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First findings of the benthic macroalgae Vaucheria cf. dichotoma (Xanthophyceae) and Punctaria tenuissima (Phaeophyceae) in Estonian coastal waters/Bentiliste makrovetikate Vaucheria cf. dichotoma ja Punctaria tenuissima esmaleiud Eesti rannikumeres.


The Baltic Sea is the world's largest brackish water body. It holds nearly 400 macroalgal species across the wide latitudinal salinity gradient, with a rapid decline in marine species richness from south to north (Middelboe et al., 1997; Larsen & Sand-Jensen, 2006). The lowest richness is predicted to be found at the horohalinicum salinity (5-8) and is reflected in relatively low macroalgal species in the NE Baltic Sea (Schubert et al., 2011).

In Estonian coastal waters, macroalgae have been recorded since the end of the 18th century (for a complete review see Martin et al., 2004). However, regular quantitative phytobenthic investigations started as late as in the 1960s (Trei, 1965), and continuous monitoring of phytobenthos started in 1995 (Martin et al., 2003). By today about 60 macroalgal taxa have been recorded in the Estonian coastal waters (according to the benthos database at the Estonian Marine Institute). Within these registered species, several algae have been found or rediscovered in the Estonian marine waters during recent decades (Martin & Kersen, 2011).

The genus Vaucheria de Candolle belongs to the class Xanthophyceae (yellow-green algae), and is one of the two benthic macroalgal genera within xanthophytes besides Tribonema (Christensen, 1987). They are mainly found on soft bottoms where they form felty growths on the mud (Snoeijs, 1999). Even though Vaucheria was long regarded as a green algae (Johnson & Merritt, 2002), molecular data show that Xanthophyceae are most closely related to Phaeophyceae (Lee, 2008).

Currently 73 species of the genus Vaucheria are known worldwide (Graham et al., 2009; Guiry & Guiry, 2011). In Europe, approximately 40 species are found, mainly in freshwater and terrestrial habitats (Rieth, 2009). In the Baltic Sea Vaucheria are mostly found sterile and therefore are difficult to identify to species level (Johnson & Merritt, 2002; Ott & Oldham-Ott, 2003; Rieth, 2009). According to the most recent macroalgal distribution index (HELCOM, 2012), 13 Vaucheria species have been recorded in the Baltic Sea, most of them found in the southern parts of the basin. Nevertheless, none of the Estonian marine phytobenthic studies have identified Vaucheria spp. previously (e.g. Trei 1983, 1986; Kukk, 1993, 1995). In the northern Baltic Sea, Vaucheria forms dense mats of loose macroalgae where it inhabits bays with soft sediment bottoms in the Baltic Proper, Gulf of Bothnia, and Gulf of Finland (Kautsky, 1992; Bergstrom & Bergstrom, 1999; Lehvo & Back, 2001).

The taxonomy of Vaucheria is based on sexual organs (oogonia and antheridia), which are observed in the field only sporadically. A widespread opinion is that Vaucheria spp. are difficult to identify to subgeneric levels (Nemjova & Kaufnerova, 2009; Schagerl & Kerschbauber, 2009). Vaucheria dichotoma (L.) Martius is a euryhaline species, preferably growing in brackish water, but it can also inhabit fresh waters (Christensen, 1988). It is believed to be of brackish or marine origin (Henschel et al., 1992) and is widespread along the European coasts, being distributed from the Black Sea to the Barents Sea (Guiry, 2010). In the Baltic Sea, V. dichotoma is the most dominant species of the genus and it occurs from the Blekinge Archipelago on the south coast of Sweden up to the northern Bothnian Bay (Snoeijs, 1999; J. Hansen, unpublished data), inhabiting also shallow bays of Finland (Holmstrom et al., 2007) and the southern part of the Baltic Sea (Nielsen et al., 1995).

