The habitat selection and invasion of a new polychaete Marenzelleria viridis is described.

Up to now the species has Iow biomass and abundance values in Estonian coastal waters. M. viridis prefers gravel substrate. The increase in the abundance of M. viridis coincides with the decline of that of another polychaete, Nereis diversicolor, in uniform habitats but not in the patchy environment. lt is likely that M. viridis may become a numerically dominant species in the zoobenthos of the region in the following years

Kotta, Ilmar

Kotta, Jonne

OceanRep

Proc. Estonian Acad. Sei. Biol. Ecol., 1998, 47, 3, 212-220 DISTRIBUTION AND INVASION ECOLOGY OF Marenzelleria viridis IN THE ESTONlAN COASTAL WATERS Jonne KOTTA and Ilmar KOTTA Estonian Marine Institute, Rävala pst. 8, EE-0001 Tallinn, Estonia, e-mail: jonne@sea.ee Received 21 May 1998 Abstract. The habitat selection and invasion of a new polychaete Marenzelleria viridis is described. Up to now the species has Iow biomass and abundance values in Estonian coastal waters. M. viridis prefers gravel substrate. The increase in the abundance of M. viridis coincides with the decline of that of another polychaete, Nereis diversicolor, in uniform habitats but not in the patchy environment. lt is likely that M. viridis may become a numerically dominant species in the zoobenthos of the region in the following years. Key words: Marenzelleria viridis, Gulf of Riga, Gulf of Finland, Väinameri Archipelago. INTRODUCTION Salinity in the greater part of the Baltic Sea is less than 8 PSU. The Baltic Sea is the largest marine area of such low salinity in the world. As compared to other brackish waters, the nurober of species in the Baltic fauna is low (Segersträle, 1957). This makes the ecosystem morevulnerable to external factors. In recent years several non-indigenous estuarine and marine species have appeared in the eastern part of the Baltic Sea, among them Hemimysis anomala G. 0. Sars and Dreissena polymorpha (Pallas). This phenomenon has been explained as the result of weakening of the physical barriers due to human activities (Salemaa & Hietalahti, 1993; Valovirta & Porkka, 1996; Kotta et al., 1998). Marenzelleria viridis (Verrill), originally inhabiting only North American estuaries, was for the first time found in the Baltic in 1985 (Bick & Burckhardt, 212 1989). Since then the species has successfully spread to the coasts of Sweden (Persson, 1991), Finland (Norkko et al., 1993), Poland (Gruszka, 1989), Lithuania (Churbanova & Olenin, 1992), and Latvia (Lagzdins & Pallo, 1994). In this paper the invasion of M. viridis to Estonian waters is described and an overview of its habitat preferences is given. MATERIAL AND METHODS The investigation area covers the main waterhodies around Estonian coast: the southern part of the Gulf of Finland, the Väinameri Archipelaga area, and the Gulf of Riga (including southern bays of Saaremaa Island and eastern andcentral parts of the Gulf of Riga). A total of 476 stations were sampled. In the present study the data from different mapping studies during 1991-97 are used. For more detailed information about sampling sites see Kotta et al., 1998. Most samples were collected by a diver (Tvärminne or suction sampler) at depths less than 15m. A van-Veen bottomgrab was used in the deeper areas. Salinity is very low in our study area: values between 3 and 7 PSU have been measured (Segersträle, 1957; Järvekülg, 1979). Sampling was performed on various bottom substrates. Depths ranged from 0.5 to 70 m. Sediments were washed through a 0.25 mm mesh, and were preserved in 4% buffered formaldehyde solution. In the laboratory the animals were counted under a stereo dissecting microscope. The total biomass of the animals in each sample was weighed to the nearest 0.5 mg and calculated for an area of 1 m2. The significance Ievel of 0.05 was adopted for all statistical tests. After testing for normality of the data (Kolmogorov-Smirnov test for goodness of fit) and homogeneity of variance (Bartlett' s and