Aquatic Ecology 32: 313322, 1998. 1998 Kluwer Academic Publishers. Printed in the Netherlands.

313

Settlement and growth of the green mussel Perna viridis (L.) in coastal waters: inuence of water velocity
S. Rajagopal1 , V. P. Venugopalan2 , K. V. K. Nair2 , G. Van der Velde1 and H. A. Jenner3
1 Department

of Ecology, Laboratory of Aquatic Ecology, University of Nijmegen, Toernooiveld, 6525 ED Nijmegen, The Netherlands (E-mail gerardv@sci.kun.nl); 2 Marine Biology Programme, Water and Steam Chemistry Laboratory, BARC Facilities, Kalpakkam 603 102, India; 3 KEMA Power Generation, P.O. Box 9035, 6800 ET Arnhem, The Netherlands
Accepted 20 November 1998

Key words: Perna viridis, ow velocity, growth rate, larval occurrence, spat settlement, spawning periods

Abstract Green mussels Perna viridis were observed to be a major foulant in the seawater intake tunnel of a coastal power station. Field experiments were carried out to ascertain what factors were responsible for the successful colonisation by mussels. Two adjacent stations (25 m apart) were selected, one representing the coastal waters and the other representing the intake screens (with higher water velocity). Gonadal activity, larval abundance, spat settlement and growth rate of the mussels were monitored at monthly intervals for a total period of two years. The results showed that the breeding activity of the mussels at the study area is inuenced largely by temporal distribution of seawater temperature. However, ensuing larval availability in the coastal waters is more dependent on food availability. On the other hand, spat settlement and growth rate are predominantly inuenced by water ow, probably as a result of increased propagule and food ux rate at higher water velocities. Higher water velocity at the intake screens also contributed to mussel dominance by preventing settlement of many potential competitors.

Introduction Green mussels, Perna viridis are widely distributed in the Indo-Pacic region; their distribution extends from Japan to New Guinea and from the Persian Gulf to South Pacic Islands (Siddall, 1980). They are a characteristic species of midlittoral and sublittoral zones where they often constitute dense populations on rocky substrata. In spite of their wide distribution and their importance in the ecology of rocky shore ecosystems, detailed works on their biology are only a few (Lee, 1985). These mussels are also important from the point of view of animal protein for human consumption and some aspects of their biology relevant to shery and culture have, therefore, been studied by other workers (Qasim et al., 1977; Sivalingam, 1977; Parulekar et al., 1982; Rivonkar et al., 1993; Rajagopal et al., 1998). However, these mussels also deserve serious attention on account of their potential to foul industrial cooling systems us-

ing seawater (Rajagopal et al., 1997). On an earlier appraisal (Rajagopal et al., 1991), it was found that out of 570 tons of fouling debris lodged inside the concrete intake tunnel of a power station, P. viridis alone constituted 411 tons. Since this was the rst time such massive colonisation of marine mussels has been observed in the cooling circuits of an Indian power station, we were interested to know what aspects of the ecology of the mussels make them such successful colonisers. This was important as earlier workers who studied fouling phenomena on the east coast had not indicated the dominance of P. viridis among the natural sessile communities (Paul, 1942; Daniel, 1954; Renganathan et al., 1982; Rao, 1990). Moreover, information regarding the breeding activity of P. viridis from different localities of the Indian peninsula was inconsistent. Larval availability and growth rate are important parameters inuencing successful colonisation by sessile species. Earlier workers had indicated the importance of water ow on the population ecol-

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Figure 1. (a) Map showing the Kalpakkam. (b) Schematic representation of the Madras Atomic Power Station seawater intake tunnel showing 2 sampling stations (not drawn to scale).

ogy of mussels (Nixon et al., 1971; Perkins, 1974; Venugopalan et al., 1991; Wildish & Kristmanson, 1997). It was possible that greater water ow experienced within the cooling water circuit could be responsible for the successful colonisation of mussels. Bearing this in mind, the population ecology of P. viridis was monitored at two locations which represented (a) their natural habitat (coastal waters) and (b) cooling intake point of the power station experiencing high water velocity. The present study investigates whether ow regimes are signicant in inuencing the population ecology viz., growth rate, breeding activity and spat settlement of P. viridis in coastal waters of Kalpakkam, east coast of India.

Figure 2. Seasonal variations in the hydrographic parameters ((a) temperature, (b) salinity, (c) dissolved oxygen and (d) chlorophyll-a) in Kalpakkam coastal waters from April 1988 to March 1990. Data are presented as mean SD.

