Oceanological and Hydrobiological Studies
International Journal of Oceanography and Hydrobiology
Volume 42, Issue 3
ISSN 1730-413X
eISSN 1897- 3191
(268–276)
2013
Received:
Accepted:
DOI: 10.2478/s13545-013-0083-x
Original research paper
Composition of protozoan communities
at two sites in the coastal zone of the
southern Baltic Sea
Krzysztof Rychert *, Katarzyna Spich, Kinga
Laskus, Michalina Pączkowska, Magdalena
Wielgat-Rychert, Gracjan Sojda
Department of Aquatic Ecology,
Pomeranian University in Słupsk,
ul. Arciszewskiego 22b, 76-200 Słupsk, Poland
Key words: ciliates, flagellates, HNF, dinoflagellates,
Baltic Sea, coastal zone
Abstract
Protozoan communities were studied in the coastal zone of
the southern Baltic Sea. Stable environmental conditions and
typical, bimodal seasonal changes in the protozoan biomass were
observed at the sampling site in Sopot (2003-2004). At the
sampling site in Ustka (2007-2008), strong benthic resuspension
and irregular impacts of fresh water resulted in atypical seasonal
changes in the protozoan biomass with a summer peak only. The
mean annual biomass had similar values at both sites: 43.2 µg C
dm-3 in Sopot and 38.6 µg C dm-3 in Ustka. The protozoan
community in Sopot was dominated by ciliates (48% of the
biomass), whereas in Ustka − by heterotrophic nanoflagellates
(53%).
*
Corresponding author e-mail: [email protected]
September 27, 2012
January 02, 2013
INTRODUCTION
In aquatic environments, protozoa, together with
bacteria, constitute the microbial food web (Azam et
al. 1983, Kirchman & Williams 2000). Smaller
protozoa, nanoflagellates feed on bacteria and
cyanobacteria, whereas larger ciliates − on other
protozoa, algae, and bacteria. In marine and brackish
environments, also heterotrophic dinoflagellates are
an important component of microbial food webs,
which ingest both algae and protozoa (Smetacek
1981, Hansen 1991, Sherr & Sherr 1994, Bralewska
&
Witek
1995).
Consequently,
microbial
communities comprise food webs with complicated
trophic
relations.
Generally,
ciliates
and
heterotrophic dinoflagellates are regarded as top
predators within the microbial food web and
nanoflagellates as the intermediate trophic level
between bacteria and cyanobacteria and the
aforementioned protozoa.
The Baltic Sea is one of the most thoroughly
studied water bodies. This includes studies of
unicellular organisms; however, studies of protozoa
are far less frequent compared to studies of bacteria
and algae. Generally, their taxonomic diversity is well
recognized (e.g. Vørs 1992, Thomsen 1992, Ikävalko
& Thomsen 1997, lkävalko 1998, Mironova et al.
2009, Grinienė et al. 2011, Rychert 2011), but there is
still a lack of studies focusing on the quantitative
importance of protozoa. In a review on planktonic
communities in the Baltic Sea, Arndt (1991)
expressed the need for studying all three of the most
important protozoan groups simultaneously –
heterotrophic
ciliates,
dinoflagellates,
and
nanoflagellates. Such comparative studies should be
focused on biomass, as abundance is misleading
because the volume of protozoa ranges over a few
orders of magnitude (e.g. Müller 1989). This kind of
studies must also cover all the seasons of the year, as
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Protozoa in the coastal zone of the Baltic Sea| 269
this permits the calculation of mean annual biomass.
Twenty years after the review by Arndt (1991), there
are still strikingly few such studies published with
reference to both marine (Garstecki et al. 2000, Yang
et al. 2008) and freshwater environments (Mathes &
Arndt 1995, Kopylov et al. 2002, Kiss et al. 2009).
Such comprehensive data are particularly important
for the calculation of the annual energy flow budgets
or construction of mechanistic ecosystem models.
