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A C. connivens specimen from the Vistula Lagoon. 1 – an internode of the main axis (stem), usually longer than branchlets; 2 – an incurved sterile branchlet. The specimen’s thalli are delicately green and lustrous as described by Dąmbska [4]. 

A C. connivens specimen from the Vistula Lagoon. 1 – an internode of the main axis (stem), usually longer than branchlets; 2 – an incurved sterile branchlet. The specimen’s thalli are delicately green and lustrous as described by Dąmbska [4]. 

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The stonewort Chara connivens was rediscovered in the Vistula Lagoon in 2011, almost 35 years after its last record. In 2012, the species was recorded for the first time in the Szczecin Lagoon. Chara connivens occurred at shallow (0.5–1.2 m) sandy-muddy and muddy bottoms of small embayments. In the Vistula Lagoon, the stonewort was represented by s...

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Context 1
... are a group of macroscopic green algae represented by 314 species [1] assigned to the family Chara- ceae within the phylum Chlorophyta; alternatively, they are treated as a separate phylum, the Charophyta [2]. The occurrence of most of the Characeae species in Europe is limited to clear, freshwater lakes of low fertility [3–8]. In response to increasing trophic status and the associated light limitation, many charophytes have therefore become rare, and are treated as sensitive bioindicators of water quality [6,9]. Although the number of sites with abundant charophyte vegetation has decreased, these macroalgae are widely distributed in different types of aquatic environments throughout the world. Due to their ability for ion regula- tion and osmotic adjustment, a set of charophyte species, commonly known from freshwaters, can occur in brackish coastal lagoons, as shown for the Baltic Sea [10]. In brackish waters, charophytes experience changes in osmotic pressure and mineral composition [7]. A brackish environment unique on a global scale is the Baltic Sea, the world’s largest brackish water body [11]. The variety of the Baltic coast types gives rise to diverse and specific biocoe- noses. Among submerged algae, a number of freshwater and brackish charophyte species were identified to represent five out of six genera within the family of Characeae [10]. Among the rarest brackish charophytes, Chara connivens P. Salzmann ex A. Braun 1835 exemplifies those species whose distribu- tion has been reported to have greatly declined [12,13]. Chara connivens is widely distributed, with records known from Europe, Africa and Northern Asia [6,12,14–16]. Within Europe, C. connivens was recorded in the western European maritime regions, along the coasts of the Mediterranean and the Baltic Sea, and in inland saline waters of central and southern Europe [4,6,12]. In Poland, historical records of the species are available for the Gulf of Gdańsk [4] and for the Vistula Lagoon [17]. Currently, C. connivens has a status of a strictly protected species. In the “Red list of plants and fungi in Poland”, published in 2006 [18], the species was assigned to the category Ex, i.e., extinct or probably extinct species. The present paper reports new records of C. connivens from the Vistula and the Szczecin Lagoons. The Vistula Lagoon observation is a rediscovery of C. connivens after almost 35 years of the previous record, while in the Szczecin Lagoon this species has never been noted so far. The occurrence of C. connivens was evidenced in the Pol- ish parts of the Vistula and Szczecin Lagoons (Fig. 1) during the vegetation studies performed on 6 July 2011 in the Vistula Lagoon and on 31 July and 14 August 2012 in the Szczecin Lagoon. The distribution and species composition of aquatic vegetation was determined by direct observations and from samples collected with an anchor. Additionally, in the Vistula Lagoon the floristic data collection was supplemented with measurements of water conductivity and temperature taken with a portable CTD probe. Chara connivens [syn. Chara globularis f. connivens (P. Salzmann ex A. Braun) R.D. Wood 1962] is a small alga, usually up to 15 cm long, rarely longer (25–50 cm) [4,12,13,15,19]. The slightly encrusted and slender thalli are delicate green and lustrous [4]. Sterile specimens of C. connivens are very similar to those of C. globularis Thuill. [15] due to presence of triplostichous stem cortex (usually isostichous or prime cells slightly larger than the secondaries) with lacking or rudimentary spine cells [4] and two rows of