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A marine coastal ecosystem is a marine ecosystem which occurs when the land meets the ocean. Marine coastal ecosystems include many different types of marine habitats, such as estuaries and lagoons, salt marshes and mangrove forests, seagrass meadows and coral reefs, kelp forests and backwaters. Directly and indirectly these provide a vast range of ecosystem services for humans, such as sequestering carbon, cycling nutrients and elements, providing nurseries and fishing grounds for commercial fisheries, preventing coastal erosion and moderating extreme events, as well as providing recreational services and supporting tourism.

Overview

Coastal seas are highly productive systems, providing an array of ecosystem services to humankind, such as processing of nutrient effluents from land and climate regulation.^[1] However, coastal ecosystems are threatened by human-induced pressures such as climate change and eutrophication. In the coastal zone, the fluxes and transformations of nutrients and carbon sustaining coastal ecosystem functions and services are strongly regulated by benthic (that is, occurring at the seafloor) biological and chemical processes.^[1]

Coastal systems also contribute to the regulation of climate and nutrient cycles, by efficiently processing anthropogenic emissions from land before they reach the ocean.^[2]^[3]^[4]^[5] The high value of these ecosystem services is obvious considering that a large proportion of the world population lives close to the coast.^[6]^[7]^[1]

Currently, coastal seas around the world are undergoing major ecological changes driven by human-induced pressures, such as climate change, anthropogenic nutrient inputs, overfishing and the spread of invasive species.^[8]^[9] In many cases, the changes alter underlying ecological functions to such an extent that new states are achieved and baselines are shifted.^[10]^[11]^[1]

In 2015, the United Nations established 17 Sustainable Development Goals with the aim of achieving certain targets by 2030. Their mission statement for their 14th goal, Life below water, is to "conserve and sustainably use the oceans, seas and marine resources for sustainable development".^[12] The United Nations has also declared 2021–2030 the UN Decade on Ecosystem Restoration, but restoration of coastal ecosystems is not receiving appropriate attention.^[13]

Intertidal zone

Intertidal zones are the areas that are visible and exposed to air during low tide and covered up by saltwater during high tide.^[14] There are four physical divisions of the intertidal zone with each one having its distinct characteristics and wildlife. These divisions are the Spray zone, High intertidal zone, Middle Intertidal zone, and Low intertidal zone. The Spray zone is a damp area that is usually only reached by the ocean and submerged only under high tides or storms. The high intertidal zone is submerged at high tide but remains dry for long periods between high tides.^[14] Due to the large variance of conditions possible in this region, it is inhabited by resilient wildlife that can withstand these changes such as barnacles, marine snails, mussels and hermit crabs.^[14] Tides flow over the middle intertidal zone two times a day and this zone has a larger variety of wildlife.^[14] The low intertidal zone is submerged nearly all the time except during the lowest tides and life is more abundant here due to the protection that the water gives.^[14]

Estuaries

Estuaries occur where there is a noticeable change in salinity between saltwater and freshwater sources. This is typically found where rivers meet the ocean or sea. The wildlife found within estuaries is unique as the water in these areas is brackish - a mix of freshwater flowing to the ocean and salty seawater.^[15] Other types of estuaries also exist and have similar characteristics as traditional brackish estuaries. The Great Lakes are a prime example. There, river water mixes with lake water and creates freshwater estuaries.^[15] Estuaries are extremely productive ecosystems that many humans and animal species rely on for various activities.^[16] This can be seen as, of the 32 largest cities in the world, 22 are located on estuaries as they provide many environmental and economic benefits such as crucial habitat for many species, and being economic hubs for many coastal communities.^[16] Estuaries also provide essential ecosystem services such as water filtration, habitat protection, erosion control, gas regulation nutrient cycling, and it even gives education, recreation and tourism opportunities to people.^[17]

Lagoons

Lagoons are areas that are separated from larger water by natural barriers such as coral reefs or sandbars. There are two types of lagoons, coastal and oceanic/atoll lagoons.^[18] A coastal lagoon is, as the definition above, simply a body of water that is separated from the ocean by a barrier. An atoll lagoon is a circular coral reef or several coral islands that surround a lagoon. Atoll lagoons are often much deeper than coastal lagoons.^[19] Most lagoons are very shallow meaning that they are greatly affected by changed in precipitation, evaporation and wind. This means that salinity and temperature are widely varied in lagoons and that they can have water that ranges from fresh to hypersaline.^[19] Lagoons can be found in on coasts all over the world, on every continent except Antarctica and is an extremely diverse habitat being home to a wide array of species including birds, fish, crabs, plankton and more.^[19] Lagoons are also important to the economy as they provide a wide array of ecosystem services in addition to being the home of so many different species. Some of these services include fisheries, nutrient cycling, flood protection, water filtration, and even human tradition.^[19]