The aim of this study was to contribute to the
knowledge about standing stocks and composition of
protozoan communities in the coastal zone of the
southern Baltic Sea. Two sampling sites differing in
their characteristics were compared. As it was
emphasized above, the studies were conducted
throughout the year to permit the description of the
biomass range and the calculation of mean annual
values. The three most important protozoan groups
in sea waters were studied: heterotrophic flagellates,
dinoflagellates, and ciliates (Arndt 1991, Sherr &
Sherr 2002). The fourth group of protozoa –
amoebae, were excluded from the analysis since their
role in the marine pelagic zone is considered to be
negligible, with the exception of estuaries where they
appear to be more relevant (Caron & Swanberg 1990,
Sherr & Sherr 2002, Rogerson et al. 2003). It should
be emphasized that large groups of amoebae, e.g.
foraminifera, are absent in the less saline parts of the
Baltic Sea (e.g. Schweizer et al. 2011).
MATERIALS AND METHODS
The sampling sites were located in the southern
Baltic Sea (Fig. 1). Water samples were collected at
the Sopot site from the terminal part of a 450 m long
wooden pier located in the Gulf of Gdańsk. Samples
were collected once a month between April 2003 and
March 2004. In Ustka (Fig. 1), sampling was done
monthly between April 2007 and April 2008 from the
terminal part of a 70 m long concrete pier; in spring
samples were collected fortnightly. The two sites
were of similar depth (6.0–6.5 m), and the sampling
events were accompanied by measurements of
temperature, salinity, oxygen concentration, Secchi
depth, and BOD5. Chlorophyll was measured only in
Sopot.
The ciliates were analyzed under an inverted
microscope with the Utermöhl (1958) method in
samples fixed with Lugol’s solution (final
concentration of 0.5%). The ciliates were identified
based on the identification keys of Marshall (1969)
and Witek (1994), a web-based key (Strüder-Kypke &
Fig. 1. Sampling sites (Ustka, Sopot) located in the
coastal zone of the Baltic Sea. Samples were collected
at the end of piers. Depths at both sites were similar
(Sopot - 6.0 m; Ustka - 6.5 m)
Montagnes 2002), and others. The cells were counted
and measured to calculate their volume. The volume
was converted into carbon units with the coefficient
of 0.11 pg C µm-3 (Edler 1979). The biomass of
loricated ciliates (carbon content, CC, pg C) was
calculated according to Verity & Langdon (1984):
𝐶𝐶 = 0.053𝑉 + 444.5
where V is the volume of lorica (µm3). The
cryptophyte-bearing ciliate Mesodinium rubrum was
considered to be generally autotrophic (Crawford
1989, Johnson & Stoecker 2005), and was
disregarded. See Rychert & Pączkowska (2012) for
the abundance of Mesodinium rubrum at the sampling
site in Ustka.
Heterotrophic dinoflagellates were analyzed in the
same samples as ciliates. The distinguishable
specimens were identified to the genus or species,
and were classified as heterotrophs according to
Thomsen (1992), Bralewska & Witek (1995), and our
own previous observations. Among unidentified
forms, heterotrophs were identified after
simultaneous observation under an epifluorescence
microscope in samples that had been pre-fixed with
Lugol’s solution, discolored with thiosulfate, and
then fixed with formalin (Sherr & Sherr 1993). Ebria
tripartita was grouped with dinoflagellates as was
done previously by Kivi (1986), WrzesińskaKwiecień & Mackiewicz (1995) and Bralewska &
Witek (1995). The biomass was calculated from the
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270 |Krzysztof Rychert, Katarzyna Spich, Kinga Laskus, Michalina Pączkowska, Magdalena Wielgat-Rychert, Gracjan Sojda
volume with the coefficients of 0.11 pg C µm-3 for
naked dinoflagellates and 0.13 pg C µm-3 for thecate
dinoflagellates (Edler 1979).
Heterotrophic flagellates larger than 8 µm were
analyzed under an inverted microscope in samples
fixed with Lugol’s solution, the same way as ciliates
and dinoflagellates. They were identified according to
Thomsen (1992), and their biomass was calculated
with the coefficient of 0.11 pg C µm-3 (Edler 1979).
