POPULATION BIOLOGY AND SECONDARY PRODUCTION OF VESICA VESICA (GMELIN, 1791) (: OLIVIDAE) ON A SANDY BEACH IN SOUTHEASTERN BRAZIL

CARLOS HENRIQUE SOARES CAETANO, VALÉRIA GOMES VELOSO AND RICARDO SILVA CARDOSO Laboratório de Dinâmica de Populações Marinhas, Departamento de Ciências Naturais, Universidade do Rio de Janeiro (UNIRIO), Av. Pasteur, no. 458, Sala 411, Urca, CEP 22290 – 240, Rio de Janeiro, Brazil

ABSTRACT vesica (Gastropoda: Olividae) is a common inhabitant of exposed sandy beaches in southern and southeastern Brazil. The population biology and secondary production of this was studied from June 1998 to July 2000 on Restinga da Marambaia beach, state of Rio de Janeiro, Brazil. Specimens were hand-collected in two sectors (10 25 m), and shell length was measured in the field. Growth and mortality were estimated from length-frequency data, using computer-based methods. Life span was estimated by an inverse von Bertalanffy growth equation, considering tmax as 95% of the asymptotic length. Mass-specific growth-rate and size-frequency methods were used to calculate secondary production. Highest population densities were observed during winter (September 1998 and 1999) and autumn (June 2000). The growth rate varied seasonally, with the slowest growth rates occurring in late summer (1st year) and spring (2nd year). Mortality rates (Z 3.12 and 2.56 year–1, for the first and second year, respectively) did not differ significantly between years. Life span was nearly 5 years. Estimated secondary production varied between 0.142 and 0.213 g AFDW m–2 year–1, while annual P/B ratio varied between 1.029 and 1.111.

INTRODUCTION from June 1998 through July 2000, always at low tide. Two rect- angular sectors (500 m apart) were established, each 10 m wide Gastropods and bivalves are among the most conspicuous with a 25-m base parallel to the waterline, located from the members of the macrofauna of exposed sandy beaches (Brown swash zone to a 50-cm water depth in the sublittoral. In each sec- & McLachlan, 1990), being generally more abundant on dissi- tor, specimens were hand-collected by two collectors until every pative and intermediate beaches than on reflective beaches snail had been removed. Shell length was measured in the field (Dexter, 1984). using a vernier caliper (0.1 mm accuracy). Afterwards, all speci- Gastropods of the family Olividae inhabit sandy shores in mens were returned to their respective sectors, except those tropical and subtropical regions of the world (Petuch & Sargent, utilized to determine the length–weight relationship (n 95). 1986), and a few species are also found in the cooler seas of In the laboratory, shells were cracked in a vice and the southern Australia and New Zealand (Smith, 1998). The removed. These were dried at 70°C for 24 h, weighed, ashed in a Olivancillaria includes about 10 species, seven of them occurring muffle furnace for 4 h at 600°C and reweighed. in Brazil (Rios, 1994), where some are important for subsistence The study period was arbitrarily divided into two sampling fishing. Nevertheless, studies of this genus are sparse and mainly years (from June 1998 to June 1999 and from July 1999 to July concern systematics (Klappenbach, 1964, 1965, 1966; Thomé, 2000) in order to analyse temporal variations in population 1966) and anatomy (Marcus & Marcus, 1959; Jurberg, 1970; parameters. Lopes, 1991; Borzone, 1995; Borzone & Vargas, 1999). Olivan- cillaria vesica (Gmelin, 1791) is a common inhabitant of beaches in southern and southeastern Brazil, living semi-buried in Environmental parameters sandy substrates from the intertidal zone to deeper areas in the Sediment samples for particle-size analysis were taken monthly sublitteral (Gianuca, 1985). According to Klappenbach (1966), with a plastic corer of 3.5 cm diameter to a depth of 10 cm, in the O. vesica occurs in two geographic forms or subspecies. One sublittoral zone. Samples were oven-dried at 70°C and sieved form with a long narrow shell, O. vesica vesica, occurs from the through graded screens in order to determine mean particle state of Rio de Janeiro south to Paraná, while another with a size (Folk & Ward, 1957). The slope of the beach was deter- smaller wider shell, O. vesica auricularia (Lamarck), ranges from mined by Emery’s profiling technique (Emery, 1961). Dean’s Santa Catarina in Brazil to the Mar del Plata in Argentina. In the dimensionless parameter (; Short & Wright, 1983) was calcu- present study we describe the growth, mortality, life span and lated for each month as a measure of beach morphodynamic secondary production patterns of O. vesica vesica, based on a two- state: Hb/Ws.T, where Hb is breaker height in cm, Ws the sand year study. settling velocity in cm s–1 (obtained from the particle size and Gibbs, Mathews & Link, 1971) and T the wave period in seconds. MATERIAL AND METHODS Water temperature and salinity in the surf zone were measured using a thermometer and a salinometer, respectively. Sampling and laboratory procedures The population of Olivancillaria vesica vesica at Restinga da Population parameters Marambaia beach (23°03 S; 43°36 W) was sampled monthly Growth and mortality functions were estimated from the monthly Correspondence: C. H. S. Caetano; e-mail: [email protected] length measurements in the population (26 length-frequency