Materials and methods Site description Kalpakkam is situated (12 32 N and 80 11 E) about 65 km south of Madras (Figure 1a). Madras Atomic Power Station (MAPS), Kalpakkam is a seawater cooled station, and uses a 468 m long sub-seabed tunnel to draw cooling water (35 m3 s1 ) for its twin (2 235 MWe) reactors (for details refer Rajagopal,

315 1997). The seawater ows by gravity from the intake (Figure 1b) via the tunnel to the forebay pump house, from where it is pumped (12 pumps) to the condensers. The coolant seawater ow in the tunnel when all the 12 pumps are running, works out to be about 3 m s1 (Madras Atomic Power Station Design Manual, 1975). The intake point is guarded by steel weld mesh screens to prevent the entry of large objects into the cooling circuit. Sampling stations Two stations were selected for the study (Figure 1b). Station 1 represents the coastal waters which is 8 m deep and experiences coastal currents of velocity in the order of 0.20.3 m s1 . Station 2 is the seawater intake point which is characterised by high water velocity (as high as 3 m s1 , depending on the ow). The distance between Sta 1 and Sta 2 is about 25 m. The physicochemical characteristics of the water are, therefore, identical at both stations, except for water velocity. Hydrographical features Hydrographical features of the study site were studied by collecting surface water samples at fortnightly intervals during the period April 1988 to March 1990. Parameters like temperature, salinity, dissolved oxygen (DO) and chlorophyll-a were monitored (Strickland & Parsons, 1972) to understand their variation and possible inuence on the breeding pattern and growth rate of the mussels. Gonad observations Mussels were collected every month (April 1988 to March 1990) from the two stations and were used for gonadal studies. About 6065 mussels (approx. 3040 mm shell length) were collected from each station. In the laboratory, the gonadal tissues were removed from the mantle lobes and xed in Bouins uid for 24 h and later transferred to 40% alcohol. Sections (1015 m) were made from wax-embedded tissues and stained with haematoxylin and eosin. The method of Seed (1969) was adopted to categorise the gonads into four groups viz., spent/resting, developing/redeveloping, ripe and spawning. Gonad index (GI) was also determined based on the method described by King et al. (1989). Larval abundance Mussel larvae were concentrated from 500 l of seawater using a 22 m mesh net (De Wolf, 1973), every month from May 1988 to May 1990. The larvae were subsequently xed in 5% buffered formalin and counted in a Sedgwick rafter counter. Spat settlement Concrete blocks (20 20 20 cm) were used to sample spat fall in coastal waters (Sta 1), as described by Nair et al. (1988) and Rajagopal et al. (1997). Three test blocks were suspended at 1 m, 4 m and 7 m using nylon ropes and retrieved after 30 d to estimate spat fall. At Sta 2, spat samples were collected from the steel intake screens at 2 m, 4 m and 6 m. Earlier trials had shown that mussel settlement on steel surfaces were comparable to those on concrete. The samples (in triplicate) were collected at each depth, and the data were averaged and presented as numbers dm2 (dm2 = 100 cm2 ). Growth rate Growth rate measurements were initiated by suspending test blocks at 1 m (Sta 1) at the beginning of spat settlement (April 1988). Every month about 30 mussels were randomly collected from the concrete blocks (1 m depth at Sta 1) and from the intake screens (2 m depth at Sta 2). Collection of mussels from test blocks involved sampling of a different subset of the mussel population every month. In order to monitor the growth increment of the same population over a period of time, mussels (11 0.6 mm shell length, n = ca. 100) were conned in cages (75 75 75 cm, 0.5 cm mesh size) and left suspended at 1 m depth at Sta 1 for 375 d (Page & Hubbard, 1987). Their shell growth increment was monitored at monthly intervals during the period October 1988 to September 1989. Statistical analysis A 2-factor analysis of variance (ANOVA) was used to examine variability in gonad index and spat settlement of P. viridis taking account of season (sampling time) and station as two independent variables (Sokal & Rohlf, 1981). Spat settlement of P. viridis at different depths was tested by 1-factor ANOVA. For post-hoc comparison of monthly means, we used student t-tests for comparison of two means and Student-NeumanKeuls (SNK) tests for comparison of multiple means

