Testing the intermittent upwelling hypothesis: upwelling, downwelling, and subsidies to the intertidal zone

The Intermittent Upwelling Hypothesis (IUH) posits that subsidies of larvae and phytoplankton to intertidal communities should vary unimodally along a gradient of upwelling from persistent upwelling to persistent downwelling with most subsidies occurring where upwelling is of intermediate strength and intermittent. Furthermore, the hypothesis states that larvae and phytoplankton are transported far offshore by strong, persistent upwelling and fail to subsidize nearshore communities, whereas weak upwelling or downwelling reduces nutrients for phytoplankton production limiting food for larvae and nearshore communities. We review studies conducted at sea and onshore and reanalyze published data to test the IUH and evaluate alternative hypotheses. To test the hypothesis, we examine five predictions that must hold if the IUH is true. (1) Larvae should inhabit the surface Ekman layer where they are transported offshore during upwelling. Larvae of many intertidal taxa occur deeper in the water column where currents flow shoreward during upwelling. (2) Larvae of nearshore species should occur farther offshore during upwelling than during relaxation or downwelling. Larvae of many nearshore species remain within several kilometers of shore during both conditions. (3) Larval settlement in intertidal communities should be lower during upwelling than relaxation or downwelling. Daily larval settlement has not observed to be higher during relaxation or downwelling events; settlement has most often been seen to vary with the fortnightly tidal cycle likely due to onshore larval transport by internal tides. (4) Larval settlement and recruitment in intertidal communities should be lower in areas of strong, persistent upwelling than where upwelling is weaker and less persistent. Recruitment of mussels and barnacles to artificial and natural substrates did not vary with the strength of upwelling, but did vary inversely with two measures of desiccation potential, and directly with indicators of surf zone hydrodynamics; larval recruitment was higher where surf zones were more dissipative with rip currents. (5) Phytoplankton subsidies to nearshore communities should be highest where upwelling is moderate and intermittent. Like larval subsidies, phytoplankton subsidies varied spatially with surf zone hydrodynamics rather than upwelling. This reconsideration of the evidence for the IUH finds the hypothesis unsupported.