Smaller flagellates (2–8 µm, and some obvious
picoflagellates of 1–2 µm diameter) were analyzed
under an epifluorescence microscope in samples
fixed with glutaraldehyde, which is considered
superior to formalin for subsequent staining with
fluorescent dyes (Bloem et al. 1986). Small flagellates
were stained with primulin (samples taken in Sopot;
Caron 1983) or, after parallel staining (separate
filters) with acridine orange and proflavin (samples
taken in Ustka, Sherr et al. 1993). Staining with
pirmulin or proflavin made it possible to observe the
red autofluorescence of chlorophyll and to
distinguish heterotrophic flagellates. These small
flagellates (<8 µm) were not identified. Their
biomass was calculated from the volume with the
coefficient of 0.22 pg C µm-3 (Børsheim and Bratbak
1987).
Some marine protozoa are mixotrophic
(Bralewska & Witek 1995, Caron 2000, Esteban et al.
2010). Only heterotrophic cells were counted among
the dinoflagellates and nanoflagellates. See Rychert
(2006) for data on the co-occurrence of pigmented
and non-pigmented flagellates at the sampling site in
Sopot. In the case of ciliates, some chloroplastsequestering forms were probably counted. They
could not be identified in samples fixed with Lugol’s
solution; it should be emphasized, however, that they
were predominantly heterotrophic.
Since the study focused on the protozoan
biomass, Lugol’s solution was used to fix the ciliates,
larger flagellates and dinoflagellates. Lugol’s solution
is known to be a good preservative for quantitative
studies (Leakey et al. 1994) but this fixative hinders
taxonomic identification. Consequently, even
thought the study produced good biomass
determinations, some of the cells could not be
identified to the species.
The authors computed the mean annual biomass
for each group of protozoa using the trapezoidal
method, which takes into account the different
periods that elapse between samplings; thus, the
mean annual values are weighted means.
RESULTS AND DISCUSSION
Environmental conditions
The range of water temperatures observed at both
sites (1–21°C) was typical of temperate waters. The
maximum temperature at the sampling site in Sopot
was recorded in August, whereas in Ustka in July.
The mean annual salinity in Sopot (Table 1) was
slightly lower compared to the southern Baltic (7.5
PSU, Majewski 1987), which is typical of enclosed
areas such as Sopot located in the Gulf of Gdańsk.
The salinity in Ustka fluctuated irregularly (Table 1)
as a result of changeable, wind-dependent impact of
fresh water from the mouth of the Słupia River,
which is located just 1.3 km to the east (discharge of
17–18 m3 s-1, HELCOM 1998). Secchi disk visibility
differed at the two studied sites. In Sopot, it ranged
from 4.5–6.5 m in winter to 2.0–2.5 m during the
spring phytoplankton bloom and in summer.
Changes in the transparency were atypical in Ustka
with higher values recorded during the growing
season (1.6–5.8 m) and the lowest values during
winter (0.5–1.3 m), which resulted from turbidity
caused by wind-induced benthic resuspension (the
strongest winds occurred in winter). The water
column at both sites was mixed and well-oxygenated.
No ice cover was observed. To sum up, the
environmental conditions at the Sopot site were
stable, whereas at the Ustka sampling site − variable
with strong benthic resuspension and irregular
impact of fresh waters.
The trophic states of both sampling sites were
estimated based on BOD5 and bacterial abundance.
The mean annual values of these parameters were
Table 1
Characteristics of sampling sites in Sopot (the Gulf of
Gdańsk, studied between April 2003 and March 2004)
and Ustka (Central Pomerania, studied between March
2007 and April 2008) both located in the coastal zone
of the southern Baltic Sea
Sopot
Salinity (mean value)
Secchi Disk (mean annual value)
BOD5 (mean annual value)
Bacterial abundance (mean annual value)
Chlorophyll (mean annual value)
Ustka
Salinity (mean value)
Secchi Disk (mean annual value)
BOD5 (mean annual value)
Bacterial abundance (mean annual value)
6.4 – 7.4 (7.1) PSU
2.0 – 6.5 (4.3) m
0.94 – 4.35 (1.59) mg O2 dm-3
4.36 – 17.1 (8.16) mln cm-3
0.99 – 15.9 (3.56) mg m-3
6.2 – 8.5 (7.2) PSU
0.5 – 5.8 (2.5) m
0.78 – 3.39 (1.44) mg O2 dm-3
1.58 – 8.16 (4.55) mln cm-3
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Protozoa in the coastal zone of the Baltic Sea| 271
higher at the Sopot sampling site (Table 1). It is also
noteworthy that the mean BOD5 value in Ustka
differ annually, as the mean value of measurements
taken by Rychert & Wielgat-Rychert (2008) between
May 2006 and April 2007 was 2.18 mg O2 dm-3 and
was significantly higher than the value of 1.44 mg O2
dm-3 obtained between April 2007 and April 2008.