Hartley' s tests), analyses of variance (ANOVA) (Sokal & Rohlf, 1981) were performed on the abundances and biomasses of M. viridis. The sedimenttype was involved as a factor. The effect of depth, abundance, and biomass of the dominant polychaete, Nereis diversicolor (0. F. Müller), was tested on the abundances and biomasses of M. viridis. RESULTS The first observation of M. viridis in Estonian waters dates from 1994. The polychaete was found in the eastern part of the Gulf of Finland near the estuary of the Pühajögi River (Narva Bay, 27° 21' E, salinity z4 PSU). Only a small individual (10 mm), probably just an occasional visitor, was recorded. Up to 1997 no other individual of the species was found at the southern coast of the Gulf of Finland. In 1995 M. viridis was caught practically in the whole littoral zone as well as in the deeper parts of the Gulf of Riga (salinity ""5.5 PSU). Inshallower areas of 213 the gulf (0-3 m), the species prefers sand or gravel bottoms. Its abundances are higher at more densely vegetated areas. At higher depths (>10m) M. viridis is confined to silty clay bottoms. In the same year M. viridis spread also north from the Gulf of Riga, into the Väinameri Archipelaga area. However, there the polychaete is restricted to the deeper parts of the archipelago (7-11 m), which corresponds to sandy clay sediments. In both regions , the communities were dominated numerically by Macoma balthica L. and Cerastaderma lamarcki (Reeve). The proportion of M. viridis in these communities was rather low, comprising 10% at the most. Figure 1 depicts the distribution of M. viridis in Estonian coastal waters. 22'00'E 24"00'E 26"00'E 2S' OO'E j;!;i!TI Dis tribu tion area of M arenzel/erin viridis GULF OF FINLAND 59' 20 'N 58'4 0'N ESTONIA 58'00'N 50 km Fig. 1. Distribution area of Marenzelleria viridis in Estoni an coastal sea. Sediment type is important in explaining the variance of abundance (ANOVA, p = 0.003) and partly the biomass of M. viridis (p = 0.066). Higher abundances were found on barder substrate (gravelly sand, pebbles rnixed with sand) and lower abundances on different types of clay bottoms ( clay and silt bottoms, clay bottoms rnixed with sand and fine sand). Biomasses werehigher on gravel bottoms and lower on clay bottoms rnixed with fine, medium, or coarse sand (Figs. 2 and 3). 214 10000 N 1000 E -'"0 c oS 100 () c CO '"0 c :;j ..D 10 <( 1 r-~ u Cl) u u + Cl) ~ u + Cl) Fig. 2. Average values of the abundance of Marenzelleria viridis at different sediment types in the Gulf of Riga and the Väinameri Archipelaga area. Sediment types: S, silt; C, clay; FS, fine sand; MS, medium sand; CS, coarse sand; ST, stone; PB, pebbles; G, gravel. 10 1 N E -0) (f) 0.1 (f) ro E .Q CD 0.01 0.001 n ~ u Cl) u Cl) + ~ u r-Cl) u + 0::) 0.., Cl) ~ + Cl) ,.. u + Cl) u + Cl) ~ Fig. 3. Average values of the biomass of Marenzelleria viridis at different sediment types in the Gulf of Riga and the Väinameri Archipelaga area. For sediment codes see Fig. 2. 215 Depth had little effect on the abundance and biomass of M. viridis (r = -0.05, p>0.05). The species was found between depths of 0.1 and 55 m, the average depth was 10.3 m. Longitude correlated weakly with the abundance of M. viridis (r = 0.16, p<0.05). During 1994-95 a significant decline of another polychaete, Nereis diversicolor, was observed in the deeper parts of the Väinameri Archipelaga area. Its mean abundance was 600 ind m-2 in 1994. In 1995 we failed to find the species in most part of the Väinameri Archipelaga area. When considering the whole study area no negative correlation between M. viridis and N. diversicolor was observed. DISCUSSION Though the first finding of M. viridis in the Latvian side of the Gulf of Riga dates from 1988 (Lagzdins & Pallo, 1994), it did not invade Estonian waters during the next five years. Extensive mapping studies of macrozoobenthos were carried out in Pärnu Bay and the surroundings in 1991 and at the coastal sea of Saaremaa Island in 1993 (Kotta & Kotta, 1995, 1997). No