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Distribution and Growth of the Keyhole Limpet <I>Fissurella Barbadensis

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Distribution and Growth of the Keyhole Limpet <I>Fissurella Barbadensis DISTRIBUTION AND GROWTH OF THE KEYHOLE LIMPET FlSSURELLA BARBADENSIS GMELIN JONET WARD The Bel/airs Research Institute of McGill University, St. James, Barbados, West Indies ABSTRACT The vertical and horizontal distributions of F. barbadensis on the rocky shores of Barbados are described. Growth rates are obtained and the effects of growth and environmental variation on some of the structural features of the shell are examined. The results are compared with the growth, distribution, and shell variations of temperate climate limpets. INTRODUCTION The genus Fissurella is found throughout temperate and tropical seas. Many species live below low tide level while others inhabit deep water or the intertidal zone (Pilsbry, 1890; Farfante, 1943; Zuniga, 1951; Gauld & Buchanan, 1959; Warmke & Abbott, 1961). F. barbadensis ranges from southeast Florida and Bermuda to the West Indies and from Mexico to British Guiana (Farfante, 1943). Lewis (1960) describes this species in Barbados living Iowan the shore from mean low water to mean sea level. There appear to be no published studies on the growth of fissurellids. However, growth rates of various Patellacea have been obtained for Patella vulgata (Russell, 1909; Orton, 1928), Acmaea dorsuosa (Abe, 1932), Patel/oida conulus (Hamai, 1937), and Patina pellucida (Graham & Fretter, 1947). The effects of growth and variation in environment on shell structure have been examined in several limpets. In A. limatula dif- ferent intertidal habitats produce variations in shell weight, in the volume of the extra-visceral space between the shell and the soft parts, in the heart rate, and in the condition of the gonads (Segal, 1956a, b). The shell height in Patella vulgata (Orton, 1928; Moore, 1934) and Acmaea dorsuosa (Abe, 1932) is greater in dry, more exposed, habitats than in damp ones. The shells are also thicker in a dry habitat. An attempt has been made to compare the tropical species, F. barbadensis Gmelin, with temperate limpets in respect to distribution, growth, and the effects of different environments on some of the structural features of the shell. The work was supported by a grant from the National Research Council of Canada. The author is very grateful for the suggestions and encourage- ment given by Dr. J. B. Lewis of the Bellairs Research Institute. My thanks also go to Dr. G. T. Ward, of the Brace Research Institute of Mc- Gill University, for discussions on some of the mathematical procedures. 300 Bulletin of Marine Science [17(2) MATERIALS AND METHODS Distribution.-The density of F. barbadensis was measured at 12 stations around the island of Barbados by counting the specimens in areas 1 m square. The average daily period of exposure to the air, allowing for wave action, was calculated geometrically from the wave amplitude and the mean tidal height as given by Lewis (1960). Direct observation confirmed these calculations. The wave amplitude was measured by recording the maximum and minimum wave heights on a vertical scale on several days of average sea conditions. Absolute Growth.-The absolute growth rates were studied over a period from June 1964 to November 1965 using three methods: 1. A group of measured limpets was placed on the concrete piles of a wharf along the west coast of Barbados. This area had previously been cleared of all other members of the species. Every month the limpets were remeasured. A small number of specimens of various sizes was used in this experiment so that individual growth rates could be obtained. 2. Other specimens were kept in aquaria with running sea water and mea- sured at monthly intervals. They fed on algae which grew on the rocks provided and on the side of the aquaria. The data from 1 and 2 (above) were employed to obtain a relation be- tween the length of the limpet and its age as follows: The data were first plotted as the regression of length at time (t + 1) on length at time t using the equation Lt+1 = mLt + i where m is the slope of the line and i is the vertical intercept (Walford, 1946; Taylor, 1959). The von Bertalanffy growth equation (1938) was then used to represent the growth curve Lt = Loo [1 - e-K(t-to)] where Loo = i/(1-m) is the asymptotic length and the constant K is given by K = (-logem). By this means, the age of any specimen of length Lt is estimated as t' = (t - to). The accuracy of the estimation depends upon the accuracy of the value of to in the von Bertalanffy equation. In