Ecological disturbance - Opportunities, Resilience, Recovery

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Although disturbances tend to negatively affect populations of resident plants, animals, and other organisms in a given ecosystem, they provide some fugitive species with opportunities to move into and gain footholds in ecosystems whose biological communities once excluded them. This process results in an increase in the biodiversity of the ecosystem. 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Although disturbances tend to negatively affect populations of resident plants, animals, and other organisms in a given ecosystem, they provide some fugitive species with opportunities to move into and gain footholds in ecosystems whose biological communities once excluded them. This process results in an increase in the biodiversity of the ecosystem. Several in-depth examples of abiotic and biotic disturbances, including the ecological opportunities they create, are"/> <meta property="og:type" content="ARTICLE"/> <meta property="og:title" content="Ecological disturbance - Opportunities, Resilience, Recovery | Britannica"/> <meta property="og:description" content="Ecological disturbance - Opportunities, Resilience, Recovery: Disturbances may be generated by abiotic, or nonliving, forces such as weather and wildfires, or they may occur as a result of biotic, or living, forces such as disease and invasions of exotic species. Although disturbances tend to negatively affect populations of resident plants, animals, and other organisms in a given ecosystem, they provide some fugitive species with opportunities to move into and gain footholds in ecosystems whose biological communities once excluded them. This process results in an increase in the biodiversity of the ecosystem. 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id="content" class="md-content"> <div class="md-article-container template-desktop infinite-pagination"> <div class="infinite-scroll-container article last"> <article class="article-content container-lg qa-content px-0 pt-0 pb-40 py-lg-20 content md-expanded" data-topic-id="1873060"> <div class="grid gx-0"> <div class="col-auto"> <div class="topic-left-rail md-article-drawer position-relative d-flex border-right-sm border-left-sm open"> <div class="drawer d-flex flex-column open"> <div class="left-rail-section-content"> <div class="topic-left-rail-header text-truncate bg-gray-50 position-relative text-right d-flex align-items-center"> <div class="tlr-title px-20 py-15 text-left"> <em class="material-icons text-gray-400 d-lg-none" data-icon="toc"></em> <a class="font-serif font-weight-bold text-black link-blue" href="https://www.britannica.com/science/ecological-disturbance">ecological disturbance</a> </div> <button aria-label="Close" class="js-sections-close-button btn-link btn-sm btn d-lg-none position-absolute top-0 p-10 right-0" > <em class="material-icons font-26" data-icon="close"></em> </button> </div> <div class="section-content pl-10 pr-20 pl-sm-50 pr-sm-60 pl-lg-5 pr-lg-10 pt-10 pt-lg-0 bg-gray-50 clear-catfish-ad"> <div class="toc mb-20"> <div class="font-serif font-14 font-weight-bold mx-15 mb-15 mt-20"> Table of Contents </div> <ul class="list-unstyled my-0" data-level="h1"><li data-target="#ref1"><div class="pl-25"><a class="link-gray-900 w-100" href="/science/ecological-disturbance">Introduction</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li><li data-target="#ref307199"><div class="d-flex align-items-center"><button class="h1-link-drawer-button btn btn-xs btn-circle d-flex rounded" type="button" aria-label="Toggle Heading"><em class="material-icons font-18" data-icon="keyboard_arrow_right"></em></button><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery">Characteristics of disturbance and recovery</a></div><div class="ml-40 toc-drawer sub-toc-drawer"><ul class="list-unstyled" data-level="h2"><li data-target="#ref307200"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery#ref307200">Disturbance intensity and the pace of recovery</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307201"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery#ref307201">Disturbance frequency and recovery</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307202"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery#ref307202">Spatial distribution</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307203"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery#ref307203">Size distribution of patches</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307204"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery#ref307204">Intermediate disturbance hypothesis</a></li></ul></div></li><li data-target="#ref307205"><div class="d-flex align-items-center"><button class="h1-link-drawer-button btn