CC BY
7t ..*:' Трансформация экосистем
V
5 \1 V-Ecosystem Transformation 1 '-у www.ecosysttrans.com
The current state and dynamics of the flora of several small lakes of the Yaroslavl Region, Russia
Eugeny A. Belyakov*, Ekaterina G. Sakharova, Alexandra S. Sokolova
I.D. Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences, Borok 109, Nekouz Distict, Yaroslavl Region, 152742 Russia
*eugenybeliakov@yandex.ru
Received: 19.05.2020 Accepted: 14.09.2020 Published online: 13.11.2020
DOI: 10.23859/estr-200519 UDC 581.93, 574.583
ISSN 2619-094X Print ISSN 2619-0931 Online
Translated by D.M. Martynova
The phytoplankton and vascular aquatic plants have been studied in small lakes Vashutinskoe, Chashnitskoe, Ryumniki, and Zaozer'e, located in the southern part of the Yaroslavl Region, Russia. The taxonomic composition of phytoplankton comprised 137 species of algae belonging to 8 divisions, 18 orders, and 76 genera. The studied water bodies were characterized by high biomass of cyanobac-teria. Aphanizomenon flos-aquae (L.) Ralfs ex Born et Flah. and the species of the genus Anabaena had a significant contribution to the phytoplankton biomass in the lakes Zaozer'e, Chashnitskoe, and Ryumniki, Limnothrixplanctonica (Wotosz.) Meff. and Microcystis aeruginosa (Kutz.) Kutz, in the Lake Vashutinskoe. In terms of phytoplankton biomass, the pelagial and shallow areas of the Lake Vashutinskoe belonged to eutrophic reservoirs, of the Lake Ryumniki, to oligotrophic. The phytoplankton biomass in the pelagial zone of the lakes Chashnitskoe and Zaozer'e indicated the mesotrophic status of these water bodies, in the littoral zone, eutrophic. The study of the phytoplankton of the lakes was carried episodically and was often performed at different periods of the growing season. Therefore, the obtained data did not allow one to assess the direction of the successional changes in the algal flora. The flora of vascular plants of lakes was represented by 118 species belonging to 4 divisions, 5 classes, 22 orders, and 37 families. Thirty-four species were common to all studied water bodies. The differences in the The differences in the flora of vascular aquatic plantsof the studied lakes were associated with differences in their trophic status, the presence of various biotopes, the nature of anthropogenic load, and the degree of eutrophication. During a hundred years of observations, some species have disappeared, but some have appeared, including adventive species (such as Elodea canadensis Michx., Bidens frondosa L., Epilobium adenocaulon Hausskn, Zizania latifo-lia Griseb., and Juncus tenuis Willd.). The study of the current state of rare and protected species of vascular plants made it possible to reveal the eventual causes and period of disappearance of some species from the lakes. The taxonomic and ecological structure of phytoplankton and vascular plants of lakes was quite diverse and usual for the water bodies of the European part of Russia.
Keywords: lakes, phytoplankton, vascular plants, flora transformation, anthropogenic influence, rare species, trophic status.
Belyakov, E.A. et al., 2020. The current state and dynamics of the flora of several small lakes of the Yaroslavl Region, Russia. Ecosystem Transforation 3 (4), 15-40.
Introduction
Small lakes are still a relatively poorly studied group of the water bodies (Izmailova et al., 2017; Lyashenko et al., 2002). They are most sensitive to climate change and are subject to large anthropogenic pressure (Belyakov and Garin, 2018; Izmailova et al., 2017; Lawniczak-Malinska and Achtenberg, 2018; Lobunicheva et al., 2013; Slukovskii et al., 2018; Strahovenko et al., 2014). The latter affects the species richness of macrophytes, which is closely related to the natural type and status of the water bodies (Maemets et al., 2010).
The studied lakes Vashutinskoe, Chashnitskoe, Ryumniki, and Zaozer'e are located in the Eastern Borisoglebsk Upland in the East European Plain. In accordance with the classification of the lakes by the depth suggested by S.P. Kitaev (1984), the Lake Vashutinskoe refers to shallow water bodies, while the Chashnitskoe, Ryumniki and Zaozer'e lakes are mid-depth water bodies. The lakes Chashnitskoe, Vashutinskoe, and Zaozer'e belong to the basin of the Klyazminskaya Nerl' River; the Lake Ryumniki is located in the Ryumniko-Osoevskaya Depression, in the watershed area of the rivers flowing southeastwards, to the Klyazminskaya Nerl' River and further to the Klyazma River, and northwards, to the Sara River and further to the Kotorosl' River (Fortunatov and Moskovskii, 1970). The studied lakes are common by their glacial origin and the peculiarities of their location, but differ by the degree of anthropogenic impact, as a result, they have different modern trophic status. In addition, according to the Decree of the Government of the Yaroslavl Region of November 2, 2017 no. 823-p "On approval of the regime of special protection of the territory of natural monuments of regional significance in the Yaroslavl Region" (Postanovlenie..., 2017), the three lakes (Ryumniki, Chashnitskoe, and Vashutinskoe) are included into the list of natural monuments of regional significance, the species of vascular plants, included into the Red Data Book of the Yaroslavl Region (Krasnaya Kniga..., 2015), are found in each lake.
The data on the ecosystem state of these water bodies are scarce. The composition and some quantitative characteristics of their flora are known only from episodic observations carried out in the XX - beginning of the XXI century. The first data on the phytoplankton of lakes were obtained in 1903 (Blokhontsev, 1903a, b). During this period, 17 species of algae were reported for the Lake Ryumniki, 20 species, for the Lake Chashnitskoe, and 13 species, for the Lake
Zaozer'e (Grigor'ev, 1903). Cyanobacteria dominated by biomass in the phytoplankton communities of the lakes. According to the results of studies performed in 1963 (Fortunatov and Moskovskii, 1970), a large number of cells of Anabaena elliptica Lemm. was registered in the Lake Ryumniki, Aphanizomenon flos-aquae (L.) Ralfs ex Born. et Flah., in the Lake Chashnitskoe, and Microcystis aeruginosa (Kütz.) Kütz., in the lakes Zaozer'e and Vashutinskoe. In the 2002, cyanobacteria from the genera Microcystis, Aphanizomenon, Oscillatoria, Aphanothece, and Woronichinia dominated (Lyashenko et al., 2002).
The first data on the flora of higher aquatic plants in the studied lakes were obtained in 1902 (Flerov, 1903). During this period, in the Lake Zaozer'e, 6 species of aquatic and coastal aquatic plants (Potamogeton perfoliatus L., Alisma plantago-aquatica L., Sagittaria sagittifolia L., Eleocharis acicularis (L.) Roem. et Schult., E. palustris (L.) Roem. et Schult., and Persicaria amfibia (L.) Delarbre) were indicated, in the Lake Chashnitskoe, 11 species, and in the Lake Ryumniki, 21 species. Further studies of these water bodies were carried out by A.P. Belavskaya only 60 years later, in 1963 (Fortunatov and Moskovskii, 1970). However, these studies were mainly focused on studying the processes of overgrowing of lakes, not the floristic composition. Apparently, it has been noted that the natural communities of aquatic plants have been greatly changed in the Lake Zaozer'e due to long-term pollution by waste from livestock farms. Significant thickets of the great manna grass Glyceria maxima (Hartm.) Holmb. were observed here, in the southwestern part of the lake, there were horsetail-calla floating mats (Fortunatov and Moskovskii, 1970). Later, in 1985, common spike-rush Eleocharis palustris, water smartweed Persicaria amphibia, and the species of the genus Carex were found in addition to the great manna grass (Trusov et al., 1988). Studying the flora of the Lake Chashnitskoe, A.P. Belavskaya reported on 21 species of vascular plants and noted the widespread association of the water horsetail Equisetum fluviatile L. and arrowheads Sagittaria sagittifolia, in the southern and southeastern parts of the lake, the association of the bottle sedge Carexrostrata Stokes and shift-bogbean communities. After 25 years, 22 plant species were indicated in the floristic lists (Trusov et al., 1988). In the Lake Ryumniki, three species were indicated for the first time by A.P. Belavskaya, bulrush Scirpus radicans Schkuhr, clasping-leaved pondweed Potamogeton perfoliatus, and bottle sedge.
New data on the flora of vascular plants appeared almost a hundred years after the first studies, when the survey of the flora of these lakes was carried out in September 2001 by G.F. Lyashenko (Lyashenko et al., 2002). As a result of these studies, 12 species of vascular plants were identified for the Lake Zaozer'e, 19 species, for the Lake Chashnitskoe, 21 species, for the Lake Ryumniki, and 14 species, for the Lake Vashutinskoe. Considering all the studies performed, the flora of vascular plants of the Lake Zaozer'e was presented by about 20 species, of the Lake Chashnitskoe, by 31 species, of the Lake Ryumniki, 40 species, and for the Lake Vashutinskoe, 34 species in total.
The study aims to assess the current state and nature of changes in phytoplankton and the floristic composition of vascular plants of small lakes Zaozer'e, Ryumniki, Chashnitskoe, and Vashutinskoe located in the Rostov District, the Yaroslavl Region, Russia, for more than 50-year period.
Materials and methods
The studies were carried out in mid-July 2018 at four lakes (Zaozer'e, Ryumniki, Chashnitskoe, and Vashutinskoe) located in the Rostov District of the Yaroslavl Region: (Fig. 1).
