Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region
Bays and estuaries of the Black Sea–Azov region are characterized by pronounced hydrological variability, which creates a wide range of environmental conditions and influences the taxonomic structure of microalgae. Despite the substantial number of studies, available data are mostly reported for sep...
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Algologia| _version_ | 1867479081454403584 |
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| author | Bren, O.G. Bren, O.A. Podorozhny, S.M. |
| author_facet | Bren, O.G. Bren, O.A. Podorozhny, S.M. |
| author_institution_txt_mv | [
{
"author": "O.G. Bren",
"institution": "Institute of Botany of the Czech Academy of Sciences, 135 Dukelska, Třeboň, Czech Republic"
},
{
"author": "O.A. Bren",
"institution": "Centre for Polar Ecology, University of South Bohemia, 3 Na Zlaté stoce, České Budějovice, Czech Republic"
},
{
"author": "S.M. Podorozhny",
"institution": "Bogdan Khmelnitsky Melitopol State Ped. Univ., 59 Naukove mistechko, Zaporizhzhia 69017, Ukraine"
}
] |
| author_sort | Bren, O.G. |
| baseUrl_str | https://algologia.co.ua/journal/oai |
| collection | OJS |
| datestamp_date | 2026-06-08T18:00:44Z |
| description | Bays and estuaries of the Black Sea–Azov region are characterized by pronounced hydrological variability, which creates a wide range of environmental conditions and influences the taxonomic structure of microalgae. Despite the substantial number of studies, available data are mostly reported for separate water bodies, which complicates their comparison. In this study, published data on the taxonomic structure of microalgae were compiled and comparatively analysed at the level of phyla and classes using multivariate statistical approaches, including ordination (NMDS) based on the Bray–Curtis index, permutational multivariate analysis of variance (PERMANOVA), and analysis of the contribution of individual taxa to differences between water body types (SIMPER). In bays, Bacillariophyta and Dinoflagellata had generally comparable species proportions, whereas in estuaries Bacillariophyta comprised substantially more species, while Dinoflagellata were represented by few species or were absent in several water bodies. These differences indicate patterns in the taxonomic structure of microalgae between bays and estuaries of the Black Sea–Azov region and could be considered in ecological monitoring. |
| doi_str_mv | 10.15407/alg36.02.107 |
| first_indexed | 2026-06-09T01:00:36Z |
| format | Article |
| fulltext |
Comparative analysis of the taxonomic structure
ISSN 2413-5984. Альгологія. 2026. 36(2) 107
BREN O.G. 1, BREN O.A. 2, PODOROZHNY S.M. 3
1 Institute of Botany of the Czech Academy of Sciences,
135 Dukelska, Třeboň, Czech Republic
2 Centre for Polar Ecology, University of South Bohemia,
3 Na Zlaté stoce, České Budějovice
3 Bogdan Khmelnitsky Melitopol State Ped. Univ.,
59 Naukove mistechko, Zaporizhzhia 69017, Ukraine
* Author for correspondence: oscillat8@gmail.com
COMPARATIVE ANALYSIS OF THE TAXONOMIC STRUCTURE OF
MICROALGAE IN BAYS AND ESTUARIES OF THE BLACK SEA–
AZOV REGION
Abstract. Bays and estuaries of the Black Sea–Azov region are characterized by pronounced
hydrological variability, which creates a wide range of environmental conditions and influences
the taxonomic structure of microalgae. Despite the substantial number of studies, available data
are mostly reported for separate water bodies, which complicates their comparison. In this
study, published data on the taxonomic structure of microalgae were compiled and
comparatively analysed at the level of phyla and classes using multivariate statistical
approaches, including ordination (NMDS) based on the Bray–Curtis index, permutational
multivariate analysis of variance (PERMANOVA), and analysis of the contribution of
individual taxa to differences between water body types (SIMPER). In bays, Bacillariophyta
and Dinoflagellata had generally comparable species proportions, whereas in estuaries
Bacillariophyta comprised substantially more species, while Dinoflagellata were represented
by few species or were absent in several water bodies. These differences indicate patterns in the
taxonomic structure of microalgae between bays and estuaries of the Black Sea–Azov region
and could be considered in ecological monitoring.