316 (Zar, 1984). Prior to the analysis the data were tested for normality and homogeneity of variance. with spawning and GI data, with two peaks in a year, viz., AprilJune and OctoberNovember. The second peak was much lower and occurred during a shorter period when compared to the rst. The spat fall also showed close correspondence with larval availability, with two peaks per annum (Figure 6). The October peak, as in the case of larval abundance, was much smaller when compared to the May-June peak. Settlement of P. viridis varied signicantly with respect to season (ANOVA, df = 23, F = 99.813, P < 0.001) and station (ANOVA, df = 1, F = 128.886, P < 0.001, Table 1). P. viridis settlement was considerably higher in 1988 when compared to 1989. The data also showed markedly higher spat settlement at Sta 2, as compared to Sta 1. The maximum spat fall recorded was 5224 numbers dm2 at Sta 2, in May 1988. Depth-wise differences in settlement were negligible (ANOVA, df = 2, F = 0.332, P = 0.72) at the intake point (Table 2), whereas at Sta 1, settlement was maximum at 4 m, followed by 1 m and least at 7 m (ANOVA, df = 2, F = 8.966, P < 0.001). Growth rate Signicant differences in growth rate were found between Sta 1 and Sta 2 (ANOVA, df = 1, F = 5.271, P < 0.05, Figure 7). At Sta 1 mussels grew at the rate of 13 0.5 mm in 30 d up to 98 0.7 mm in 370 d. At the same time, mussels growing at Sta 2 exhibited maximum growth of 27 1.1 mm in 49 d up to 119 2.1 mm in 375 d. The difference in size of the mussels at the end of the period at the two stations was significant (t-tests, P < 0.001). A statistical comparison of the cage data (11 mm size group) with test block data, indicated that at the end of one year the size of caged mussels was not signicantly different from that of test block mussels (t-tests, P > 0.05).

Results Hydrographical features Surface water temperature ranged from 25.9 C (December 1989) to 31.3 C (October 1989) during the study period (Figure 2a). It was characterised by two well dened maxima, one occurring during May/June (30.9 C) and the other in October (31.3 C). Salinity ranged from 27.0% to 35.5%, with a maximum in May/June and a minimum in November (Figure 2b). Dissolved oxygen of surface water varied from 4.19 mg l1 (June 1989) to 6.24 mg l1 (September 1988) (Figure 2c). A denite pattern of seasonal change in dissolved oxygen concentrations was not discernible. Chlorophyll-a ranged from 1.06 mg m3 to 9.99 mg m3 (Figure 2d). Maxima were found in May and minima in November. The chlorophyll values peaked during MayJune months, indicating greater availability of phytoplankton in the coastal waters. However, NovemberDecember months saw a general decrease in chlorophyll values; there was no chlorophyll increase corresponding with the second temperature peak observed in October. Spawning, larval abundance and spat settlement Histological data showed that spawning of P. viridis at Kalpakkam started by March and peaked in May (Figure 3). Extensive spawning during this period was indicated by a large proportion of spawning mussels in May and presence of spent gonads in June and July samples. The latter also corresponds with a decrease in GI in July (Figure 3). The GI values increased in August and September, due to the large proportion of redeveloping gonads. The second spawning season peaked around October and was followed by a short cessation of spawning activity in the population. Gonad index was positively correlated with the temperature (Sta 1: y = 839.1 + 34.4x , r2 = 0.82, P < 0.001; Sta 2: y = 1100.2 + 43.4x , r2 = 0.90, P < 0.001, Figure 4). A comparison of GI did not show any signicant difference between Sta 1 and Sta 2 (ANOVA, df = 1, F = 2.787, P = 0.10, Table 1). Larval abundance in the coastal water uctuated widely during the study period (Figure 5). The maximum (39532 larvae m3 ) was recorded in May 1988. The data, in general, showed good correspondence

Discussion Prior to our studies on the massive colonisation of the seawater intake system of MAPS (Rajagopal et al., 1991), there are no published reports of large-scale fouling by P. viridis in cooling water circuits of the tropical Indo-Pacic. Huang et al. (1983) have reported P. viridis to be an important fouler of ship hulls, piers, buoys and rafts in Hong Kong waters (see also Lee, 1985; Cheung, 1993). Most of the earlier studies on P. viridis from Indian waters (Rao et al., 1975; Qasim et al., 1977; Narasimhan, 1980; Parulekar et al.,

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Table 1. Results of ANOVA testing whether gonad index and spat settlement of the green mussel, Perna viridis were different at Sta 1 (coastal waters) and Sta 2 (intake screens) during April 1988 to March 1990 Source of variation Gonad index Station Time Station Time Error Spat settlement Station Time Station Time Error df SS MS F P

1 23 23 48

173.34 467003.99 16551.91 2985.50

173.34 20304.52 719.65 62.20

2.78 326.45 1.57

0.102 0.001 0.001

1 23 23 96

5134000.69 91446540.64 23705459.97 3824044.67

5134000.69 3975936.55 1030672.17 39833.80

128.89 99.81 25.87

0.001 0.001 0.001

Figure 3. Seasonal variations of gonadal maturity stages (%) and gonadal index of Perna viridis in coastal waters and intake screens of power station in Kalpakkam.