The Lake Zaozer'e (other names: Zaozerskoe, Pokrovskoe, Monastyrskoe) is located in a closed depression. The lake is drainless. The area of the water surface is 32 hectares, the maximum depth is 10 m, the average depth, 5 m. During the phytoplankton bloom in the lake, the water transparency reaches 10-30 cm, pH, 9.0-9.2, and the electrical conductivity, 27.2-30.0 ^S. The lake has a rounded shape, the shores are gentle. The increased content of alkali metal ions associated with water pollution with organic matter was noted (Fortunatov and Moskovskii, 1970; Grigor'ev, 1903). Regard must be paid to a smaller amount of organic matter having been entered to the lake for a long time, which is primarily due to the closure of livestock farms located along the lake shores. The soils near the coast are sandy, the silt layer over the sandy bottom increases along with the depth increase, the silt has a pungent smell of hydrogen sulfide. The sapropel deposits in the lake are not found (Lyashenko et al., 2002).
The Lake Ryumniki (other names: Ryumnikov-skoe, Ryumnikovo). The area of the water surface is 153 hectares, the maximum depth is 10 m, the average depth, 3 m. The water in the lake is weakly mineralized (16-17 mg/L), sulfate class, pH 6.0-6.9. The water transparency is 1.4-1.6 m. In the years of
Fig. 1. Location of the studied lakes in the Rostov District of the Yaroslavl Region, Russia.
mean moisture regime, the lake is drainless; in high-water years, part of the water flows out to the swamps adjacent to the southwestern coast. The thickness of the sapropel layer in the lake basin ranges from 0.5 to 2.0 m; in the southern part, the sapropel is overlain by a layer of peaty silt up to 3-m thick. The coastal areas along the northern coast and partially along the southern coast are sandy. In the southern part of the lake, the shores are steep, formed by loams and boulders, in the southeastern part of the lake, they are low, gentle. In the northeastern part, there is a swampy backwater; the northwestern, western and southwestern coasts are highland moors with birch forests or wet alder groves (Fortunatov and Moskovskii, 1970; Kuzmichev et al., 1990).
The Lake Chashnitskoe (other names: Chashnitsy, Chashnikovo, Chashniche). The area of the water surface is 54 hectares, the maximum depth of the lake is 11 m, the average depth is 4-5 m. The waters of the lake are slightly mineralized (19.0-23.5 m/L) and are characterized by high transparency. The electrical conductivity of the surface water layer ranges from 36.1 to 43.0 ^S, pH, 6.7-7.6. In dry years, the lake has no drainage; in high-water years, the water from the lake flows through the ravine towards the Lake Karash and further, through the Pashma River, into the. Klyazminskaya Nerl' River. The shores and most of the littoral zone of the lake are composed by sandy-pebble or sandy soils and, to a lesser extent, by peat. The sediments forming the bottom of the lake are also heterogeneous. Thus, in the eastern part of the lake, sandy-silty deposits are widespread, sometimes with an admixture of pebbles and boulders. Near the western and southern coasts, mainly peaty silts are found, overlapping gray sapropel (thickness from 0.10 to 4.5 m) in some areas. In the central part of the lake, clayey sapropel dominates, while in the southern part, the sapropel stratum is covered by a 2-m layer of peaty silt (Fortunatov and Moskovskii, 1970; Grigor'ev, 1903; Kuzmichev et al., 1990; Lyashenko et al., 2002).
The Lake Vashutinskoe (other names: Vashutino). The area of the water surface is 310 hectares, the maximum depth is 6.5 m, the average depth, 3.5 m. The waters of the lake are characterized by sufficiently high mineralization of 100 m/L. The water transparency ranges from 40-60 cm near the shores to 80 cm in the center of the lake; pH is 6.6-7.9; the electrical conductivity of the surface water layer is 58-59 ^S. The lake is open; the Sutyaga River inflows to the lake from the north-west, the Dubets Brook, from the north; the River Kamenka (a tributary of the Klyazminskaya Nerl River) outflows from the southeastern part of the lake. The shores of the lake are swampy in the northern and southern parts. The bottom is formed mainly by peaty silts, overlapping sapropel in some areas (Fortunatov and Moskovskii, 1970; Kuzmichev et al., 1990; Lyashenko et al., 2002).
Hydrophysical and hydrochemical characteristics ofwater (temperature, electrical conductivity, dissolved oxygen) were measured with a multiparameter hand-held portable probe YSI-ProPlus (YSI, USA) in the surface layer (10 cm), the pH was determined using a portable analyzer Combo HI 98129 (Hanna Instruments, Germany), the water transparency was assessed using the Secchi disk.
Integral samples were taken in the pelagial zone and in the shallow waters covered with thickets ofhigher aquatic vegetation in order to determine the biomass and species composition of algae. The collection and processing of phytoplankton samples was carried out according to standard methods (Metodika..., 1975). Taxa with biomass equal to or exceeding 10% of the total biomass were considered dominants. When analyzing the phytoplankton communities of the lakes, the microalgae biomass and the species richness of phytoplankton were considered. The trophic status of the lakes was determined by the phytoplankton biomass in accordance with the classification by S.P. Kitaev (1984, 2007).
The vegetation cover of the lakes was studied according to the methods generally accepted in hydrobotany (Katanskaya, 1981; Shennikov, 1964; etc.). The material was collected by the route method using a rowing boat and going round the lake along the shore. During the field surveys, the taxonomic composition of the flora of the vascular plants of the lakes was taken into account (Papchenkov, 2001), the depth of plant growth and the nature of the soil were noted. When compiling a plant species list of the lake, materials from our own research were used. The species of vascular plants were determined using the taxonomic keys of local flora (Lisitsyna et al., 2009; Maevskii, 2014). Latin names of taxa were verified in accordance with the summary of P.F. Maevskii (2014), recent taxonomy was also taken into account for a number of taxa. For example, the nomenclature of ferns is given here in accordance to M.M. Christenhusz and M.W. Chase (2014), the nomenclature of the genera Ranunculus L. and Lysimachia L., by the APG IV system (Byng et al., 2016; Emadzade et al., 2010; Hao et al., 2004). Taxonomic, ecological, and ecobiomorphological analyzes of the flora were performed in accordance with the recommendations of V.G. Papchenkov (2001).
The plant species lists are presented for each lake (Appendices 1 and 2), these species were those observed in the lake area in mid-July 2018.
Results and discussion
The greatest depth and the lowest water transparency during the study period were registered in the Lake Zaozer'e (Table 1). The highest temperature, dissolved oxygen content, and pH were also recorded here, which was apparently associated with a high biomass of cyanobacteria. It should be noted that the
Table 1. Some abiotic characteristics of small lakes in the Rostov District of the Yaroslavl Region in July 2018. P - pelagial zone, OL - overgrown littoral zone, «-» - no data.
Parameter
Sampling depth, m Water transparency, m Water temperature, °C Dissolved oxygen, mg/L
Oxygen saturation, %
Total dissolved solids, mg/L
Lake
Chashnitskoe Zaozer'e Ryumniki Vashutinskoe
P OL P OL P OL P OL
7 1
21.6
100
1
0.9 21.8
99
9
0.3 22.3
0.4 0.1 23.4
139.7 132
31.9 31.2 29.9 27.3
3.5 1.4 22.2
8.89 8.62 12.40 11.90 8.36
96
0.9
down to the bottom
22.0
7.83
91
17.6 17.6
2.6 0.6 21.4
1.2 0.4 22.1
рН
6.8
6.8
9.0
9.2
6.5
6.9
6.6
7.0
active photosynthesis exactly causes an increase in water pH and an increase in the oxygen concentration in the reservoir during the phytoplankton bloom (Kellar and Paerl, 1980; Paerl and Ustach, 1982). The lowest water transparency was characteristic of the shallowest of the studied lakes, the Lake Vashutinskoe, the highest water transparency in the pelagial and littoral zones was observed in the lakes Ryumniki and Chashnitskoe. The lakes Vashutinskoe, Ryumniki, and Chashnitskoe did not differ significantly in terms of pH and dissolved oxygen content. The lowest mineralization was observed in the Lake Ryumniki, the highest, in the Lake Chashnitskoe.
Algal flora
In total, 137 species of algae belonging to 8 divisions, 18 orders, and 76 genera were identified in the studied lakes of the Rostov District of the Yaroslavl Region. The largest number of species was found in the Lake Vashutinskoe (81 species), followed by the lakes Chashnitskoe and Zaozer'e (69 species each), in the Lake Ryumniki, 65 species were found (Appendix 1). The largest number of species in the pelagial zone was found in the Lake Vashutinskoe (68 species), the lowest, in the Lake Chashnitskoe (50 species). The number of phytoplankton species in the pelagial zone in the lakes Zaozer'e and Ryumniki was 53 and 54 species, respectively. In the shallow waters of the lakes, the number of algal species was lower than that in the deep-water part: 29 species in the Lake Zaozer'e, 37, in the Lake Ryumniki, and 46 species each in the lakes Chashnitskoe and Vashutinskoe.