Keywords: Bacillariophyta, Dinoflagellata, bays, estuaries, phyla, classes, regional patterns,
Black Sea–Azov region
Надійшла до редакції 01.05.2026. Після доопрацювання 19.05.2026. Опублікована 20.06.2026
Ц и т у в а н н я . Bren O.G., Bren O.A., Podorozhny S.M. 2026. Comparative analysis of the taxonomic
structure of microalgae in bays and estuaries of the Black Sea–Azov region. Альгологія. 36(2): 107–117.
https://doi.org/10.15407/alg36.02.107
This is open access article under the CC BY license (https://creativecommons.org/licenses/by/4.0/)
© Інститут ботаніки ім. М.Г. Холодного НАН України, 2026
© Видавець ВД «Академперіодика» НАН України, 2026
Bren O.G., Bren O.A., Podorozhny S.M.
108 ISSN 2413-5984. Algologia. 2026. 36(2)
Introduction
Bays and estuaries are characterized by fluctuations in hydrological and
physicochemical parameters, which also affect aquatic biota. Microalgae
represent an important and integral component of coastal aquatic ecosystems and
respond rapidly to environmental changes. Their species composition and
taxonomic structure reflect the ecological state of a water body and allow the
detection of changes that are not always clearly captured by other indicators
(Barinova et al., 2019; Barsanti, Birindelli, 2021; Shevchenko et al., 2023).
Despite the availability of numerous studies on microalgae in the Black Sea-
Azov region, the results are typically reported for individual water bodies
(Dolinska, 2016; Terenko, Terenko, 2000; Gerasimyuk, Ennan et al., 2023) or
limited to a single type of water body, particularly estuaries (Gerasimyuk, 2018).
This approach complicates comparative analysis of microalgal taxonomic
composition across different types of water bodies and limits generalization at
the regional scale.
As a result, it remains unclear whether the taxonomic structure of
microalgae differs between different types of water bodies, particularly between
bays and estuaries, and which taxa contribute to these differences, if present. The
Black Sea–Azov region, characterized by a high number of bays and estuaries,
their spatial proximity, and the availability of accumulated phycological data,
provides a suitable setting for such a comparative analysis.
The aim of this study is to perform a comparative analysis of the taxonomic
structure of microalgae in bays and estuaries of the Black Sea–Azov region.
We hypothesize that the taxonomic structure of microalgae at the level of
phyla and classes differs between bays and estuaries in this region.
Materials and Methods
The study is based on a synthesis of published data on the taxonomic structure of
microalgae in bays and estuaries (estuarine water bodies) of the northern part
of the Black Sea–Azov region (Fig. 1). Data were compiled for estuaries of the
northwestern Black Sea region (Gerasimyuk, 2018), water bodies of the eastern
Azov region (Kovaleva, 2008), bays of Crimea (Tokareva et al., 2008; Nevrova,
2015), and the Odesa Bay (Terenko, Terenko, 2000). In total, 22 water bodies
were analysed, including 6 bays and 16 estuaries.
Taxonomic assignment of microalgal taxa followed the classification
adopted in AlgaeBase (Guiry, Guiry, 2025).
Similarity among water bodies was assessed using the Bray–Curtis index
calculated based on the relative proportions of phyla. Ordination was performed
using non-metric multidimensional scaling (NMDS). Differences in taxonomic
Comparative analysis of the taxonomic structure
ISSN 2413-5984. Альгологія. 2026. 36(2) 109
structure between water body types (bays and estuaries) were tested using
permutational multivariate analysis of variance (PERMANOVA) with 9,999
permutations. Homogeneity of multivariate dispersion was assessed using
PERMDISP. To evaluate the contribution of individual taxonomic groups to
differences between bays and estuaries, similarity percentage analysis (SIMPER)
based on the Bray–Curtis index was applied. Differences in the distribution of
relative proportions of microalgal phyla between (bays and estuaries were tested
using the non-parametric Mann–Whitney test. Effect size was estimated using
the Vargha–Delaney A statistic. All calculations were performed in PAST 5.3
(Hammer et al., 2001).