rudimentary stipulodes (uppers slightly larger than lowers or all imperceptible). The features important in distinguishing the two species include the branchlets (6–10 in a whorl composed of 6–13 segments each, out of which the terminal 1–2 celled segment is ecorticate) which in C. connivens are strongly incurved in fertile male specimens, often also in fertile female individuals [4,6,12,15]. This feature is marked on the photographs of individuals from the Szczecin (Fig. 2) and Vistula (Fig. 3) Lagoons. Additionally, the axial internodes are very long, up to 6 times as long as the branchlets [12]; this was observed in our collections (Fig. 3). The most important distinguishing trait is that C. connivens is dioecious, while C. globularis is monoecious. In C. connivens , antheridia and oogonia develop at the lowermost 1–4 and 1–3 branchlet nodes, respectively [4]. The red solitary antheridia of up to 1 mm in diameter in C. connivens are larger than those in C. globularis [12,16]. The oogonia are up to 1.1 mm long and the oospores are dark. The specimens collected in the two lagoons showed the characters described above, typical of the vegetative thallus parts of C. connivens . In addition, besides sterile specimens, both lagoons yielded fertile male specimens which were more abundant in the Szczecin Lagoon (Fig. 2), where fertile female individuals were found as well. On 6 July 2011, C. connivens was found in the western part of the Vistula Lagoon (Fig. 1), on sandy mud at a depth of 1.1 m. The area was partially isolated by abundant mono- specific assemblages of emergent vegetation, particularly Phragmites australis (Cav.) Steud. and Schoenoplectus lacustris (L.) Palla. As the site is shallow and the water dynamics is low, the water column temperature was high (19.5°C). The site is located 2 km away from the nearest river mouth; the riverine inflow resulted in low salinity (0.8 psu). The water was transparent down to the bottom; however, it was not possible to examine the submerged vegetation distribution from the shipboard as the bottom was densely covered by the filamentous green algae Cladophora sp., Oedogonium sp. and Rhizoclonium riparium (Roth) Harvey (those accounted for 90% of the biological material collected). The site sup- ported a high diversity of macrophyte species occurring as single and small-sized specimens. They were dominated ...
Context 2
... are a group of macroscopic green algae represented by 314 species [1] assigned to the family Chara- ceae within the phylum Chlorophyta; alternatively, they are treated as a separate phylum, the Charophyta [2]. The occurrence of most of the Characeae species in Europe is limited to clear, freshwater lakes of low fertility [3–8]. In response to increasing trophic status and the associated light limitation, many charophytes have therefore become rare, and are treated as sensitive bioindicators of water quality [6,9]. Although the number of sites with abundant charophyte vegetation has decreased, these macroalgae are widely distributed in different types of aquatic environments throughout the world. Due to their ability for ion regula- tion and osmotic adjustment, a set of charophyte species, commonly known from freshwaters, can occur in brackish coastal lagoons, as shown for the Baltic Sea [10]. In brackish waters, charophytes experience changes in osmotic pressure and mineral composition [7]. A brackish environment unique on a global scale is the Baltic Sea, the world’s largest brackish water body [11]. The variety of the Baltic coast types gives rise to diverse and specific biocoe- noses. Among submerged algae, a number of freshwater and brackish charophyte species were identified to represent five out of six genera within the family of Characeae [10]. Among the rarest brackish charophytes, Chara connivens P. Salzmann ex A. Braun 1835 exemplifies those species whose distribu- tion has been reported to have greatly declined [12,13]. Chara connivens is widely distributed, with records known from Europe, Africa and Northern Asia [6,12,14–16]. Within Europe, C. connivens was recorded in the western European maritime regions, along the coasts of the Mediterranean and the Baltic Sea, and in inland saline waters of central and southern Europe [4,6,12]. In Poland, historical records of the species are available for the Gulf of Gdańsk [4] and for the Vistula Lagoon [17]. Currently, C. connivens has a status of a strictly protected species. In the “Red list of plants and fungi in Poland”, published in 2006 [18], the species was assigned to the category Ex, i.e., extinct or probably extinct