The marine benthic alga Punctaria tenuissima (C. Agardh) Greville (= Despotrichum undulatum (J. Agardh) Reinke) is a member of the family Chordariaceae. It is characterized by an erect flattened blade arising from the discoid holdfast, having 'hairs' from the surface edge and surface cells that are quadrate or rectangular in rows (Fletcher, 1987). Punctaria tenuissima originates from the Atlantic region and has been found in the North Atlantic, the Mediterranean, the Black Sea, and even the Indian Ocean (Guiry & Guiry, 2011). This species has been reported as a non-indigenous benthic species in the Mediterranean (Ambrogi, 2000), where, due to its epiphytic life strategy, it was introduced probably by oyster culture (Boudouresque & Verlaque, 2010). In the Baltic Sea, P. tenuissima has been recorded from the Kattegat to the southern Bothnian Sea (Nielsen et al., 1995), inhabiting a salinity gradient from 5 to 24 (Snoeijs, 1999). It is rare in the outer archipelago and is often found as an epiphyte on other algae, Zostera marina or Ruppia maritima (Tolstoy & Osterlund, 2003). Punctaria tenuissima has been found also in littoral communities of the Finnish Archipelago Sea (Hallfors et al., 1975; Bostrom et al., 2004) up to the Gulf of Finland (Nielsen et al., 1995); however, it is absent from the Gulf of Finland nowadays (HELCOM, 2012).

The aim of this paper is (1) to report on the recent findings of Vaucheria and Punctaria, present the morphological characteristics of the collected material, and (2) to give a review of potential reasons why the algae have not been recorded previously in the Estonian marine flora.


The study material includes data of a marine benthic habitat inventory (EU LIFE project: Marine Protected Areas in the Eastern Baltic Sea) in the Estonian coastal waters (Fig. 1) in 2005-2006, wrack monitoring in the Gulf of Riga (MARMONI project 'Innovative approaches for marine biodiversity monitoring and assessment of conservation status of nature values in the Baltic Sea'), and Fucus/Furcellaria epiphyte sampling during 2011. Therefore, specimens have been collected by different methods and devices--bottom grab, SCUBA diving, frame sampler, and underwater videography.


Benthos sampling was performed in summer. Three replicated quantitative samples were collected by an Ekman-type bottom grab (0.02 [m.sup.2]), a plant frame (0.04 [m.sup.2]), or by thallus removal (Kersen et al., 2011). Sediment samples were sieved in the field and proportions of different sediment types were determined (i.e. silt, sand, gravel, shingle, stone, and decay). Algal material was stored in a deep freezer at -20[degrees]C until analysing. Simplified Wave Model (SWM) method was used to calculate the wave exposure for mean wind conditions represented by the ten-year period from 1 January 1997 to 31 December 2006 (Isaus, 2004). SWM exposure metric ([m.sup.2] [s.sup.-1]) indicates the following exposure classes: from 4000 to 10 000--very sheltered, from 10 000 to 100 000--sheltered, and from 100 000 to 500 000--moderately exposed. These exposure levels follow the EUNIS classification modified for the Baltic conditions (e.g. HELCOM, 2008; Nikolopoulos & Isaus, 2008).

Wrack samples (sensu Orr et al., 2005) were collected from three transects, all located parallel to the shoreline (e.g. Kersen & Martin, 2007). Transect lengths were 5 m and samples were collected with a 20 cm x 20 cm frame at each metre.

Species identification was done after Christensen (1987) and Fletcher (1987) by means of compound microscope (magnification x40-100) with independent opinions of Swedish algologists used for confirmation (J. Hansen and L. Kautsky, pers. comm.). Measurements of cells and other structure were made using a micrometre eyepiece. Thereafter algae were weighed (dry weight, 60 [degrees]C for a week) or visual abundance of Vaucheria was estimated (from wrack samples) as the ratio to total wrack biomass.