J. Moll. Stud. (2003) 69: 67–73 © The Malacological Society of London 2003 C. H. S. CAETANO, V. G. VELOSO & R. S. CARDOSO samples with individual lengths grouped into 2-mm size classes). Statistical analysis The routine Electronic Length Frequency Analysis (ELEFAN) of the package FAO-ICLARM Stock Assessment Tools (FISAT) Simple linear regressions were performed to assess the relation- was used to fit a seasonal von Bertalanffy model (Gayanilo, ships between the density of O. vesica vesica and the density of Sparre & Pauly, 1996) to the set of restructured length-frequency their main food item, the intertidal mole crab Emerita brasiliensis samples. This curve has the form: Schmitt (data obtained from Cardoso, Veloso & Caetano, 2002); sand particle size; beach slope; and water temperature π π π π [–K(t–t0) + (KC/2 )sin 2 (t–ts)–(KC/2 )sin 2 (t0–ts)] Lt L[1–e ] and salinity. Both O. vesica vesica and E. brasiliensis were collected during the same sampling period. An analysis of covariance where Lt is length (mm) at time t; L is the theoretical maximum (ANCOVA) was used to compare mortality rates between years, length that the species would reach if it lived indefinitely; K is using age as the covariate. The Mann–Whitney U-test was used to the curvature parameter; C is a constant for the amplitude of compare biomass and secondary production between years. In oscillation in seasonal growth; to is age at zero length; and ts is the all statistical analyses, a significance level of 5% was adopted. initial point of seasonal oscillation in relation to t 0 and WP (winter-point, i.e. the period of growth reduction, expressed as a decimal fraction of the year). The graphical representation of this equation produces a curve that is evaluated through the RESULTS goodness of fit index Rn (Gayanilo et al., 1996). The instantaneous mortality rate (Z) was calculated by the Environmental parameters single negative exponential model, using the length-converted Restinga da Marambaia beach showed a modal intermediate catch curve method (Pauly, Moreau & Abad, 1995) of the FISAT morphodynamic state with a mean omega () value of 1.8 program (Gayanilo et al., 1996). (SD 0.6). The beach had a gentle slope (1/11.73–1/85.71). Life span was estimated by an inverse von Bertalanffy growth The sediment was composed of fine to medium sands, ranging equation, considering maximum length as 95% of the asymp- from 0.20 to 0.32 mm (x¯ 0.23; SD 0.03). The mean tem- totic length (King, 1995). perature in the water of the surf zone was 22.7ºC (SD 2.6), with the lowest temperatures of 17.5 and 16.6ºC occurring in Length–weight relationships spring (November 1998 and 1999, respectively), and the highest temperature of 28.7ºC in February 1999. The salinity of the surf The relationship between shell length and ash-free dry weight waters was nearly constant (x¯ 34.3; SD 0.5). of the soft tissue was estimated by linear regression analysis, with the data converted to natural logarithms in the equation: lnW ln a b ln L, where W is the mean ash-free dry weight per Population abundance individual (g); L is the length of the size class (mm); and a is the intercept on the Y axis and b is the slope of the regression line. Densities of O. vesica vesica varied considerably with the greatest densities observed during winter (September 1998 and 1999) and autumn (June 2000; Fig. 1). No significant correlations Secondary production between the density of O. vesica vesica and the abiotic (sand par- Secondary production was calculated by two methods. The ticle size, water temperature and salinity) and biotic (density first was the mass specific growth rate (MSGR) of Crisp (1984). of Emerita brasiliensis) factors analysed were observed, except for In this method production is given by the equation: beach slope (r 0.45; P 0.05). P fi *G i *wi * t, where fi is the mean number of individuals in size class i, Gi is the mass specific growth rate of size class i, Growth, life span and mortality wi the mean weight of the size class i and t is the interval of time. Mass specific growth rate (Gi) can be obtained by During the study period, 1594 snails were collected and meas- Gi ln wi + 1 – lnwi/ t(i + 1 – i). The second was the size-frequency ured: 720 in the first year (June 1998–June 1999) and 874 in the method (SF; Hynes & Coleman, 1968; Hamilton, 1969; Benke, second year (July 1999–July 2000). The smallest individual was 1979; Menzie, 1980) estimated by: 21 mm and the largest 51 mm. Estimates of growth indicated Σ 1/2 P [ (Nj – Nj + 1)(Wj * Wj + 1) ] i/CPI, where Nj is the number moderate seasonal oscillations (C 0.6 and 0.4), with slowest if individuals that grow to the mean size of the size class during growth rates occurring in late summer (WP 0.3*12 3.6 – 1/2 1/i of the year (Hamilton, 1969), (Wj*Wj + 1) is the mean geo- March) for the first year, and in spring (WP 0.9*12 10.8 – metric weight between two sucessive length classes, i is the October/November) for the second year (Table 1, Fig. 2). The number of size classes and CPI is the time from hatching to estimated life span (tmax) was 4.28 and 4.99 years, for the first death of the largest size class. Annual mean biomass was calcu- and second years, respectively. ΣΣ ∆ –1 lated as: B f i * w i * t. The instantaneous mortality (Z) was 3.12 year in the first The turnover rates (P/B), derived of the ratio between annual year and 2.56 year–1 in the second year (Fig. 3). Mortality rates secondary production (P) and annual mean biomass (B), was did not differ significantly between years (ANCOVA F 2.78, also calculated. df 1/12, P 0.05).

