Diatom research has been conducted in the Gulf of Gdańsk for nearly 100 years. The first researcher to document the occurrence of diatoms in the bay area was Schulz (1926). The modern and fossil diatom flora of the Gulf of Gdańsk was described by, among others, Rumek (1948), Pliński (1987, 1988, 1990, 1995), Witkowski (1994), Witkowski & Pempkowiak (1995), Pliński & Kwiatkowski (1996), Stachura & Witkowski (1997), Witak (2000, 2002, 2010, 2013), Bogaczewicz-Adamczak et al. (2001), Witak et al. (2006, 2011), Witak & Dunder (2007), Leśniewska & Witak (2008, 2011), Pliński & Witkowski (2009, 2011, 2013a,b), Witak & Pędziński (2018), Pędziński & Witak (2019). Previous studies usually focused on diatoms preserved in the sediments in different parts of the Gulf of Gdańsk. The diatom research in the coastal zone of the Gulf of Gdańsk was used to determine the degree of water organic pollution (Bogaczewicz-Adamczak et al. 2001; Zgrundo & Bogaczewicz-Adamczak 2004) and to investigate diatom preferences with respect to habitat (Witak et al. 2020). An important aspect of many diatom studies is to determine the degree of anthropogenic eutrophication. A multifaceted discussion on the relationship between diatoms and eutrophication in the Gulf of Gdańsk was included in the works of Witkowski (1994), Stachura & Witkowski (1997), Witak (2010, 2013), Leśniewska & Witak (2011), Witak & Pędziński (2018) and Pędziński & Witak (2019). Consequently, the so-called anthropogenic assemblage dominated by small planktic diatoms tolerating high levels of nitrogen, phosphorous and organic matter was defined, represented by
The latest research on species diversity of epilithon, epipsammon and epiphyton was conducted in the inner coastal zone of the Hel Peninsula (Witak et al. 2020). That study has shown that diatom taphocoenoses are represented by species that prefer one type of microhabitat: epilithon is characterized by the presence of
The study area is located in the coastal zone of the south-western part of the Gulf of Gdańsk, the southern Baltic Sea. This region includes shallows with a depth not exceeding 30 cm b.s.l. along the shore between Gdynia and Sopot, as well as an area of deeper water in the Port of Gdynia dredged to ca. 16 m b.s.l (Fig. 1). Three types of coasts occur in the area, i.e. accumulative, cliffs and anthropogenic. The northern section represents the anthropogenic coast and comprises the Port of Gdynia and the Seaside Boulevard with a concrete revetment. The central part of the analyzed fragment of the coastal zone has the form of a cliff. This part of the shore is exposed to the intense erosive action of waves, which results in the presence of the Orłowo Cliff with the most seaward jutting Orłowo Headland, which constitutes a conventional border between Puck Bay and the open part of the Gulf of Gdańsk. To the south, the cliff is cut off by the Kacza River. This part of the shore features anthropogenic elements in the form of hydrotechnical structures, i.e. a pier and spurs. The southern part, located between the mouth of the Kolibianka River and the mouth of the Vistula River outside the city of Gdańsk, has an accumulative character. In the Sopot area there is a wide sand beach. Nevertheless, hydrotechnical structures such as the pier and spurs largely affect the hydrodynamics of this part of the bay (Basiński et al. 1993).
Figure 1
Location of the study area

The hydrological regime of the coastal zone of the Gulf of Gdańsk is determined by shallow water depth, climatic conditions, and the inflow of saline waters from the open sea. In addition, freshwater discharge from the surrounding land is an important factor influencing the hydrology of the study area. The deeper part of the Gulf of Gdańsk is controlled by the strong runoff from the Vistula mouth (Kravtsov et al. 2002). The influence of the Vistula River in the coast between Gdynia and Sopot is strongly limited due to prevailing winds blowing from the W, WNW and WSW directions, which distribute river water in an easterly direction. On the other hand, winds blowing from the E, ENE and ESE directions cause the Vistula runoff to move westward. The impact of the Vistula waters in the study area increases during extreme floods when the flood wave can reach up to 27 km from the mouth of the Vistula (Wielgat-Rychert et al. 2013). Moreover, the waters of the Vistula have the greatest impact on the nutrient load in the Gulf of Gdańsk. They introduce 90% of nitrogen and 81% of phosphorus (Andurlewicz & Witek 2002). However, the content of nutrients in the coastal zone of the Gulf of Gdańsk is significantly affected by industry (0.3% N, 0.9% P) and wastewater treatment plants (3% N, 3% P; Andurlewicz & Witek 2002). The impact of other rivers and streams, including the Kacza, the Chylonka, the Kolibianka, the Swelinia and the Karlikowski Stream is limited to the proximity of their outlets (Majewski 1972). The Baltic Sea monitoring studies conducted in 2010–2020 showed that the concentrations of total phosphorus and total nitrogen were lower in 2020 than in previous years (Drgas 2021).
