The Importance of Small Planktonic Copepods and Their Roles in Pelagic Marine Food Webs

Small planktonic marine copepods (＜ 1 mm in length) are the most abundant metazoans on Earth. Included are adults and copepodites of calanoid genera such as Paracalanus, Clausocalanus, and Acartia; cyclopoid genera such as Oithona, Oncaea, and Corycaeus; planktonic harpacticoids of the genus Microsetella; and nauplii of almost all copepod species. Despite the abundance of small copepods, they have historically been undersampled due to the use of nets with meshes ＞ 200- 333 μm. Recent studies have shown, however, that when appropriate net meshes of 100 μm or less are used, small copepods vastly exceed the abundance and sometimes the biomass of larger ones. Failure to adequately account for small copepods may cause serious underestimations of zooplankton abundance and biomass, the copepod grazing impact on phytoplankton primary production, zooplankton-mediated fluxes of chemicals and materials, and trophic interactions in the sea. The feeding ecology of small copepods is less well-known than that of adults of larger copepod species, such as members of the genus Calanus. Further, most feeding information for small copepods is for coastal genera such as Acartia, rather than for offshore taxa. Although it is generally assumed that small copepods, including nauplii, feed primarily upon small-sized phytoplankton cells, most such information comes from rearing or feeding studies on limited laboratory diets. There have been few examinations of actual copepod feeding on mixed diets of natural phytoplankton and microzooplankton found in the sea, but some of those have produced surprises. For instance, some species of Oithona and Paracalanus and even nauplii of Arctic Calanus spp. may feed primarily as predators upon heterotrophic protists, rather than as grazers of phytoplankton. Also, nauplii of various tropical copepod species have been shown to feed upon bacterioplankton. Thus, numerous basic questions remain as to the feeding ecology and grazing/predation impact of small copepods in the sea. Despite limited knowledge of what small copepods eat, it is clear that many higher-trophic-level consumers eat them. Numerous studies have shown that copepod nauplii, Oithona spp. , and other small copepods are important prey of fish larvae and other planktivores. Small copepods exhibit a variety of reproductive strategies to compensate for losses to their populations due to predation. These include having high fecundity and growth rates, when not limited by insufficient food; having high reproduction and growth rates at warmer temperatures; having limited motion and low respiration rates, allowing the investment of more energy in reproduction; and having extended longevity to maximize lifetime reproductive output. Thus, small copepods are important links in marine food webs, serving as major grazers of phytoplankton, as components of the microbial loop, and as prey for ichthyoplankton and other larger pelagic carnivores. Our present inadequate understanding of the true abundance, biomass, trophic ecology, and role of small copepods in biogenic fluxes precludes proper understanding of the ecology of the sea.