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large and severe as a hurricane, tornado, or volcanic eruption.
                        Some disturbances are sudden, such as landslides or floods,
                        while others are more gradual, such as climate change. Some
                        disturbances  regularly reoccur  and are  considered  normal
                        aspects of a system (such as periodic fire, seasonal storms, or
                        cyclic insect outbreaks). Organisms may, in fact, adapt to regu-
                        lar and predictable types of disturbance. For instance, many
                        plants that grow in fire-prone regions have evolved ways of
                        surviving fire and have seeds that depend on fire to germinate
                        in the nutrient-rich soil that fire leaves behind. Today, human
                        impacts are major sources of disturbance for ecological com-
                        munities worldwide—from habitat alteration to pollution to the
                        introduction of non-native species such as the zebra mussel.
                            Communities are dynamic systems and may respond
                        to  disturbance  in  several  ways.  A  community  that  resists
                        change and remains stable despite disturbance is said to show
                        resistance to the disturbance. Alternatively, a community may
                        show resilience, meaning that it changes in response to distur-
                        bance but later returns to its original state. Or, a community
                        may be modified by disturbance permanently and never return   Figure 4.13 Primary succession begins as organisms
                        to its original state.                               colonize a lifeless new surface. Here, plants establish on a
                                                                             volcanic lava flow.

                        Succession follows severe disturbance                landscape (Figure 4.13). Species that arrive first and colonize
                                                                             the new substrate are referred to as pioneer species. Pioneer
                        If a disturbance is severe enough to eliminate all or most   species are well adapted for colonization, having traits such as
                        of the species in a community, the affected site may then   spores or seeds that can travel long distances.
                        undergo a somewhat predictable series of changes that ecolo-  The pioneers best suited to colonizing bare rock are the
                        gists have traditionally called succession. In the conventional   mutualistic aggregates of fungi and algae known as lichens.
                        view of this process, there are two types of succession.   In lichens, the algal component provides food and energy via
                        Primary succession follows a disturbance so severe that no   photosynthesis while the fungal component grips the rock and
                        vegetation or soil life remains from the community that had   captures the moisture that both organisms need. As lichens
                        occupied the site. In primary succession, a biotic commu-  grow, they  secrete acids  that break  down the rock  surface,
                        nity is built essentially from scratch. In contrast, secondary   beginning the formation of soil. Small plants and insects
                        succession begins when a disturbance dramatically alters an   arrive, providing more nutrients and habitat. As time passes,
                        existing community but does not destroy all living things   larger  plants  and animals  establish  themselves,  vegetation
                        or all organic matter in the soil. In secondary succession,   increases, and species diversity rises.
                        vestiges of the previous community remain, and these build-  Secondary succession begins when a fire, a hurricane, log-
                        ing blocks help shape the process.                   ging, or farming removes much of the biotic community. Consider
                            At terrestrial sites, primary succession takes place after   a farmed field in eastern North America that has been abandoned
                        a bare expanse of rock, sand, or sediment becomes newly   (Figure 4.14). After farming ends, the site will be colonized by
                        exposed to the atmosphere.  This can occur when glaciers   pioneer species of grasses, herbs, and forbs that disperse well or
                        retreat, lakes dry up, or volcanic lava or ash spreads across the   were already in the vicinity. Soon, shrubs and fast-growing trees   CHAPTER 4 • S PEC i ES   i n TERA CT i on S   A nd Co mmuni T y E C ology




                                                                                     Hardwoods
                                                                        Pines
                                                         Saplings,
                                                        poplar trees
                                                                                                   Figure 4.14 Secondary
                                                                                                   succession occurs after a
                                           Shrubs,                                                 disturbance (such as a fire,
                                           seedlings                                               landslide, or farming) removes
                            Grasses,
                           herbs, forbs                                                            most vegetation from an
                                                                                                   area. Shown is a typical series of
                                                                                                   changes in a plant community of
                                                                                                   eastern North America following
                        Time                                                                       abandonment of a farmed field.  103







           M04_WITH7428_05_SE_C04.indd   103                                                                                    12/12/14   2:55 PM
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