Rivers provide warm water in summer and cool water in winter. On the other hand, seawater from the Gdańsk Basin have the opposite effect and cause temperature increase in winter and decrease in summer (Nowacki 1993a). Moreover, thermal conditions of the waters in the study area are strictly dependent on changes in air temperature in particular seasons. Long-term studies show the advantage of higher water temperatures compared to air temperature from August to January. The opposite trend is observed from March to June. Moreover, two compensations for temperature differences are noticeable. The first one occurs in February and the second one in July (Kwiecień 1990). The lowest air temperatures in the coastal zone of the south-western Gulf of Gdańsk are recorded in January and range from –1°C to –2°C, while the highest extreme values occur in August (ca. 28°C; Kwiecień 1990). At the same time, the surface water temperature reaches the lowest values in February (ca. 0°C) and the highest values in August (ca. 24°C; Herman 2021). The average water temperature in the coastal zone between Gdynia and Sopot from December to April (cold season) is 2.5–3.0°C. In the warm season, the temperature increases from 15.6–16.0°C in the vicinity of river mouths to 15.1–15.5°C in the area of the Port of Gdynia (Urbański et al. 2007). Seasonal changes in salinity are also observed. The average salinity in the cold season near the port and the Orłowo Headland is 7.59–7.75 PSU. In the coastal zone of Sopot, the salinity increases to 7.76–7.78 PSU. In the warm season, the salinity of 7.38–7.49 PSU is observed in almost all parts of the coastal zone of Gdynia and Sopot. Only in the area of the Orłowo Headland, the salinity drops to the value of 7.18–7.37 PSU due to the inflow of the Kacza River (Urbański et al. 2007).
The water circulation in the Gulf of Gdańsk is determined by the shape of the coastal zone and the frequent occurrence of winds from the western sector (Kowalik 1990). The basin is dominated by clockwise currents with an average speed of about 10 cm s-1 (Nowacki 1993b). The westerly winds cause the formation of currents that bring water from the outer side of the Hel Peninsula to the eastern side of the Gulf of Gdańsk. The coastal zone is characterized by the propagation of currents parallel to the shore (Kowalik 1990). There are also bottom currents, the direction of which is opposite to the surface currents and their speed is about 4 cm s-1 (Nowacki 1993b).
The bottom of the shallow-water zone of the Gulf of Gdańsk is mainly composed of sandy sediments (Kramarska 1995). In the Port of Gdynia, there are fine-grained sands and muds. In the marina, on the other hand, there are silty sands (Urbański et al. 2007). The remaining part of the studied coastal zone is covered with very well-sorted fine-grained sand. Less sorted sediments, represented by coarse sand, gravel and stones, occur only in the area of the Orłowo Cliff (Majewski 1990). The content of organic matter in sandy sediments is low and does not exceed 1%. However, in the mouth of the Kacza River, it increases to 5% (Majewski 1990).
The flora near the seabed of the coastal zone of the Gulf of Gdańsk is not rich and usually occurs to a depth of 8 m (Ringer 1990). However, high biodiversity is typical of the shallow-water region along a coastal cliff moraine plateau called Kępa Redłowska, where 17 species of macroalgae were identified (Pliński & Florczyk 1993). Numerous species of red algae –
The material studied was collected in July 2016 along the coastal zone of the Gulf of Gdańsk from the Port of Gdynia up to the city limits of Sopot. A total of 19 sites were surveyed, including six sites in the Port of Gdynia (sites GaH1–6), eight sites along the coastal zone of Gdynia (sites Ga1–8) and five sites along the coastal zone of Sopot (sites So1–5; Fig. 1). Four samples were collected at site Ga2 (Ga2a,b,c,d), and two samples each were collected at sites Ga3 (Ga3a,b) and So1 (So1a,b; Table 1). At most sites, the diatom flora was analyzed from several microhabitats. At five sites (Ga2–3, Ga6–8), the study material was collected from stones. Sand was collected at five sites in Gdynia (Ga1–2, Ga5, Ga7–8) and four sites in Sopot (So1–2, So4–5). Muddy sediments were collected only in the Port of Gdynia (GaH1–6). Seagrasses were collected at 13 sites (Ga1–8, So1–5). A total of 37 samples were analyzed.