Table 1. Growth, mortality and life span estimates for Olivancillaria vesica vesica at Restinga da Marambaia beach.

Period L KCWPRnZ tmax

1st year: June 1998/June 1999 59.0 0.7 0.6 0.3 0.252 3.12 4.28 2nd year: July 1999/July 2000 59.5 0.6 0.4 0.9 0.246 2.56 4.99

–1 L, asymptotic length (mm); K, curvature parameter (yr ); C, intensity of seasonal oscillation; WP, point of –1 slowest growth rate in the year; Rn, goodness of fit index; Z, mortality rates (yr ); tmax, life span (yr).

68 POPULATION BIOLOGY AND PRODUCTIVITY OF OLIVANCILLARIA

1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 20 1 Figure 1. Monthly variations in density (ind m–2: mean SD) of Olivancillaria vesica vesica at Restinga da Marambaia beach. 2 3 4 5 6 7 8 9 30 1 2 3 4 5 6 7 8 9 40 1 2 3 4 5 6 7 8 9 50 1 Figure 2. Seasonal growth curve fitted for Olivancillaria vesica vesica at Restinga da Marambaia beach. First year: June/1998–June/1999 (upper), Second year: 2 July/1999–July/2000 (lower). 3 4 5 Secondary production DISCUSSION 6 7 The regression equation between length-ash free dry weight for Most species on sandy beaches show strong fluctuations in 8 O. vesica vesica was: lnW –12.52 3.43 lnL (n 95, r 0.92, population abundance (Veloso, Cardoso & Fonseca, 1997). The 9 P 0.0001). factors tested (density of the principal prey Emerita brasiliensis, 60 Estimates of mean annual biomass and secondary production sand particle size, water temperature and salinity), except for 1 by the mass-specific growth rate and size-frequency methods beach slope, did not contribute significantly to explaining the 2 were slightly higher for the second year, however they were fluctuations in density of O. vesica vesica. According to this 3 not significantly different between years (U 66.00, P 0.05; relationship (density vs beach slope), the highest abundances 4 U 65.24, P 0.05, for biomass and secondary production, of O. vesica vesica were observed in months when the slope was 5 respectively). The P/B ratio showed no well-defined pattern, gentler (Fig. 4). This indicates that the observed population 6 with higher values occurring in the first year as estimated by peaks may be associated with greater ease of migration to the 7 the size-frequency method and in the second year by the mass- shallower areas, since these animals use the swash, which is influ- 8 specific growth rate method (Table 2). enced by the beach slope, to move up and down the beach

69 C. H. S. CAETANO, V. G. VELOSO & R. S. CARDOSO

Figure 3. Length-converted catch curves for Olivancillaria vesica vesica during the first year (June 1998–June 1999) and second year (July 1999–July 2000).

Figure 4. Olivancillaria vesica vesica. A. Monthly variations in beach slope (——) and density (- - - -). B. Scatter diagram of densities plotted against beach slope, with regression line (r 0.45).