The lowest phytoplankton biomass in the pelagial zone was recorded in the Lake Ryumniki (0.65 mg/L),
the highest, in the Lake Vashutinskoe (7.13 mg/L) (Fig. 2A). According to the algae biomass, the pelagial zone of the lakes Chashnitskoe and Zaozer'e was characterized by mean values (2.18 mg/L and 3.38 mg/L, respectively). The phytoplankton biomass in the pelagial zone of the lakes turned out to be generally lower than that in the littoral zone overgrown with higher aquatic vegetation (Fig. 2A). The greatest difference was observed in the Lake Zaozer'e, where biomass in the littoral zone was 6 times greater comparing to the pelagial zone, the lowest, in the Lake Ryumniki (1.5 times more in the littoral zone than in the pelagial zone).
Cyanobacteria dominated by biomass in the Lake Chashnitskoe (~ 50% of the total biomass), followed by green algae (15-25%) (Fig. 2B). Bacillariophyta (14%) and Dinophyta (10%) also prevailed in the pelagial zone of the lake, Chrysophyta (13%), in the littoral zone. In the deep-water part of the Lake Chashnitskoe, the dominating species were Snowella lacustris (Chod.) Kom. et Hind. (21%), Anabaena sp. (17%), and Aulacoseira ambigua (Grun.) Sim. (12%); in shallow waters, these were Anabaena sp. (37%), Dinobryon divergens Imhof (12%), and the species of the genus Chlamydomonas Ehrenb. (10%) (Appendix 1).
In the Lake Zaozer'e, cyanobacteria dominated also (91-99%) in the studied biotopes (Fig. 2B). Aphanizomenon flos-aquae prevailed in the pelagial and littoral zones of this water body (84% and 78%, respectively). In shallow waters, Microcystis aeruginosa (18%) was a subdominant (Appendix 1).
The ratio of the dominant algae divisions was different in the pelagial and littoral zones of the Lake Ryumniki (Fig. 2B). In the deep-water part of the lake,
Fig. 2. Phytoplankton biomass (A) and the share of different groups of planktonic algae in regard to the total biomass (B) in different parts of the studied lakes of the Rostov District of the Yaroslavl Region, Russia, in July 2018.
the biomass was dominated by Dynophyta (48%), Chlorophyta (17%), and Cyanobacteria (16%), while in shallow water, Cyanobacteria (63%) and Cryptophyta (19%) prevailed. Ceratium hirundinella (O.F. Müll.) Schrank dominated by biomass (43%) in the pelagial zone, in the littoral zone, these were Aphanizomenon flos-aquae (34%), Anabaena sp. (25%), and Cryptomonas marssonii Skuja (17%) (Appendix 1).
In the Lake Vashutinskoe, the dominant algae divisions were cyanobacteria (53% in the pelagial and 37% in the littoral zones) and diatoms (26% and 52%, respectively) (Fig. 2B). Limnothrix planctonica (Wotosz.) Meff. (30%), Aulacoseira ambigua (19%), and Microcystis aeruginosa (13%) prevailed in the deep-water area, Aulacoseira ambigua (51%) and Limnothrix planctonica (26%), in shallow waters.
Based on the phytoplankton biomass data obtained for the pelagial zone, the Lake Vashutinskoe belongs to the eutrophic water bodies, the lakes Chashnitskoe and Lake Zaozer'e, to mesotrophic, and the Lake Ryumniki, to oligotrophic (Kitaev, 1984, 2007). Shallow areas of the reservoir are characterized by a higher phytoplankton biomass than deep-water ones; according to the trophicity scale, the lakes Chashnitskoe, Zaozer'e, and Vashutinskoe are classified as eutrophic reservoirs. Shallow waters of the Lake Ryumniki have an oligotrophic status. In should be noted that the coastal areas are characterized by higher enrichment with biogenic elements, which probably contributes to the active development of phytoplankton in these zones (Korneva, 1983; Solov'eva and Korneva,
2006). The lowest species richness of phytoplankton has been noted in the shallow waters, which also confirms the advantage of cyanobacteria (whose biomass is higher in the littoral zone than in the pelagial zone) in high rates of nutrient consumption, so they are responsible for the light limitation for other phytoplankton species (Paerl and Huisman, 2009). The study of the processes occurring in the shallow waters is especially important in regard to their role in the ecosystem of the entire water body. It is known that the eutrophication processes beginning in the littoral areas are then observed in the deep-water zones of the reservoir (Sakharova and Korneva, 2018).
The study of the phytoplankton of these lakes was very episodic, the sampling was carried out only in September (Fortunatov and Moskovskii, 1970; Lyashenko et al., 2002); therefore, the available data do not allow us to conclude on the vector of successional changes in the algal community. The predominance of cyanobacteria in all lakes was noted both in our studies and earlier (Fortunatov and Moskovskii, 1970; Grigor'ev, 1903; Lyashenko et al., 2002). Decreasing role of Aphanizomenon flos-aquae and species of the genus Anabaena was reported earlier (Lyashenko et al., 2002), but these algae made a significant contribution to biomass (up to 80%) during our studies. As it was observed in September 2001 (Lyashenko et al., 2002), during our studies, phytoplankton of the Lake Vashutinskoe was similar in composition of the dominant complex (genera Microcystis and Limnothrix) with other shallow eutrophic lakes, for example, the Lake Nero (Babanazarova et al., 2011).
Vascular plants
Currently, the flora of vascular plants of all studied lakes includes 118 species belonging to 4 divisions, 5 classes, 22 orders, and 37 families. A full list of vascular plants is presented in Appendix 2.
The Lake Zaozer'e was characterized by the poorest species richness (50 species; Table 2). This was caused by the anthropogenic disturbance of its vegetation cover, associated with the income of the organic-rich wastewater from two livestock farms, which were previously located on the shores of the lake (Fortunatov and Moskovskii, 1970). It should be noted that the farms have not been operating for a long time; however, the lake still "blooms" strongly in the summer months, and so the water transparency drops sharply and thus inhibits the development of submerged aquatic plants. The oligotrophic Lake Ryumniki was characterized by the richest species composition (82 species; Table 2); apparently, this was due to a significant number of different biotopes (both natural and anthropogenically disturbed).
Cyperaceae Juss and Poaceae Barnhart were the dominant families of the flora of all studied lakes (Table 3). Their share in the total composition of the flora reached 22.0% in the Lake Zaozer'e, about 24.0%, in the Lake Chashnitskoe, 23.1%, in Lake Ryumniki, and 27.3% in the Lake Vashutinskoe. Redard must be paid to a significant share of grasses in the flora of the studied lakes, which can serve as an indicator of an increase of the anthropogenic load (Teteryuk, 2012). In the Lake Ryumniki, Ranunculaceae Juss may also be included into the list of dominant families. The floras of the lakes Chashnitskoe, Ryumniki, and Vashutinskoe were characterized by a pronounced distinctness along with the common leading genera Carex L., Persicaria (L.) Mill., and Potamogeton L. (Table 3). Thus, Juncus L. and Sparganium L. were the leading genera in the Lake Chashnitskoe, in addition to the genera listed above, Salix L. and Ranunculus L., in the Lake Ryumniki, and Calamagrostis Adans, in the Lake Vashutinskoe.
When considering ecological and coenotic structure, hygrophytes were the leading group of
Table 2. The taxonomic structure of the flora of small lakes in the Rostov District of the Yaroslavl Region in July 2018.
Number of taxonomic groups Chashnitskoe Zaozer'e Lake Ryumniki Vashutinskoe
Species 71 50 82 76
Genera 51 37 59 56
Families 30 24 37 31
Orders 21 16 23 23
Classes 4 4 5 3
Divisions 2 2 3 2
Table 3. Contribution of the dominant families and genera of vascular plants to the species richness of the flora of small lakes in the Rostov District of the Yaroslavl Region in July 2018.
Lake Family Contribution to the species richness, % Genus Contribution to the species richness, %
Cyperaceae 12.6 Carex 5.6
Poaceae 11.3
Chashnitskoe Araceae, Lamiaceae, Polygonaceae, Typhaceae 5.6 each Juncus, Persicaria, Potamogeton, Sparganium 4.2 each
Cyperaceae 12.0
Poaceae 10.0
Zaozer'e Typhaceae Araceae, Lamiaceae, Rubiaceae 8.0 6.0 each Sparganium, Carex, Galium 6.0 each
Poaceae 13.4 Carex 6.1
Cyperaceae 9.7 Salix, Ranunculus 5.0 each
Ryumniki Ranunculaceae Araceae, Lamiaceae, Salicaceae 6.1 5.0 each Potamogeton, Persicaria 3.6 each
Poaceae 14.3 Carex 7.8
Vashutinskoe Cyperaceae Araceae, Lamiaceae, Polygonaceae 13.0 5.2 each Calamagrostis, Potamogeton, Persicaria ~ 4.0 each
the flora of each of the studied lakes: 44.0% in the Lake Zaozer'e, 39.4% in the Lake Chashnitskoe, 39.0% in the Lake Ryumniki, and 36.8% in the Lake Vashutinskoe. The second place belonged to hygrohelophytes (24.0% each in the lakes Zaozer'e and Chashnitskoe, 25.0% in the Lake Vashutinskoe), followed by hydrophytes (14.0%, in the Lake Zaozer'e, 17.0% in the Lake Chashnitskoe, and 19.7% in the Lake Vashutinskoe). It should be noted that in the Lake Ryumniki, in contrast to the other lakes, hydrophytes (22.0%) dominated over hygrohelophytes (20.7%). The share of helophytes, as usual, was much lower: 16.0% in the Lake Zaozer'e, 15.6% in the Lake Chashnitskoe, 12.0% in the Lake Vashutinskoe, and 7.4% in the Lake Ryumniki. Mesophytes were either absent (the Lake Zaozer'e), or are present by no more than one or two species (2.4% in the Lake Ryumniki, 1.4% in the Lake Chashnitskoe, and 1.3% in the Lake Vashutinskoe).