Characteristics of the analyzed samples. Types of microhabitats: st – stone, sa – sand, mu – mud, sg – seagrass. Black dots represent single samples collected
Sites | Samples | Φ | λ | Location | Depth [m] | Types of microhabitats | ||||
---|---|---|---|---|---|---|---|---|---|---|
st | sa | mu | sg | |||||||
GaH1 | GaHl | 54°32.523N | 18°30.32E | GDYNIA PORT | port channel | 13.6 | • | |||
GaH2 | GaH2 | 54°32.30N | 18°31.20E | Dock VII | 15.6 | • | ||||
GaH3 | GaH3 | 54°32.10N | 18°31.85E | port channel | 12.3 | • | ||||
GaH4 | GaH4 | 54°31.85N | 18°33.27E | Dock III | 14.4 | • | ||||
GaH5 | GaH5 | 54°31.70N | 18°33.62E | south channel | 8.7 | • | ||||
GaH6 | GaH6 | 54°31.26N | 18°33.50E | Dock I | 9.2 | • | ||||
Ga1 | Ga1 | 54°30.999N | 18°33.138E | GDYNIA | beach | 0.25 | • | • | ||
Ga2 | Ga2a | 54°30.774N | 18°33.062E | 0.25 | • | • | • | |||
Ga2b | • | |||||||||
Ga2c | • | |||||||||
Ga2d | • | |||||||||
Ga3 | Ga3a | 54°30.573N | 18°33.203E | Seaside Boulevard | 0.3 | • | ||||
Ga3b | • | |||||||||
Ga4 | Ga4 | 54°30.499N | 18°33.258E | 0.3 | • | |||||
Ga5 | Ga5 | 54°30.333N | 18°33.412E | 0.3 | • | • | ||||
Ga6 | Ga6 | 54°30.147N | 18°33.527E | 0.3 | • | • | ||||
Ga7 | Ga7 | 54°29.101N | 18°34.118E | beach | 0.3 | • | • | • | ||
Ga8 | Ga8 | 54°28.788N | 18°33.834E | pier | 0.2 | • | • | • | ||
So1 | So1a | 54°27.057N | 18°34.062E | SOPOT | beach | 0.3 | • | • | ||
So1b | • | |||||||||
So2 | So2 | 54°26.811N | 18°34.345E | pier | 0.25 | • | • | |||
So3 | So3 | 54°26.447N | 18°34.636E | beach | 0.3 | • | ||||
So4 | So4 | 54°26.115N | 18°35.051E | 0.24 | • | • | ||||
So5 | So5 | 54°25.666N | 18°35.751E | 0.27 | • | • |
Diatom samples containing live cells were prepared according to the standard procedure of Battarbee (1986). All samples were treated with 30% H2O2 to remove organic matter. However, sediment samples (ca. 0.5–1 g of dry sediment) were previously treated with 10% HCl to remove calcium carbonate. Quantitative diatom analysis was carried out on all samples. Moreover, qualitative analysis was performed on sandy and muddy sediments. The random settling technique was used to estimate the concentration of diatom valves per unit weight of dry sediment (Bodén 1991). Permanent diatom preparations were mounted in Naphrax with a refractive index nD of 1.73. The analysis was performed under a Nikon ECLIPSE E200 light microscope at a magnification of ×100, using oil immersion. Only whole, undamaged diatom valves were considered for counting. The counting method of Schrader and Gersonde (1978) was applied, and ca. 300–500 valves in each sample were counted to estimate relative abundance of each taxon. Row counts were converted into relative abundance of all valves counted. The following identification keys were used to identify diatoms: Hustedt (1927–1966), Krammer & Lange-Bertalot (1986, 1988, 1991a,b), Pankow (1990), Krammer (2000), Lange-Bertalot (2001), Bąk et al. (2012). In addition, the identified diatoms were classified with respect to their autoecological preferences, including microhabitat, salinity, trophic and saprobic status, which was completed based on OMNIDIA 6.08 software. All details of ecological groupings were presented by Witak et al. (2020). In the case of some marine and brackish water species, trophic and saprobic preferences are irrelevant. The percentage content of all ecological groups was estimated in each sample. Only species with frequency exceeding 3% in at least one sample were selected for diatom diagrams prepared using TILIA 2.0.37 (Grimm 2011).