Fig. 1. Map of bays and estuaries in the northern part of the Black Sea-Azov region included in the
analysis
Results
Based on published data, a compiled dataset of relative proportions (%) of the
main phyla and classes of microalgae was obtained (Fig. 2). In each analysed
water body, regardless of type, Bacillariophyta prevailed in the taxonomic
structure in terms of the species proportion (31.2–97.5%). Representatives of
Cyanobacteriophyta also contributed substantially, with proportions ranging
from 0.9 to 23.1%.
At the same time, differences between water body types were observed. In
particular, Dinoflagellata and Haptophyta showed relatively high proportions in
bays (up to 43.5% and 10.2% respectively). In estuaries they generally had lower
proportions of the total number of recorded species or were absent:
Dinoflagellata — up to 5.4% (with exception in the Budak Liman — 26.9%) and
Haptophyta 3.8%. A similar pattern was observed for Chrysophyceae,
Cryptophyceae, and Dictyochophyta, which were mainly recorded in bays and
Bren O.G., Bren O.A., Podorozhny S.M.
110 ISSN 2413-5984. Algologia. 2026. 36(2)
were nearly absent in estuaries, although their proportions were low in all
studied water bodies.
Fig. 2. Representation of microalgal taxa (phyla and classes) in the studied bays and estuaries of
the Black and Azov Seas (%)
Non-metric multidimensional scaling (NMDS; stress = 0.09) revealed a
clear separation of the studied water bodies according to their type (Fig. 3). The
first NMDS axis represents the main gradient in the data and distinctly separates
marine bays from estuaries. All bays are located in the negative part of axis 1,
whereas estuaries consistently cluster in its positive part, indicating differences
in the taxonomic structure of microalgae between these water bodies. The
second axis explains a smaller portion of the variation and does not play a
substantial role in separating bays and estuaries.
The distribution of taxonomic groups in the ordination space showed that
some taxa (Dinoflagellata, Haptophyta and Dictyochophyta) were positioned in
the negative part of axis 1 with bays, whereas others occupied its positive
part and were characteristic of estuaries. The separation of water bodies
along axis 1 is primarily driven by taxa located at its extreme positions.
It should be noted that the Taganrog Bay and Dinska Bay do not group with
the other bays, instead occupying positions among the estuaries.
Comparative analysis of the taxonomic structure
ISSN 2413-5984. Альгологія. 2026. 36(2) 111
Fig. 3. NMDS ordination of coastal water bodies based on the Bray–Curtis index calculated from
the relative proportions of microalgal taxa. Black points represent water bodies, and purple points
represent microalgal taxonomic groups
PERMANOVA results indicated statistically significant differences in the
taxonomic structure of microalgae between bays and estuaries (F = 16.16,
p = 0.0002), with water body type explaining approximately 45% of the total
variation. The test for homogeneity of multivariate dispersion (PERMDISP)
showed no statistically significant differences between bays and estuaries
(p = 0.06), allowing reliable interpretation of the PERMANOVA results.
SIMPER analysis (Table) showed that differences in the proportions of
Bacillariophyta and Dinoflagellata accounted for approximately 70% of the
overall dissimilarity between bays and estuaries. Bacillariophyta had
substantially higher mean relative proportions in estuaries (80.6%) compared to
bays (49.1%), whereas Dinoflagellata were characteristic of bays (26.9%) and
were much less represented in estuaries (3.1%). Additional differences were
contributed by Cyanobacteriophyta, which were more frequently recorded in
estuaries than in bays (9.2% and 4.1%, respectively).
To assess differences between bays and estuaries in the distribution of major
microalgal taxa, the non-parametric Mann–Whitney test was applied. The
analysis showed that the distribution of Bacillariophyta proportions was shifted
towards higher values in estuaries compared to bays (p < 0.01; A = 0.10). In
contrast, Dinoflagellata exhibited the opposite pattern, with higher proportions
in bays (p < 0.01; A = 0.94). Similarly, Haptophyta were also characterized by
higher values in bays (p < 0.01; A = 0.86), although their overall representation
was lower than that of Dinoflagellata.
