Life cycle strategies of copepods in coastal upwelling zones

Life cycles of copepods of coastal upwelling zones are of the multigenerational type–as many as 10 or more generations may be produced each year, depending upon water temperature, food concentration and length of the upwelling season. Abundant food resources and moderate temperature convey advantages to those copepods living in coastal upwelling zones, however, there is a clear disadvantage in that coastal upwelling zones are highly advective environments. Typically, water circulation patterns are such that surface waters are carried offshore, deeper waters carried onshore and most of the water column over the continental shelf is moving equatorward. The challenge to copepod species that inhabit upwelling systems is life cycle closure–how do eggs, nauplii, juveniles and adults avoid being swept out of these ecosystems in the face of persistent transport out of the system? In this review, I first list the species which dominate coastal upwelling ecosystems then discuss three variations on the multigenerational life cycle scheme that are observed in upwelling systems. The latter part of the review is devoted to discussion of how individuals are retained in the productive continental shelf waters within coastal upwelling ecosystems. The suggestion is made that the only copepod species that successfully achieve life cycle closure in such systems are those that are preadapted to upwelling circulation patterns. Our quantitative understanding of the relative importance of physical factors (such as advection) and biological factors (birth, growth, and mortality) on life cycle strategies and population dynamics is quite rudimentary. It would help our understanding if there were more field studies and more computer modeling studies that focused on seasonal cycles of abundance, development times and vertical distribution of life cycle stages, and measurements of water circulation patterns.