the case of the limpets Patelloida conulus (Hamai, 1937) and Acmaea dorsuosa (Abe, 1932), the age, t, can be determined independently from the annual rings on the shell and the constant, to, the theoretical age at length zero, thus evaluated. The age of F. barbadensis cannot be obtained from annual rings because of the relatively steady growth of this species throughout the year and the 1967] Ward: Distribution and Growth of Fissurella 301 3 <141m2 < 201m 2 <71m2 o 2 4 MILES I I I II FIGURE1. Map of Barbados showing density of Fissurella barbadensis at 12 stations and the annual wind direction. Wind rose based on 50-year means (U. S. Navy, 1949). ], Half Moon Fort; 2, Six Men's Bay; 3, Paynes Bay; 4, Deep Water Harbour; 5, St. Lawrence; 6, Oistins Bay; 7, Silver Sands; 8, Long Bay; 9, Conset Bay; 10, Bath; 11, Bathsheba; 12, River Bay. accuracy of the estimated age, t', therefore depends upon the accuracy of a separate estimate of to inferred from the study of specimens in the labo- ratory. From observations on newly settled limpets in the aquaria it has been determined that to is probably negligible and is unlikely to be greater than two weeks. 3. From June 1964 to June 1965, a monthly sample of approximately 100 limpets was collected along transect lines perpendicular to the shore in an area of beach rock at Six Men's Bay (Fig. 1). All sizes of animals above 6 mm long were represented in the sample from their upper limit in the pink zone to their lower limit in the surf zone. The length, width, and height of each animal were measured to the nearest 0.5 mm with vernier calipers. All animals were replaced within the same transect area from which they had been collected to ensure that any particular size group was not depleted from the population. The transect was made in a different position each month. Specimens below 6 mm long were not in- cluded in the samples because they are easily overlooked. 302 Bulletin of Marine Science fl7(2) TABLE 1 THE VERTICAL INTERTIDAL ZONES OF THE ROCKY SHORES IN BARBADOS, AFTER LEWIS (1960) Tidal height above zero datum Zone Range (ft) (m) Surf Mean low water springs-mean low water 0-1.2 0-0.37 Pink Mean low water-just below mean high water 1.2-2.8 0.37-0.85 Green Just below mean high water-6 inches above mean high water 2.8-3.8 0.85-1.16 Relative Growth.- The horizontal beach rock platforms at Six Men's Bay were selected as a sheltered area and the cliff station at Oistins (Fig. 1) was chosen as an exposed area to study the variations in shell dimensions, weight, and volume with distribution and growth. At each station high level specimens were collected from the upper pink and lower green zones and the low level animals were obtained from the lower pink and upper surf zones. These zones have already been described by Lewis (1960) and are summarized in Table 1. The shell of Fissurella is approximately the shape of an elliptical cone. The volume of an elliptical cone is a constant, 7T/12, times the product of the three leading dimensions which, in this case, are the major and minor diameters of the base and the height of the shell. The volume is thus ex- pressed as [17T2 . D2H] where H is the height and D is the geometrical mean diameter of the shell base. The shells are not exactly the shape of an elliptical cone, as they are somewhat flattened at the apex due to the apical hole and have the anterior end slightly tapered; the differences, however, would have a negligible effect on the volume. For convenience, the con- stant, 7T/12, was omitted from the calculations. The values for D~H must therefore be multiplied by the constant to obtain the true volumes. The ratio H/D is used as an index to show the variation in the relative height of the shell. The shell length/breadth, L/B, ratio is used to indi- cate the variation in the shape of the base of the shell. The shell thickness was measured by a vernier gauge to 0.1 mm using a point about 2 mm posterior to the apical hole. The shell and soft parts were separated by cutting the muscles of at- tachment. Both parts were damp dried on paper towelling and weighed to the nearest 0.001 g. The volume under the shell was measured to the nearest 0.01 cc by inverting the shell in a plasticine cup. Water, to which soap had been added to reduce surface tension, was introduced through a 1967J Ward: Distribution and Growth of Fissurella 303 + W HAR F 30 o AQUARIUM 25 E E .•...+ 20 UJ e> <t I- 15 « :I: l- e> z 1 0 UJ ...J 5 o o 5 10 15 20 25 30 LENGTH AT AGE t (mm) FIGURE 2. Growth data for individual F. harhadensis. 1 cc graduated pipette until the meniscus was level with the edge of the shell.
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