Bren O.G., Bren O.A., Podorozhny S.M.
112 ISSN 2413-5984. Algologia. 2026. 36(2)
Table. Microalgal taxa contributing to differences between bays and estuaries (SIMPER
analysis)
Taxon Contributin
g, %
Mean proportion in
bays, %
Mean proportion in
estuaries, %
Bacillariophyta 39.7 47.4 80.6
Dinoflagellata 30.1 28.4 3.1
Cyanobacteriophyta 7.9 4.1 9.2
Chlorophyta 6.3 6.3 4.7
Haptophyta 5.5 5.0 0.4
Cryptophyceae 2.6 24 0.2
Euglenophyta 2.5 2.3 0.1
Chrysophyceae 2.0 1.9 0.8
Dictyochophyta 1.4 1.2 0.1
Charophyta 0.8 0.0 0.7
Xanthophyceae 0.7 0.6 0.1
Raphidophyceae 0.4 0.3 0.0
Olisthodiscophyceae 0.1 0.1 0.0
For other taxonomic groups, including Chlorophyta and Cyano-
bacteriophyta, despite their occurrence in nearly all analysed water bodies, no
statistically significant differences in the distribution of proportions between
bays and estuaries were detected (p > 0.05).
Thus, the results of the comparative analysis indicate differences in the
taxonomic structure of microalgae between bays and estuaries of the Black Sea–
Azov region, primarily reflected in the relative proportions of Bacillariophyta
and Dinoflagellata.
Discussion
The results obtained in this study support the proposed hypothesis that the
taxonomic structure of microalgae at the level of phyla and classes differs
between bays and estuaries of the Black Sea–Azov coast.
The high representation of Bacillariophyta and Dinoflagellata in coastal
water bodies worldwide, as well as their sensitivity to environmental changes,
has been reported by other studies (Carstensen et al., 2015; Garrido et al., 2016;
Miao et al., 2024). At the same time, the dominance of these groups in terms of
species proportions varies among studies. Some reports are consistent with our
findings (Cardoso, Torgan, 2007; Saifullah et al., 2014; Telesh et al., 2025),
whereas others indicate a prevalence of diatoms in bays (Chan, Hamilton, 2001;
Comparative analysis of the taxonomic structure
ISSN 2413-5984. Альгологія. 2026. 36(2) 113
Souza et al., 2013; Stanca et al., 2013) or dinoflagellates in estuaries (Sellner et
al., 2001; Prabhudessai, Rivonker, 2020).
This variability likely reflects the primary role of local environmental
conditions, particularly salinity, water exchange intensity, and turbulence
(Elferink et al., 2020; Bi et al., 2021; Xu et al., 2024), while the water body type
represents a generalized expression of these factors. In this context, the results
obtained in this study should be interpreted primarily as reflecting regional
features of the taxonomic structure of microalgae in coastal water bodies of the
Black Sea-Azov region.
Along with the observed general patterns in the distribution of taxonomic
groups, some deviations were identified. In particular, in Taganrog Bay and
Dinska Bay, the proportion of Bacillariophyta substantially exceeds that of
Dinoflagellata, which may be related to specific salinity conditions as well as
trophic and hydrochemical regimes of these water bodies (Belova et al., 2026;
Sukhinov, Belova, 2026).
Another example is Budak Liman, which, in contrast, shows a taxonomic
structure closer to that of bays, particularly Odesa Bay, with comparable
proportions of these two major groups. Limited hydrological and hydrochemical
data for this water body do not allow a clear explanation of these differences.
It should be noted that the dataset for coastal water bodies of the Azov Sea
in this study is smaller and includes fewer sites compared to those from the
Black Sea coast. For some water bodies, available phycological data are limited,
particularly for Zapadensky and Shahaivsky estuaries (Kovaleva, 2008). In
addition, some strongly transformed water bodies (e.g., Molochny and Utliuk
estuaries) (Bren et al., 2022), characterized by unstable and significantly altered
hydrological conditions, were not included in the analysis.