species. The present paper reports new records of C. connivens from the Vistula and the Szczecin Lagoons. The Vistula Lagoon observation is a rediscovery of C. connivens after almost 35 years of the previous record, while in the Szczecin Lagoon this species has never been noted so far. The occurrence of C. connivens was evidenced in the Pol- ish parts of the Vistula and Szczecin Lagoons (Fig. 1) during the vegetation studies performed on 6 July 2011 in the Vistula Lagoon and on 31 July and 14 August 2012 in the Szczecin Lagoon. The distribution and species composition of aquatic vegetation was determined by direct observations and from samples collected with an anchor. Additionally, in the Vistula Lagoon the floristic data collection was supplemented with measurements of water conductivity and temperature taken with a portable CTD probe. Chara connivens [syn. Chara globularis f. connivens (P. Salzmann ex A. Braun) R.D. Wood 1962] is a small alga, usually up to 15 cm long, rarely longer (25–50 cm) [4,12,13,15,19]. The slightly encrusted and slender thalli are delicate green and lustrous [4]. Sterile specimens of C. connivens are very similar to those of C. globularis Thuill. [15] due to presence of triplostichous stem cortex (usually isostichous or prime cells slightly larger than the secondaries) with lacking or rudimentary spine cells [4] and two rows of rudimentary stipulodes (uppers slightly larger than lowers or all imperceptible). The features important in distinguishing the two species include the branchlets (6–10 in a whorl composed of 6–13 segments each, out of which the terminal 1–2 celled segment is ecorticate) which in C. connivens are strongly incurved in fertile male specimens, often also in fertile female individuals [4,6,12,15]. This feature is marked on the photographs of individuals from the Szczecin (Fig. 2) and Vistula (Fig. 3) Lagoons. Additionally, the axial internodes are very long, up to 6 times as long as the branchlets [12]; this was observed in our collections (Fig. 3). The most important distinguishing trait is that C. connivens is dioecious, while C. globularis is monoecious. In C. connivens , antheridia and oogonia develop at the lowermost 1–4 and 1–3 branchlet nodes, respectively [4]. The red solitary antheridia of up to 1 mm in diameter in C. connivens are larger than those in C. globularis [12,16]. The oogonia are up to 1.1 mm long and the oospores are dark. The specimens collected in the two lagoons showed the characters described above, typical of the vegetative thallus parts of C. connivens . In addition, besides sterile specimens, both lagoons yielded fertile male specimens which were more abundant in the Szczecin Lagoon (Fig. 2), where fertile female individuals were found as well. On 6 July 2011, C. connivens was found in the western part of the Vistula Lagoon (Fig. 1), on sandy mud at a depth of 1.1 m. The area was partially isolated by abundant mono- specific assemblages of emergent vegetation, particularly Phragmites australis (Cav.) Steud. and Schoenoplectus lacustris (L.) Palla. As the site is shallow and the water dynamics is low, the water column temperature was high (19.5°C). The site is located 2 km away from the nearest river mouth; the riverine inflow resulted in low salinity (0.8 psu). The water was transparent down to the bottom; however, it was not possible to examine the submerged vegetation distribution from the shipboard as the bottom was densely covered by the filamentous green algae Cladophora sp., Oedogonium sp. and Rhizoclonium riparium (Roth) Harvey (those accounted for 90% of the biological material collected). The site sup- ported a high diversity of macrophyte species occurring as single and small-sized specimens. They were dominated ...

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... Recent field data of the Wielki Zalew already indicated Potamogeton perfoliatus and Myriophyllum spicatum at a depth of 2-2.2 m (Wozniczka, pers. com.) and the re-occurance of charophytes is reported by Brzeska et al. (2015). Charophytes were recently observed in the Kleines Haff, as well. ...
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... Out of the three above mentioned estuarine regions from which C. connivens was recorded by the end of 19th, two still support populations of this species: the Vistula and the Oder estuaries, both with major lagoons there constituting refugia for the charophytes identified in more recent times (Brzeska et al., 2015;Garbacik-Wesołowska, 1969;Pliński et al., 1978;Szarejko, 1955;Urbaniak & Gąbka, 2014;Volodina & Gerb, 2018, this study). ...