Morphological description

The Vaucheria specimen collected from Someri can be characterized as follows: macroscopic thallus of siphonous (i.e. without cell walls) cylindrical filaments were 90-110 [micro]m wide, branched in a few cases, forming green felt-like patches (Fig. 2). In the cytoplasm numerous elliptical chloroplasts were found. No pyrenoids were observed in the specimen separately from the chloroplast due to the limitations of light microscopy.



Although I did not find sexual organs of this specimen (on which the taxonomy of the genera is mostly based), I concluded that it was most likely Vaucheria cf. dichotoma (L.) Martius. This conclusion was made by following an authoritative identification key of Xanthophyceae (Christensen, 1987) and also considering the environmental tolerance (mainly salinity preference, e.g. Christensen, 1988; Henschel et al., 1992) and distribution records around the Baltic Sea. This was the first find of Vaucheria cf. dichotoma in Estonian marine waters. The species was found within a wrack community on the shore of the Someri Peninsula, in the Gulf of Riga, on 19 May 2011. It was found at a moderately exposed site on a moraine/sandy shore (Fig. 1) where it formed up to 22% of the total wrack biomass (DW [g.sup.-2]). A month later we found the species again at that site, but its abundance had decreased to 2% of the total wrack biomass (Table 1).

Occurrence of Vaucheria in wrack is due to its siphonous thallus structure, which enables to effectively overcome the water transport problem and enables the alga to grow outside truly aquatic habitats (Christensen, 1987). I suspect that the species had started to grow inside the wrack instead of having been snatched by storm waves and landed on the shore because we could not find any sublittoral Vaucheria in May-June. Vaucheria is also known to be ecologically important as a substrate stabilizer if strong currents occur (Stevenson, 1996; Calvo & Barbara, 2004). Therefore it is very likely that Vaucheria can enhance the diversity and abundance of soft bottom macrophytes and macrofauna in the coastal area (Bolam & Fernandes, 2002). The genus is found also in Atlantic salt marshes, where species of Vaucheria are pioneers, settling over the bare mud and facilitating decomposition of silt (Calvo et al., 1999).

Two months later Vaucheria was found for the first time in sublittoral habitats near Haapsi in the Gulf of Riga (Fig. 1). Vaucheria was the dominant phytobenthic species on the soft bottom, which was covered also with Cladophora glomerata, Chara aspera, Zannichellia palustris, and Fucus vesiculosus. This was the first time to observe Xanthophyceae directly in the Estonian marine waters.

This study is the first to report a yellow-green alga of macroscopic form in the Gulf of Riga (after Nielsen et al., 1995; HELCOM, 2012). Vaucheria was previously registered in phytoplankton samples in Parnu Bay (Tenson, 1995), where it probably occurred as zoospores. I am only aware of one previous record adjacent to Estonia, namely Skuja (1924) described occurrence of Vaucheria in the southern Gulf of Riga on a sandy bottom (cited in Kukk, 1993, 1995). We probably registered Vaucheria sp. quantitatively on sublittoral soft bottoms in Saaremaa bays as early as in 2005 and 2006 and again in 2010 by video observation (Table 1), but without reference specimen identification.

I argue that it is possible to find Vaucheria in many sheltered semi-enclosed bays with river inflows. Vaucheria is the most frequent macroalga together with Cladophora and Ulothrix in Estonian running waters (Trei, 2001; Trei & Paal, 2004). Growing together with higher plants, Vaucheria has been observed even in oxidation ponds in Estonia (Viik, 1999) and presumably it belongs to the lacustrine macroflora in Estonia (see Leppik, 1922; Molder, 1944).


Morphological description

The P. tenuissima specimen collected from Kakumae can be characterized as follows: spirally twisted erect blades, dimensions 0.8 mm x 40 mm, numerous plurilocular sporangia with several hairs from the edge of the thallus (Fig. 3). In surface view the cells are quadrate or rectangular, regularly ordered, 20 [micro]m in diameter, each containing discoid chloroplasts.


