of a particular lake?
The concentrations of chlorophyll a, TN, TP
and TOC in water were higher in the Chara beds
compared with the Typha stations in most cases.
The Shannon-Weaver biodiversity index
of periphytic algae revealed very high values in
all lakes and amounted in the Typha and Chara
stands to the mean values 4.68 and 3.87 respectively in Wielkowiejskie Lake, 3.32 and 4.39 in
Budzyńskie Lake and 3.08 and 3.91 in Dębiniec
Lake. Furthermore, the standardized index of
evenness exhibited the greatest differentiation
in the Typha station with the highest (0.76) in
Wielkowiejskie and the lowest (0.52) in Dębiniec
Lake.
The analysis of the growth-forms of periphytic
communities showed slightly higher diversity of
slowly moving and stalked diatoms in most cases
at the Chara stations of all studied lakes.
The similarity between periphytic communities in the Typha and Chara stands of the examined lakes, compared using the Ward method and
Euclidean distance measure, revealed a stronger
relationship within a particular habitat but not
a lake. The clearest pattern of similarity grouping
a particular habitat was obtained for the summer
period, when macrophytes reached their optimum.
The Jaccard similarity index confirmed the
differentiation of both examined habitats, reaching the mean value of 0.34.
Based on the biomass of single periphytic
species, habitat selectivity for seven species was
found. Significantly higher biomass was obtained
in the case of the rush zone for only one algae
species – Eunotia lunaris (Ehr.) Grun. In the Chara bed six species revealed significantly higher
biomasses – Cosmarium regnelli Wille, Oocystis marssonii Lemm., Ulotrix zonata (Weber et
16
Beata Messyasz, Natalia Kuczyńska-Kippen
Moor) Kütz., Scenedesmus acuminatus (Lagerh.)
Chod., Merismopedia elegans A. Braun and Phacus orbicularis Hübner.
The habitat preference of periphytic communities to different macrophyte species may
be due to the environmental factors, including
the specific architecture of a particular macrophyte substrate such as density or texture of the
plant surface. Furthermore, seasonality, as well
as physical-chemical parameters may structure
periphytic communities within the littoral zone
of lakes.
KEY WORDS: periphyton, algae, shallow
lakes, Chara, Typha, similarity, biodiversity
1. INTRODUCTION
The presence of aquatic vegetation in
lakes relates to their role in providing zooplankton with a refuge against planktivorous
fish and invertebrate predation (S chr iver et
al. 1995, Kaires a lo et al. 1998). However,
macrophytes constitute a vast substrate for
the growth of periphytic communities especially epiphytic (attached) algae (G ons
1979), as well as plant-associated invertebrates (Paters on 1993, D ug gan 2001).
The phenomenon of the influence of expanding macrophyte density, through the enlargement of the possible substrata surface, on the
increase of the total periphyton biomass is
well known (Pie czy ńska 1976).
In turn, periphyton overgrowing the underwater parts of aquatic plants may have
a negative impact on the growth of macrophytes as it can reduce light attenuation. It
has been proved that periphyton may restrict
the degree of the light that reaches the plant
by as much as 80% (Ondok 1978) and may
also limit the diffusion of some nutrients, including carbon (S and-Jens en and B or um
1984, S chef fer 2001). Moreover, periphyton may serve as a source of food for organisms inhabiting the littoral zone. The main
components of this zone are usually epiphytic (i.e. sessile, attached) algae, which can
be accompanied by bacteria and protozoans
(Jürgens et al. 1994, Thei l-Niels en and
S øndergaard 1999). However, periphyton
may also contain great amounts of detritus
which is built up in the periphyton coverage (Van Dijk 1993). The grazing activity of animals associated with macrophytes
may cause the removal of periphytic communities from plant surfaces and thus enable better plant growth (Jones et al. 1999,
2000, James et al. 2000). On the other hand,
macrophytes competing with periphyton for
light and carbon develop an adaptive strategy
called allelopathy (G op a l and G o el 1993).
Aquatic plants are known to suppress algal
communities through the excretion of chemical substances that inhibit phytoplankton
growth (Wiu m - Anders en et al. 1982, Na kai et al. 1999) or zooplankton development
(D orgel lo and He yc o op 1985, Bu r ns and
D o dds 1999). Chara in particular has been
the object of experiments on allelopathic activity (Fors b erg et al. 1990, K leiven and
S z c z e p ańska 1988). Stonewort beds are
often surrounded by remarkably clear water compared to other macrophyte stands in
a particular lake (Bl i ndow 1987).