individuals of M. viridis were detected in either studies. Similarly, M. viridis was found in the Finnish (northern) side of the Gulf of Finland already in 1990 (Stigzelius et al., 1997). The sampling in the Estonian side of the gulf in 1990-96 proved that the polychaete had not succeeded in colonizing its southern coast. Some rare individuals of M. viridis were found in deeper areas of the western side of the Gulf of Finland and Tallinn Bay in 1997. The single individual found on the southern coast of the eastern part of the gulf in 1994 suggests that no stable community of Marenzelleria has formed in this region. As previously suggested by other authors (Essink & Kleef, 1993) M. viridis may be transported in the bailast water, which could explain its presence far off its dispersion limits. Besides, anti-clockwise circulation of the currents would not permit M. viridis to spread from the northern side of the Gulf of Finland towards its southern side. The larvae are unable to complete their development at salinities below 5 PSU (George, 1966), which may frequently occur in the easternmost part of the Gulf of Finland. Despite five years of "stagnation" the polychaete successfully spread over the Gulf of Riga in 1994-95. Specimens were found equally in the deeper part of the gulf (near Ruhnu Island) and in the transects of the southern coast of Saaremaa Island, the opening area of Pärnu Bay, and the borderline between Estonia and Latvia. Why did M. viridis keep waiting till 1994? The salinity of the Gulf of Riga did not vary much during 1992-94. Figure 4 depicts the temperature fluctuations of the Gulf of Riga in 1992-95. One may notice higher water temperatures in summer 1994, which might have led to a better condition and higher reproductive potential of M. viridis. A high reproduction rate of the polychaete is needed to 216 5, ~e lS lS ~0 rs re V!. of :ie :1g as ~le =>4 lis >ff tOt ds at he he he aa nd ga >nS m ve to 20 u 0 ~ 15 ~ s 10 (!) E-< 5 IV ,o- .• -. /-+-:: ;-. c· IJ- • .,....Y--·..-; :+ •• ' ' ' ' ,; ,; :.. t) ... /~· V VI --:+- 1992 - -o - 1993 n.---·- ..... ·.0 •• "...0·-··-··()."') - •• • :- • • -fJr- 1994 ' . ...... . vn VIll .... '.- -o - 1995 'c.._ • • "o IX ". ' "' '.. ... ' .. ·,'o. ... , .. ... . .. ..... X Fig. 4. Seasonal fluctuation in temperature in the Gulf of Riga in 1992-95. XI exceed the critical abundance of juveniles in order to successfully invade new areas. The reproduction of M. viridis occurs in the Baltic in September-October (Zettler et al., 1995). Dominant wind directions at the gulf during this time of the year are from south-west or south (Berzinsh, 1995) assuring the dispersion of the myriad of larvae of M. viridis into Estonian coast and the establishment of a permanent community. Already at the beginning of the 1990s the larvae of a polychaete (probably M. viridis) were observed in the plankton samples from the northeastern part of the Gulf of Riga (M. Simm, pers. comm.). This finding is in disagreement with our data, as not a single individual of M. viridis was found in the benthic communities before 1995. As salinity did not limit the expansion of the distribution area of M. viridis in the 1990s it is possible that lower temperatures had a negative impact on the development and settlement of the larvae of M. viridis or the abundance values of the larvae were too low to colonize the , northern part of the Gulf of Riga. As compared to the figures presented by Lagzdins & Pallo (1994) for the Latvian side of the Gulf of Riga in 1990, the abundances here are rather low, hardly surpassing 100 ind m-2. Though, one may assume that the carrying capacity of an environment is far higher than the abundances recorded during the first year of colonization (i.e. in 1995). Therefore higher abundances may be recorded in the following years. In Estonian waters M. viridis inhabits similar bottom types (i.e., muddy, sandy, or gravelly sediments) as previously reported in the Baltic and North seas (Atkins et al., 1987; Bick & Burckhardt, 1989; Essink & Kleef, 1993; Lagzdins & Pallo, 1994). As a difference to Bick & Burckhardt (1989), the abundance and biomass values of M. viridis differed significantly between sediment types. 