The same study (Rychert & Wielgat-Rychert 2008)
also reported unstable conditions at the sampling site
in Ustka.
Heterotrophic ciliates
The ciliates observed at both studied sites were
reported previously in the Baltic plankton by
Leppänen & Bruun (1988), Wrzesińska-Kwiecień &
Mackiewicz (1995), Witek (1998), and Setälä & Kivi
(2003). The ciliates observed most frequently at both
sampling sites were: Strombidium spp., Strobilidium
spp., Tintinnopsis spp., Urotricha spp., Holophrya spp.,
and Balanion comatum. Common ciliates included also
Lohmanniella oviformis and the heterotrophic
Mesodinium sp. These were accompanied by predatory
ciliates from the genera Askenasia, Monodinium,
Didinium, and Lacrymaria, which were of particular
importance in spring and autumn. Some large ciliates
like Strobilidium spiralis, Strobilidium sphaericum,
Strombidium mirabile, Strombidium stylifer, and Prorodon
spp. were observed except in summer. Additionally,
two distinct ciliate communities were observed at the
sampling site in Sopot. The first one was observed in
late spring and was dominated by Euplotes sp., which
accounted for 80% of the ciliate biomass; it was
observed after the spring peak of ciliates comprised
of strombidiids, strobilidiids, and predatory ciliates.
During the same season in Ustka, Euplotes sp. was
also observed, but in small numbers. The second
distinct community was found in Sopot during the
second peak that occurred in September and
consisted mainly of Helicostomella subulata. This
community was not observed in Ustka. The mass
occurrences of Euplotes sp. in late spring and
Helicostomella subulata in late summer were observed
previously in the offshore waters of the Gulf of
Gdańsk (Witek 1998, Wasik & Mikołajczyk 1996),
the Gulf of Riga (Boikova 1984), and the Gulf of
Finland (Kivi 1986). Van Beusekom et al. (2009)
reported a peak of H. subulata in late summer in the
Bornholm Basin (Baltic Sea) as well. Witek (1998)
reported high Vorticella spp. abundance in the Gulf of
Gdańsk in summer. Vorticella spp. was also observed
at both studied sites but in small numbers. Other
ciliates, like scuticociliates and benthic migrants
(karyorelictids and hypotrichs), were of little
importance. The highest ciliate biomass of 91.8 µg C
dm-3 (Fig. 2) was observed in Sopot in May 2003. In
Ustka, the seasonal changes in the ciliate biomass
were irregular, and the highest biomass was observed
in July (Fig. 2, 67.8 µg C dm-3). The seasonal changes
in the ciliate biomass at the Sopot sampling site were
typical, and similar bimodal patterns were described
previously in many regions of the Baltic Sea
(Smetacek 1981, Arndt 1991, Witek 1998, van
Beusekom et al. 2009). Seasonal changes in Ustka
resembled patterns observed in estuaries such as
Southampton Water (Leakey et al. 1992). Typical
characteristic of estuaries are higher contributions of
tintinnids (Leakey et al. 1993, Suzuki & Taniguchi
1998, Urrutxurtu et al. 2003, Rychert 2011). Indeed,
tintinnids in Ustka accounted for 23% of the mean
annual ciliate biomass, whereas in Sopot, the
Solid lines refer to estimates in the first year of the study, and dashed
lines refer to values in the second year of the study.