Macrophytes, which differ in architecture, the fineness of leaves and also in the
texture of the plant surface, may affect the
structure of organisms inhabiting the littoral
zone of lakes (R af f ael l i et al. 2000, We t z el 2001). The chemical substances released
by particular aquatic plant species may suppress particular groups or species of algae to
varying extents (Gross and Süt feld 1994,
Gross et al. 2003). E mi ns on and Moss
(1980) as well as Bl i ndow (1987) have given
evidence that different plant species may possess various periphytic algae communities.
More detailed studies have been made on the
colonization and development of periphyton on artificial substrates (Hame e d 2003,
Ác s et al. 2003) or rush vegetation (A lb ay
and A kc a a l an 2003), however, there is still
insufficient data concerning the changes in
periphytic communities on different macrophyte species. There is also some uncertainty
as to whether the kind of macrophyte habitat
will determine the algal structure or whether it is the physical-chemical parameters of
a particular lake, which will have a stronger
impact.
The initial analysis of the densities of
periphytic communities in selected habitats
showed that there was a trend for higher
abundances of particular algae groups in the
Chara beds of the examined lakes, with the
exception of Xantophyceae, which dominated in Typha stands. However, statistically
Periphytic algal communities
significant differences were only recorded
for Chlorophyta, both in the case of the total
numbers, as well as biomass (Kuczy ńskaKipp en et al. 2005). This allowed a more
specific analysis to be conducted. Thus, the
aim of the study was to find out whether
there is an influence of macrophyte species,
differing morphologically and representing
two various ecological types, on the development of particular periphytic algae species.
An attempt was also undertaken towards
answering the question of what is more important in the structuring of the periphytic
communities – the specific architecture of
a particular macrophyte substratum or the
physical-chemical features of a particular
lake? More detailed aims included the habitat preferences of selected periphytic species,
the analysis of growth-forms and similarity
as well as the species diversity of periphytic
communities in two macrophyte habitats.
2. STUDY AREA
The study was conducted on three lakes of
the Wielkopolska region, in the western part
of Poland (Lake Wielkowiejskie – 52o17,8’N
–16o40,0’E; Lake Budzyńskie – 52o14,8’N
–16o49,5’E; Lake Dębiniec – 52o28,7’N–
17o13,6’E). These lakes were selected due to
their similar morphometry – they are not
large and all belong to shallow polymictic
(Jańcza k et al. 1996) water bodies (Table 1).
They are typical macrophyte-dominated lakes
with an extensive cover of various aquatic
vegetation associations overgrowing large
areas of the lakes. The whole basin of each
lake is surrounded by a well-developed belt
of emergent macrophytes mainly with Typha
angustifolia L. and also partly accompanied
by Phragmites australis (Cav.) Steud. Beds of
Chara tomentosa L. occupied the shallower
parts compared with other submerged mac-
17
rophyte species and there were overwintering plants in each lake. In Wielkowiejskie
Lake the Chara meadows covered extensive
parts of the lake bottom, consisting of separated patches of two species: Ch. tomentosa
L. and Ch. hispida L. However, the Chara tomentosa beds were smaller compared to the
second stonewort species. In the remaining
lakes stoneworts created single and separated
one-species beds.
Based on the dominating species among
the phytoplankton in the open water (Chroomonas acuta Utermöhl, Crucigenia tetrapedia (Kirchner) W. et G.S. West, Kirchneriella contorta (Schmidle) Bohlin, Scenedesmus
ecornis (Ehrenb.) Chodat, Gymnodinium albulum Lindemann in Budzyńskie Lake; Ankistrodesmus falcatus (Corda) Ralfs, Cyclotella
distinguenda Hustedt, Dinobryon divergens
Imhoff in Dębiniec Lake; Chroomonas acuta,
Cryptomonas erosa Ehr., C. marssonii Skuja,
Dinobryon divergens, Kirchneriella contorta
in Wielkowiejskie Lake) it was found that all
the investigated lakes may be considered as
eutrophic (C ele w ic z et al. 2004).
