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Annual Growth of the Cockle Clinocardium Ciliatum in the Norwegian Arctic (Svalbard Area)

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Annual Growth of the Cockle Clinocardium Ciliatum in the Norwegian Arctic (Svalbard Area) ... Hydrobiologia 440: 331-338, 2000. 331 M.B. Jones, J.M.N Azevedo, A.!. Neta, A.C. Costa & A.M. Frias Martins (eds), Island, Ocean and Deep-Sea Biology. '' © 2000 Kluwer Academic Publishers. Annual growth of the cockle Clinocardium ciliatum in the Norwegian Arctic (Svalbard area) Minna Elisabeth Tallqvist1 & Jan Henry Sundet2 1Department ofBiology & Husă Biologica! Station, Ăbo Akademi University, FIN-20500 Ăbo, Finland 2Norwegian Institute of Fisheries and Aquaculture, N-9002 TromsrjJ, Norway Key words: Bivalvia, growth, Arctic, Clinocardium ciliatum, Svalbard Abstract The Svalbard Islands are influenced by warm Atlantic water in the south and west, and cold Arctic water in the east. lce cover, and hence the location of the highly productive marginal ice zone, varies both intra and interannually. Part of the primary production accumulates on the bottom and is utilized by the benthos. In this study, the annual growth of the cockle Clinocardium ciliatum (Fabricius, 1780) from three sites in Svalbard waters is reported. Moffen, the site in the north (80° Ol' N, 13° 48' E) is located in the northernmost areas influenced by Atlantic water. The Storfjorden site (77° 10' N, 20° 09' E) is situated in cold Arctic water masses, and the Bear Island site (74° 50' N, 18° 54' E) is in the Polar front area where Atlantic and Arctic water masses meet. Annual growth of cockles was ana1ysed retrospectively by measuring externa! growth increments, which gave annual growth records from the 1970s to 1996. Shell height for age for different year classes was highest at the Storfjorden site, and lowest at Bear Island. Periods of high growth occurred at Storfjorden and Bear Island during the 1980s while the beginning of 1990s was characterized by low growth. At Moffen, growth was more variable between single years. Severa! factors are influencing the growth of C. ciliatum in the Svalbard area and growth cannot be coupled to only one environmental factor like ice cover. Introduction In the Arctic, suspension feeders are subjected to large seasonal and annual variations in food supply, Generally, bivalve growth is dependent upon age and and the marginal ice zone is considered to have a size, but is affected also by environmental factors such great impact on the primary production in these areas as temperature, current velocity and food availability (Wassman et al., 1991). In our study, we sampled (Richardson et al., 1980; Appeldoorn, 1981; Mac Don­ cockles from three sites with different characteristics ald & Bourne, 1987; Jones et al., 1989; Navarro et al., of ice coverage and water masses. Moffen, in the north 1992; Arsenault et al. 1997; Witbaard et al., 1997), and of Spitsbergen Island, is located in the marginal ice biotic factors such as competition (Peterson & Andre, zone where ice conditions vary even over short peri­ 1980). In Arctic areas, food supply is probably one of ods of time (days/weeks); Storfjorden, on the east of the most important factors affecting growth of benthic Spitsbergen Island, has the longest period of ice cover animals and species are forced to adjust to a short and is situated in Arctic water masses; Bear Island growth season (Picken, 1979; Sagar, 1980; Clarke, is situated in the Polar front area where Atlantic and 1988). Polar bivalves grow slower than similar spe­ Arctic water masses meet (Fig. 1). We hypothesized cies in more temperate areas (Ralph & Maxwell, 1977; that long periods of annual ice may decrease growth Clark, 1983) and the growth in winter can be close to in benthic organisms due to depressed primary produc­ zero (Richardson, 1979). In a study of the bivalve Ma­ tion compared with areas with less ice. Consequently, coma balthica, growth was more dependent on food sites influenced by Atlantic water will show higher supply than on temperature (Beukema et al., 1985) and growth rates than sites in Arctic water masses. It has the same has been shown for other species living in been suggested that there is a strong coupling between environments with marked seasonality (Clarke, 1988). the pelagic and the benthic environment in the Arctic 332 below 5 o C. Together with another common cockle species in this area, Serripes groenlandicus, it is im­ portant as food for walrus and seals (Fay & Burnes, 1988). 