Fig. 2. Seasonal changes in the biomass of
heterotrophic
ciliates
(HC),
heterotrophic
dinoflagellates
(HDF),
and
heterotrophic
nanoflagellates (HNF) at sampling sites in Sopot (upper
graph, 2003–2004) and Ustka (lower graph, 2007–
2008)
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272 |Krzysztof Rychert, Katarzyna Spich, Kinga Laskus, Michalina Pączkowska, Magdalena Wielgat-Rychert, Gracjan Sojda
corresponding value was lower (15%). The mean
annual ciliate biomass in Sopot and Ustka was 20.5
µg C dm-3 and 13.3 µg C dm-3, respectively, and both
values were higher compared to the Baltic Proper
(4.8 µg C dm-3, Samuelsson et al. 2006), the inner
Gulf of Gdańsk (Baltic Sea, 5.4 µg C dm-3, Witek
1998), the Kiel Bight (Baltic Sea, 7.1 µg C dm-3,
Smetacek 1981), similar to the previous estimates in
the coastal zone of the Gulf of Gdańsk (~20 µg C
dm-3, Witek et al. 1997), and much lower than in the
Darss-Zingst estuary (southern Baltic), where it was
100 µg C dm-3 (Arndt et al. 1990). Thus, the biomass
observed at the study sites corresponded to other
values reported from the Baltic Sea.
Heterotrophic dinoflagellates
The most frequently observed (all year round)
heterotrophic dinoflagellates at the Sopot site were
small gymnodinoids and gyrodinoids (<15 µm).
Thecate dinoflagellates from the Diplopsalis group,
Protoperidinium bipes, Protoperidinium brevipes, Cladopyxis
sp., Katodinium glaucum, Amphidinium sphenoides,
Gyrodinium spirale and others, were less frequent. Two
peaks were observed: (i) the peak in spring (May-June
2003) mainly comprised thecate dinoflagellates from
the Diplopsalis group, and (ii) the autumn peak
(October 2003) comprised small gymnodinoids and
gyrodinoids (Fig. 2). At the site in Ustka, Ebria
tripartita was most common. Small gymnodinoids and
gyrodinoids, dinoflagellates from the Diplopsalis
group, and Protoperidinium spp. were less common. A
clear peak of Protoperidinium spp. and Ebria tripartita
was observed in June (Fig. 2). It is noteworthy that
the increased Ebria tripartita abundance is typical of
estuaries (Pollehne et al. 1995, Wrzesińska-Kwiecień
& Mackiewicz 1995). The organisms recorded at the
two sites were previously described from the Baltic
Sea (Hansen 1991, Bralewska & Witek 1995, Witek &
Pliński 2005, HELCOM 2006).
In terms of the occurrence time and composition,
the heterotrophic dinoflagellate peaks observed at the
two study sites corresponded well with observations
from other regions of the Baltic Sea (Smetacek 1981,
Hansen 1991, Bralewska & Witek 1995). The mean
annual biomass of heterotrophic dinoflagellates
observed in Sopot (13.0 µg C dm-3) was similar to
those reported by Witek from the open waters of the
inner part of the Gulf of Gdańsk (11.2 µg C dm-3,
Witek 1995). In Ustka, heterotrophic dinoflagellates
were less abundant (4.8 µg C dm-3), and their biomass
was similar to values observed in the coastal zone of
the Gulf of Gdańsk in Orłowo (6.6 µg C dm-3,
Kwiatkowska 1999), the Gdańsk Deep (4.1 µg C
dm-3, Witek 1995), the open waters of the southern
Baltic (5.6 µg C dm-3, Witek 1995), and the Kiel Bight
(5.1 µg C dm-3, Smetacek 1981). All the cited biomass
values are mean annual values calculated for surface
waters.
Heterotrophic nanoflagellates
The majority of heterotrophic nanoflagellates of
the 2–8 µm fraction could not be analyzed
taxonomically. Larger identifiable flagellates (8–20
µm) accounted for on average 10% of the biomass of
all heterotrophic flagellates (2–20 µm) in Ustka and
7% in Sopot. Kathablepharis remigera Clay and Kugrens
1999, Telonema spp., and Leucocryptos marina (Braarud)
Butcher 1967 were most frequent and all of them
were reported previously in the Baltic Sea
(Mackiewicz 1991, lkävalko 1998, Vørs 1992,
Wrzesińska-Kwiecień
&
Mackiewicz
1995).