3. MATERIAL AND METHODS
Periphyton was collected from the rush
vegetation (Typha angustifolia, Phragmites
australis) and stonewort bed (Chara tomentosa) of every examined lake. Research was
carried out in the shallow part (0.5–1.0 m
depth) of each lake. Even though it is known
that periphyton differs vertically on the macrophyte stem (A lb ay and A kc a a l an 2003)
it was not possible to collect it as separate
vertical sections from the macrophyte stems
but only as one section (ca. 0.2 m; macrophyte stems were cut out from a depth of
0.1–0.3 m) due to the very shallow depth of
the examined lakes, especially in the rush
zone where the depth never exceeded 0.5 m.
Table 1. Main morphometric characteristics of the studied lakes (Jańcza k et al. 1996).
Parameters
L. Budzyńskie
L. Dębiniec
L. Wielkowiejskie
Surface area (ha)
Max depth (m)
Mean depth (m)
Shoreline length (m)
Special status
11.0
2.7
1.4
2900
National Park
15.0
7.4
3.4
1600
Landscape Park
13.3
2.8
1.4
2300
National Park
18
Beata Messyasz, Natalia Kuczyńska-Kippen
Fig. 1. The sampling area of periphytic communities (for epiphytic algae and chemical components)
inside Typha and Chara stands.
After cutting the plant stems of the length of
0.2 m each from the area of 0.25 × 0.25 m
(Fig. 1), the periphyton was firstly rinsed out
in distilled water from each stem and then
it was removed manually using a knife and
a small brush. The periphyton was collected
from the known average biomass of plant
growing per unit lake area. The obtained results of periphyton biomass adequate to 12.5 L
of lake water were later calculated into one
litre. The method of collecting periphyton
from the underwater stems of vegetated substratum comprising the known volume unit
of lake water (0.0125 m3) was applied in order
to compare the structure of periphytic communities overgrowing two different types of
macrophyte habitats that differed in their architecture – morphologically and spatially. In
the case of Ch. tomentosa, stems of this macrophyte species were understood as the main
stem with branchlets, while the underwater
Typha stem together with its adherent leaves
were treated as a single length unit.
The periphyton material was collected
three times. Each of the three subsamples
was randomly selected from each examina-
tion station and the three results were pooled
together, so they could later be used as
a mean value during the interpretation stage
(n = 54). The biomass of periphyton was collected from the known average plant biomass
growing per lake area unit, where the biomass
of the periphyton that was obtained from the
volume of 12.5 l (0.25 × 0.25 × 0.2 m) (Fig. 1)
was later calculated for one litre.
The samples were collected three times
in 2003, including the spring (April), summer (July) and autumn (September) seasons,
from the same parts of the lakes. Samples
were first fixed in Lugol solution and then
preserved in formaldehyde.
Periphyton matter was collected also
in order to analyse the chemical content.
A chemical analysis of periphyton was conducted so as to evaluate the concentration of
TN, TP and TOC within the periphyton. The
method of periphyton sampling for chemical
analysis was the same as that used for periphyton collection for taxonomical and quantity
analysis (Fig. 1). The chemical analyses were
conducted according to Standard Methods for
Examination of Water and Wastwater (1992).
Periphytic algal communities
Chlorophyll a concentration (corrected
for pheopigments) from water of a particular
plant station was collected using a plexiglass
core sampler (Ø 50 mm) and determined fluorometrically according to the procedures described by Strickland and Parsons (1972).
Periphyton algae in samples were counted
using the Uter möh l (1958) sedimentation
method. Cells were the main counted units.
For filamentous blue-greens and greens the
length unit of 100 µm was taken for one individual. The dimensions of thirty individuals
from each species were measured according
to the shape of a standard geometrical figure.
Biovolumes were calculated using the formula for the appropriate geometric shape according to R ott (1981). The abundance and
biomass of periphyton species were related to
volume of water (Fig. 1) and expressed per
water volume unit (g·m–3).
The diatom growth-forms including
slowly moving, moving and stalked diatoms
were distinguished (Ku hn et al. 1981) and
their analysis within particular stations was
conducted.
The diversity index H’ was expressed
with the Shannon-Weaver method (Marga lef 1957):
H’ = Σ (ni/N) log2 (ni/N)
where: N – total biomass of periphytic
community,
ni – biomass of (i)-species.