Vegetated coastal ecosystems

Coastal wetlands

Mangrove forests

Salt marshes

Coastal wetlands are among the most productive ecosystems on Earth and generate vital services that benefit human societies around the world. Sediment-stabilization by wetlands such as salt marshes and mangroves serves to protect coastal communities from storm-waves, flooding, and land erosion.^[24] Coastal wetlands also reduce pollution from human waste,^[25]^[26] remove excess nutrients from the water column,^[27] trap pollutants,^[28] and sequester carbon.^[29] Further, near-shore wetlands act as both essential nursery habitats and feeding grounds for game fish, supporting a diverse group of economically important species.^[30]^[31]^[32]^[33]^[34]

Mangrove forests

Mangroves are trees or shrubs that grow in low-oxygen soil near coastlines in tropical or subtropical latitudes.^[35] They are an extremely productive and complex ecosystem that connects the land and sea. Mangroves consist of species that are not necessarily related to each other and are often grouped for the characteristics they share rather than genetic similarity.^[36] Because of their proximity to the coast, they have all developed adaptions such as salt excretion and root aeration to live in salty, oxygen-depleted water.^[36] Mangroves can often be recognized by their dense tangle of roots that act to protect the coast by reducing erosion from storm surges, currents, wave, and tides.^[35] The mangrove ecosystem is also an important source of food for many species as well as excellent at sequestering carbon dioxide from the atmosphere with global mangrove carbon storage is estimated at 34 million metric tons per year.^[36]

Salt marshes

Salt marshes are a transition from the ocean to the land, where fresh and saltwater mix.^[37] The soil in these marshes is often made up of mud and a layer of organic material called peat. Peat is characterized as waterlogged and root-filled decomposing plant matter that often causes low oxygen levels (hypoxia). These hypoxic conditions causes growth of the bacteria that also gives salt marshes the sulfurous smell they are often known for.^[38] Salt marshes exist around the world and are needed for healthy ecosystems and a healthy economy. They are extremely productive ecosystems and they provide essential services for more than 75 percent of fishery species and protect shorelines from erosion and flooding.^[38] Salt marshes can be generally divided into the high marsh, low marsh, and the upland border. The low marsh is closer to the ocean, with it being flooded at nearly every tide except low tide.^[37] The high marsh is located between the low marsh and the upland border and it usually only flooded when higher than usual tides are present.^[37] The upland border is the freshwater edge of the marsh and is usually located at elevations slightly higher than the high marsh. This region is usually only flooded under extreme weather conditions and experiences much less waterlogged conditions and salt stress than other areas of the marsh.^[37]

Seagrass meadows

Seagrasses form dense underwater meadows which are among the most productive ecosystems in the world. They provide habitats and food for a diversity of marine life comparable to coral reefs. This includes invertebrates like shrimp and crabs, cod and flatfish, marine mammals and birds. They provide refuges for endangered species such as seahorses, turtles, and dugongs. They function as nursery habitats for shrimps, scallops and many commercial fish species. Seagrass meadows provide coastal storm protection by the way their leaves absorb energy from waves as they hit the coast. They keep coastal waters healthy by absorbing bacteria and nutrients, and slow the speed of climate change by sequestering carbon dioxide into the sediment of the ocean floor.

Seagrasses evolved from marine algae which colonized land and became land plants, and then returned to the ocean about 100 million years ago. However, today seagrass meadows are being damaged by human activities such as pollution from land runoff, fishing boats that drag dredges or trawls across the meadows uprooting the grass, and overfishing which unbalances the ecosystem. Seagrass meadows are currently being destroyed at a rate of about two football fields every hour.

Kelp forests

Kelp forests occur worldwide throughout temperate and polar coastal oceans.^[39] In 2007, kelp forests were also discovered in tropical waters near Ecuador.^[40]

Physically formed by brown macroalgae, kelp forests provide a unique habitat for marine organisms^[41] and are a source for understanding many ecological processes. Over the last century, they have been the focus of extensive research, particularly in trophic ecology, and continue to provoke important ideas that are relevant beyond this unique ecosystem. For example, kelp forests can influence coastal oceanographic patterns^[42] and provide many ecosystem services.^[43]

However, the influence of humans has often contributed to kelp forest degradation. Of particular concern are the effects of overfishing nearshore ecosystems, which can release herbivores from their normal population regulation and result in the overgrazing of kelp and other algae.^[44] This can rapidly result in transitions to barren landscapes where relatively few species persist.^[45]^[46] Already due to the combined effects of overfishing and climate change, kelp forests have all but disappeared in many especially vulnerable places, such as Tasmania's east coast and the coast of Northern California.^[47]^[48] The implementation of marine protected areas is one management strategy useful for addressing such issues, since it may limit the impacts of fishing and buffer the ecosystem from additive effects of other environmental stressors.

