ABSTRACT
Many species of subtidal fishes are found transiently in shallow water in the high intertidal zone, typically during a high tide (Gibson, 1982). Subtidal fishes that spawn on beaches in the intertidal zone come from a variety of different lineages, including smelts (Osmeridae), New World silversides (Atherinopsidae), herring (Clupeidae), toadfish (Batrachoididae), fugu puffer (Tetraodontidae), sandfish (Trichodontidae), sand lance (Ammodytidae), and even flatfish (Pleuronectidae). In most cases the spawning is timed with the higher tide levels or a higher shore water level caused by large wind waves (DeMartini, 1999). In all cases spawning in each species is associated with a particular type of substrate for the eggs, whether sand, rock, or vegetation, that is submerged and becomes available for oviposition with the highest tides (Conover & Kynard, 1984). As for the resident intertidal fishes, in general the eggs deposited by subtidal migrants in the upper beach are exposed to air for some or all of their incubation period (Martin et al., 2004).