The evenness index J (the range from
0 to 1) concerning all identified species and
their biomass (g·m–3) was calculated based on
the Shannon-Weaver diversity index (Pielou
1975). This was used to prevent the weighting of the H’ index by rare species:
J = H’/Hmax
where: Hmax = log2 s
s – number of species of the community.
The similarity of epiphytic communities
between Chara and Typha stands was calculated using two different methods (Jaccard
index; Ward method and Euclidean distance
measure).
The Jaccard index was used in order to
analyse the % similarity of periphytic com-
19
munities in a particular habitat. The Jaccard
index varies from 0 to 1 and is calculated
from the following equation:
Sj = a/(a + b + c)
where: a = number of species in sample
A and sample B (joint occurences),
b = number of species in sample B but
not in sample A,
c = number of species in sample A but
not in sample B.
Furthermore, the similarity between
periphytic communities on Typha and Chara
of different lakes was compared using the
Ward method and Euclidean distance measure (S oka l 1961). The graphic form of the
similarity was presented using a tree-diagram
(Krebs 1989).
The Mann-Whitney U-test or Student’s
t-test were used in order to determine the effect of site on the biomass of particular algae
species and groups (n = 54). For statistical
analysis only those species of periphytic algae
which had a high level of frequency in the examined material were selected. Species of low
frequency (below 30%) were not included in
the analysis in order to avoid the effect of accidentality of the final calculations.
4. RESULTS
The concentration of chlorophyll a was
higher in the Chara bed compared with the
Typha bed in all the lakes (Z = –5.68; P <0.00).
The highest concentrations were obtained in
Lake Dębiniec among stoneworts (Table 2).
Significantly higher concentrations of
TN (Z = –6.31; P <0.00), TP (Z = –6.31;
P <0.00) and TOC (Z = –4.59; P <0.00) were
found in periphyton area collected from the
Chara stands of all the examined lakes compared to the Typha stands. Moreover, when
analysing particular lakes – the same pattern
was also found, with statistically significant
higher values of TN, TP and TOC for Chara
bed (Table 2).
The Shannon-Weaver biodiversity
index was very high in all the lakes and
reached at the Typha and Chara stations the
mean values of 4.68 and 3.87 respectively
in Wielkowiejskie Lake, 3.32 and 4.39 in
Budzyńskie Lake and finally 3.08 and 3.91
20
Beata Messyasz, Natalia Kuczyńska-Kippen
Table 2. Average (all data, three seasons) values (±SD) of chlorophyll a (mg·m–3), TP, TN, TOC (g·m–3)
for two macrophyte beds and lakes and significance of difference Typha values versus Chara values.
Typha
Chara
P
Chlorophyll a
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
19.50 (5.29)
18.71 (5.36)
30.59 (18.53)
94.08 (55.79)
66.58 (37.23)
104.92 (38.65)
<0.00
<0.00
<0.00
TP
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
0.07 (0.03)
0.05 (0.01)
0.09 (0.02)
0.29 (0.02)
1.42 (1.18)
0.60 (0.20)
<0.00
<0.00
<0.00
TN
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
0.43 (0.33)
0.18 (0.06)
0.45 (0.22)
1.67 (0.74)
2.14 (0.82)
2.25 (0.97)
<0.00
<0.00
<0.00
TOC
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
4.58 (5.48)
6.00 (7.17)
8.10 (6.80)
28.73 (41.45)
66.04 (3.88)
19.83 (10.52)
<0.00
<0.00
<0.00
Parameters
Table 3. Average (all data, three seasons) values (±SD) of the evenness index (J)
JJ) and diversity index (H’)
of periphytic algae for two macrophyte beds and lakes and significance of difference Typha values versus
Chara values.
Typha
Chara
P
Evenness, J
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
0.76 (0.08)
0.53 (0.20)
0.52 (0.18)
0.61 (0.08)
0.68 (0.03)
0.62 (0.06)
0.00
0.05
0.23
Diversity, H’
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
4.68 (0.32)
3.32 (1.20)
3.08 (1.01)
3.87 (0.58)
4.39 (0.20)
3.91 (0.30)
0.01
0.01
0.07
Parameters
Typha
%
Chara
100
80
60
40
20
0
W
B
D
slowly moving
moving
W
B
stalked
others
D
Fig. 2. The percent participation of diatom growth-forms in the abundance of periphytic algae in the Typha and Chara stands of each lake (W – Lake Wielkowiejskie, B – Lake Budzyńskie, D – Lake Dębiniec)
Fig. 2
in particular season.
