Abstract
The effects of tides on littoral marine habitats are so ubiquitous that shorelines are commonly described as ‘intertidal’, whereas waves are considered a secondary factor that simply modifies the intertidal habitat. However mean significant wave height exceeds tidal range at many locations worldwide. Here we construct a simple sinusoidal model of coastal water level based on both tidal range and wave height. From the patterns of emergence and submergence predicted by the model, we derive four vertical shoreline benchmarks which bracket up to three novel, spatially distinct, and physically defined zones. The (1) emergent tidal zone is characterized by tidally driven emergence in air; the (2) wave zone is characterized by constant (not periodic) wave wash; and the (3) submergent tidal zone is characterized by tidally driven submergence. The decoupling of tidally driven emergence and submergence made possible by wave action is a critical prediction of the model. On wave-dominated shores (wave height ≫ tidal range), all three zones are predicted to exist separately, but on tide-dominated shores (tidal range ≫ wave height) the wave zone is absent and the emergent and submergent tidal zones overlap substantially, forming the traditional “intertidal zone”. We conclude by incorporating time and space in the model to illustrate variability in the physical conditions and zonation on littoral shores. The wave:tide physical zonation model is a unifying framework that can facilitate our understanding of physical conditions on littoral shores whether tropical or temperate, marine or lentic.
Highlights
Littoral habitats, those lying between the low-tide line and the upper limit of aquatic species on the shore, are among the most studied and well-known aquatic habitats
A third zone, bracketed by benchmarks two and three, is sandwiched between the emergent and submergent tidal zones when wave height is greater than tidal range and is termed the wave zone because it is washed by waves at both high and low tide (Figs. 2F and 2I)
Our model of coastal water level indicates that waves do not expand and elevate physical littoral zones that exist in the absence of waves
Summary
Those lying between the low-tide line and the upper limit of aquatic species on the shore, are among the most studied and well-known aquatic habitats. Much attention has been devoted to the study of organisms on rocky shores - in particular their vertical zonation, the upper and lower limits of species, and distribution along gradients of wave exposure. Stephenson & Stephenson (1949) and Stephenson & Stephenson (1972) proposed their “universal features of zonation between tide-marks on rocky coasts” after conducting surveys of vertical biotic zonation on littoral shores world-wide. More recent research continues to demonstrate that biotic zones and species distribution limits do not consistently occur at the same shore levels, even within shores (Benedetti-Cecchi & Cinelli, 1997)
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