The flora of small lakes was traditionally dominated by herbaceous plants; their contribution to the species richness of each particular lake ranged as 90.2-96.0%. Perennials and annual / biennial plants formed the basis of the vegetation cover of lakes (89.4-94.7%). This group also included polycarpous and sporulating species (Isoetes echinospora Durieu,
Equisetum fluviatile, and Thelypteris palustris Schott). Annuals made up to 5.3-7.8%. It is interesting that in the Lake Ryumniki annual plants were represented not by 3-4 species (as in other lakes), but by 8 species (9.7% of the lake flora).
The rare and protected species of the Yaroslavl Region (Krasnaya Kniga..., 2015) were represented by Nuphar pumila (Timm) DC. in the Lake Vashutinskoe and Isoetes echinospora in the Lake Ryumniki. Sparganium gramineum Georgi was recorded in all studied water bodies.
Thirty-four plant species were found in all the studied water bodies, of which 4 were hydrophytes: Elodea canadensis Michx., Lemna minor L., Spirodela polyrhiza (L.) Schleid., Sparganium gramineum; six species were helophytes (Equisetum fluviatile L., Alisma plantago-aquatica, Sagittaria sagittifolia, Glyceria maxima, Phragmites australis (Cav.) Trin. ex Steud., and Typha latifolia L.), and two species were hydrohelophytes (Eleocharis acicularis and E. palustris).
Since 1950s, I. echinospora, Elatine hydropiper L., and E. triandra Schkuhr. have disappeared from the flora of the Lake Chashnitskoe; in the Lake Zaozer'e, these are Persicaria amphibia, and Potamogeton
perfoliatus; in the Lake Rymniki - Elatine callitrichoides (W. Hyl.) Kauffm., Nupharpumila, Carex elongata L., Sparganium natans L., and Potamogeton lucens L.; in the Lake Vashutinskoe - I. echinospora, Potamogeton berchtoldii Fieb., P. natans L., Caulinia flexilis Willd., Zizaniapalustris L., Carexcaespitosa L., Ranunculus trichophyllum Chaix, R. reptans L., Elatine hydropiper, Callitriche hermaphroditica L., and Galium trifidum L.
The disappearance of Isoetes echinospora from the flora of the lakes Chashnitskoe and Vashutinskoe could have occurred during the period from 19651972 to 1985; apparently, it is associated with an increase in the trophic status of these water bodies (Fortunatov and Moskovskii, 1970; Lyashenko et al., 2002). Currently, this species has survived only in the oligotrophic Lake Ryumniki, where the state of its population can be considered satisfactory. I. echinospora grows here along the southern and southeastern coasts at the depths from 0.15 to 1.0 m; the sediments are sandy, less often sandy with thin deposit of spring floods. It is found as single specimens and does not form clusters or thickets. It should be noted that the data on the growth of I. lacustris L. in the Yaroslavl Region, given earlier for the lakes Ryumniki, Vashutinskoe, and Chashnitskoe, are erroneous (Garin, 2015).
Sparganium x longifolium Turcz. ex Ledeb., which is a hybrid between the boreal-montane Eurasian species Sparganium gramineum, preferring ultra-fresh water bodies with a nearly neutral pH, and S. emersum Rehm., a plurizonal Holarctic species with a wide ecological amplitude (Belyakov et al., 2017), registered during our studies in the lakes Chashnitskoe and Zaozer'e, is an indicator of an increase in the degree of trophicity as a result of vigorous economic activity (Fortunatov and Moskovskii, 1970; Lyashenko et al., 2002).
Active water blooming (transparency of 10 cm and less) in the summer in the Lake Zaozer'e was apparently a consequence of the extremely low number of submerged hydrophytes found here. Thus, the Potamogeton perfoliatus and Persicaria amphibia previously noted here were not found in the lake in 2001 (Lyashenko et al., 2002), although in the early summer period, when the water in the lake is still clear, Elatine hydropiper, Potamogeton crispus L., and Ceratophyllum demersum L. are found here. It should be noted that the last two species are the indicators of the eutrophication of the reservoir. In addition, both parental species of bur-reed and their hybrid co-grow on this lake. The alternating groups of Sparganium gramineum and S. x longifolium are located in a narrow strip along the northern and northwestern shores of the lake, while S. emersum is distributed exclusively along the southern coast. In the future, this situation may lead to the loss of S. gramineum from the flora of the lake.
Due to eutrophication and active overgrowth of the north-eastern shore of the Lake Vashutinskoe, Nuphar pumila (Timm) DC are endangered, as well as a single small clump of Sparganium gramineum (no more than 1 m in diameter); in addition, the appearance of Stratiotes aloides L., an indicator species of eutrophication, in this lake raises concerns. Absence of Chara strigosa A. Br. in 2018, a relict species listed in the Red Data Book of the Yaroslavl Region (2015), is another indicator of the eutrophication processes in the Lake Vashutinskoe. Chara strigosa is a stenothermal cold-water freshwater, predominantly light-loving plant that is considered an indicator of oligotrophic waters (Langangen and Mjelde, 2010; Romanov et al., 2014). For the first time, this species was noted in the Lake Vashutinskoe in 1972, and the last time it was found here in 2017 in the form of a few and strongly suppressed shoots.
The saturation of the flora of lakes with adventive species is extremely low. Elodea canadensis, Bidens frondosa L., and Epilobium adenocaulon Hausskn were noted in all studied lakes. Zizania latifolia Griseb. and Juncus tenuis Willd. were also registered in the lake Vashutinskoe. The high adaptive potential of these plants allows them to colonize quickly and hold successfully their niches in phytocenoses. It should be noted that E. canadensis, apparently, could appear in the studied lakes not earlier than in the 1980s and currently does not form significant thickets in the water bodies. Cultivation of the Far Eastern perennial Zizania latifolia and the Canadian annual plant Z. palustris in the Lake Vashutinskoe was associated with the presence of a hunting farm on its shore, which is still functioning. Apparently, these species were planted along the Lake Vashutinskoe as early as in 1960s and 1970s in order to increase the food supply and improve the habitat conditions for semiaquatic animals (Lisitsyna et al., 2009; Maltseva and Bobrov, 2017). Two more North American invasive species, Epilobium adenocaulon and Bidens frondosa, are now widely distributed in all areas of the middle zone of the European part of Russia (Lisitsyna et al., 2009; Maevskii, 2014).
Conclusions
In the studied water bodies, 137 species of algae belonging to 8 divisions, 18 orders, and 76 genera were identified. The algoflora of the Lake Vashutinskoe is the richest (81 species), followed by the lakes Chashnitskoe and Zaozer'e (69 species each), 65 species inhabit the Lake Ryumniki. Despite the fact that phytoplankton of the studied water bodies is known only from episodic observations, cyanobacteria predominate in the lakes. A significant contribution to the biomass of Aphanizomenon flos-aquae and species of the genus Anabaena was noted in the lakes Zaozer'e, Chashnitskoe, and Ryumniki, in the Lake Vashutinskoe, they were Limnothrix planctonica and Microcystis aeruginosa. According to the phytoplankton biomass of
the pelagial zone, the Lake Vashutinskoe is classified as eutrophic water body, the lakes Chashnitskoe and Zaozer'e, as mesotrophic, and the Lake Ryumniki, as oligotrophic. By trophicity, shallow waters of the lakes Chashnitskoe, Zaozer'e, and Vashutinskoe belong to the eutrophic water bodies, while the littoral zone of the Lake Ryumniki has an oligotrophic status. The irregular nature of the observations carried out in different years and periods of the growing season does not allow us to conclude on the vector of the successional changes in the algal flora.
The total list of vascular plants of the studied lakes includes 118 species belonging to 4 divisions, 5 classes, 22 orders, and 37 families; 34 species are common for all studied lakes. The Lake Ryumniki is the richest by the species composition, the Lake Zaozer'e, the poorest. In general, the differences in the taxonomic composition of the flora of certain lakes are associated with both the diversity of biotopes and the different level of conservation of the vegetation cover, as well as with the degree of eutrophication of a particular water body. The low species richness of the flora of the Lake Zaozer'e is mainly due to the monotonous nature of the coastline and the active blooming of water in summer. In accordance with the ecological-coenotic characteristics of the flora of the lakes, the group of hygrophytes is traditionally the largest in number of species, followed by hygrohelophytes, and then hydrophytes. In the oligotrophic Lake Ryumniki, the representatives of the ecological group of hydrophytes prevail over hygrohelophytes. The prevalence of herbaceous plants found in the studied lakes is natural and characteristic of aquatic flora.After 1950s, two species have disappeared from the Lake Zaozer'e, three species, from the Lake Chashnitskoe, five species, from the Lake Ryumniki, and eleven species, from the Lake Vashutinskoe. For example, I. echinospora was not found after 1965 in the Lake Chashnitskoe and after 1972 in the Lake Vashutinskoe. Caulinia flexilis was not found in the Lake Vashutinskoe since 1972. Potamogeton crispus, Sparganium x longifolium, Ceratophyllum demersum, Elodea canadensis, Stratiotes aloides and some other species are among the main indicators of eutrophic waters observed in the studied lakes. The saturation of the flora of lakes with adventive species is low: only two adventive species was registered in the Lake Chashnitskoe over the entire period of research, five species, in the Lake Ryumniki, and ten species, in the Lake Vashutinskoe.