The brown alga Punctaria tenuissima (C. Agardh) Greville was found for the first time in marine waters of Estonia close to the Kopu Peninsula, Hiiumaa Island, on 5 June 2011. It was found at a moderately exposed site (Fig. 1) at a depth of 4.2 m on a hard substrate with a high vegetation coverage (100%). The algal community at the site was dominated by Cladophora glomerata, Polysiphonia fucoides, Chorda filum, Stictyosiphon foeniculaceus, Ceramium tenuicorne, and Furcellaria lumbricalis. The new species was found in a frame sample with a low biomass (Table 2). It is most likely the first record of P. tenuissima in Estonian marine waters, although Nielsen et al. (1995) mentioned that the species was reported in coastal waters of western Estonia before 1970. However, we are not aware of any reliable original records yet. In June 2011, P. tenuissima was found epiphytically in Kakumae Bay, the Gulf of Finland (Fig. 1). It was found in low biomass growing on Furcellaria and Fucus at depths from 1 to 4.1 m in a sheltered site (Table 2). According to a recent check-list of HELCOM (2012), this is also the first time P. tenuissima was found in the entire Gulf of Finland.


One of the reasons why the brown alga P. tenuissima has not been found in the Estonian waters before is obviously its complex life history. The alga grows both microthalli and macrothalli. The different morphological phases of the thallus are mainly modified by temperature (Rietema & van den Hoek, 1981) and pluriseriate ribbon-like thalli form under lower water temperatures (4-16 [degrees]C). Even then, the alga appears with low coverage (Eriksson et al., 2002). Moreover, according to Parente et al. (2010), comparison of the life histories of P. tenuissima and P. latifolia suggests that they are conspecific.

Another reason behind this distribution pattern of P. tenuissima is the lack of a regular seasonal monitoring of phytobenthos to register also epiphytic flora on the macroalgae. This is an important shortcoming since the species is known as a spring-summer annual in the boreal Atlantic, and if epiphytic then it grows mainly on overwintering leaves of Zostera marina (Novaczek, 1987; Bostrom et al., 2004). This kind of methodological problem was also noted for the British Isles (Hardy & Guiry, 2003) and the Mediterranean (Bellemo et al., 1999).

doi: 10.3176/eco.2012.2.05


I am very grateful to Joakim Hansen, Gustav Johansson, and Lena Kautsky for helping to identify algal specimens. I thank Anastasiia Kovtun-Kante, Agnes Siiber, Liis Rostin, and Remi Treier for analysing algal samples. This study was financed by the Estonian Science Foundation under grant ETF8775, EU Life+ project MARMONI, EU Interreg IVA project HISPARES, and Estonian target financed project SF0180013s08.


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Priit Kersen

Estonian Marine Institute, University of Tartu, Maealuse 14, 12618 Tallinn, Estonia

Institute of Mathematics and Natural Sciences, Tallinn University, Narva mnt. 25, 10120 Tallinn, Estonia;

Received 19 December 2011, revised 7 March 2012, accepted 8 March 2012
Table 1. New records of Vaucheria cf. dichotoma in Estonian marine
waters with basic environmental characteristics. Sediment % indicates
the proportion of bottom coverage

Sublittoral survey

Site                  Sampler        Date      Depth,    Station

Saastna             Birge grab    5.08.2005      0.75    SLSAAST4
Saastna             Birge grab    5.08.2005      0.5    SL33
Kihelkonna Diving                 1.08.2006      1.5    LF409
Kihelkonna          Video         1.09.2010      2.1    VILH471
Kihelkonna          Video         1.09.2010      1.1    VILH472
Haapsi              Plant frame   5.08.2011      1.0    Marmoni 4

Site                  Lat        Lon             Wave
                    (WGS-84)   (WGS-84)        exposure,
                                          [m.sup.2] [s.sup-1]