btn-xs btn-circle d-flex rounded" type="button" aria-label="Toggle Heading"><em class="material-icons font-18" data-icon="keyboard_arrow_right"></em></button><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Ecological-opportunities-provided-by-disturbances">Ecological opportunities provided by disturbances</a></div><div class="ml-40 toc-drawer sub-toc-drawer"><ul class="list-unstyled" data-level="h2"><li data-target="#ref307206"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Ecological-opportunities-provided-by-disturbances#ref307206">Fire legacy in Yellowstone</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307207"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Ecological-opportunities-provided-by-disturbances#ref307207">Disturbance and biodiversity in intertidal zones</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307208"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Ecological-opportunities-provided-by-disturbances#ref307208">Disturbance and biodiversity in prairie landscapes</a></li></ul><ul class="list-unstyled" data-level="h2"><li data-target="#ref307209"><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Ecological-opportunities-provided-by-disturbances#ref307209">Depression creation by marine animals</a></li></ul></div></li><li data-target="#ref307210"><div class="d-flex align-items-center"><div class="ml-25"></div><a class="w-100 link-gray-900" href="/science/ecological-disturbance/Ecological-opportunities-provided-by-disturbances#ref307210">Management implications</a></div><div class="ml-40 toc-drawer sub-toc-drawer"></div></li></ul> <a class="toc-extra-link link-gray-900" href="https://www.britannica.com/science/ecological-disturbance/additional-info">References & Edit History</a> <a class="toc-extra-link link-gray-900" href="/facts/ecological-disturbance">Quick Facts & Related Topics</a> </div> <div class="tlr-media-slider pb-10 mb-30"> <a class="section-header link-gray-900 font-serif font-14 font-weight-bold mb-10 mx-10" href="https://www.britannica.com/science/ecological-disturbance/images-videos">Images</a> <div class="slider js-slider position-relative d-inline-flex align-items-center mw-100 "> <div class="slider-container js-slider-container overflow-hidden d-flex overflow-hidden text-nowrap ml-15"> <a href="https://cdn.britannica.com/11/154611-050-1222C65C/view-remains-trees-hillside-wildfires-forest-fire.jpg" data-href="/media/1/1873060/170896" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/11/154611-004-17A3E23C/view-remains-trees-hillside-wildfires-forest-fire.jpg" alt="ecological disturbance caused by forest fire" height="50" /> </a> <a href="https://cdn.britannica.com/97/95197-004-7F9B8F09/climax-community-succession-rock-areas-plants-hundreds.jpg" data-href="/media/1/1873060/125658" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/97/95197-004-7F9B8F09/climax-community-succession-rock-areas-plants-hundreds.jpg" alt="primary ecological succession" height="50" /> </a> <a href="https://cdn.britannica.com/98/95198-004-2619E3FA/succession-disturbance-fire-flood-environments-primary-stages.jpg" data-href="/media/1/1873060/125659" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/98/95198-004-2619E3FA/succession-disturbance-fire-flood-environments-primary-stages.jpg" alt="secondary ecological succession" height="50" /> </a> <a href="https://cdn.britannica.com/22/115822-004-8D237E70/tree-lava-patch-Ecosystem-recovery-eruption-California.jpg" data-href="/media/1/1873060/170897" class="media-overlay-link d-inline-block mr-5"> <img loading="lazy" src="https://cdn.britannica.com/22/115822-004-8D237E70/tree-lava-patch-Ecosystem-recovery-eruption-California.jpg" alt="forest recovery" height="50" /> </a> </div> <button disabled class="prev-button js-prev-button position-absolute btn btn-circle shadow btn-blue " aria-label="Previous"> <span class="material-icons md-24" data-icon="keyboard_arrow_left"></span> </button> <button disabled class="next-button js-next-button position-absolute btn btn-circle shadow btn-blue " aria-label="Next"> <span class="material-icons md-24" data-icon="keyboard_arrow_right"></span> 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Paine</div> <div class="editor-description font-12 font-serif mt-5 clamp-description text-black">Professor emeritus, Department of Biology, University of Washington, Seattle, Washington.</div> </a> <div data-popper-arrow></div> </div> <span class="btn btn-link editor-link p-0 qa-byline-link gtm-byline font-12 byline-contributor text-decoration-underline"> Robert T. Paine</span></div> <div class="font-serif font-12 text-gray-700"> <span class="qa-fact-checked-by">Fact-checked by</span> <div class="editor-popover popover p-0"> <a class="d-block p-20 font-12" href="/editor/The-Editors-of-Encyclopaedia-Britannica/4419" > <div class="editor-title font-16 font-weight-bold">The Editors of Encyclopaedia Britannica</div> <div class="editor-description font-12 font-serif mt-5 text-black">Encyclopaedia Britannica's editors oversee subject areas in which they have extensive knowledge, whether from years of experience gained by working on that content or via study for an advanced degree. They write new content and verify and edit content received from contributors.