Acknowledgments
The work was carried out within the framework of the State Tasks nos. AAAA-A18-118012690096-1 and AAAA-A18-118012690099-2. Special thanks go to I.P. Samakov, the Head of the Pereslavl hunting farm, V.S. Gal'tsov, chief hunting expert, and other employees of the hunting farm for invaluable support during the studies on the Lake Vashutinskoe.
References
Babanazarova, O.V., Sidelev, S.I., Aleksandri-na, E.M., Sakharova, E.G., Kurmayer, R., 2011. Phytoplankton structure and microcystine concentration in the highly eutrophic Nero Lake. Water Resources 2, 229-236. https://doi.org/10.1134/ S0097807811020023
Belyakov, E.A., Garin, E.V., 2018. Long-term dynamics of flora of karst lakes: Changes and current state. Biosystems Diversity 26 (2), 160-169. https://doi.org/10.15421/011825
Belyakov, E.A., Shcherbakov, A.V., Lapirov, A.G., Shilov, M.P., 2017. Morfologiya i ekologicheskie osobennosti Sparganium x longifolium (Typhace-ae) v centre evropeiskoy chasti Rossii [Morphology and ecological characteristics of Sparganium x longifolium (Typhaceae) in the central part of the European Russia]. Biosystems Diversity 25 (2), 154-161. doi:10.15421/011723. (In Russian).
Blokhontsev, E.N., 1903a. O fitoplanktone nekoto-rykh ozer Rostovskogo uezda Yaroslavskoi gu-bernii i dvukh ozer Vladimirskoi gubernii [About phytoplankton of some lakes of the Rostov Uezd (parish) of the Yaroslavl Guberniya (government) and two lakes of the Vladimir Guberniya]. Trudy Saratovskogo obshchestva estestvoispytatelej i lyubitelej prirody [Proceedings of the Saratov Society of Naturalists and Nature Enthusiasts] 4 (2), 253-269. (In Russian).
Blokhontsev, E.N., 1903b. Fitoplankton Rostovskikh ozer [Phytoplankton of the lakes Rostovskie]. Zem-levedenie [Earth Sciences] 10 (4), 47-54. (In Russian).
Byng, J.W., Chase, M.W., Christenhusz, M.J.M., Fay, M.F., Judd, W.S. et al., 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181 (1), 1-20. https://doi.org/10.1111/boj.12385
Christenhusz, M.M., Chase, M.W., 2014. Trends and concepts in fern classification. Annals of Botany 113 (4), 571-594. https://doi.org/10.1093/aob/mct299
Emadzade, K., Lehnebach, C., Lockhart, P., Horandl, E., 2010. A molecular phylogeny, morphology and classification of genera of Ranuncu-leae (Ranunculaceae). Taxon 59 (3), 809-828. https://doi.org/10.2307/25677670
Flerov, A.F., 1903. Botaniko-geograficheskie ocher-ki. III. Rostovskii krai [Botanical and geographical
essays. III. Rostov Region]. Zemlevedenie [Earth Sciences] 10 (2-3), 193-218. (In Russian).
Fortunatov, M.A., Moskovskii, B.D., 1970. Ozera Yaroslavskoi oblasti: Kadastrovoe opisanie i krat-kie limnologicheskie kharakteristiki [Lakes of the Yaroslavl Region: Cadastral description and brief limnological characteristics]. In: Fortunatov, M.A. (ed.), Ozera Yaroslavskoi oblasti i perspektivy ikh khozyaistvennogo ispol'zovaniya [Lakes of the Yaroslavl Region and prospects for their economic use]. Yaroslavl State Pedagogical University, Yaroslavl, USSR, 3-177. (In Russian).
Garin, E.V., 2015. Sosudistye sporovye rasteniya flory Yaroslavskoi oblasti [Vascular cryptogames of the flora of the Yaroslavl Region]. In: Lapi-rov, A.G. (ed.), Horizons of Hydrobotany. Trudy IBVV RAN. Vyp. 71 (74) [Transactions of the I.D. Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences. Issue 71 (74)]. Filigran', Yaroslavl', Russia, 48-59. (In Russian).
Grigor'ev, S., 1903. Ozera Rostovskogo uezda [Lakes of the Rostov Uezd]. Zemlevedenie [Earth Sciences] 10 (2-3), 163-192. (In Russian).
Hao, G., Yuan, Y.-M., Hu, C.-M., Ge, X.-J., Zhao, N.-X., 2004. Molecular phylogeny of Lysimachia (Myrsinaceae) based on chloroplast trnL - F and nuclear ribosomal ITS sequences. Molecular Phy-logenetics and Evolution 31, 323-339. https://doi. org/10.1016/S1055-7903(03)00286-0
Izmailova, A.V., Rumyantsev, V.A., Drabkova, V.G., 2017. Limnologicheskaya izuchennost evropeiskoy chasti Rossii v svete problem antropogennoi modi-fikatsii ozernykh ekosistem [Limnological study of the European part of Russia in regard to the issue of anthropogenic modification of lake ecosystems]. Izvestiya Russkogo geograficheskogo obchshest-va [Bulletin of the Russian Geographical Society] 149 (6), 1-13. (In Russian).
Katanskaya, V.M., 1981. Vysshaya vodnaya rasti-tel'nost' kontinental'nyh vodoemov SSSR. Metody izucheniya [Higher aquatic vegetation of the continental water bodies of the USSR. Study methods]. Nauka, Leningrad, USSR, 187 p. (In Russian).
Kellar, P.E., Paerl, H.W., 1980. Physiological adaptations in response to environmental stress during an N2-fixing Anabaena bloom. Applied and Environmental Microbiology 40, 587-595.
Kitaev, S.P., 1984. Ekologicheskie osnovy bioproduk-tivnosti ozer razlichnykh pririodnykh zon [Ecological
basis of bioproductivity of lakes of various natural zones]. Nauka, Moscow, USSR, 208 p. (In Russian).
Kitaev, S.P., 2007. Osnovy limnologii dlya gidrobi-ologov i ikhtiologov [Fundamentals of Limnology for Hydrobiologists and Ichthyologists]. Karelian Scientific Center, Russian Academy of Sciences, Petrozavodsk, Russia, 395 p. (In Russian)
Korneva, L.G., 1983. Planktonnye algotsenozy pribrezhiya Rybinskogo vodokhranilichsha [Plankton algocoenoses of the coastal areas of the Rybinsk Reservoir]. In: Monakov , A.V., Presnovod-nye gidrobionti i ikh biologiya [Freshwater aquatic organisms and their biology]. Nauka, Leningrad, Russia, 38-51. (In Russian).
Krasnaya kniga Yaroslavskoy oblasti [Red Data book of the Yaroslavl Region], 2015. Academiya-76, Yaroslavl, Russia, 472 p. (In Russian).
Kuz'michev, A.I., Ekzertsev, V.A., Lisitsyna, L.I., Dovb-nya, I.V., Trusov, B.A. el al., 1990. Flora i rasti-tel'nost' ozer Yaroslavskoi oblasti [Flora and vegetation of the lakes in the Yaroslavl Region]. In: Ekzertsev, V.A. (ed.), Flora i produktivnost' pelagicheskikh i litoral'nykh fitotsenozov vodoemov basseina Volgi [Flora and productivity of pelagic and littoral phyto-cenoses of the water bodies of the Volga River basin]. Nauka, Leninrgad, USSR, 272 p. (In Russian).
Langangen, A., Mjelde, M., 2010. Handlingsplan for kalksj0er - Faktaark for viktige arter av kransalger og tj0nnaks. Milj0vernavdelingen, Fylkesmannen i Oppland, Lillehammer, Norway, 39 p.
Lawniczak-Malinska, A.E., Achtenberg, K., 2018. On the use of macrophytes to maintain functionality of overgrown lowland lakes. Ecological Engineering 113, 52-60. https://doi.org/10.1016Zj.eco-leng.2018.02.003
Lisitsyna, L.I., Papchenkov, V.G., Artemenko, V.I., 2009. Flora vodoemov Volzhskogo basseina. Opre-delitel' sosudistykh rastenii [Flora of the water bodies of the Volga River basin. Taxonimic keys for vascular plants]. KMK, Moscow, Russia, 219 p. (In Russian).
Lobunicheva, E.V., Borisov, M.Ya., Filonenko, I.V., Filippov, D.A., 2013. Otsenka ekologicheskogo sostoyaniya malykh vodoemov. Uchebnoe posobie [Environmental assessment of small water bodies. A tutorial]. State Research Institute on Lake and River Fisheries (FGBNU GosNIORKh), Vologda, Russia, 218 p. (In Russian).