80.J Description of sampling sites Generally, Arctic waters are characterized by high sea­ sonality (due to the light regime) with a short growth season and annual variation in ice coverage. The polar night (i.e. when the sun is constantly under the ho­ rizon) is from 22nd October to 20th February at the northernmost site and from 6th November to 5th Feb­ ruary at the southernmost site. The midnight sun (i.e. when the sun is constantly above the horizon) is from 15th April to 27th August at the northernmost site and from 30th April to 12th August at the southernmost site. In the Svalbard area, the maximum ice extent is in March and the minimum in September. Moffen (80° OI' N, 13° 48' E) was sampled on 7th September 1997 at 112-130 m water depth (Fig. l ). This site is dominated by gravei and scallop beds, and C. ciliatum were found only in hauls where clay was the domin­ ant sediment. Of the 64 individuals sampled, 63 were intact and used in the analyses. Probability for ice in March is about 70-80% (Vinje & Kvambekk, 1991). The Storfjorden site (77° 1O' N, 20° 09' E) was 700 sampled on 14th September 1997 at a sampling depth of 80-118 m. This site was dominated by clay and 162 cockles were sampled, of which 154 were intact Figure 1. A map of the study area. Ali sites (indicated with black and used in the analyses. The probability of ice cover dots) are situated in Svalbard lsland waters with Moffen in the in March at Storfjorden is 100% (Vinje & Kvambekk, north, Storfjorden in the east of the island Spitsbergen, and the 1991). southernmost site north of Bear Island. Bear Island site (74° 50' N, 18° 54' E) was sampled on 1lth November 1997 at a depth range of 100- due to a slow response of zooplankton to increased 130 m. The substratum was clay with a fraction of primary production in the spring (Grebmeier et al., empty cockles; 146 cockles were sampled, of which 1988; Peinert et al., 1989). 143 were included in the analyses. The probability of In this paper, the annual growth of the com­ ice cover in March at Bear Island is about 30% (Vinje mon Arctic cockle, Clinocardium ciliatum (Fabricius, & Kvambekk, 1991). 1780), is compared between three Arctic sites in the Svalbard area. At each sampling site, bivalve growth patterns are linked to ice coverage. Clinocardium cili­ Materials and methods atum is one of the common larger bivalve species in the Svalbard area (Rozycki, 1987). It prefers clay Bivalves were sampled with a triangulardredge (open­ sediment (Ockelmann, 1958) and is found in a wide ing 1 m*1 m*1 m, inner net size 1 cm) from the depth range from the intertidal down to about 500 m research vessel 'Jan Mayen' (Troms0 University) dur­ (Bernard, 1979). Maximum size may be up to 70 mm ing two cruises in 1997. Sediment was sieved through (Bernard, 1979), but cockles of up to 40 mm are most a 5 mm mesh and C. ciliatum were picked out. Cockles common in our study area. It is a circumpolar species were washed and shell height was measured with ver­ of Pacific origin and is found at water temperatures nier callipers to the nearest 0.01 mm. The body was 333 mean. In this way, different year classes of bivalves a) 30 can be analysed together and growth at different calen­ dar years can be compared. Standardization of growth 20 rings is a method widely used in dendrochronology 10 (Fritts, 1976; Schweingruber, 1989) and has also been applied in growth analysis of marine bivalves (Wit­ o ~ baard, 1996a,b ). Standardization is do ne by comparing b) ~ 30 expected values with actual values. Expected values v" of growth (increment size) can be calculated by dif­ '- 20 o ferent methods, most of them based on curve fitting. .,~ .o 10 The time series of indices describe higher or lower E z" growth than expected for different years. The stand­ o ardization was done only from the 6th increment (i.e. c) 30 the largest increment). Therefore, the standardization was done only for ages with decreasing increment size 20 and the increase in growth in young ages (0- 6) was 10 excluded from the analyses. Studies in dendrochrono­ logy show that the period of years with increasing o growth at younger ages is usually not very sensit­ 7 9 Il 13 15 17 19 21 23 25 27 29 31 33 ive to environmental factors and these years are often Age (years) excluded to facilitate the standardization calculations (Fritts, 1976). We present indices only for the time Figure 2. Age distribution of Clinocardium ciliatum from (a) Mof­ fen, (b) Storfjorden and (c) Bear lsland. period 1980- 1994, excluding the 1970s because of the low number of cockles representing this time period. The ice data are presented as the number of days separated from the shell and the shells were stored with more than 70% ice coverage at the sites between for further analyses. Age determination was done by 15th March to 1st October (1980-1994). counting externa! growth rings on the shells. This species has very marked externa! growth rings, and these were assumed to be formed annually due to the Results pronounced seasonality in the area and on the basis of results from other polar mollusc species (Ralph The age distribution of the bivalves sampled varied & Maxwell, 1977; Richardson, 1979; Picken, 1980; between sites (Fig.
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