Saastna             58.42577   23.07843           3 400
Saastna             58.42477   23.08277           5 600
Kihelkonna Diving   58.35869   21.98711          33 600
Kihelkonna          58.35582   21.97116          26 000
Kihelkonna          58.36015   21.96862          28 000
Haapsi              58.39705   23.71138         307 000

Site                   Biomass,      Silt,   Sand,   Gravel,
                    DWg [m.sup.-2]     %       %        %

Saastna                 0.6345        60      16
Saastna                 3.3229        60       0
Kihelkonna Diving                      5      80       10
Kihelkonna                                    95        5
Kihelkonna                                    95        5
Haapsi                 281.515                60

Site                Shingle,   Decay,   Stone,
                       %         %        %

Saastna                          24
Saastna                40
Kihelkonna Diving       5
Haapsi                 30                 10

Wrack survey

Site     Sampler      Date      Depth   Section     Lat        Lon
                                                  (WGS-84)   (WGS-84)

Someri    Frame    19.05.2011    n.a.      1      58.35483   23.7463
Someri    Frame    19.05.2011    n.a.      3      58.35373   23.7462
Someri    Frame    14.06.2011    n.a.      2      58.35437   23.74642

Site            Wave             Occurrence       Biomass
              exposure,          in samples    proportion, %
         [m.sup.2] [s.sup-1]

Someri         200 000         1 sample of 5       < 1
Someri         200 000         4 samples of 5       12
Someri         200 000         2 samples of 5        2

n.a., not applicable.

Table 2. New records of the Punctaria tenuissima in Estonian marine
waters with basic environmental characteristics. Sediment % indicates
the proportion of bottom coverage

Punctaria in epiphytic survey

Host alga                 Depth,     Lat        Lon         Date
                            m      (WGS-84)   (WGS-84)

Furcellaria lumbricalis    4.1     59.45805   24.56707   22.06.2011
Furcellaria lumbricalis    4.1     59.45805   24.56707   22.06.2011
Furcellaria lumbricalis    4.1     59.45805   24.56707   22.06.2011
Fucus vesiculosus          1       59.45833   24.56894   22.06.2011
Fucus vesiculosus          2.4     59.45833   24.56844   22.06.2011
Fucus vesiculosus          2.4     59.45833   24.56844   22.06.2011
Fucus vesiculosus          2.4     59.45833   24.56844   22.06.2011

Host alga                 Location           Wave
                                     [m.sup.2] [s.sup.-1]

Furcellaria lumbricalis   Kakumae           70 000
Furcellaria lumbricalis   Kakumae           70 000
Furcellaria lumbricalis   Kakumae           70 000
Fucus vesiculosus         Kakumae           70 000
Fucus vesiculosus         Kakumae           70 000
Fucus vesiculosus         Kakumae           70 000
Fucus vesiculosus         Kakumae           70 000

Host alga                   Punctaria,     Host alga,   Epiphytic
                               DWg            DWg         load,
                             on host                        %

Furcellaria lumbricalis       0.0001         4.6672       0.002
Furcellaria lumbricalis       0.0001         2.9183       0.003
Furcellaria lumbricalis       0.0001         3.7622       0.003
Fucus vesiculosus             0.0002        26.663        0.001
Fucus vesiculosus             0.0001        12.4422       0.001
Fucus vesiculosus             0.0001         7.3967       0.001
Fucus vesiculosus             0.0004        22.9225       0.002

Punctaria in habitat mapping survey

Station   Depth,     Lat        Lon         Date     Location
            m      (WGS-84)   (WGS-84)

KOPU130    4.2     58.90903   22.1001    5.06.2011     Kopu

Station           Wave              Biomass,      Shingle,   Boulder,
               exposure,         DWg [m.sup.-2]      %           %
          [m.sup.2] [s.sup.-1]

KOPU130         427 000              0.0025          60         40
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Author:Kersen, Priit
Publication:Estonian Journal of Ecology
Article Type:Report
Geographic Code:4EXES
Date:Jun 1, 2012
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