Lyashenko, G.F., Lazareva, V.I., Lyashenko, O.A., 2002. Dinamika vysshei vodnoi rastitel'nosti i plank-
tona v malykh ozerakh basseina Verkhnei Volgi [Dynamics of higher aquatic vegetation and plankton in small lakes of the upper Volga River basin]. In: Papchenkov, V.G. (ed.), Dinamika raznoobrazi-ya gidrobiontov vo vnutrennikh vodoemakh Rossii [Dynamics of diversity of aquatic organisms in the inland waters of Russia]. Yaroslavl State Technical University, Yaroslavl, Russia, 34-57. (In Russian).
Maevskii, P.F., 2014. Flora srednei polosy evropeiskoi chasti Rossii [Flora of the temperate climatic zone of the European part of Russia]. 11th edition. KMK, Moscow, Russia, 635 p. (In Russian).
Mal'tseva, S.Y., Bobrov, A.A., 2017. Alien species of vascular plants in the Rybinsk Reservoir (Upper Volga, Russia). Russian Journal of Biological Invasions 8 (4), 321-326. https://doi.org/10.1134/ S2075111717040063
Maemets, H., Palmik, K., Haldna, M., Sudnitsyna, D., Melnik, M., 2010. Eutrophication and macrophyte species richness in the large shallow North-European Lake Peipsi. Aquatic Botany 92 (4), 273-280. https://doi.org/10.1016Zj.aquabot.2010.01.008
Metodika izucheniya biogeotsenozov vnutrennikh vodoemov [Method of Analysis of Biogeocenosises of Inland Water Bodies], 1975. Mordukhai-Boltovskoy, F.D. (ed.), Nauka, Moscow, USSR, 240 p. (In Russian).
Paerl, H.W., Huisman, J., 2009. Climate change: a catalyst for global expansion of harmful cyano-bacterial blooms. Environmental Microbiology Reports 1 (1), 27-37. https://doi.org/10.1111/j.1758-2229.2008.00004.x
Paerl, H.W., Ustach, J.F., 1982. Blue-green algal scums: an explanation for their occurrence during freshwater blooms. Limnology and Oceanography 27, 212-217.
Papchenkov, V.G., 2001. Rastitelnyi pokrovvodoemov
1 vodotokov Srednego Povolzh'ya [Vegetation cover of the water bodies and watercourses of the middle Volga River region]. International Academy of Business and New Technologies, Yaroslavl, Russia, 214 p. (In Russian).
Postanovlenie pravitel'stva Yaroslavskoi oblasti ot
2 noyabrya 2017 goda N 823-p "Ob utverzhdenii rezhima osoboy okhrany territorii pamyatnikov prirody regionalnogo znacheniya Yaroslavskoi oblasti" [Decree of the Government of the Yaroslavl Region of November 2, 2017, no. 823-p "On approval of the special protection regime
for the territory of nature monuments of regional significance of the Yaroslavl Region"]. (In Russian).
Romanov, R.E., Chemeris, E.V., Vishnyakov, V.S., Chepinoga, V.V., Azovskii, M.G., Kuklin, A.P., Timofeeva, V.V., 2014. Chara strigosa (Streptophyta: Charales) v Rossii [Chara strigosa (Streptophyta: Charales) in Russia]. Botanicheskiy Jurnal [Botanical Journal] 99 (10), 1148-1161. (In Russian). https:// doi.org/10.1134/S1234567814100097
Sakharova, E.G., Korneva, L.G., 2018. Phytoplank-ton in the littoral and pelagial zones of the rybinsk reservoir in years with different temperature and water-level regimes. Inland Water Biology 1, 6-12. https://doi.org/10.1134/S1995082918010157
Shennikov, A.P., 1964, Vvedenie v geobotaniku [Introduction to geobotany]. Leningrad State University, Leningrad, USSR, 447 p. (In Russian).
Slukovskii, Z.I., Shelekhova, T.S., Syroezhko, E.V., 2018. Otklik diatomovoy flory malogo ozera na vozdeistviye tyazhelykh metallov v usloviyakh urbanizirovannoi sredy Respubliki Karelii [A response of diatoms from the small lake on heavy metals effect in an urban environment, Republic of Karelia]. Vestnik Sankt-Peterburgskogo universiteta. Nauki o Zemle. [Vestnik of Saint Petersburg University. Earth Sciences] 63 (1), 103-123. (In Russian). https://www.doi. org/10.21638/11701/spbu07.2018.106
Solov'eva, V.V., Korneva, L.G., 2006. Struktura i dinamika fitoplanktona melkovodii i pelagiali Volzhs-kogo plesa Rybinskogo vodokhranilishcha [Structure and dynamics of phytoplankton of shallow waters and pelagic zone of the Volga Reach of the Rybinsk Reservoir]. Biologiya vnutrennikh vod [Inland Water Biology] 4, 34-41. (In Russian).
Strakhovenko, V.D., Kabannik, V.G., Malikova, I.N., 2014. Geochemical features of the ecosystem of the Lake Kolyvanskoe (Altai Krai) and the effect of the technogenic impact. Lithology and Mineral Resources 3, 213-227.
Teteryuk, B.Y., 2012. Flora i rastitelnost' drevnikh ozer evropeiskogo Severo-Vostoka Rossii [Flora and vegetation of ancient lakes of the European north-east part of Russia]. Nauka, Saint-Petersburg, Russia, 237 p. (In Russian).
Trusov, B.A., Lapirov, A.G., Lyashenko, G.F., 1988. O zarastanii nekotorykh ozer Yaroslavskoi oblasti [About the overgrowth of some lakes in the Yaroslavl Region]. Biologiya vnutrennikh vod [Inland Water Biology] 78, 17-21. (In Russian).
SUPPLEMENTARY MATERIALS
Appendix 1
Taxonomic composition of phytopiankton of small lakes of the Rostov District of the Yaroslavl Region in July 2018 . 1 - presence of the taxon, 0 - absence P - pelagial zone,
OL - overgrown littoral zone
Lake
Taxon Chashnitskoe Zaozer'e Ryumniki Vashutinskoe
P OL P OL P OL P OL
DIVISION CYANOBACTERIA
Order Chroococcales
Aphanocapsa holsatica (Lemmermann) Cronberg et Komârek 1 1 1 0 1 1 1 1
A. incerta (Lemmermann) Cronberg et Komârek 0 0 1 0 1 0 1 0
Aphanothece clathrata W. et G.S. West f. Clathrata 0 0 0 0 1 0 0
A. clathrata f. brevis (Bachmann) Elenkin 1 1 0 1 1 1 1 1
Chroococcus limneticus Lemmermann 1 1 1 1 1 1 1 1
Merismopedia marssonii Lemmermann 0 0 0 0 0 1 1
M. tenuissima Lemmermann 0 0 0 0 1 1 0 0
Microcystis aeruginosa (Kützing) Kützing 1 1 1 1 1 1 1 1
M. flos-aquae (Wittrock) Kirchner 0 0 0 1 0 0 0 0
M. pulverea (H.C. Wood) Forti 1 0 0 0 0 0 1 0
M. smithii Komârek et Anagnostidis 0 1 0 1 0 0 0 0
M. wesenbergii (Komârek) Komârek 1 1 1 1 0 0 0 0
Snowella lacustris (Chodat) Komârek et Hindâk 1 1 0 0 0 0 1 1
DD CD
Qi TT
o
_<
m
>
CD
ro o ro o
m
o
0
1
Co
cB-
S1
Co
S1
o
CO
cn I
o
Order Nostocales
Anabaena flos-aquae Brébisson ex Bornet et Flauhault 0 1 1 0 0 0 0 0
/A. scheremetievi Elenkin 0 0 0 0 0 0 1 0
Anabaena sp. 1 1 1 1 1 1 0 1
Aphanizomenon flos-aquae ( Linnaeus) Ralfs ex Bornet et Flahault 1 0 1 1 1 1 0 0
/A. klebahnii Elenkin ex Pechar 0 0 0 0 0 0 1 1
ro
Lake ro 00
Taxon Chashnitskoe Zaozer'e Ryumniki Vashutinskoe
P OL P OL P OL P OL
Order Oscillatoriales
Limnothrix planctónica (Wotoszyñska) Meffert 0 0 1 0 0 0 1 1
L. redekeiiVan Goor 0 0 0 0 0 0 1 1 DD CD
Lyngbya limnetica Lemmermann 0 1 0 0 0 0 1 1 03
Oscillatoria sp. 1 0 0 0 0 0 0 0 O
Planktothrix agardhii Gomont 0 0 0 0 0 0 1 0 m >