A total of 147 species, subspecies, varieties and forms belonging to 56 genera were identified in the material studied (Fig. 2). Epipelic diatoms were the most diverse group, represented by 86 species. Epilithic and epiphytic diatoms were less diverse, with 68 and 71 species identified in these groups, respectively. The lowest diversity, 68 species, was observed in the epipsammic community.
Figure 2
The number of diatom taxa (species, subspecies, varieties, forms) versus ecological preferences: *eh-euhalobous, mh-mesohalobous, oh–oligohalobous halophilous, oi–oligohalobous indifferent; **et–eutraphentic, emt–eumesotraphentic, mt–mesotraphentic, mot-meso-oligotraphentic, ot–oligotraphentic, edt–eurydystrophic, ir-irrelevant; ***ps–polysaprobous, ams–α-mesosaprobous, abms–α-β-mesosaprobous, bms–β-mesosaprobous, os– oligosaprobous, x-xenosaprobous, ir-irrelevant

Diatom taphocoenoses of the shallow coastal zone of the Gulf of Gdańsk are usually dominated by benthic species. Planktic species constituted the allochthonous element of the studied material. The exception is muddy sediments in the Port of Gdynia, where epipelon is mainly represented by planktic species with frequency reaching 80%. This group was predominantly represented by eutraphentic diatoms, including
In the Gdynia region, oligohalobous halophilous, eutraphentic and β-mesosaprobic diatoms predominate in the epilithon. Their frequency usually exceeds 80% (Fig. 3). However, at the mouth of the Kacza River (Ga8), the frequency of β-mesosaprobionts drops to ca. 40%. At the same time, the percentage of α-mesosaprobionts increases to ca. 40%. Diatoms growing on stones (Gdynia sites) were usually dominated by the species
Figure 3
Percentage content of the diatom ecological groups in epilithon (stone), epipsammon (sand) and epipelon (mud)

Figure 4
Frequency of the main diatom taxa in epilithon (stone), epipsammon (sand) and epipelon (mud)

The epipsammic assemblage of the Gdynia and Sopot zone is dominated by eutraphentic diatoms with a maximum frequency of 78% at site Ga8 (Fig. 3). They were accompanied by eu-mesotraphentic species (7–35%). The percentage of saline groups indicates a slightly higher frequency of mesohalobous diatoms at the Sopot sites than in Gdynia. The brackish species
The epipelic assemblage found only in the Port of Gdynia (GH1–8) was dominated by eutraphentic (49–82%) and α- and β-mesosaprobic diatoms (10–30% and 10–35%, respectively; Fig. 3). The concentration of diatom valves is clearly higher in the port muds than in the sandy sediments of Gdynia and Sopot and ranges between 25 x 105 valves/g (GaH5) and 75 x 105 valves/g (GaH1). The marine species
The epiphytic assemblage occurring at all sites in Gdynia and Sopot, except the port, is dominated by diatoms preferring nutrient-rich waters (70–98%) and tolerating large amounts of organic matter, i.e. α- and β-mesosaprobionts (33–91%, 7–62% respectively; Fig. 5). Between sites Ga1 and So1, the epiphytic assemblage is dominated by oligohalobous halophilous species, i.e.
Figure 5
Percentage content of the diatom ecological groups in epiphyton (seagrass)

Figure 6
Frequency of the main diatom taxa in epiphyton (seagrass)

Our results have shown significant differences in species spectra in relation to microhabitats. Changes in saline conditions caused by the inflow of river waters as well as anthropopressure had a significant impact on the species diversity of the diatom flora. The observed differences in individual diatom communities (epilithic, epipsammic, epipelic and epiphytic) allowed us to distinguish four groups: (1) species characteristic of one microhabitat, (2) species living in two types of microhabitats, (3) species observed in three microhabitats, and (4) species occurring in all types of microhabitats.
Some diatom species were found in only one of the analyzed communities (epipsammon, epipelon, epiphyton).