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... The rediscovered and newly discovered populations of C. connivens in the Vistula and Szczecin Lagoons presented in this article confirm the predictions of Torn et al. (2020), at least in case of Lake Wicko Wielkie (northernmost part of the Szczecin Lagoon). Previously, no charophytes were recorded from this area (Garbacik-Wesołowska, 1969), but in 2012 this species turned out to be abundant locally, forming a dense assemblage without any other chrophytes (Brzeska et al., 2015). Along in the Vistula Spit, the species was found to be still frequent, but in scarcity or rarely in small and not dense patches, co-occurring with C. aspera, C. contraria, C. globularis, C. tomentosa and N. obtusa (Krajewski et al., 2015;Brzeska-Roszczyk, 2020;this study). ...
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... Modeltest3.7 [34] was used to determine the parameters and the best surrogate model for each marker (Table 3). Diagnosis: This species is similar to Chara connivens, but the cortex of the latter is triplotichous and obviously different from this species [35][36][37][38]. It is also related to Chara virgata (=C. ...
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... They are widely distributed in freshwater as well as brackish and marine habitats, from tropical to polar regions (Wood 1965). The occurrence of Characeae taxa is mostly limited to clear, freshwater lakes with low fertility (Dąmbska 1964;Krause 1969;1981;1997;Martin et al. 2003;Brzeska et al. 2015). Many stonewort species are characteristic of hard, oligo-mesotrophic water ecosystems (Krause 1981;1997). ...
... Chara connivens P. Salzmann ex A. Braun, known as a convergent stonewort, belongs to the group of brackish charophytes. To date, C. connivens has been reported mainly from Western Europe, the Mediterranean Sea basin region and the Baltic Sea (Krause 1981;1997;Torn et al. 2004;Urbaniak & Gąbka 2014;Brzeska et al. 2015). Several stands of C. connivens were also located in the inland saline waters of Central and Southern Europe (Dąmbska 1964;Krause 1997;. ...
... Communities with C. connivens are mostly pioneer in ephemeral shallow waterbodies (Felzines & Lambert 2012). In Europe, C. connivens stands were recorded in Albania (Zeneli & Kashta 2016), Bulgaria (Temniskova et al. 2008), Estonia Kotta et al. 2004;Torn et al. 2015), Finland (Appelgren et al. 2004), France (Hy 1913;Corillion 1957), Georgia (Barinova & Kukhaleishvili 2014), Germany (Wood 1965;Luther 1979;Ludwig & Schnittler 1996), Great Britain (Stewart & Church 1992;Bryant & Stewart 2002;John et al. 2011), Greece (Langangen 2010;, Ireland (Stewart & Church 1992), Latvia (Skuja 1928;Kostkevičienė & Sinkevičiene 2008), the Netherlands Bruinsma 2000;Bruinsma et al. 2018), Poland (Urbaniak & Blaženčić 2012;Urbaniak & Gąbka 2014;Brzeska et al. 2015;Krajewski et al. 2015), Portugal (Corillion 1957;Cambra Sánchez et al. 1998), Romania (Caraus 2002;2017), Russia (Kaliningrad Oblast) (Romanov et al. 2018a,b), Spain (Reyes Prósper 1910;Aboal 1985;Cambra Sánchez et al. 1998;Cirujano et al. 2008;del Pozo et al. 2011), and Sweden (Luther 1979;Blindow 1988;Wallström & Persson 1999;Tolstoy & Österlund 2003;Torn et al. 2004;Langangen 2007). In Africa, populations of C. connivens were found in Algeria (Wood 1965;Muller et al. 2017), Egypt (Corillion 1957), Morocco (Wood 1965;Muller et al. 2017), and Tunisia (Corillion 1957;Wood 1965;Muller et al. 2017). ...