70 POPULATION BIOLOGY AND PRODUCTIVITY OF OLIVANCILLARIA

1 (‘swash-riders’). In general, variations in population density Scolelepis gaucha (Orensanz & Gianuca). Contreras, Defeo & 2 of intertidal macrofauna have been attributed more to biotic Jaramillo (1999) associated the variations in growth rate of the 3 factors, mainly recruitment, than to abiotic factors (Dexter, crustacean Emerita analoga (Stimpson) with seasons of low tem- 4 1984; Bamber, 1993; Borzone & Souza, 1997). peratures. Curtis, Kinley & Tanner (2000) studied the growth of 5 The largest individual of O. vesica vesica collected measured the gastropod Ilyanassa obsoleta (Say) and observed that infec- 6 51 mm. This is shorter than maximum lengths found along the tion by trematodes retarded its growth. A decrease in the growth 7 Brazilian coast by Marcus & Marcus (1959) in São Paulo, 60 mm, rate of the polychaete Laeonereis acuta (Treadwell) near the 8 and Gofferjés (1950) in Paraná, 68 mm. It is generally assumed breeding season was observed by Omena & Amaral (2000); the 9 that marine poikilotherms that inhabit the colder waters of decrease attributed to the large allocation of energy to the pro- 10 higher latitudes grow more slowly, attain larger sizes, and live duction of gametes. 1 longer than do individuals of the same or closely related species The life span estimated for O. vesica vesica, between 4.28 2 from warmer waters (Dehnel, 1955; Ray, 1960; Fonseca, Veloso and 4.99 years, is much shorter than that observed for two 3 & Cardoso, 2000). This being so, the observed variation in size closely-related species: biplicata, between 8 and 12 years 4 may be explained by the differences in temperature and latitude (Stohler, 1969) and Oliva oliva Linnaeus, nearly 10 years 5 of those localities where these investigators obtained their (Tursch, Ouin & Bouillon, 1995). However, the estimates for 6 material. We found no individuals with a shell length less than Olivella biplicata and Oliva oliva should be treated with caution, 7 21 mm. The absence of smaller individuals could be explained since they appear to have been proposed by simple inference, 8 by an alternative hypothesis of a size-stratified distribution, in because the authors did not use specific methods for estimating 9 which those individuals with shells shorter than 21 mm were life span. 20 located beyond the arbitrary sampling limits established in this The values of secondary production obtained by size-frequency 1 study. Edwards (1968) noted that the juveniles of Olivella biplica- and mass-specific growth rates methods ranged from 0.142 to 2 ta (Sowerby) live preferentially in deeper areas, while the adults 0.213 g AFDW m–2 yr–1. These values are low compared to esti- 3 live in shallower water. According to Greifeneder (1981), this mates for other gastropods, such as melanoides Deshayes, 4 pattern seems to be a widespread ecological strategy among 1.06 g AFDW m–2 yr–1 (Ansell, McLusky, Stirling & Trevallion, 5 gastropods, being recorded in some species of the family 1978); Bullia rhodostoma Reeve, 0.59 g AFDW m–2 yr–1 (McLachlan 6 Olividae and in Conus litteratus Linnaeus (Conidae). It