217 Several authors have suggested the presence of interspecific competltwn between two polychaetes, Nereis diversicolor and M. viridis (Essink & Kleef, 1993; Lagzdins & Pallo, 1994), though the relationship was not found to be statistically significant. In the Gulf of Riga the increase in the abundance of M. viridis did not coincide with the decline of N. diversicolor. The region inhabited by M. viridis in the Väinameri Archipelaga area is characterized as a very homogeneaus in sediment type and algal community (i.e. sandy clay sediments are covered with a layer of red algae, Furcellaria lumbricalis (Huds.) Lamour). Among macrozoobenthos practically only Macoma balthica, Cerastaderma lamarcki, and N. diversicolor are present in these sediments, while Mytilus edulis L. is more abundant in the Furcellaria layer. Because of the uniformity and low number of species, this community is more vulnerable to an intruder such as M. viridis. In the shallow, patehier environment of the Väinameri Archipelaga area, M. viridis has not yet been observed, and as usual, N. diversicolor dominates numerically among polychaetes. As the benthic primary production of the Väinameri Archipelaga area is relatively high due to the shallowness of the area, and the growth rate of M. viridis is strongly influenced by food quality and quantity (Bochert & Bick, 1995), supposedly this region will be the most densely populated by M. viridis in the next couple of years. One may already notice higher biomasses (though not significantly higher yet) in the Väinameri Archipelaga area. Consequently, this region may be regarded as the most probable source of M. viridis for the southern side of the Gulf of Finland. As a conclusion, we can say that M. viridis has not yet reached its stable biomass values in the study area. Up to now M. viridis has colonized practically the whole possible substrate in the Gulf of Riga and the Väinameri Archipelaga area. At the sametime the species was found only at some places in the Gulf of Finland, including the deeper areas in the western part of the gulf and Tallinn Bay. M. viridis has invaded the former area possibly via the Väinameri Archipelaga area, whereas the occurrence of this polychaete in Tallinn Bay is probably connected with the shipping activity. We predict that M. viridis will form a permanent community in the western and central parts of the Gulf of Finland in the next two years. ACKNOWLEDGEMENTS This study was partly financed by the Nordic Council of Ministers and EU project MAST-III. We would like to thank Mrs. Tiia Rosenberg for her valuable help in performing the Iabaratory analysis. We owe special thanks to Mrs. Anne Timm, who kindly provided us the temperature data of the Gulf of Riga. 218 011 ef, be of on s a ay s.) --:a, ile he an he al, IS of :k, m tot llS :rn >le lly go of [111 IS ill of u le REFERENCES Atkins, S. M., Jones, A. M. & Garwood, P. R. 1987. The ecology and reproductive cycle of a population of Marenzelleria viridis (Annelida: Polychaeta: Spionidae) in the Tay Estuary. Proc. R. Soc. Edinb., 92B, 311-322. 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Marenzelleria viridis'e LEVIK JA INV ASIOONIÖKOLOOGIA EESTI RANNIKUMERES Jonne KOTTAja Ilmar KOTTA On käsitletud Marenzelleria viridis' e elukeskkonna eelistusi ja invasiooni-ökoloogiat. Liigi arvukus ja biomass on Eesti rannikumeres veel väikesed. M. viridis eelistab kruusaseid p5hju. Homogeensetel merealadel kaasneb M. viridis' e arvukuse suurenemisega teise hulkharjasussi Nereis diversicolor arvukuse vähenemine. Heterogeensetel aladel seilist seost ei täheldatud. Lähi-tulevikus v5ib M. viridis muutuda domineerivaks liigiks Eesti rannikumeres. 220 

Distribution and invasion ecology of Marenzelleria viridis in the Estonian Coastal Waters

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