Multiscale patterns of spatial and temporal variation in density and population structure were used to evaluate the generality of a three—trophic—level cascade among sea otters (Enhydra lutris), invertebrate herbivores, and macroalgae in Alaska. The paradigm holds that where sea otters occur herbivores are rare and plants are abundant, whereas when sea otters are absent herbivores are relatively common and plants are rare. Spatial patterns were based on 20 randomly placed quadrats at 153 randomly selected sites distributed among five locations with and four locations without sea otters. Both sea urchin and kelp abundance differed significantly among locations with vs. without sea otters in the Aleutian Islands and southeast Alaska. There was little (Aleutian Islands) or no (southeast Alaska) overlap between sites with and without sea otters, in plots of kelp density against urchin biomass. Despite intersite variation in the abundance of kelps and herbivores, these analyses demonstrate that sea otter predation has a predictable and broadly generalizable influence on the structure of Alaskan kelp forests. The percent cover of algal turf and suspension feeder assemblages also differed significantly (although less dramatically) between locations with and without sea otters. Temporal variation in community structure was assessed over periods of from 3 to 15 yr at sites in the Aleutian Islands and southeast Alaska where sea otters were 1) continuously present, 2) continuously absent, or 3) becoming reestablished because of natural range expansion. Kelp and sea urchin abundance remained largely unchanged at most sites where sea otters were continuously present or absent, the one exception being at Torch Bay (southeast Alaska), where kelp abundance varied significantly through time and urchin abundance varied significantly among sites because of episodic and patchy disturbances. In contrast, kelp and sea urchin abundances changed significantly, and in the expected directions, at sites that were being recolonized by sea otters. Sea urchin biomass declined by 50% in the Aleutian Islands and by nearly 100% in southeast Alaska following the spread of sea otters into previously unoccupied habitats. In response to these different rates and magnitudes of urchin reduction by sea otter predation, increases in kelp abundance were abrupt and highly significant in southeast Alaska but much smaller and slower over similar time periods in the Aleutian Islands. The different kelp colonization rates between southeast Alaska and the Aleutian Islands appear to be caused by large—scale differences in echinoid recruitment coupled with size—selective predation by sea otters for larger urchins. The length of urchin jaws (correlated with test diameter, r2 = 0.968) in sea otter scats indicates that sea urchins <15—20 mm test diameter are rarely eaten by foraging sea otters. Sea urchin populations in the Aleutian Islands included high densities of small individuals (<20 mm test diameter) at all sites and during all years sampled, whereas in south...

Kelps are highly productive seaweeds found along most temperate latitude coastlines, but the fate and importance of kelp production to nearshore ecosystems are largely unknown. The trophic role of kelp-derived carbon in a wide range of marine organisms was assessed by a natural experiment. Growth rates of benthic suspension feeders were greatly increased in the presence of organic detritus (particulate and dissolved) originating from large benthic seaweeds (kelps). Stable carbon isotope analysis confirmed that kelp-derived carbon is found throughout the nearshore food web.

In Torch Bay, Alaska, USA, sea urchins structure shallow subtidal communities by consuming most macroscopic algae. Experimental removal of urchins leads to the development, within 1 yr, of a kelp association of high biomass and diversity. In the 2nd yr, however, these attributes decrease as a single kelp species, Laminaria groenlandica, becomes dominant. Additional field experiments confirm the competitive superiority of this kelp. These manipulations lead to predictions regarding the ecological role of sea otters, a once common (but now absent) sea urchin predator. The predictions are tested and verified by examining areas into which sea otters have recently been transplanted. As expected, a rapid and extensive modification of algal species composition and a dramatic increase in kelp biomass follow the return of sea otters.

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