Generally, no differences were found between the
community composition at the two sites studied. The
abundance of heterotrophic flagellates in Sopot was
similar to that reported by Piwosz & Pernthaler
(2010) who studied nanoflagellates at the same site in
Sopot in 2007. The mean annual biomass of
nanoflagellates in Sopot (9.7 µg C dm-3) was much
higher than the corresponding values reported by
Samuelsson et al. (2006) for the Bothnian Bay and
the Bothnian Sea (1.7–3.5 µg C dm-3) and slightly
higher than in the Baltic Proper (Baltic Sea, 5.1–7.9
µg C dm-3, Samuelsson et al. 2006). Seasonal changes
with one peak in spring (Fig. 2) resembled those
observed in the open Gulf of Gdańsk (Baltic Sea,
Mackiewicz 1991), the Gulf of Riga (Baltic Sea,
Boikova 1984), and in the Bay of Biscay (Atlantic
Ocean, temperate zone, Granda & Álvarez 2008).
The biomass of heterotrophic nanoflagellates was
higher and their seasonal changes were different at
the sampling site in Ustka. The mean annual biomass
of heterotrophic nanoflagellates (20.5 µg C dm-3) was
twice as high as the corresponding value in Sopot.
Although this is a high mean value, even higher mean
biomass was reported previously from the pelagic
zone of the Baltic Sea, e.g. in shallow inlets of the
southwestern Baltic where the mean, combined
biomass of dino- and nanoflagellates was 180 µg C
dm-3 (Garstecki et al. 2000). Such high biomass of
flagellates was attributed to the resuspension of
benthic organisms. Garstecki et al. (2000) reported
that flagellates are more easily resuspended in the
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Protozoa in the coastal zone of the Baltic Sea| 273
water column than larger protozoa. Most probably a
similar mechanism resulted in the high biomass of
nanoflagellates at the Ustka sampling site because of
strong sediment resuspension, as it was described
previously. Seasonal changes in the biomass with one
summer peak are similar to the seasonal trends
observed in the Rhine River (Weitere & Arndt 2002),
the Danube River (Kiss et al. 2009), the estuarine
Southampton Water (Brandt & Sleigh 2000), or the
estuary of Biscay Bay (Urrutxurtu et al. 2003).
Therefore, the seasonal occurrence of heterotrophic
flagellates in Ustka could also be driven by fresh
water flowing from the Słupia River.
Composition of protozoan communities
At the sampling site in Sopot, seasonal changes in
the protozoan biomass (heterotrophic ciliates,
dinoflagellates, and nanoflagellates together) were
bimodal with two distinct peaks: the higher one of
138.8 µg C dm-3 in spring, represented mainly by
ciliates, and the smaller one of 61.3 µg C dm-3 in
autumn dominated by heterotrophic dinoflagellates.
The mean annual biomass of protozoa in Sopot
(calculated as a weighted mean) was 43.2 µg C dm-3.
The unstable conditions in Ustka resulted in atypical
seasonal changes in the protozoan biomass. The
maximum biomass of all three groups of protozoa,
i.e. 151.6 µg C dm-3, was recorded in spring and early
summer, in July, whereas no autumn peak was
observed. The mean annual protozoan biomass at
the Ustka sampling site was 38.6 µg C dm-3.
As mentioned in the Introduction, there are only
few studies that provide data on the biomass of all
three main groups of protozoa for the entire year.
Generally, the biomass is equally distributed between
heterotrophic ciliates and dinoflagellates in open and
coastal oceanic waters (Lessard & Swift 1985, Sherr
& Sherr 2002, Montagnes et al. 2010), in high latitude
ecosystems (Levinsen et al. 2000), and in the Baltic
Sea (Hansen 1991, Smetacek 1981). This is also the
case for heterotrophic ciliates and nanoflagellates in
the Pacific Ocean (Suzuki et al. 1998). Arndt (1991)
reported that heterotrophic ciliates, dinoflagellates,
and nanoflagellates each contribute about one third
of the mean annual protozooplankton biomass in the
Baltic Sea. The available data on the mean annual
biomass in the Baltic Sea are presented in Table 2,
which indicates that the biomass of the three main
components are roughly comparable.