</div> </a> <div data-popper-arrow></div> </div> <span class="btn btn-link editor-link p-0 qa-byline-link font-12 "> The Editors of Encyclopaedia Britannica</span></div> <div class="last-updated font-12 font-serif"> <a class="byline-edit-history" href="https://www.britannica.com/science/ecological-disturbance/additional-info#history" rel="nofollow">Article History</a> </div></div> </div> <button class="d-flex d-lg-none btn btn-outline-blue border rounded-sm shadow-sm mobile-toc-button gtm-mobile-toc-inline-button d-none d-sm-block js-sections-inline-button module-spacing btn d-lg-none"> <em class="material-icons mr-5 ml-n10 my-n5 md-icon" data-icon="toc"></em> Table of Contents </button> <div class="d-flex d-sm-none flex-row"> <button class="d-flex d-lg-none btn btn-outline-blue border rounded-sm shadow-sm mobile-toc-button gtm-mobile-toc-inline-button js-sections-inline-button module-spacing"> <em class="material-icons mr-5 ml-n10 my-n5 md-icon" data-icon="toc"></em> Table of Contents </button> <button class="ai-ask-button btn border-2 ai-ask-button btn border-2 module-spacing btn-sm js-inline-ai-ask-button btn-outline-red-400 border-red-400 p-10 ml-5"> Ask the Chatbot a Question </button> </div> <div class="js-qf-module qf-module px-40 px-sm-20 py-15 mx-auto module-spacing font-14 bg-gray-50 rounded"> <div class="facts-list mt-10"> <div class=""> <div class="js-fact mb-10 line-clamp clamp-3"> <dl> <dt>Related Topics: </dt> <dd><a href="/science/ecosystem" topicid="178597">ecosystem</a></dd> <dd><a href="/science/habitat-loss" topicid="1377188">habitat loss</a></dd> <dd><a href="/science/secondary-succession" topicid="531823">secondary succession</a></dd> </dl> <button class="js-more-btn d-none btn btn-unstyled font-12 bg-gray-50" aria-label="Toggle more/less fact data"> <em class="js-content link-blue">(Show more)</em> </button> </div> <div class="text-center"> <a class="btn btn-sm btn-link p-0" href="/facts/ecological-disturbance"> See all related content </a> </div> </div> </div> </div><span class="marker before-article"></span><span class="marker h3"></span><section data-level="1" id="ref307205"> <span class="marker PREMOD1 mod-inline"></span><p class="topic-paragraph">Disturbances may be generated by abiotic, or nonliving, forces such as weather and wildfires, or they may occur as a result of biotic, or living, forces such as disease and invasions of exotic species. Although disturbances tend to negatively affect populations of resident plants, animals, and other organisms in a given <a href="https://www.britannica.com/science/ecosystem" class="md-crosslink autoxref " data-show-preview="true">ecosystem</a>, they provide some fugitive species with opportunities to move into and gain footholds in ecosystems whose biological <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="communities" href="https://www.merriam-webster.com/dictionary/communities" data-type="MW">communities</a> once excluded them. This process results in an increase in the <a href="https://www.britannica.com/science/biodiversity" class="md-crosslink autoxref " data-show-preview="true">biodiversity</a> of the ecosystem. Several in-depth examples of abiotic and biotic disturbances, including the ecological opportunities they create, are provided below.</p><span class="marker MOD1 mod-inline"></span> <section data-level="2" id="ref307206"><h2 class="h2">Fire legacy in Yellowstone</h2> <span class="marker PREMOD2 mod-inline"></span><p class="topic-paragraph">In 1988 <span id="ref1175429"></span><a href="https://www.britannica.com/place/Yellowstone-National-Park" class="md-crosslink " data-show-preview="true">Yellowstone National Park</a> was the site of an immense <a href="https://www.britannica.com/science/forest-fire" class="md-crosslink autoxref " data-show-preview="true">forest fire</a>. Of the park’s 9,000 square km (about 3,500 square miles), much of it dominated by lodgepole pine (<em>Pinus contorta</em>), more than half were scorched, and approximately 36 percent of the park was burned. The fire generated a landscape mosaic of incinerated and unburned pine stands. Fifteen years later tree density varied from 566 to 545,000 trees per hectare, depending on the severity of the local burn and the proximity of the scorched areas to surviving and reproducing trees. The <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="legacy" href="https://www.merriam-webster.com/dictionary/legacy" data-type="MW">legacy</a> of each site had important <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="implications" href="https://www.merriam-webster.com/dictionary/implications" data-type="MW">implications</a> for the recovery process.