Highly advective upwelling systems along the western margins of continents are widely believed to transport larvae far offshore in surface currents resulting in larval wastage, limited recruitment, and increased population connectivity. However, suites of larval behaviors effectively mediate interspecific differences in the extent of cross-shelf migrations between nearshore adult habitats and offshore larval habitats. Interspecific differences in behavior determining whether larvae complete development in estuaries or migrate to the continental shelf are evident in large estuaries, but they sometimes may be disrupted by turbulent tidal flow or the absence of a low-salinity cue in shallow, low-flow estuaries, which are widespread in upwelling systems. Larvae of most species on the continental shelf complete development in the coastal boundary layer of reduced flow, whereas other species migrate to the mid- or outer shelf depending on how much time is spent in surface currents. These migrations are maintained across latitudinal differences in the strength and persistence of upwelling, in upwelling jets at headlands, over upwelling-relaxation cycles, and among years of varying upwelling intensity. Incorporating larval behaviors into numerical models demonstrates that larvae recruit closer to home and in higher numbers than when larvae disperse passively or remain in surface currents. 1. Introduction Eastern boundary upwelling systems have been studied intensively, because they are one of the most productive marine ecosystems producing ~20% of the fish catch from less than 1% of the global ocean [1]. Strong equatorward winds drive broad, slow eastern boundary currents that attain maximum velocities 50 to 200？km from shore (Figure 1). Wind together with rotation of the earth (Coriolis) along the coastal boundary generates a shallow offshore flow at the surface (Ekman transport) and a drop in sea level, which draws cold, nutrient-rich, bottom water to the surface where sunlight fuels high primary production and fisheries for sardines and anchovies. Spatially varying wind velocity (wind stress curl) and increasing wind offshore also generate upwelling (Ekman pumping). Figure 1: Schematic block diagram of generalized circulation. Prevailing winds blow equatorward toward the south with brief periods of relaxation or reversal. Surface waters flow offshore in the Ekman layer, which is weak and a few meters deep nearshore and stronger and about 15 to 30？m deep offshore. Cold, deep waters flow onshore and upwell to the surface often forming a front with warmer
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