Although knowledge on dispersal patterns is essential for predicting long-term population dynamics, critical information on the modalities of passive dispersal and potential interactions between vectors is often missing. Here, we use mangrove propagules with a wide variety of morphologies to investigate the interaction between water and wind as a driver of passive dispersal. We imposed 16 combinations of wind and hydrodynamic conditions in a flume tank, using propagules of six important mangrove species (and genera), resulting in a set of dispersal morphologies that covers most variation present in mangrove propagules worldwide. Additionally, we discussed the broader implications of the outcome of this flume study on the potential of long distance dispersal for mangrove propagules in nature, applying a conceptual model to a natural mangrove system in Gazi Bay (Kenya). Overall, the effect of wind on dispersal depended on propagule density (g l-1). The low-density Heritiera littoralis propagules were most affected by wind, while the high-density vertically floating propagules of Ceriops tagal and Bruguiera gymnorrhiza were least affected. Avicennia marina, and horizontally floating Rhizophora mucronata and C. tagal propagules behaved similarly. Morphological propagule traits, such as the dorsal sail of H. littoralis, explained another part of the interspecific differences. Within species, differences in dispersal velocities can be explained by differences in density and for H. littoralis also by variations in the shape of the dorsal sail. Our conceptual model illustrates that different propagule types have a different likelihood of reaching the open ocean depending on prevailing water and wind currents. Results suggest that in open water, propagule traits (density, morphology, and floating orientation) appear to determine the effect of water and wind currents on dispersal dynamics. This has important implications for inter- and intraspecific variation in dispersal patterns and the likelihood of reaching suitable habitat patches within a propagule's viable period.

Abstract. Although wind has been recognized to be an important factor in the dispersal of hydrochorous mangrove propagules, and hence in the quantification of (meta)population dynamics, the species-specific sensitivity to wind effects has not been studied. We combined observations from a controlled experiment (flume tank) and in situ experiments to understand wind and water current contributions to dispersal potential as well as to estimate real dispersal ranges due to immediate response to tidal currents (two outgoing tides). This was done for 4 species with propagules differing in morphological and buoyancy properties (i.e. Rhizophora mucronata, Ceriops tagal, Heritiera littoralis and Xylocarpus granatum). The flume experiments revealed that the influence of wind depends on the density of a propagule (and hence its buoyancy characteristics) and that typical morphological characteristics of the dispersal unit are additionally important. H. littoralis propagules were influenced most, because on the one hand their low density (613.58 g L−1; n =10) enables them to float on top of the water surface, and on the other hand their "sailboat-like" structure provides a relatively large surface area. The X. granatum fruits appeared to be the least influenced by ambient wind conditions, explained by the smooth surface and spherical shape of which, because of the fruit's high density (890.05 g L−1; n = 1), only a small part sticks above the water surface. Although the seeds of X. granatum are of a similar size class than H. littoralis propagules, they are (like the X. granatum fruits) largely submerged due to their high density (870.66 g L−1; n = 8), hence catching less wind than H. littoralis propagules. The influence of wind on the dispersal of the horizontally floating C. tagal and R. mucronata dispersal units was strong, comparable to that of H. littoralis propagules. A differential effect of wind was found within elongated propagules, which directly follows from the floating orientation of the propagules. While the dispersal path of vertically floating propagules was influenced by the strength and direction of the water currents and to a lesser extent by ambient wind conditions, the dispersal path of horizontally floating propagules was influenced by both strength and direction of the water currents and prevailing wind forces. To validate the flume results, propagules of C. tagal and R. mucronata were released during outgoing tide in a tidal creek in Gazi Bay (Kenya), followed by observation of their dispersal distance and direction, while knowing the actual dominant wind direction. In line with the flume results, this study showed that wind plays an important role in the dispersal distance of the propagules. The present study provides important mechanistic insight into the effect of wind on hydrochorous mangrove propagule dispersal, thereby yielding an essential step towards the construction and optimization of (particle-based) hydrodynamic dispersal models.

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