</p><span class="marker MOD2 mod-inline"></span> <span class="marker PREMOD3 mod-inline"></span><p class="topic-paragraph">The fire history of large terrestrial ecosystems such as Yellowstone National Park can be obtained from historical records or by determining the ages of the resident trees. In Yellowstone, seven sites that began to recover from 1 to about 300 years ago following burning events were sampled for their <a href="https://www.britannica.com/science/species-richness" class="md-crosslink autoxref " data-show-preview="true">species richness</a>. The sites with greatest biological species richness were some 13 to 25 years into their recovery. This data supports the notion that disturbance separated in space and time yields a mosaic of habitats in various stages of recovery and that such landscape heterogeneity makes the <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="collective" href="https://www.merriam-webster.com/dictionary/collective" data-type="MW">collective</a> region more biologically <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="diverse" href="https://www.merriam-webster.com/dictionary/diverse" data-type="MW">diverse</a> than any individual site found within it.</p><span class="marker MOD3 mod-inline"></span></section> <section data-level="2" id="ref307207"><h2 class="h2">Disturbance and biodiversity in intertidal zones</h2> <span class="marker PREMOD4 mod-inline"></span><p class="topic-paragraph">On marine rocky shores, disturbance is commonplace, and the size of the disturbed area varies depending on the physical forces involved and the degree of exposure to waves. For instance, floating logs and other debris in the <a href="https://www.britannica.com/science/littoral-zone" class="md-crosslink autoxref " data-show-preview="true">intertidal zone</a> can batter the resident organisms, producing localized mortality. In addition, boulders in intertidal zones are overturned as a function of their mass and the severity of the wave stress. If the boulders are too large to be overturned, the surface is stable and a characteristic algal <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="community" href="https://www.merriam-webster.com/dictionary/community" data-type="MW">community</a> develops on each boulder’s surface. When the boulders are overturned, however, the residents are killed, and the surface is invaded by a suite of different species. As a result, the size of the boulders, combined with the strength of the wave forces, influences the local patterns of biodiversity in the intertidal zone.</p><span class="marker MOD4 mod-inline"></span> <span class="marker PREMOD5 mod-inline"></span><p class="topic-paragraph">The <span id="ref1175501"></span><a href="https://www.britannica.com/animal/California-mussel" class="md-crosslink ">California mussel</a>, which dominates exposed rocky shores in the western <a href="https://www.britannica.com/place/United-States" class="md-crosslink autoxref " data-show-preview="true">United States</a> and Canada, serves as another example. Winter storms scour mussels off the rock surfaces, producing patches or gaps that vary in initial size from less than 100 square cm (15.5 square inches) to 35 square metres (about 377 square feet). Recovery to the original mussel-dominated space can take 8 to 35 years, depending on many factors but especially on the tidal height of the mussel bed and the productivity of the surrounding waters. These events can be modeled demographically, permitting a view of the <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="dynamics" href="https://www.merriam-webster.com/dictionary/dynamics" data-type="MW">dynamics</a> of spatial pattern as it varies with gap size, disturbance intensity, and rate of recovery.</p><div class="module-spacing"> </div><span class="marker MOD5 mod-inline"></span></section> <section data-level="2" id="ref307208"><h2 class="h2">Disturbance and biodiversity in prairie landscapes</h2> <span class="marker PREMOD6 mod-inline"></span><p class="topic-paragraph">Biologically based disturbances also provide opportunities for previously excluded species to invade and occupy a disrupted ecosystem. The earliest invaders are fugitive species, and disturbance is a basic requisite of this common ecological strategy. For instance, the common <span id="ref1175502"></span><a href="https://www.britannica.com/plant/teasel" class="md-crosslink " data-show-preview="true">teasel</a> (<em>Dipsacus sylvestris</em>) often functions as a biennial plant that inhabits disturbed sites. It has a broad-leafed basal rosette (a cluster of leaves forming a crowded circle), up to about 100 cm (about 39 inches) in diameter, that dies back as the plant matures, usually in its second year. This space can be invaded by <span id="ref1175504"></span><a href="https://www.britannica.com/plant/common-winter-cress" class="md-crosslink ">common winter cress</a> (<em>Barbarea vulgaris</em>), an invasive winter annual, as well as the seeds of later successional species that enrich the local biological <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="diversity" href="https://www.merriam-webster.com/dictionary/diversity" data-type="MW">diversity</a>.