Periphytic algal communities
in Dębiniec Lake. The highest value of the
standardized index of evenness was recorded for the Typha station at Wielkowiejskie
Lake and the lowest for Typha at Dębiniec
Lake (Table 3). When analysing the results
for each lake the Shannon-Weaver biodiversity index and the evenness index show the
significant differences between the stands
21
of Typha and Chara only in the case of Lake
Wielkowiejskie where both indexes for Typha are higher then for Chara (Table 3). In
the remaining lakes the differences are not
statistically significant.
During the analysis of the growth-forms
of periphytic communities it was found that
in most cases in the Chara stand of all the
Fig. 3. The habitat preference of
particular periphytic species towards the Chara or Typha stand.
Algae mean values (SD) of biomass
expressed per g·m–3 (see Methods).
22
Beata Messyasz, Natalia Kuczyńska-Kippen
examined lakes a slightly higher, although
not significant, diversity of various forms,
including stalked and slowly moving, was
found when compared with the Typha stations (Fig. 2).
The biomass of single periphytic species
was also found to differ for both beds. Significantly higher biomasses were found in the
Typha bed for one diatom species Eunotia lunaris (Ehr.) Grun. (Z = 2.18; P = 0.03) and in
the Chara stand for chlorophytes: Cosmarium
regnelli Wille (Z = –2.34; P = 0.02), Oocystis
marssonii Lemm. (Z = –1.99; P = 0.05), Ulotrix
zonata (Weber et. Moor) Kűtz. (Z = –2.21;
P = 0.03), Scenedesmus acuminatus (Lagerh.)
Chod. (Z = –2.16; P = 0.03), for cyanoprokaryota Merismopedia elegans A. Braun
(Z = –2.18; P = 0.03) and for one euglenoid Phacus orbicularis Hűbner (Z = –2.18;
P = 0.03) (Fig. 3).
The Jaccard similarity index reached the
mean value of 0.34 for all three lakes together. The similarity of periphytic communities
between both macrophyte stands was lowest
in Dębiniec Lake (0.28) and the highest in
Budzyńskie Lake (0.42) but the difference is
not statistically significant (Table 4).
The analysis of the periphytic communities among Chara and Typha stands in all
three investigated lakes revealed that a higher
similarity, using the Ward method and Euclidean distance measure, was found in most cases
among a particular habitat (Chara or Typha)
but not among a particular lake (Fig. 4).
Table 4. Jaccard index of similarity of periphytic algae between Typha and Chara habitats in particular
lakes and all lakes together (total).
Lakes
Lake Wielkowiejskie
Lake Budzyńskie
Lake Dębiniec
Total
Mean (±SD)
Min
Max
0.31 (0.06)
0.42 (0.11)
0.28 (0.35)
0.34 (0.10)
0.16
0.31
0.19
0.16
0.45
0.53
0.42
0.53
W-sp-Typha
D-sp-Typha
W-sp-Ch
B-sp-Ch
D-au-Typha
W-su-Typha
W-au-Typha
B-sp-Typha
B-su-Typha
D-su-Typha
D-sp-Ch
W-su-Ch
B-au-Ch
B-su-Ch
B-au-Typha
D-su-Ch
D-au-Ch
W-au-Ch
0
10
20
30
40
50
60
70
Distance measure
Fig. 4. The average value of the periphytic algal community similarity in all studied lakes and seasons
(the Ward method and Euclidean distance measure).
(W – Wielkowiejskie Lake, B – Budzyńskie Lake, D – Dębiniec Lake; Ch – Chara; sp – spring, su – summer, au – autumn).
Periphytic algal communities
5. DISCUSSION
Parameters such as chlorophyll a, biomass and taxonomic composition are the
most commonly measured periphyton algae
components (Z imb a and Hops on 1997).
When describing the community structure
these indices are often taken into consideration.
It was found that in the water between
stems of the Typha bed the concentration
of chlorophyll a was lower due to the dominance of diatoms in periphyton of this zone,
where the fucoxanthine pigment prevails.
