<I>Cenchritis Muricatus</I>
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BULLETIN OF MARINE SCIENCE, 84(3): 307–313, 2009 E FFECTS of DISTURBANCE on CENCHRITIS MURICATUS (BEADED PERIWINKLE) pOPULATIONS ON small islanDS in THE BAHAMAS Jonah Piovia-Scott ABSTRACT D isturbance can have multiple impacts on shoreline gastropods. This study compares populations of a common supralittoral snail on islands in exposed and protected areas; the former are subject to much more disturbance from wave ac- tion and storm surges. Cenchritis muricatus (Linnaeus, 1758) density was six times higher on protected islands than on exposed islands, representing 2.5 times more biomass. Contrary to expectation, individuals were larger on exposed islands than on protected islands (mean lengths were 26.1 mm and 20.0 mm, respectively); this difference was primarily explained by a significant negative relationship between body size and density coupled with the fact that exposed islands had lower densities. I suggest that periodic large disturbances and oceanographic processes associated with dispersal limit the abundance of C. muricatus on exposed islands, and that larger sizes on exposed islands were probably due to enhanced growth caused by reduced intraspecific competition. D isturbance can have profound impacts on ecological communities (Sousa, 1984; Pickett and White, 1985). These impacts have been particularly well-studied in shoreline ecosystems where disturbance, usually in the form of wave action, affects a wide variety of taxa (Dayton, 1971; Paine and Levin, 1981; Underwood, 1999; Walker et al., 2008). Gastropods are a common component of most shoreline ecosystems, and disturbance can affect gastropods both directly, by inhibiting settlement (Crisp, 1955; Sousa, 1979; Bushek, 1988; Pawlik and Butman, 1993) or dislodging individuals from the substrate (Boulding and van Alstyne, 1993; Trussell, 1997), and indirectly, by removing predators (Menge and Sutherland, 1976; Menge, 1978), competitors (Steffani and Branch, 2003a,b), or facilitators (Underwood, 1999). These direct and indirect effects can lead to either increased (Underwood and McFadyen, 1983; Brown and Quinn, 1988) or decreased (Menge and Sutherland, 1976; Boulding and van Al- styne, 1993) abundance in exposed sites. In contrast, there is a relatively consistent tendency for conspecific individuals to be smaller at exposed sites than at protected sites (e.g., Emson and Faller-Fritsch, 1976; Etter, 1989; Richardson and Brown, 1990; Trussell, 1997), a pattern usually attributed to increased susceptibility to dislodge- ment with size (Denny et al., 1985; Trussell et al., 1993) or changes in foraging be- havior that limit growth rate at exposed sites (Etter, 1996). This study examined the effects of disturbance on the abundance, size, and biomass of a common supralittoral gastropod by surveying populations on exposed and protected islands. The Exuma Cays in the central Bahamas (Fig. 1A) represent an ideal system for studying the effects of disturbance. Small islands in close physical proximity experi- ence very different disturbance regimes. Islands located in “creek” areas are sheltered from high wind and wave action, whereas islands outside of these sheltered areas are not. Exposed islands are also more affected by periodic large storms (Spiller et al., 1998), and the eyes of four hurricanes have passed within 130 km of the study sites in Bulletin of Marine Science 307 © 2009 Rosenstiel School of Marine and Atmospheric Science of the University of Miami 308 BULLETIN OF MARINE SCIENCE, VOL. 84, NO. 3, 2009 Figure 1. Map of the study location in (A) Exuma Cays, the Bahamas indicating protected islands P and exposed islands E, (B) near Staniel Cay and (C) Great Exuma. the decade prior to this study (Hurricanes Lili 1996, Floyd 1999, Michelle 2001, and Frances 2004). The littorinid snail Cenchritis muricatus (Linnaeus, 1758) is the most abundant supralittoral gastropod on small islands in the Exuma Cays (pers. obs.). Cenchritis muricatus is common throughout the Caribbean, South Florida, and the Bahamas (Clench and Abbott, 1942; Abbott, 1954; Trussell, 1997), where it is distributed from the waterline up to at least 3.6 m in vertical height and 28 m in horizontal distance from the water’s edge (Lang et al., 1998; Emson et al., 2002). Organisms in this supra- littoral fringe habitat are thought to be regulated chiefly by abiotic factors (Vermeij, 1972a; Lang et al., 1998). I explored the effect of disturbance on C. muricatus by examining its abundance, size, and biomass on exposed and protected islands at two study sites in the Exuma Cays. Materials and Methods Study System.