Pseudonabaena limnetica Lemmermann 1 0 1 1 0 0 1 1 CD <—H
DIVISION CHRYSOPHYTA Order Chromulinales
Chromulina sp. 0 0 0 0 1 0 1 0
Chrysococcus rufescens G.A. Klebs 1 1 1 0 1 1 1 1
Chrysococcus sp. 0 0 0 1 0 0 0 0
Dinobryon bavaricum Imhof 0 0 0 0 0 0 1 1
D. divergens Imhof 1 1 0 0 0 0 0 0
Order Synurales
Mallomonas sp. 0 0 0 0 0 0 0 1
Synura sp. 1 1 0 0 1 0 1 0
DIVISION BACILLARIOPHYTA
Order Thalassiosirales
Cyclotella comta Kützing 0 0 0 1 0 0 0 0
C. meneghiniana Kützing 0 1 0 0 0 0 0 1
Cyclotella sp. 1 0 1 0 1 0 1 0
Stephanodiscus invisitatus Hohn et Hellerman 0 0 0 0 0 0 1 0
Order Melosirales
Aulacoseira ambigua (Grunow) Simonsen 1 1 0 0 1 0 1 1
ro o ro o
m
o
0
1
Co
cB-S
31
Co
S1
i
cu
o'
CJ
cn I
o
Order Araphales
Asterionella formosa Hassal 0 0 1 0 1 0 1 0
Diatom a tenuis Aga rd h 0 0 0 0 0 0 1 0 DD CD
Flagilaria capucina Desmazières 1 0 0 0 1 0 0 1 03
F. crotonensis Kitton 0 1 0 0 0 0 1 0 o _<
Tabellaría fenestrata (Lyngbye) Kützing 1 0 0 0 0 0 0 0 m >
Ulnaria acus (Kützing) M. Aboal 0 1 0 0 0 0 1 1 CD <—H ro o ro o
U. ulna (Nitzsch) P. Compère 0 0 0 0 0 0 1 1
Order Raphales
Achnanthes sp. 0 0 0 0 1 0 0 0 m
Amphora sp. 0 1 1 0 0 0 0 0 o o %
Cymbella sp. 0 0 0 0 0 0 1 0 CO cB- s
Navícula sp. 0 0 1 0 1 0 1 0 3 ^
Nitzschia acicularis (Kützing) W. Smith 1 0 1 0 0 0 1 0 QJ 3 Co
N. palea (Kützing) W. Smith 0 0 1 0 0 0 1 1 S1
Nitzschia sp. 0 0 0 0 0 1 0 0 cu o'
DIVISION XANTHOPHYTA 3 M
Order Heterococcales ^
Goniochloris fallax Fott 0 0 0 0 0 0 0 1 en 1
G. parvula Pascher 0 0 0 0 0 1 0 0 o
Ophiocytium capitatum Wolle 0 1 0 0 0 0 0 0
Tetraedriella spinigera Skuja 1 0 0 0 0 0 1 0
DIVISION CRYPTOPHYTA
Order Cryptomonadales
Chroomonas acuta Utermöhl 1 1 0 1 1 1 1 1 ro
CO
Chroomonas sp. 0 1 0 1 0 1 1 0
Cryptomonas marssonii Skuja 1 1 0 1 1 1 1 1
C. ovata Ehrenberg 1 1 0 1 1 1 1 1
C. curvata Ehrenberg 1 1 1 0 0 1 1 1
Cryptomonas sp. 1 0 0 0 0 0 1 1
Rhodomonas lens Pascher 0 1 1 0 1 0 1 0
DIVISION DINOPHYTA
Order Peridiniales
Ceratium hirundinella (O.F. Müller) Schrank 0 0 0 0 1 0 1 0
Gymnodinium sp. 1 0 1 0 0 0 1 0
Peridiniopsis kevei Grigorszky et Vasas 1 0 0 0 0 0 0 0
Perediniopsis sp. 0 1 0 0 1 1 0 0
DIVISION CHLOROPHYTA
Order Chlamydomonadales
Chlamydomonas monadina Stein 1 1 0 0 1 0 0 0
Chlamydomonas spp. 1 1 1 1 1 1 1 1
Order Chlorococcales
Ankistrodesmus bibraianus (Reinsch) Korschikov 1 1 1 0 1 1 1 0
A. falcatus (Corda) Ralfs 0 0 1 0 0 1 0 1
A. fusiformis Corda 0 0 0 0 1 0 1 0
Ankyra viridis (Massjuk) Tzarenko 0 0 1 1 0 0 0 0
Closteriopsis acicularis (G.M. Smith) Belcher et Swale 0 1 0 0 1 1 1 1
Coelastrum microporum Nägeli 0 1 0 0 0 0 0 0
Coenococcus planctonicus Korshikov 0 0 0 0 0 1 0 0
Crucigenia tetrapedia (Kirchner) W. et G.S. West 1 0 0 0 1 1 0 0
Taxon Chashnitskoe Zaozer'e Ryumniki Vashutinskoe
P OL P OL P OL P OL
Crucigeniella apiculata (Lemmermann) Komárek 0 0 0 1 0 0 1 0
C. rectangularis (Nägeli) Komárek 0 0 0 0 1 0 0 0
Dictyosphaerium chlorelloides (Naumann) Komárek et Perman 0 0 1 0 0 0 0 0
D. pulchellum Wood 1 1 1 0 0 1 1 0
D. tetrachotomum Printz 1 1 0 0 1 1 0 1
Didymocystis planctónica Korshikov 1 1 1 0 0 0 1 1
Golenkinia radiata C hod at 0 0 0 0 0 0 1 0
Golenkiniopsis parvula (Woronichin) Korshikov 0 0 0 0 0 0 0 1
Kirchneriella contorta (Schmidle) Bohlin var. elegans (Playfair) Komárek 0 0 0 0 1 1 0 0
K. irregularis (G.S. Smith) Korschikov 0 0 0 0 0 0 1 0
K. obesa (W. West) Schmidle 0 0 1 0 0 0 0 0
Lagerheimia ciliata (Lagerheim) C hod at 1 0 0 0 0 0 0 0
L. genevensis (Chodat) Chodat 0 0 1 0 0 0 0 0
L. subsalsa Lemmermann 0 0 0 0 0 0 0 1
Monoraphidium arcuatum (Korschikov) Hindák 1 0 1 0 0 0 1 1
M. contortum (Thuret) Komárkova-Legnerová 0 0 1 1 1 0 1 1
M. komarkovae Nygaard 1 1 0 0 1 1 0 0
M. minutum (Nägeli) Komárkova-Legnerová 0 0 1 0 0 0 0
M. tortile (W. et G.S. West) Komárkova-Legnerová 1 1 0 0 1 1 0 0
Oocytis borgei Snow 0 1 0 0 1 0 0 0
0. elliptica West 0 0 0 0 1 0 0
0. lacustris Chodat 1 1 0 0 1 1 0 0
Oocytis sp. 0 0 1 0 1 0 1 1
Pediastrum boryanum (Turpin) Meneghini var. boryanum 0 0 1 1 0 0 0 0
P. duplex Meyen 0 0 0 1 0 0 1 0
R tetras (Ehrenberg) Ralfs 0 1 1 1 1 0 0 0
Scenedesmus arcuatus Meyen 0 0 1 1 1 0 1 0
S. bicaudatus Dedussenko 0 0 1 0 1 0 0 0
S. denticulatus Lagerheim var. linearis Hansgirg 0 0 1 0 0 0 0 0
S. ellipticus (W. et G.S. West) Chodat 1 1 1 0 1 1 0 0
S. gutwinskii Chodat 0 0 1 0 0 0 0 1
S. incrassatulus Bohlin 1 0 1 0 0 0 0 0
S. magnus Meyen 0 1 0 1 0 0 0 1
S. obliquus (Turpin) Kützing 0 1 0 1 1 1 0 0
S. opoliensis P. Richter var. carinatus (Lemmermann) Chodat 0 0 1 0 0 0 0 0
S. parvus (G.M. Smith) Bourrelly et Manguin 0 0 1 0 0 0 0 0
S. quadricauda (Turpin) Brébisson 1 0 1 1 1 0 0 1
S. sempervirens Chodat 0 0 1 0 0 0 1 0
Schroederia robusta Korschikov 0 0 1 1 0 0 0 0
Spermatozopsis exsultans Korshikov 0 0 1 0 0 0 0 0
Sphaerocystis planctónica (Korschikov) Bourrelly 1 0 0 0 0 1 0 0
Tetraedron caudatum (Corda) Hansgirg 1 0 0 0 0 0 1 1
T. minimum (A. Braun) Hansgirg 0 1 1 0 1 0 1 1
Tetrastrum triangulare (Chodat) Komárek 1 0 1 0 1 1 1 0
Treubaria triappendiculata Bernard 0 0 1 0 1 0 0 0
Westella botryoides (W. West) De-Willdeman 1 0 0 0 0 0 1 0
Order Ulotrichales
Elakatothrix genevensis (Reverdin) Hindák 0 0 0 0 1 0 0 0
Koliella longiseta (Vischer) Hindák 0 0 1 0 1 1 1 0
Ulothrix subtilissima Raben hörst 0 0 0 0 1 0 0 0
Lake
Zaozer'e Ryumniki P OL P OL
Order Oedogoniales
Oedogonium sp. 0 0 0 0 0 0 1 0
Order Desmidiales
Cosmarium bioculatum Brebisson 1 1 0 0 0 1 0 0
C. impressulum Elfving 0 0 0 1 0 0 0 0
C. meneghinii Breb. ex Ralfs 0 1 0 0 0 0 0 0
Cosmarium sp. 0 0 0 1 1 0 1 0
Staurastrum sp. 1 0 1 0 0 0 1 1
Staurodesmus sp. 0 0 0 0 1 0 0 0
DIVISION EUGLENOPHYTA
Order Euglenales
Cryptoglena australis Playfair 0 0 0 0 0 0 0 1
C. pigra Ehren berg 1 0 0 0 0 0 0 0
Euglena sp. 1 1 1 0 1 1 1 0
Lepocinclis ovum (Ehrenberg) Lemmermann 0 0 0 0 0 0 1 0
Trachelomonas cylindrica Ehrenberg 0 0 0 0 0 0 1 1
T. volvocina Ehren berg 0 0 1 0 1 0 1 1
TOTAL 50 46 53 29 54 37 68 46
от
CD Qi
о
_<
m
>
CD (—b
ro о ro о
m
о
0
1
Co
cB-
Co
S1
о
CJ
СЛ I
о
w w
Appendix 2
Taxonomic composition of the vascular plants of small lakes of the Rostov District of the Yaroslavl Region in July 2018. 1 - presence of the taxon, 0 - absence.
w
Lake
Taxon
Chashnitskoe
Zaozer'e
Ryumniki
Vashutinskoe
DIVISION LYCOPODIOPHYTA Class Lycopodiopsida
FAMILY ISOETACEAE RCHB. Isoetes echinospora Durieu
DIVISION EQUISETOPHYTA Class Equisetopsida
FAMILY EQUISETACEAE Michx. ex DC. Equisetum fluviatile L.