1982; Rajagopal et al., 1998) have been related to the shery aspects. This necessitated a study on important aspects of the population ecology of P. viridis, which

are relevant to its potential to colonise cooling water circuits.

318 Cheung (1993), working on the population dynamics of P. viridis in the Tolo Harbour, Hong Kong, reported two recruitment periods per year: one from July to September and another from November to March. Lee (1985), who worked in the Victoria Harbour (Hong Kong) population, reported a single recruitment period for P. viridis which extended from June to September. Our data, which includes gonadal observations (Figure 3), larval abundance (Figure 5) and spat settlement (Figure 6), clearly indicate two recruitment periods for P. viridis at Kalpakkam. The rst one extends from April to June and the second from September to October, the latter being less intensite. A comparison of the present data with the data from other parts of the Indian coast (Figure 8) indicate that breeding activity of this species could vary substantially within narrow geographical regions, as observed in Hong Kong waters by Lee (1985 and 1988) and Cheung (1993). Many of the published literature regarding P. viridis reproductive behaviour on the east coast of India (refer Figure 8), are based on spat settlement data (Godwin, 1980; Karande et al., 1983; Nair et al., 1988), which may not give a correct picture regarding gonadal activity. Some of the nearby areas (Edaiyur backwaters and Kovalam, Figure 1a) experience hydrographic conditions (especially salinity) signicantly different from those at Kalpakkam, which is an open coast experiencing typical marine conditions for most part of the year. This could have been a probable cause of the differences in reproductive behaviour of P. viridis at these sites. In the case of the blue mussel Mytilus edulis (L.) Bayne and Worrall (1980) have also reported two spawning periods at Lynher and a single one at Cattewater in the UK. Obviously, local hydrographical conditions (including pollution levels) and food availability may act key factors for a reproductive strategy (Lee, 1985). Though the spawning activity of the mussels growing at Sta 1 and Sta 2 exhibit two clear peaks per annum, which are roughly comparable to each other in terms of intensity, the ensuing larval concentration in the water shows only one prominent peak, corresponding to the rst spawning activity. A possible explanation for this is as follows. The gonadal development and spawning activity of P. viridis at Kalpakkam is linked with the water temperature which shows two clear peaks, one in April-June and another in October. Gonadal index matches with seasonal uctuations of temperature (Figure 4). However, larval abundance in the coastal waters is probably decided largely by food availability. Chlorophyll-a, which is used here

Figure 4. The relationship between the gonad index of Perna viridis and water temperature in Kalpakkam coastal waters. Table 2. Data on spat settlement of Perna viridis at different depths at Sta 1 and Sta 2 during April 1988 to March 1990. One-way ANOVA followed by multiple comparison tests (SNK tests) were used to determine whether depthwise spat settlement of mussels differed signicantly at Sta 1 and Sta 2 Coastal waters Depth 1m 4m Mean Maximum Minimum Median SD SE Lower 95% Cl Upper 95% Cl ANOVA 314 1734 0 37 524 112 82 546 517 2486 0 95 783 167 170 865 Intake screens Depth 2m 4m 641 5225 0 138 1172 234 158 1125 601 4512 0 120 1191 238 109 1092

7m 68 529 0 14 129 28 13 127

6m 687 4779 0 150 1258 252 168 1207

F = 8.966, P < 0.001

F = 0.332, P > 0.05

as an indicator of phytoplanktonic food availability, shows only a single summer peak in AprilJune. A rise in water temperature in October is not associated with a phytoplankton bloom. We hypothesise that food acts as a limiting factor for the growing larvae, resulting in low numbers of mussel larvae in the coastal waters following the second spawning in October. However, this would need conrmation by additional experimentation. The larval densities observed during the April June peak were comparable with data reported from elsewhere. The highest density recorded in the present study was 39 532 larvae m3 . Schram (1970) recorded a similar (40 000 larvae m3 ) gure for M. edulis from the Oslofjord. Hopkins (1977) showed that lamellibranch larvae in Tampa Bay ranged from 1200 to 15 500 larvae m3 , with an annual mean of 8000

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Figure 5. Seasonal occurrence of mussel larvae in Kalpakkam coastal waters.

Figure 7. Growth rate of Perna viridis in coastal waters, intake screens of power station and experimental cages (initial shell length 11 0.6 mm, n = 100, suspended in coastal waters at 1 m depth) in Kalpakkam. Data are presented as mean SD.