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The paper presents the first record of a Chara connivens (Characeae) stand from Fuerteventura (Canary Islands, Spain). The species was previously recorded only on Tenerife and Lanzarote, mostly in marine and artificial habitats. Physicochemical parameters of water and morphological features of the thalli and plant community were analyzed. General habitat requirements of C. connivens populations located in Europe, North Africa, and South-West Asia were reviewed. The newly described stand was located in a natural rock crevice that was part of a riverbed in El Barranco de las Peñitas (Penitas Canyon). This habitat was unshaded and filled with brackish water. C. connivens co-occurred with Cladophora glomerata and Ruppia maritima . Our report on the presence of C. connivens in the Spanish territory of the Atlantic Ocean is the first in almost 40 years. The distribution of C. connivens and its plant associations on the Canary Archipelago are poorly explored, which is why this topic requires further research.
... Our field surveys and a literature study proves that different submerse macrophyte species are still present in the western lagoon and single macrophyte stands were found in a water depth of up to 1.8 m. This is true for the eastern, Polish part as well (Brzeska et al. 2015). Recent studies by Nowak et al. (2008) and Blindow et al. (2016) documented that germinable diaspores of several species are present in the sediments of all observed German Baltic coastal water. ...
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The Systems Approach Framework with an integrated Ecological-Social-Economic assessment was applied to address the issue of zebra mussel (Dreissena polymorpha) farming in the large Oder (Szczecin) Lagoon, southern Baltic Sea. Heavy eutrophication hampers the use of the lagoon and zebra mussel farming is considered as new use and potential measure to improve water quality. Three alternative scenarios were developed in interaction with local stakeholders: 1) the production of mussels as fresh feed and meal on a commercial basis seemed not profitable, because of a limited market for fresh mussels (zoos, aquaculture) and low prices for organic feed. 2) Mussel cultivation to improve transparency and attractiveness of bathing waters near beaches had only a limited potential (0.2 m improvement of Secchi depth). A higher mussel biomass would increase the risk of temporary hypoxia. 3) Mussels farms for improving the environmental status (according to EU Water Framework Directive) by supporting macrophyte restoration were considered as the most promising scenario. Our model simulations suggested that as soon as a compensation for nutrient removal is considered, all mussel farm scenarios could cover the costs. Experiments and literature confirm that the conditions for an environmental friendly farming approach in the lagoon are suitable. Steps towards and problems associated with an implementation, e.g. invasion of Dreissena bugensis (quagga mussel), are discussed. Each step of the Ecological-Social-Economic assessment and major lessons learnt are documented in detail. Altogether, the approach turned out to be very suitable for this issue.
... Despite the narrowest breadth of environmental niche space compared to other studied species , Chara connivens will benefit the most from climate change. During the last 20 years the distribution of C. connivens has been increasing in the Estonian coastal waters (Torn et al. 2004(Torn et al. , 2015 as well as in the Baltic Sea (Appelgren et al. 2004;HELCOM 2013b;Brzeska et al. 2015). The predicted occurrence of the marine species Zostera marina decreased under lower salinity conditions. ...
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The potential response of occurrence of charophyte and angiosperm communities to climate change in the brackish environment of the Baltic Sea was investigated using the boosted regression trees (BRT) modelling method. The aim of the study was to analyse sensitivity of various species to climate change and to predict the changes in species’ distribution under projected changes of environmental variables. The results showed that depth was the most influential environmental variable in predicting the spatial distribution of charophytes and angiosperms. Other environmental variables had notably lower importance in determining community structure and the order of importance was species specific. The exceptions were the charophytes Chara horrida and C. tomentosa, for which the influence of wave exposure was stronger than the influence of depth. The studied species could be divided into three groups based on the predicted effect of climate change: (1) species that benefit from change (Chara connivens, Potamogeton perfoliatus, Myriophyllum spicatum, C. aspera, C. baltica, C. canescens, Ruppia maritima), (2) species with no notable change (Chara tomentosa, C. horrida, Stuckenia pectinata), and (3) species that decline (Zostera marina, Zannichellia palustris, Tolypella nidifica). Currently, the shallow and sheltered West Estonian Archipelago Sea hosts favourable habitat conditions for most of the charophyte and angiosperm species. The most significant predicted change was the decline or disappearance of brackish and marine species and the increase of freshwater species due to the expected decrease of salinity in the Baltic Sea.