is sug- & Van der Horst, 1979); sarmaticus Linnaeus, 1.18 and 7 gested that the ecological meaning of this distribution pattern is 3.66 g AFDW m–2 yr–1 (McLachlan & Lombard, 1980); Melanoides 8 that, first, small juveniles at depth are better protected against tuberculata (Müller), 12.09 g AFDW m–2 yr–1 (Dudgeon, 1986); 9 desiccation and visual predators, and secondly, that mate-find- Chilina gibbosa Sowerby, 14.18 g AFDW m–2 yr–1 (Bosnia, Kaisin 30 ing is enhanced by a higher density of adult sexually mature & Tablado, 1990) and Hydrobia ulvae (Pennant), 8.0 g AFDW 1 snails in shallower waters. Future research on population bio- m–2 yr–1 (Sola, 1996). 2 logy of O. vesica vesica should include experiments to test these The P/B values of the above-mentioned species show a similar 3 hypotheses. pattern to that observed for secondary production. The estim- 4 The growth rate in O. vesica vesica showed moderate seasonal ated values for O. vesica vesica are, again, much lower than for the 5 variation, with the lowest rates (WP) occurring in late summer other species, except for B. rhodostoma and T. sarmaticus, which 6 and spring, for the first and second year, respectively. The in spite of having high secondary production show a low P/B 7 decrease in growth rate during the summer can be explained by (Table 3). 8 the low densities of E. brasiliensis, its chief prey, on this beach The low P/B values observed for O. vesica vesica, B. rhodostoma 9 (Cardoso et al., 2002). For the second year, low temperatures and T. sarmaticus appear to be related to differences in body 40 recorded in November (16.6°C) may explain the lower growth dimensions (Table 3). The negative relationship between the 1 rate during that period. Santos (1994) observed that variations turnover rate (P/B) and individual body mass (Brey & Clarke, 2 in beach morphodynamics influenced growth of the polychaete 1993; Tumbiolo & Downing, 1994) and longevity (Waters, 1977; 3 4 Table 2. Mean annual biomass (B) in g AFDW m–2, secondary production (P) in g AFDW m–2 yr–1 and turnover 5 rates (P/B) in yr–1 of Olivancillaria vesica vesica at Restinga da Marambaia beach according to size-frequency (SF) 6 and mass specific growth rate (MSGR) methods. 7 8 SF MSGR 9 Period PBP/B PBP/B 50 1 1st year: June 1998/June 1999 0.142 0.128 1.111 0.174 0.169 1.029 2 2nd year: July 1999/July 2000 0.149 0.142 1.048 0.213 0.201 1.063 3 4 5 Table 3. Comparison of values of secondary production (P) in g AFDW m–2 yr–1, turnover rates (P/B) in yr–1 and maximum shell 6 length (Lmax) in mm among gastropod species. 7

8 Species PP/BLmax Source 9 60 Olivancillaria vesica vesica 0.14 – 0.21 1.02–1.11 51 This study 1 Bullia melanoides 1.06 13.50 * Ansell et al. (1978) 2 Bullia rhodostoma 0.59 0.90 55 McLachlan & Van der Horst (1979) 3 Turbo sarmaticus 1.18 – 3.66 0.48–0.69 97 McLachlan & Lombard (1980) 4 Melanoides tuberculata 12.09 4.81 10 Dudgeon (1986) 5 Chilina gibbosa 14.18 3.39 32 Bosnia et al. (1990) 6 Hydrobia ulvae 8.00 2.90 7 Sola (1996) 7 8 *Data not provided by authors.

71 C. H. S. CAETANO, V. G. VELOSO & R. S. CARDOSO

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