DIVISION PTERIDOPHYTA Class Polypodiopsida
FAMILY THELYPTERIDACEAE Pic. Serm. Thelypteris palustris Schott
DIVISION SPERMATOPHYTA Class Magnoliopsida
FAMILY NYMPHAEACEAE Salisb. Nuphar lutea (L.) Smith N. pumila (Timm) DC. Nymphaea Candida J. Pres I FAMILY POTAMOGETONACEAE Rchb. Potamogeton berchtoldii Fieb. P. compressus L. P. crispus L. P. gramineus L.
0 0 0
0 0 1 0
1 0 0
1 0 0 0
от
CD Qi O
m
>
CD <—b
£U
ro о ro о
m
о
0
1 I
Co
S1
о
M
сл I
о
P. natans L. 1
P. perfoliatus L. 1 FAMILY HYDROCHARITACEAE Juss.
Elodea canadensis Michx. 1
Hydrocharis morsus-ranae L. 0
Stratiotes aloides L. 0 FAMILY ALISMATACEAE Vent.
Alisma plantago-aquatica L. 1
Sagittaria sagittifolia L. 1 FAMILY ARACEAE Juss.
Calla palustris L. 1
Lemna minor L. 1
L. trisulea L. 1
Spirodela polyrhiza (L.) Schleid. 1 FAMILY IRIDACEAE Juss.
Iris pseudacorus L. 1 FAMILY POACEAE Barnhart
Agrostis stolonifera L. 1
Calamagrostis canescens (Web.) Roth 1
C. epigeios (L.) Roth 0
C. neglecta (Ehrh.) Gaertn., B. Mey. et Schreb. 0
C. purpurea (Trin.) Trin. 1
Festuca pratensis Huds. 0
Glyceria fluitans (L.) R. Br. 1
G. maxima (C. Hartm.) Holmb. 1
Leersia oryzoides (L.) Sw. 0
Lake
Zaozer'e Ryumniki Vashutinskoe
0 1
0 1 1
1 1 1
1 1 1
0 1
1 1 1
1 1 1
1 1 1
1 1 1
0 1 1
1 1 1
0 1 1
1 1 1
1 1
0 1
0 1 1
0 1
0 1
1 1 1
1 1 1
0 0 1
го
CD
03 TT
о
_<
m
>
(D
<—ь Q3_
ro о ro о
m
о
0
1 I
s §■
Co
S1
i
аз ^
о' со
сл I
о
СлЭ
сл
Molinia caerulea (L.) Moench 0
Phalaroides arundinacea (L.) Rauschert 1
Phleum pratense L. 1
Phragmites australis (Cav.) Trin. ex Steud. 1
Poa annua L. 0
P. pratensis L. 0
Scolochloa festucacea (Willd.) Link 0
Zizania latifolia (Griseb.) Stapf 0 FAMILY JUNCACEAE Juss.
J uncus conglomeratus L. 0
J. articulatus L. 1
J. effusus L. 1
J. filiformis L. 1
J. compressus Jacq. 0
J. tenuis Willd. 0 FAMILY TYPHACEAE Juss.
Sparganium emersum Rehm. 1
S. gramineum Georgi 1
S. x longifolium Turcz. ex Ledeb. 1
Typha latifolia L. 1 FAMILY CYPERACEAE Juss.
Carex acuta L. 1
C. canescens L. 0
C. echinata Murray 0
C. hirta L. 0
C. lasiocarpa Ehrh. 0
Lake °>
Zaozer'e Ryumniki Vashutinskoe
0 1 0
1 1 1
0 0 0
0 1 1 от
CD
0 1 0 0 1 0 I
0 0 1
1 о 0
0 0 0
0 1 1
0 0 0
0 1 0
m
>
0 0 12.
ro о ro о
m
о
0
1
со cB-S
0 0 1 §
Co
S1 i
1 0 1a-
o'
1 1 1 3 1 1 1 w
1 0 0 s
1 1 1
1 1 1 1 0 1 0 1 0 1 1 1 0 0 1
СЛ I
о
C. leporina L. C. pseudocyperus L. C. rostrata Stokes C. vesicaria L.
Eleocharis acicularis (L.) Roem. et Schult. E. palustris (L.) Roem. et Schult. Schoenoplectus lacustris (L.) Palla Scirpus radicans Schkuhr S. sylvaticus L.
FAMILY CERATOPHYLLACEAE S.F. Gray
Ceratophyllum demersum L.
FAMILY RANUNCULACEAE Adans.
Caltha palustris L.
Ranunculus lingua L.
R. repens L.
R. reptans L.
R. sceleratus L.
R. trichophyllum Chaix ex Vi II.
FAMILY GROSSULARIACEAE DC.
Ribes nigrum L.
FAMILY HALORAGACEAE R.Br. Myriophyllum verticil latum L. FAMILY ROSACEAE Juss. Comarum palustre L. Filipéndula ulmaria (L.) Maxim. Potentilla anserina L.
Zaozer'e Ryumniki Vashutinskoe
10 0 0 10 0 0 1111 0 0 11 1111 1111 10 0 1 10 11 10 0 0
1111 0 0 0 1 0 0 10 10 10 0 110 0 0 10
1 0 1
1 0 o
1
0
1
1 1 o
FAMILY BETULACEAE S.F. Gray Alnus glutinosa (L.) Gaertn. FAMILY SALICACEAE Juss. Salix cinerea L. S. x tragi lis L. S. pentandra L. S. triandra L.
FAMILY ELATINACEAE Dumort. El atine hydropiper L. E. triandra Schkuhr FAMILY ONAGRACEAE Juss. Epilobium adenocaulon Hausskn. Epilobium palustre L. Epilobium parviflorum Schreb. FAMILY LYTHRACEAE J.St.-Hil. Lythrum salicaria L. FAMILY BRASSICACEAE Burnett Cardamine amara L. Rorippa amphibia (L.) Bess. R. palustris (L.) Bess. FAMILY POLYGONACEAE Juss. Persicaria amphibia (L.) Delarbre P. hydropiper (L.) Delarbre P. lapathifolia (L.) S.F. Gray Rumex aquaticus L.
Chashnitskoe Zaozer'e Ryumniki Vashutinskoe
1 0 1 0
1 0
1 1 0
0
1 0 0 1
0 0
1 1 0
от
CD QÍ О
m
>
CD <—b
£U
ro о ro о
m
о
0
1
со cB-
Co
S1
О
0
1
1
0
1 1
0
1
0
1 1 1 1
о со
сл I
о
FAMILY CARYOPHYLLACEAE Juss. Stellaria aquatica (L.) Scop. S. palustris Ehrh. ex Retz. FAMILY PRIMULACEAE Vent. Lysimachia thyrsi flora L. L. vulgaris L.
FAMILY RUBIACEAE Juss. Galium palustre L. G. rivale (Sibth. et Sm.) Griseb. G. tri fid um L.
FAMILY BORAGINACEAE Juss.
Myosotis cespitosa Schultz
M. palustris (L.) Nathh.
FAMILY SOLANACEAE Juss.
Solanum dulcamara L.
FAMILY LENTIBULARIACEAE Rich.
Utricularia vulgaris L.
FAMILY LAMIACEAE Lindl.
Lycopus europaeus L.
Mentha arvensis L.
Scutellaria galericulata L.
Stachys palustris L.
FAMILY PLANTAGINACEAE Juss.
Callitriche palustris L.
FAMILY SCROPHULARIACEAE Juss.
Scrophularia nodosa L.
Chashnitskoe Zaozer'e Ryumniki Vashutinskoe
0 0
1 1
1 1 1
0 0
1 1
1 1
1 0 1
0 1
0 1
1 1
1 0 0
1 0
от
CD QÍ О
m
>
CD
<—ь £ü
Ю О
ю о
m
о
0
1
Со
З1
о со
сл I
о
W CD
FAMILY ASTERACEAE Dumort. Achillea ptarmica L. Bidens cernua L. B. frondosa L.
FAMILY MENYANTHACEAE Dum. Menyanthes trifoliata L. FAMILY APIACEAE Lindl. Cicuta virosa L. Oenanthe aquatica (L.) Poir. Sium latifolium L. Thyselium palustre (L.) Raf.
TOTAL
0 1
71
1 0 0 1
50
1 0 1 1 82
1 0 1
1 0 0 1
76