Among other limitations, the dataset includes both planktonic and benthic
microalgal communities, and for some water bodies, information is available for
only one of these components, which may affect the completeness of the
taxonomic representation. Despite these limitations, the analysis allowed
generalization of the main features of microalgal taxonomic structure in coastal
water bodies and revealed differences between their types at the level of higher
taxa.
The results of this study may be useful for monitoring the ecological status
of coastal water bodies. Future research could focus on expanding spatial
coverage and testing the observed patterns in other regions, as well as applying
molecular, non-culture-based approaches to improve data comparability and
reduce the time required for data acquisition.
Bren O.G., Bren O.A., Podorozhny S.M.
114 ISSN 2413-5984. Algologia. 2026. 36(2)
Conclusions
This study synthesizes published data on the proportions of microalgal
taxonomic groups in bays and estuaries of the Black Sea-Azov coast and
provides their comparative analysis at the level of phyla and classes. In bays,
Bacillariophyta and Dinoflagellata generally showed comparable proportions,
whereas in estuaries Bacillariophyta substantially prevailed. These differences
reflect patterns in the taxonomic structure of microalgae between the studied
water body types and reflect features of the Black Sea–Azov region.
ETHICS DECLARATION
The authors declare no conflict of interst.
ORCID
O.G. Bren: https://orcid.org/0000-0001-7423-9258
O.A. Bren: https://orcid.org/0009-0003-7632-6285
S.M. Podorozhny: https://orcid.org/0000-0002-7702-7602
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https://doi.org/10.1016/j.ocecoaman.2023.106982
Comparative analysis of the taxonomic structure
ISSN 2413-5984. Альгологія. 2026. 36(2) 117
Брен O.Г. 1, Брен O.А. 2, Подорожний С.M. 3
1 Інститут ботаніки Академії наук Чеської Республіки,
вул. Дукельська, 135, Тршебонь, Чеська Республіка
2 Центр полярної екології Південночеського університет
вул. На Злате Стоце, 3, Чеські Будейовиці, Чеська Республіка
3 Мелітопольський державний педагогічний університет імені Богдана Хмельницького,
вул. Наукове містечко, 59, Запоріжжя 69017, Україна
Порівняльний аналіз таксономічної структури мікроводоростей заток та лиманів
Чорноморсько-Азовського регіону
Затоки та лимани Чорноморсько-Азовського регіону характеризуються вираженою
гідрологічною мінливістю, що формує широкий спектр умов середовища та впливає на
таксономічну структуру мікроводоростей. Попри значну кількість досліджень, наявні дані
переважно наводяться для окремих водойм, що ускладнює їхнє порівняння. У роботі
узагальнено літературні дані про таксономічну структуру мікроводоростей та проведено
їхній порівняльний аналіз на рівні відділів і класів із застосуванням багатовимірних
статистичних підходів, включаючи ординацію (NMDS) на основі індексу Брея–Куртіса,
пермутаційний аналіз варіації (PERMANOVA) та SIMPER-аналіз. У затоках Bacillariophyta
та Dinoflagellata характеризувалися загалом співставними пропорціями видів, тоді як у
лиманах Bacillariophyta включали значно більшу кількість видів, а Dinoflagellata були
представлені незначною кількістю видів або були відсутні в окремих водоймах. Виявлені
відмінності свідчать про наявність закономірностей у таксономічній структурі
мікроводоростей між затоками та лиманами Чорноморсько-Азовського регіону та можуть
бути перспективними для використання в екологічному моніторингу.