Figure 6. Monthly variations of mussel spat settlement at different depths in coastal waters and intake screens of power station in Kalpakkam.

larvae m3 . Fish and Johnson (1937) reported peak abundance of more than 25 000 larvae m3 in the Bay of Fundy. The present data indicate signicant variations in the settlement intensity of P. viridis at the two stations

(Table 2). Maximum settlement was observed at Sta 2. The high settlement intensity on the intake screens (Sta 2) is explicable on account of the high ow rate, which would enhance the propagule ux rate to the substratum. Though the water velocity is high at this station, mussel larvae are capable of settling at such high water velocities (Neitzel et al., 1984; Rajagopal, 1997). Neitzel et al. (1984) reported that at velocities as high as 3.5 m s1 , mussels could settle and colonise new surfaces. Depth-related variation in intensity, which is obvious at Sta 1 is lacking at Sta 2. At Sta 1, the settling plantigrades clearly prefer the intermediate depth (4 m), when compared to the near-

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Figure 8. Breeding periodicity of green mussel, Perna viridis (L.) reported from Indian waters. Thick lines represent intense reproductive activity ( gonadal observations; spat settlement data).

surface (1 m) (SNK tests, P < 0.05) or near-bottom (SNK tests, P < 0.001). This behaviour is probably due to the subtidal habitat of these mussels. Depthwise differences in spat fall are likely if the larvae are non-uniformly distributed in the water column or alternatively, the settling larvae prefer discrete light regimes. However, this depth-related feature was not apparent at Sta 2, owing probably to the turbulent water ow which would disturb any vertical distribution of larvae. An analysis of the data presented in Figure 9 shows that, apart from P. viridis, not many species were able to colonise the intake screens, apparently due to the high velocity regime. The high velocity would permit settlement of only those larval forms, such as mussels, which have the ability to withstand a high shear force (Neitzel et al., 1984). This would have increased mussel dominance at Sta 2 not only directly, but also indirectly, by reducing competition. In our study, a maximum growth size of 119 2.1 mm shell length was recorded in the rst year for P. viridis growing on the intake screens. This value is much higher than the values reported from elsewhere in India: 93 mm from Kakinada on the east coast

(Narasimhan, 1980) and 96 mm from Goa on the west coast (Rao et al., 1975; Rivonker et al., 1993). We also observed that mussels kept conned in cages in the coastal waters reached the same size as those on test blocks after a period of one year growth (Figure 7). The increase in growth rate at Sta 2 is, probably due to increased food ux rate caused by the high ow rate. The overriding importance of food supply in mussel growth rate has been emphasised by Seed (1976), Seed and Suchanek (1992) and Wildish and Kristmanson (1997). Other authors who studied growth rates of P. viridis include Lee (1985, 1986) and Cheung (1993). Lee (1986) recorded low (5 mm month1) growth rates in the polluted Victoria Harbour, Hong Kong. Similarly, Cheung (1993) reported a growth of 49.7 mm in one year from Tolo Harbour, another polluted harbour in Hong Kong. When compared to values reported earlier (Vakily, 1989), the growth rate recorded at Sta 2, in comparison, stands apart as the gure (10 mm month1) is the highest ever reported for P. viridis.

321 recorded at Sta 2 (high water velocity) is the highest ever reported for P. viridis.

Acknowledgements We express our sincere thanks to S. V. Narasimhan and S. B. Kuppuraju for the facilities and support. Thanks are due to J. M. van Groenendael and B. Kelleher for suggestions. Thanks are also due to C. van der Rijt for his assistance with the statistics. This work was supported by a grant from the Board of Research in Nuclear Sciences, Department of Atomic Energy, Government of India.

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Figure 9. (a) Species diversity of the sessile community colonising concrete test blocks at Sta 1 (initiated on 9th April 1988 at 1 m, 4 m and 7 m) and on intake screens at Sta 2 (initiated on 9th April 1988 at 2 m, 4 m and 6 m). Monthly mean values were calculated from the three sampling depths at each station. (b) Major taxa are those whose percentage cover exceed more than 10%.

Conclusions (1) Temperature plays a key role in inuencing the reproductive activity of P. viridis in Kalpakkam coastal waters. (2) Though intense spawning of the green mussel population occurs two times a year, ensuing larval availability is high only during one season (summer) and subdued during the second season. Reduced availability of food during second spawning season is a probable reason for this low larval availability. (3) Higher water velocity has no signicant effect on reproductive activity, but it profoundly enhances spat settlement on surfaces. (4) Higher water velocity prevented many potential colonisers (competitors for mussels) from settling at Sta 2, thereby increasing mussel dominance. (5) Increased velocity signicantly increases mussel growth rate. The growth rate (10 mm month1)

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