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Keystone                     Figure 4.12 Sea otters are a
                                                                   absent                      keystone species.  A keystone (a)
                                                                                               is the wedge-shaped stone at the
                                                                                               top of an arch that holds its struc-
                                                                                               ture together. A keystone species
                                                                                               (b) is a species that exerts great
                                Keystone                                                       influence on a community’s com-
                                                                                               position and structure. Sea otters
                                                                                               consume sea urchins that eat kelp in
                                                                                               coastal waters of the Pacific. Otters
                     (a) A keystone
                                                                                               keep urchin numbers down, allow-
                                                                                               ing lush underwater forests of kelp
                                                                                               to grow, providing habitat for many
                                                                Sea otter absent               species. When otters are absent,
                                                                                               urchins increase and devour the kelp,
                                                                                               destroying habitat and depressing
                           Sea otter                                                           species diversity.
                           (keystone
                           species)               Kelp






                                                                                     Overgrazed
                                                                                     kelp




                                                                                     Explosion of
                                                  Sea                                sea urchin
                                                  urchin                             population




                     (b) A keystone species


                        The removal of top predators in the United States was   redistribute nutrients, and selectively protect or destroy differ-
                     an uncontrolled large-scale experiment with unintended con-  ent insects and plants within the radius of their colonies. And
                     sequences, but ecologists have verified the keystone species   zebra and quagga mussels alter the communities they invade
                     concept in controlled scientific experiments. Classic research   by filtering plankton out of the water.
                     by marine biologist Robert Paine established that the preda-  Less  conspicuous  organisms  toward  the  bottoms  of
                     tory sea star Pisaster ochraceus shapes the community com-  food chains may exert still more impact. Remove the fungi
                     position of intertidal organisms (p. 445) on North America’s   that decompose dead matter, or the insects that control plant
                     Pacific  coast. When  Pisaster is present in this community,   growth, or the phytoplankton that are the base of the marine
                     species diversity is high, with various types of barnacles,   food chain, and a community may change very rapidly indeed.
                     mussels, and algae. When Pisaster is removed, the mussels   However, because there are usually more species at lower
                     it preys on become numerous and displace other species, sup-  trophic levels, it is less likely that any single one of them alone
                     pressing species diversity.                          has wide influence. Often if one species is removed, other spe-
                        Research off the U.S. Atlantic coast published in 2007   cies that remain may be able to perform many of its functions.
                     suggests that the reduction of shark populations by commer-
                     cial fishing has allowed populations of certain skates and rays
                     to increase, which in turn has depressed numbers of bay scal-  Communities respond to disturbance
                     lops and other bivalves they eat.                    in various ways
                        Animals at high trophic levels—such as wolves, sea stars,
                     sharks, and sea otters (see Figure 4.12)—are often viewed as   The removal of a keystone species is just one type of distur-
                     keystone species that can trigger trophic cascades. However,   bance that can modify the composition, structure, or function
                     other types of organisms also exert strong community-wide   of an ecological community. In ecological terms, a disturbance
                     effects. “Ecosystem engineers” physically modify the envi-  is an event that affects environmental conditions rapidly and
                     ronment shared by community members. Beavers build dams   drastically, resulting in changes to the community and ecosys-
                     across streams, creating ponds and swamps by flooding large   tem. A disturbance can be as localized as a tree falling in a
                     areas of land. Prairie dogs dig burrows that aerate the soil   forest, creating a gap in the canopy that lets in light and alters
             102     and serve as homes for other animals. Ants disperse seeds,   conditions for plants and animals in the gap. Or it can be as







           M04_WITH7428_05_SE_C04.indd   102                                                                                    12/12/14   2:55 PM
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