Dispersal Dynamics of the Bivalve Gemma Gemma in a Patchy Environment

Abstract

The purpose of this study was to analyze the dispersal dynamics of the ovoviviparous bivalve Gemma gemma (hereafter referred to as Gemma) in an environment disturbed by the pit—digging activities of horseshoe crabs, Limulus polyphemus. Gemma broods its young and has no planktonic larval stage, so all dispersal is the result of juvenile and adult movement. Animal movement was measured using natural crab pits, hand—dug simulated crab pits, and cylindrical bottom traps in the intertidal zone at Tom's Cove, Virginia, USA. This study demonstrated that horseshoe crabs create localized patches with reduced densities of Gemma, that all sizes and ages of Gemma quickly disperse into these low density patches, and that the mechanism of dispersal is passive bedload and suspended load transport. Freshly excavated natural pits had significantly lower Gemma densities than did undisturbed background sediment, but there were no significant differences in total density of other species, number of species, and species diversity (H'). Equitability (J') was greater in pits than in controls because of the reduced abundance of Gemma, the numerically dominant species. Newly dug simulated crab pits also had significantly lower Gemma densities than controls and returned to control levels by the next day. Density recovery trajectories for individually marked pits showed consistent responses in summer and fall, but not in winter when low Gemma abundance resulted in greater variability among pits. Significant positive correlations between the volume of sediment and the number of Gemma collected per bottom trap support the hypothesis that Gemma dispersal is a passive transport phenomenon. Assuming no active, density—dependent movement, the product of the Gemma density frequency distribution in undisturbed background sediment and the frequency distribution of sediment volume collected per trap created a predicted Gemma frequency distribution in traps that matched the actual distribution. Absolute dispersal rates and relative dispersal rates (absolute dispersal rate divided by background density in undisturbed sediment) into pits and traps were greater in summer than winter. Dispersal rate results suggest that increased horseshoe crab disturbance in summer may cause an increase in Gemma transport. Because Gemma individuals are dispersed by hydrodynamic action, it was expected that small, young individuals would be most easily transported in the bedload. There was, however, little evidence that movement into pits and traps was size— or age—selective. Most recent benthic dispersal research has focused on the large—scale movement and settlement patterns of invertebrate larvae. The results from this study illustrate that dispersal of bottom—dwelling juveniles and adults plays an important role in regulating the local distribution and abundance of Gemma. Previous workers have shown that young Gemma live in dense aggregations and that growth and fecundity are reduced at such high densities, leading to population crashes. This study demonstrated a mechanism by which Gemma disperses into low—density patches where intraspecific competition may be mitigated, possibly resulting in enhanced individual reproductive success and population fitness.