Similar results were obtained by B asu et
al. (2000) who found that chlorophyll a was
higher at dense vegetation sites than in those
which are sparsely vegetated. In all the lakes
the Chara stand was much denser compared
to the Typha bed where the stems were less
densely situated to each other in space (C ele w icz et al. 2004). However, the higher values
of chlorophyll a within Chara bed reflect the
dominance of chlorophytes, which contain
chlorophyll a as a pigment. Moreover, green
algae are known to be attached to the substratum in a less coherent way than diatoms and
some of them only accompany the overgrowing community (D elb e c que 1983, We t z el
1983), thus their presence in the interstem
water may increase and as a consequence the
concentration of chlorophyll a reaches higher values. Representatives of this group of
algae may more often be unattached due to
e.g. water waving, so the presence of chlorophytes may increase in the water between the
macrophyte stems. However, when analysing
the concentrations of nutrients and TOC in
the periphyton overgrowing particular macrophyte species it was found that in the case
of the Chara stand all the values were much
higher compared to the Typha station, which
influenced the structure of periphytic communities in both habitats.
When analysing the Shannon-Weaver
biodiversity of periphytic algae it was found
that in most cases the values reached a very
high level in all the examined lakes, with an
average value of 4.68 recorded inside the Typha stand in Wielkowiejskie Lake. Stevenson
and Yangdong (2001) proved that this index may exceed the 4.5 value when the taxa
richness is higher than 70. Also in the case
23
of all the studied lakes the number of periphytic algal species was very high with 191
taxa in total in Wielkowiejskie Lake, 183 in
Budzyńskie Lake and 172 in Dębiniec Lake
(Kuc z y ńska - Kipp en et al. 2005). The
evenness index revealed the highest species
diversity among Typha vegetation. The values in the Chara beds were similar for all the
investigated lakes. In the case of Budzyńskie
and Dębiniec lakes higher diversity was
found in the bed of Chara. However, the
sustained high diversity in the Typha bed of
Wielkowiejskie Lake throughout the whole
year may possibly be explained by the allelopathic effect revealed by the Chara meadows
in this lake. Stoneworts here cover extensive
areas of the lake bottom, thus creating compact and continuous plant beds. Within this
vegetation cover individual Chara tomentosa
beds are separated and surrounded by Ch.
hispida. It can then be presumed that Chara
hispida, which was not found in Budzyńskie
or Dębiniec lakes, releases substances which
can selectively influence algal development.
Several authors, e.g. Bl i ndow (1987) or
Na kai et al. (1999) have observed a similar phenomenon where the Chara meadows
are inhibited by the algal communities. The
analysis of both biodiversity indexes revealed
significant differences between the Typha
and Chara stands in Budzyńskie and Wielkowiejskie lakes, which confirms the great
differentiation between those two habitats.
The analysis of the biomass for seven
periphytic algal species indicated their habitat preferences. The diatom Eunotia lunaris
is a typical periphytic species that colonizes
large and smooth surfaces associated with
Typha stands. The cells of this species attach
singularly or in bunches to the macrophyte
stem. So, the long-stalked species compete
successfully with those that create gelatinous
tubes or matrix masses along with the development of periphyton (R o os 1983, We t z el
1983). In the Typha zone diatoms are more
abundant due to the fact of extensive overshading, which they prefer (Ondok 1978).
Light is an essential factor for photosynthesis, so particular groups of algal communities may differ in their light requirements.
Furthermore, it was also proved in situ, that
Chara may greatly restrict periphytic algae
development, especially epiphytic diatoms
24
Beata Messyasz, Natalia Kuczyńska-Kippen
(Wium-Anders en et al. 1982, Mű l ler
1999), which is why no diatom species were
found to choose selectively the Chara habitat
in the examined lakes.
However, in the Chara bed six species of
significantly higher biomass in this zone were
found, representing mostly chlorophytes
(Cosmarium regnellii, Oocystis marssonii,
Ulotrix zonata, Scenedesmus acuminatus),
and also cyanoprokaryota (Merismopedia elegans) and euglenophyta (Phacus orbicularis).