—This study was conducted in the Exuma island chain in the central Baha- mas (Fig. 1A) in September and October 2005. I surveyed 16 small islands (480–3033 m2) for C. muricatus, which inhabited all parts of each island. Eight islands were located near Staniel Cay, five of these were in “creek” areas protected on both sides by larger islands and three were on the exposed west side of the island chain (Fig. 1B). One of the exposed islands was a narrow peninsula of a slightly larger island. The remaining eight islands were located 100 km to the southwest, near Great Exuma. Four of these were on the exposed south side of the island while the other four were in a protected harbor on the north side (Fig. 1C). Sampling.—I surveyed C. muricatus populations on each island by counting and measur- ing all C. muricatus in circular plots with an area of 0.1963 m2 (0.25 m radius). Between 25 and 50 plots were distributed systematically on each study island using transects. First, I es- P IOVIA-SCOTT: EFFECT OF DISTURBANCE ON BEADED PERIWINKLES 309 tablished a primary transect across the longest axis of the island, beginning and ending at the high water line; I then established secondary transects perpendicular to the primary transect at regular intervals. Sampling plots were established at 2 m intervals along the secondary transects, starting 1 m from the high water line. I adjusted the spacing between secondary transects in order to sample a similar number of plots on each island. Cenchritis muricatus density was calculated as the mean number of individuals per plot on each island. Body size was measured as the length along the longest axis of the shell; I used the average length of sampled individuals on an island for analyses. The relative biomass of each individual was the cube of the height, which is proportional to the dry tissue mass (Borkowski, 1974); I calculated biomass as the mean relative biomass per plot on each island. Analysis.—Differences between exposed and protected islands in C. muricatus density, mean length, and biomass were analyzed using ANOVA, with exposure and study site as predictors. Protected islands and exposed islands did not differ in mean area (t-test:P = 0.77) or mean perimeter-to-area ratio (t-test: P = 0.63). I analyzed the effect of density on size using linear regression. All models were consistent with assumptions of homogeneity of variances and normality of residuals. Analyses were conducted using PROC GLM in SAS v 8.0 (SAS Institute, 1999). Results The number of snails counted and measured per island ranged from 2 to 432. There were significant differences inC. muricatus density, average length, and biomass be- tween exposed and protected islands. Protected islands had six times more individu- als per plot than exposed islands (ANOVA: F1,13 = 5.95, P = 0.030; Fig. 2A). Average C. muricatus length was greater on exposed islands (26.1 mm) than on protected islands (20.0 mm) (ANOVA: F1,13 = 11.65, P = 0.005; Fig. 2B). Average biomass per plot was 2.5 times larger on protected islands (ANOVA: F1,13 = 6.97, P = 0.020; Fig. 2C). There was a negative relationship between mean shell length and mean density (F1,14 = 38.36, P < 0.001; Fig. 3). This pattern was also evident on protected islands alone (F1,7 = 33.46, P < 0.001), but was not significant on exposed islands (F1,5 = 4.56, P = 0.086), although the trend was in the same direction. Discussion Cenchritis muricatus had higher densities and higher biomass on protected islands than on exposed islands, indicating that exposure to disturbance limits snail density and biomass. Individuals on exposed islands were larger than those on protected Figure 2. Cenchritis muricatus on protected (n = 9) and exposed (n = 7) islands: (A) Mean (± SE) number of individuals per plot, (B) mean (± SE) shell length of all measured individuals on an island, and (C) mean (± SE) biomass per plot. 310 BULLETIN OF MARINE SCIENCE, VOL. 84, NO. 3, 2009 Figure 3. Mean Cenchritis muricatus shell length and mean density per plot on protected (closed symbols) and exposed (open symbols) islands. Squares represent islands near Staniel Cay and circles represent islands near Great Exuma. Linear regression line is shown (R2 = 0.73). islands, but this effect may be explained primarily by the fact that these islands had lower densities. Since exposed islands have lower densities but larger individuals, the difference in biomass between protected and exposed islands is not as great as the difference in density. Four processes could contribute to the observed differences in C. muricatus den- sity, size, and biomass between exposed and protected islands: (1) removal by dis- turbance, (2) recruitment limitation, (3) intraspecific competition, and (4) predation. Interspecific competition is unlikely to be important, as C. muricatus is the only mollusc that regularly occurs over 1 m inland from the high water line on the study islands (pers. obs.). While other processes, such as parasitism, could be important, I do not have sufficient evidence to address them here. Exposed islands experience higher levels of erosive wind and wave action in the supralittoral zone (L. Yang, University of California-Davis, pers. comm.) and tend to be more affected by periodic large storms (Spiller et al., 1998).