</p><span class="marker MOD6 mod-inline"></span> <span class="marker PREMOD7 mod-inline"></span><p class="topic-paragraph">American <a href="https://www.britannica.com/animal/badger" class="md-crosslink " data-show-preview="true">badgers</a> (<em><span id="ref1175505"></span><a href="https://www.britannica.com/animal/American-badger" class="md-crosslink ">Taxidea taxus</a></em>) create localized disturbances in tallgrass prairies by digging for their rodent prey; digging produces mounds of dirt 0.2–0.3 square metre (2.2–3.2 square feet) in size. These holes and dirt mounds function as localized disturbances that enrich the field’s spatial patterning and provide a necessary resource (i.e., bare ground) for a number of fugitive plant species—for example, the stiff goldenrod (<em>Solidago rigida</em>).</p><span class="marker MOD7 mod-inline"></span></section> <section data-level="2" id="ref307209"><h2 class="h2">Depression creation by marine animals</h2> <span class="marker PREMOD8 mod-inline"></span><p class="topic-paragraph">On unconsolidated marine sediments, the most <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="conspicuous" href="https://www.merriam-webster.com/dictionary/conspicuous" data-type="MW">conspicuous</a> agents of disturbance are foraging <a href="https://www.britannica.com/animal/vertebrate" class="md-crosslink " data-show-preview="true">vertebrates</a>, such as <a href="https://www.britannica.com/animal/whale" class="md-crosslink " data-show-preview="true">whales</a>, <a href="https://www.britannica.com/animal/walrus" class="md-crosslink " data-show-preview="true">walrus</a>, <a href="https://www.britannica.com/animal/sea-otter" class="md-crosslink " data-show-preview="true">sea otters</a>, and <span id="ref1175506"></span><a href="https://www.britannica.com/animal/ray-fish" class="md-crosslink " data-show-preview="true">rays</a>. These consumers dig pits or make depressions in which <a href="https://www.britannica.com/science/detritus" class="md-crosslink " data-show-preview="true">detritus</a> accumulates, attracting secondary consumers. Such pits and depressions can range from less than 1 metre (3.3 feet) to more than 20 metres (about 66 feet) in diameter. The depths of these disturbances vary; some are more than 40 cm (about 16 inches) deep. Collectively, they can cover over 30 percent of a bay’s bottom. In California the pits dug by feeding rays cause increased <a href="https://www.britannica.com/science/predation" class="md-crosslink autoxref " data-show-preview="true">prey</a> mortality, but they also serve as receptacles of organic materials, a resource <a class="md-dictionary-link md-dictionary-tt-off eb" data-term="consumed" href="https://www.britannica.com/dictionary/consumed" data-type="EB">consumed</a> by many members of the <a href="https://www.britannica.com/science/community-biology" class="md-crosslink autoxref " data-show-preview="true">biological community</a>. The disturbance of the sediment by rays yields a spatial mosaic that varies by degrees of recolonization.</p><span class="marker MOD8 mod-inline"></span></section></section> <span class="marker h4"></span><section data-level="1" id="ref307210"><h2 class="h1">Management implications</h2> <span class="marker PREMOD9 mod-inline"></span><p class="topic-paragraph">Limiting the impact that a disturbance will have on an ecosystem is difficult because the timing of the disturbance, its location, its intensity, and its spatial patterning are unpredictable. However, a future disturbance event may be forecast based on time series data (i.e., records of past occurrences) and the event’s probability of occurrence across a long interval. For example, geologists have assigned probabilities to <a href="https://www.britannica.com/science/earthquake-geology" class="md-crosslink " data-show-preview="true">earthquake</a> occurrences along the <a href="https://www.britannica.com/place/Ring-of-Fire" class="md-crosslink " data-show-preview="true">Pacific Ring of Fire</a>, and meteorologists have attempted to calculate the probability of a 100-year snow event (an uncommonly large snowfall that has a 1 