They are all common species, tending to live
in small water bodies as well as in large lakes,
and preferring eutrophic waters. These species are free-moving, non-sessile forms and
they can often be found in the pelagic zone of
lakes, especially Cosmarium regnellii, Oocystis marssonii, Scenedesmus acuminatus. They
avoided the Typha macrophyte sites, where
the diatoms dominated (Kuczy ńska-Kipp en et al. 2005), creating a dense and compact layer of periphytic communities. Ulotrix
zonata, like other filamentous species, can
often be found among macrophytes of complicated spatial and morphological structure
due to the possibilites of attachment to the
furcated shoots with rough surface (St armach 1972, B er r y and L embi 2000).
Particular plant species differ markedly in
shape and these different forms are therefore
important with respect to their own typical
associated organisms. Various factors are responsible for spatial habitat segregation in
shallow lakes, including tolerance to changes
of chemical environment and physical parameters which relate to the architecture of
macrophyte habitat (R os enzweig 1991).
Chemical factors are likely to vary with different types of macrophytes (C onde-Porc una 2000), which has also been proved
in the case of the investigated water bodies
where significant differences in the chemical
composition were recorded when comparing Chara and Typha stands. The content of
TN, TP and TOC in periphyton, which differed significantly between both macrophyte
stands, may then affect the habitat preferences of particular algae species as they require
specific nutrient conditions. For instance
Ulotrix zonata prefers habitats rich in organic matter (St ar mach 1972), as found in
the case of Chara beds. Habitat selectivity of
organisms is also influenced by space hetero-
geneity, which is connected with the number
of available sites in relation to a particular
surface area (Krebs 2001). Morphological
structure, especially the plant surface structure and density of macrophytes may also affect the creation of specific conditions within
particular vegetation stands (Pie c z y ńska
and Sp o d nie wska 1963, Cy r and D ow n i ng 1988, v an den B erg et al. 1997).
Both vegetated habitats in the studied lakes
differed in their biometric measurements,
which modified the habitat conditions for
inhabiting organisms and thus affected their
habitat preferences. The rush vegetation is
characterised by sparse stem spatial structure and vertically orientated shoots (C ele w ic z et al. 2004, Kuczyńska-Kippen and
Nagengast in press), while stoneworts create
a much denser macrophyte habitat due to
their much longer stems. With the increase
of the mosaic structure of the vegetated
substratum the creation of more ecological
niches is also observed, thus favouring the
development and habitat selectivity of inhabiting organisms (Kuczyńska-Kippen and
Nagengast, in press).
The Jaccard index of similarity of the
periphyton algae community indicated
a great differentiation between the examined
macrophyte stands due to very low values of
similarity. Additional analysis of similarity
carried out with the use of the Ward method
and Euclidean distance measure between
both examined plant stations in all three
lakes revealed a higher relation among the
examined habitats. The morphological and
spatial structure of a particular macrophyte
bed seems to be the factor responsible for
the similarity of periphytic communities in
the examined lakes. Many authors who have
dealt with invertebrates inhabiting the surface of macrophytes have also asserted this
fact. They have emphasized the influence of
the plant biomass and its morphological differentiation on the structure of animal communities (Cy r and D ow ni ng 1988, Paters on 1993).
The differentiation of both macrophyte
habitats was also confirmed in the case of the
analysis of the growth-forms of periphytic
communities. It was recorded that in most
cases the stonewort stands of all the studied
lakes had a slightly higher diversity of vari-
Periphytic algal communities
ous forms, in particular the stalked and slowly moving.
The changes in the periphytic structure
on different macrophyte species may be due
to the variation of environmental factors,
including the specific architecture of macrophyte stands, seasonality, physical-chemical
parameters as well as biotic relations, e.g. allelopathy or physical parameters of a particular macrophyte substrate (D ug gan 2001).
The obtained results revealed that not
a particular lake but the specific architecture of a particular macrophyte substratum
determined the structure of the periphytic
algal communities. In the case of the Chara stand that was characterised by higher
concentrations of nutrients and TOC in the
periphyton overgrowing its surface, higher
concentration of chlorophyll a in the interstem spaces as well as by more spatially and
morphologically complicated structure, the
biocoenotical parameters of epiphytic algae
reached higher values compared to the Typha stand. In most of cases the periphytic
algal communities of the Chara bed were
more diverse. Also the stonewort stand possessed a much larger group of habitat-related algae species. Moreover, the similarity
indices also confirmed the distinctiveness
between both the examined macrophyte
habitats.
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