К л ю ч о в і с л о в а : Bacillariophyta, Dinoflagellata, затоки, лимани, відділи, класи,
регіональні закономірності, Чорноморсько-Азовський регіон
Цитування : Брен О.Г., Брен О.А., Подорожний С.М. 2026. Порівняльний аналіз таксономічної
структури мікроводоростей заток і лиманів Чорноморсько-Азовського регіону. Альгологія. 36(2): 107–
117. https://doi.org/10.15407/alg36.02.107
|
| id | oai:algologia.co.ua:article-122 |
| institution | Algologia |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2026-06-09T01:00:36Z |
| publishDate | 2026 |
| publisher | M.G. Kholodny Institute of Botany, NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | algologiacoua/3e/9c32f3af760fa5dfbccb1b671df7d13e.pdf |
| spelling | oai:algologia.co.ua:article-1222026-06-08T18:00:44Z Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region Порівняльний аналіз таксономічної структури мікроводоростей заток та лиманів Чорноморсько-Азовського регіону Bren, O.G. Bren, O.A. Podorozhny, S.M. Bacillariophyta Dinoflagellata затоки лимани відділи класи регіональні закономірності Чорноморсько-Азовський регіон Dinoflagellata bays estuaries phyla classes regional patterns Black Sea–Azov region Bays and estuaries of the Black Sea–Azov region are characterized by pronounced hydrological variability, which creates a wide range of environmental conditions and influences the taxonomic structure of microalgae. Despite the substantial number of studies, available data are mostly reported for separate water bodies, which complicates their comparison. In this study, published data on the taxonomic structure of microalgae were compiled and comparatively analysed at the level of phyla and classes using multivariate statistical approaches, including ordination (NMDS) based on the Bray–Curtis index, permutational multivariate analysis of variance (PERMANOVA), and analysis of the contribution of individual taxa to differences between water body types (SIMPER). In bays, Bacillariophyta and Dinoflagellata had generally comparable species proportions, whereas in estuaries Bacillariophyta comprised substantially more species, while Dinoflagellata were represented by few species or were absent in several water bodies. These differences indicate patterns in the taxonomic structure of microalgae between bays and estuaries of the Black Sea–Azov region and could be considered in ecological monitoring. Затоки та лимани Чорноморсько-Азовського регіону характеризуються вираженою гідрологічною мінливістю, що формує широкий спектр умов середовища та впливає на таксономічну структуру мікроводоростей. Попри значну кількість досліджень, наявні дані переважно наводяться для окремих водойм, що ускладнює їхнє порівняння. У роботі узагальнено літературні дані про таксономічну структуру мікроводоростей та проведено їхній порівняльний аналіз на рівні відділів і класів із застосуванням багатовимірних статистичних підходів, включаючи ординацію (NMDS) на основі індексу Брея–Куртіса, пермутаційний аналіз варіації (PERMANOVA) та SIMPER-аналіз. У затоках Bacillariophyta та Dinoflagellata характеризувалися загалом співставними пропорціями видів, тоді як у лиманах Bacillariophyta включали значно більшу кількість видів, а Dinoflagellata були представлені незначною кількістю видів або були відсутні в окремих водоймах. Виявлені відмінності свідчать про наявність закономірностей у таксономічній структурі мікроводоростей між затоками та лиманами Чорноморсько-Азовського регіону та можуть бути перспективними для використання в екологічному моніторингу. M.G. Kholodny Institute of Botany, NAS of Ukraine 2026-06-20 Article Article application/pdf https://algologia.co.ua/journal/article/view/36.2.107 10.15407/alg36.02.107 Algologia; Vol. 36 No. 2 (2026); 107-117 Альгологiя; Том 36 № 2 (2026); 107-117 Альгология; Том 36 № 2 (2026); 107-117 2413-5984 0868-8540 10.15407/alg36.02 en https://algologia.co.ua/journal/article/view/36.2.107/142 Copyright (c) 2026 O.G. Bren, O.A. Bren, S.M. Podorozhny (Author) https://creativecommons.org/licenses/by-nc/4.0 |
| spellingShingle | Bren, O.G. Bren, O.A. Podorozhny, S.M. Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region |
| title | Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region |
| title_alt | Порівняльний аналіз таксономічної структури мікроводоростей заток та лиманів Чорноморсько-Азовського регіону |
| title_full | Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region |
| title_fullStr | Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region |
| title_full_unstemmed | Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region |
| title_short | Comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the Black Sea–Azov region |
| title_sort | comparative analysis of the taxonomic structure of microalgae in bays and estuaries of the black sea–azov region |
| topic_facet | Bacillariophyta Dinoflagellata затоки лимани відділи класи регіональні закономірності Чорноморсько-Азовський регіон Dinoflagellata bays estuaries phyla classes regional patterns Black Sea–Azov region |
| url | https://algologia.co.ua/journal/article/view/36.2.107 |
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