The composition of the protozoan community at
the Sopot sampling site revealed the elevated
relevance of ciliates (48% of the protozoan biomass)
and the lower importance of heterotrophic
nanoflagellates (22% of the protozoan biomass). The
most likely explanation for this is the higher trophic
state of this site. The high abundances of bacteria
(the annual mean of 8.16 million cells cm-3, Table 1)
indicated that ciliates could compete with
nanoflagellates for bacteria as prey. It is well
documented that the importance of ciliates as
bacterivores in the freshwater environments
increases along the trophic gradient (e.g. Beaver &
Crisman 1989, Šimek et al. 2000). The contribution
of heterotrophic dinoflagellates to the protozoan
community appeared to be typical (30%, Table 2).
Table 2
Mean annual biomass of the protozoan community components in different Baltic Sea regions (data arranged
according to increasing heterotrophic ciliate biomass)
Area
Heterotrophic ciliates
(µg C dm-3)
Bothnian Bay (offshore)
1.5–1.6
Bothnian Sea (offshore)
1.8–1.9
Southern Baltic (offshore)
3.0
Baltic Proper (offshore)
4.2–4.8
Gdańsk Deep (offshore)
4.7
Gulf of Finland (coastal zone)
5.1
Inner Gulf of Gdańsk (offshore)
5.4
Kiel Bight
7.1
Southern Baltic (coastal zone, Ustka)
13.3
Inner Gulf of Gdańsk (coastal zone, Sopot)
20.5
Southern Baltic (coastal zone, Rassower Strom)
40
Southern Baltic (coastal zone, Kirrbucht)
43
1
Combined biomass of heterotrophic nanoflagellates and dinoflagellates
Heterotrophic
dinoflagellates
(µg C dm-3)
no data
no data
5.6
no data
4.1
1.2
11.2
5.1
4.8
13.0
Heterotrophic
nanoflagellates
(µg C dm-3)
1.7–2.6
3.1–3.5
fragmentary data
5.1–7.9
fragmentary data
no data
fragmentary data
fragmentary data
20.5
9.7
131
1801
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Witek 1995
Samuelsson et al. 2006
Witek 1995
Kivi 1986
Witek 1995
Smetacek 1981
This study
This study
Garstecki et al. 2000
Garstecki et al. 2000
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274 |Krzysztof Rychert, Katarzyna Spich, Kinga Laskus, Michalina Pączkowska, Magdalena Wielgat-Rychert, Gracjan Sojda
In Ustka, heterotrophic flagellates accounted for
most of the biomass (53%, Table 2), whereas ciliates
were less important (35%) and heterotrophic
dinoflagellates contributed only 12% of the biomass.
The low contribution of heterotrophic dinoflagellates
to the protozoan biomass could be explained by the
impact of fresh water. As mentioned in the
Introduction, dinoflagellates are an important
component of the microbial food web in the marine
ecosystems, but not in the freshwater ones. The
reduced contribution of dinoflagellates to protozoan
communities was also observed in the coastal zone of
the Gulf of Finland (Kivi 1986). The elevated
relevance of nanoflagellates, similar to that observed
at a sampling site in Kirrbucht (Garstecki et al. 2000),
most likely resulted from the benthic resuspension.
The current study provides a comprehensive
description of the protozoan community inhabiting
the coastal waters of the southern Baltic Sea. Because
the sampling site in Ustka proved to be of estuarine
character, further research is necessary to estimate
the biomass of amoebae occurring there, which are
generally
unimportant
in
pelagic
marine
environments but can be abundant in estuaries
(reviewed in the Introduction). Lesen et al. (2010)
reported that in the Hudson River Estuary, amoebae
accounted for one fourth of the total protozoan
biomass.
ACKNOWLEDGEMENT
This study was partially supported by the Polish
Ministry of Science and Higher Education (Grant No
2 P04F 074 27).
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