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Sea urchins are major components of marine communities. Theirgrazing limits algal biomass, and they are preyed upon by manypredators. Purple sea urchins (Strongylocentrotus purpuratus)are among the best studied species. They live in environmentsthat alternate between two stable states: luxuriant, species-richkelp forests and sea urchindominated "barrens." The transitionfrom one state to the other can be initiated by several factors,including the abundance of algal food, predators, storm intensities,and incidence of disease. Purple sea urchins compete with othergrazers, some of which are important fishery resources (suchas abalones and red sea urchins), and they are harvested forscientific research. Revelations from their genome will leadto a better understanding of how they maintain their ecologicalimportance, and may in turn enhance their economic potential.
Long Marine Laboratory, University of California, Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, USA.
E-mail: pearse{at}biology.ucsc.edu
Sea urchins are major grazers in shallow seas worldwide (1).Purple sea urchins (S. purpuratus) and other strongylocentrotidsea urchins of the Northern Hemisphere are particularly importantand are the most intensively studied (2). A delicate balancebetween sea urchin grazing and kelp forest productivity leadsto stable states that alternate between luxuriant kelp forestsand relatively species-depauperate sea urchin "barrens" (3).Curiously, the densities of sea urchins are often similar withinkelp forests and sea urchin barrens. Within kelp forests, seaurchins are nearly stationary, feeding on captured pieces ofkelp litter ("drift kelp") that are produced and shed in highquantities from the kelp plants. However, when the kelps areremoved by storms or El Niño events, the remaining seaurchins actively forage on young kelp recruits and on driftkelp brought in from elsewhere, preventing the reestablishmentof the kelp forest. The sea urchins can be decimated by storms(4) or diseases (5), allowing the kelp forest to return to thearea.
In addition to the alternation of kelp forests and sea urchinbarrens, temporal and spatial variation in recruitment of seaurchins can greatly influence their abundance, and thereforetheir ecological role, within kelp forests. Infrequent peaksin recruitment can lead to spurts in sea urchin densities (6),sometimes triggering intense grazing that removes the kelp,transforming a kelp forest into a sea urchin barren (7) (Fig. 1).Recruitment intensity is determined mainly by the supply ofsea urchin larvae, which in turn depends on the oceanographicconditions that bring the larvae to suitable areas to settle(8).
Fig. 1. Purple sea urchins (S. purpuratus) grazing on the remains of a giant kelp hold fast after an unusually heavy recruitment in Carmel Bay, California (7). [Photo courtesy of J. M. Watanabe]
[View Larger Version of this Image (137K GIF file)]
Predators almost certainly limited the abundance of sea urchinsin the past, and the presence of sea urchin barrens around theworld is due at least in part to decimation of predators byhumans (9). In southern California, where sea urchindominatedbarrens are common, major predators such as spiny lobsters,sheep-head fish, and sea otters have been greatly reduced duringthe past century (10). Sea otters, in particular, are effectivein keeping sea urchin densities low throughout their range (11),which often tips the balance toward kelp forests and enhanceskelp forest productivity and biodiversity (12). Other factorsalso can favor kelp forests or barrens, even in the absenceof major sea urchin predators (13). For example, sea urchinscompete with other grazers. Purple sea urchins compete in particularwith abalones and red sea urchins (14). However, because bothabalones and red sea urchins have experienced intensive fishingpressure for the past several decades, competitive interactionsamong these species have probably decreased, perhaps relievingpurple sea urchins from adverse competition. On the other hand,juvenile purple sea urchins and abalones often are shelteredfrom predators under the spines of red sea urchins (15), andboth may be negatively affected by heavy fishing of red seaurchins. The strengths of these interactions remain poorly knownand are complicated by variation in fishing pressures. Red seaurchins have been the basis of a major fishery in the easternPacific for the past several decades. However, the U.S. seaurchin fishery has been in decline, mainly because of a declinein demand by the Japanese market ($110 million worth of seaurchins were exported from the United States to Japan in 1993compared with about $36 million in 2003) (16). The smaller,less valuable, purple sea urchins have been a minor componentof the fishery but continue to play an extensive role in scientificresearch (2).
Sea urchins are central in structuring marine benthic communities,both as grazers and prey, and are economically valuable in fisheries.There are important differences among the different species:Some are more effective as grazers than others, and they varyin their diets, growth rates, longevities, and importance infisheries. Some show no sign of senility and live for well overa century (17). Elucidation of their genomes will open new avenuesof research into the underlying genetic and evolutionary basesof these variations.
References and Notes
1. C. Harrold, J. S. Pearse, Echinoderm Studies2, 137 (1987).
2. L. Rogers-Bennett, in Edible Sea Urchins: Biology and Ecology, J. M. Lawrence, Ed. (Elsevier, Amsterdam, Netherlands, 2007), pp. 393425.
3. P. K. Dayton et al., Ecol. Monogr.62, 421 (1992). [CrossRef]
14. M. J. Tegner, P. K. Dayton, ICES J. Mar. Sci.57, 579 (2000).
15. L. Rogers-Bennett, J. S. Pearse, Conserv. Biol.15, 642 (2001). [CrossRef]
16. D. Sweetnam et al., Calif. Coop. Oceanic Fish. Invest. Rep.46, 10 (2005).
17. T. A. Ebert, J. R. Southon, Fish. Bull.101, 915 (2003).
18. I thank J. Watanabe for providing the photo used in Fig. 1, L. Rogers-Bennett for sharing her manuscript with me, V. Pearse and an anonymous reviewer for providing comments on the manuscript, and R. A. Cameron for inviting me to prepare it.
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