percent probability of occurring in any given year). In contrast, large disturbances of biological origin are much more difficult to predict, because of the <a class="md-dictionary-link md-dictionary-tt-off eb" data-term="vast" href="https://www.britannica.com/dictionary/vast" data-type="EB">vast</a> numbers of species and species’ interactions involved. In fact, the most-devastating biological disturbances are caused by poorly known or unknown microorganisms, such as those that caused the rapid, near-total die-off of the <a href="https://www.britannica.com/animal/sea-urchin" class="md-crosslink autoxref " data-show-preview="true">sea urchin</a> <em>D. antillarum</em> in the 1980s (<a href="https://www.britannica.com/science/ecological-disturbance/Characteristics-of-disturbance-and-recovery#ref307201" class="md-crosslink " data-show-preview="true"><em>see above</em></a>) or the introduced fungal pathogen <em>Cryphonectria parasitica</em> that destroyed American <a href="https://www.britannica.com/plant/chestnut" class="md-crosslink " data-show-preview="true">chestnut</a> trees (<em>Castanea dentata</em>) during the first half of the 20th century.</p><span class="marker MOD9 mod-inline"></span> <span class="marker PREMOD10 mod-inline"></span><p class="topic-paragraph">Large, long-lasting ecological disturbances that stress natural ecosystems on a global (rather than local) scale pose significant challenges to management and restoration efforts. For example, the global impact on marine ecosystems of <a href="https://www.britannica.com/science/ocean-acidification" class="md-crosslink " data-show-preview="true">ocean acidification</a>, which stems from the absorption of <a href="https://www.britannica.com/science/carbon-dioxide" class="md-crosslink " data-show-preview="true">carbon dioxide</a> by seawater, is not well understood. Likewise, while the rise of average global near-surface air temperatures (i.e., <a href="https://www.britannica.com/science/global-warming" class="md-crosslink " data-show-preview="true">global warming</a>) has been found to increase <a class="md-dictionary-link md-dictionary-tt-off eb" data-term="drought" href="https://www.britannica.com/dictionary/drought" data-type="EB">drought</a> frequency and severity, thereby modifying patterns of ecosystem productivity and the geographic ranges of many species, how these changes could impact ecosystems globally remains uncertain. In the case of overfishing, which is a persistent global problem—in 2010, for example, 53 percent of the world’s fish stocks were fully exploited (i.e., operating at maximum sustainable yield) and another 17 percent were overexploited—marine ecologists have documented changes in the structure of fish populations and that of their prey that ultimately could change the structure of the ecosystems they inhabit. The consequences of the <a href="https://www.britannica.com/science/trophic-cascade" class="md-crosslink " data-show-preview="true">trophic cascades</a> that are likely to follow such changes are yet to be fully explored.</p><span class="marker MOD10 mod-inline"></span> <span class="marker PREMOD11 mod-inline"></span><p class="topic-paragraph">Two additional concerns complicate management and restoration in the <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="context" href="https://www.merriam-webster.com/dictionary/context" data-type="MW">context</a> of ecological disturbance. The first is <a href="https://www.britannica.com/science/climate-change" class="md-crosslink " data-show-preview="true">climate change</a>. It is not known whether the imposition of more general but gradual disturbances associated with climate change will magnify the consequences of natural disturbances and alter the rates of recovery in unanticipated ways. The second is the involvement of <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="compounding" href="https://www.merriam-webster.com/dictionary/compounding" data-type="MW">compounding</a> factors. If ecosystems collapse when disturbances are <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="compounded" href="https://www.merriam-webster.com/dictionary/compounded" data-type="MW">compounded</a> (that is, occur within the system’s expected recovery time), <a class="md-dictionary-link md-dictionary-tt-off mw" data-term="alternative" href="https://www.merriam-webster.com/dictionary/alternative" data-type="MW">alternative</a> states may be formed that present novel combinations of species and unknown dynamics, challenging both management and restoration efforts.</p><span class="marker MOD11 mod-inline"></span><span class="md-signature"><a href="/contributor/Robert-T-Paine/9197251">Robert T. 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Ecological disturbance - Opportunities, Resilience, Recovery | Britannica