Differentiation between true mangroves and mangrove associates based on leaf traits and salt contents

Mangrove species are classified as true mangroves and mangrove associates. However, as for some fringe species found mainly on the landward transitional zones of mangroves, no consensus among scientists could be reached in favor of this classification and much debate arises. We hypothesized that true mangroves differ from mangrove associates physiologically and ecologically in their ability to survive in mangrove environment. To test this hypothesis, leaf structural traits and osmotic properties were used to describe variation in 33 mangrove species (17 true mangroves, 6 mangrove associates and 10 controversial species). Specific leaf area (SLA) of true mangroves as well as leaf nitrogen concentration on a leaf mass (Nmass) were lower than that of mangrove associates; leaf succulence was, in general, twice as high in true mangroves compared to mangrove associates; true mangroves accumulated 8–9 times more Na and Cl than mangrove associates and the former had K/Na ratios <0.5, but the latter had K/Na ratios >0.5. These results indicated that true mangroves differed reliably from mangrove associates in leaf traits and osmotic properties. True mangroves are true halophytes and mangrove associates are glycophytes with certain salt tolerance. Combining distribution pattern information, the 10 controversial species were reclassified.