Three groups were distinguished among the diatoms occurring in two microhabitats. The first group represented by
The species
Diatoms occurring in all types of microhabitats are represented by the species reported by Snoeijs (1993) in the epilithon (
The port of Gdynia is impacted by seawater of the Gulf of Gdańsk, which is well documented by the dominance of marine and brackish diatoms in the muddy sediments. Due to ship traffic and intensive port activity, the water column and bottom sediments contain a large number of organic pollutants. Diatom taphocoenoses in its north-western part (GaH1–2) are rich in polysaprobionts (≤ 44%) and β-mesosaprobionts (≤ 37%). Moreover, a higher frequency of the α-mesosaprobiont
The inflow of water from the Kacza River has a significant impact on the salinity conditions in the this part of the coastal zone. The Kacza River is the second largest river flowing through the Tri-City (Augustowska et al. 1994). Between the marina (Ga1) and the Orłowo Headland (Ga8), brackish diatoms (
The hydrological regime of the coastal zone in Sopot is influenced by saline waters of the Gulf of Gdańsk. This is evidenced by the dominance of eutraphentic mesohalobous, represented mostly by
The results of the diatom study in the coastal zone of the Gulf of Gdańsk, between Gdynia and Sopot, indicate the presence of a diverse flora belonging to epilithic, epipsammic, epipelic and epiphytic assemblages. Most of the identified benthic species occurred in two or three microhabitats:
Epilithon and epiphyton were represented by The species
However, considering the diatom–microhabitat relationships, two specific groups were distinguished:
Epipsammon, epipelon and epiphyton were characterized by the presence of at least one species that was not observed in other communities. The species Some diatom species, i.e.
Based on the ecological preferences of the identified diatom flora, certain differences in the quality of water were observed in the Port of Gdynia and in the coastal zone of Gdynia and Sopot:
The high frequency of α-mesosaprobionts dominated by The high degree of eutrophication of the waters of the study area was evidenced by the abundance of species with high trophic requirements, represented by The change in salinity in the study area is related to the distance from the mouths of nearby rivers and streams. Due to the limited water dynamics associated with the breakwater, the area of the Port of Gdynia is dominated by marine species, of which
Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Characteristics of the analyzed samples. Types of microhabitats: st – stone, sa – sand, mu – mud, sg – seagrass. Black dots represent single samples collected
Sites | Samples | Φ | λ | Location | Depth [m] | Types of microhabitats | ||||
---|---|---|---|---|---|---|---|---|---|---|
st | sa | mu | sg | |||||||
GaH1 | GaHl | 54°32.523N | 18°30.32E | GDYNIA PORT | port channel | 13.6 | • | |||
GaH2 | GaH2 | 54°32.30N | 18°31.20E | Dock VII | 15.6 | • | ||||
GaH3 | GaH3 | 54°32.10N | 18°31.85E | port channel | 12.3 | • | ||||
GaH4 | GaH4 | 54°31.85N | 18°33.27E | Dock III | 14.4 | • | ||||
GaH5 | GaH5 | 54°31.70N | 18°33.62E | south channel | 8.7 | • | ||||
GaH6 | GaH6 | 54°31.26N | 18°33.50E | Dock I | 9.2 | • | ||||
Ga1 | Ga1 | 54°30.999N | 18°33.138E | GDYNIA | beach | 0.25 | • | • | ||
Ga2 | Ga2a | 54°30.774N | 18°33.062E | 0.25 | • | • | • | |||
Ga2b | • | |||||||||
Ga2c | • | |||||||||
Ga2d | • | |||||||||
Ga3 | Ga3a | 54°30.573N | 18°33.203E | Seaside Boulevard | 0.3 | • | ||||
Ga3b | • | |||||||||
Ga4 | Ga4 | 54°30.499N | 18°33.258E | 0.3 | • | |||||
Ga5 | Ga5 | 54°30.333N | 18°33.412E | 0.3 | • | • | ||||
Ga6 | Ga6 | 54°30.147N | 18°33.527E | 0.3 | • | • | ||||
Ga7 | Ga7 | 54°29.101N | 18°34.118E | beach | 0.3 | • | • | • | ||
Ga8 | Ga8 | 54°28.788N | 18°33.834E | pier | 0.2 | • | • | • | ||
So1 | So1a | 54°27.057N | 18°34.062E | SOPOT | beach | 0.3 | • | • | ||
So1b | • | |||||||||
So2 | So2 | 54°26.811N | 18°34.345E | pier | 0.25 | • | • | |||
So3 | So3 | 54°26.447N | 18°34.636E | beach | 0.3 | • | ||||
So4 | So4 | 54°26.115N | 18°35.051E | 0.24 | • | • | ||||
So5 | So5 | 54°25.666N | 18°35.751E | 0.27 | • | • |
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