Coral reefs

Coral reefs are one of the most well-known marine ecosystems in the world, with the largest being the Great Barrier Reef. These reefs are composed of large coral colonies of a variety of species living together. The corals from multiple symbiotic relationships with the organisms around them.^[49]

Coral reef
Global distribution of coral, mangrove, and seagrass diversity
A gentle but threatened dugong grazes a seagrass meadow, encouraging regrowth

Bivalve reefs

Bivalve reefs provide coastal protection through erosion control and shoreline stabilization, and modify the physical landscape by ecosystem engineering, thereby providing habitat for species by facilitative interactions with other habitats such as tidal flat benthic communities, seagrasses and marshes.^[50]

Coastal predators

Food web theory predicts that current global declines in marine predators could generate unwanted consequences for many marine ecosystems. In coastal plant communities, such as kelp, seagrass meadows, mangrove forests and salt marshes, several studies have documented the far-reaching effects of changing predator populations. Across coastal ecosystems, the loss of marine predators appears to negatively affect coastal plant communities and the ecosystem services they provide.^[51]

The green world hypothesis predicts loss of predator control on herbivores could result in runaway consumption that would eventually denude a landscape or seascape of vegetation.^[52] Since the inception of the green world hypothesis, ecologists have tried to understand the prevalence of indirect and alternating effects of predators on lower trophic levels (trophic cascades), and their overall impact on ecosystems.^[53] Multiple lines of evidence now suggest that top predators are key to the persistence of some ecosystems.^[53]^[51]

With an estimated habitat loss greater than 50 percent, coastal plant communities are among the world’s most endangered ecosystems.^[54]^[55]^[56] As bleak as this number is, the predators that patrol coastal systems have fared far worse. Several predatory taxa including species of marine mammals, elasmobranchs, and seabirds have declined by 90 to 100 percent compared to historical populations.^[57]^[58] Predator declines pre-date habitat declines,^[57] suggesting alterations to predator populations may be a major driver of change for coastal systems.^[59]^[60]^[51]

There is little doubt that collapsing marine predator populations results from overharvesting by humans. Localized declines and extinctions of coastal predators by humans began over 40,000 years ago with subsistence harvesting.^[61] However, for most large bodied, marine predators (toothed whales, large pelagic fish, sea birds, pinnipeds, and otters) the beginning of their sharp global declines occurred over the last century, coinciding with the expansion of coastal human populations and advances in industrial fishing.^[57]^[62] Following global declines in marine predators, evidence of trophic cascades in coastal ecosystems started to emerge,^[63]^[64]^[65]^[66] with the disturbing realisation that they affected more than just populations of lower trophic levels.^[53]^[51]

Understanding the importance of predators in coastal plant communities has been bolstered by their documented ability to influence ecosystem services. Multiple examples have shown that changes to the strength or direction of predator effects on lower trophic levels can influence coastal erosion,^[67] carbon sequestration,^[68]^[69] and ecosystem resilience.^[70] The idea that the extirpation of predators can have far-reaching effects on the persistence of coastal plants and their ecosystem services has become a major motivation for their conservation in coastal systems.^[53]^[69]^[51]

Coastal biogeochemistry

Globally, eutrophication is one of the major environmental problems in coastal ecosystems. Over the last century the annual riverine inputs of nitrogen and phosphorus to the oceans have increased from 19 to 37 megatonnes of nitrogen and from 2 to 4 megatonnes of phosphorus.^[71] Regionally, these increases were even more substantial as observed in the United States, Europe and China. In the Baltic Sea nitrogen and phosphorus loads increased by roughly a factor of three and six, respectively.^[72] The riverine nitrogen flux has increased by an order of magnitude to coastal waters of China within thirty years, while phosphorus export has tripled between 1970 and 2000.^[73]^[74]^[1]

Efforts to mitigate eutrophication through nutrient load reductions are hampered by the effects of climate change.^[9] Changes in precipitation increase the runoff of N, P and carbon (C) from land, which together with warming and increased CO₂ dissolution alter the coupled marine nutrient and carbon cycles.^[75]^[76]^[1]

In contrast to the open ocean where biogeochemical cycling is largely dominated by pelagic processes driven primarily by ocean circulation, in the coastal zone, pelagic and benthic processes interact strongly and are driven by a complex and dynamic physical environment.^[77] Eutrophication in coastal areas leads to shifts toward rapidly growing opportunistic algae, and generally to a decline in benthic macrovegetation because of decreased light penetration, substrate change and more reducing sediments.^[78]^[79] Increased production and warming waters have caused expanding hypoxia at the seafloor with a consequent loss of benthic fauna.^[80]^[81] Hypoxic systems tend to lose many long-lived higher organisms and biogeochemical cycles typically become dominated by benthic bacterial processes and rapid pelagic turnover.^[82] However, if hypoxia does not occur, benthic fauna tends to increase in biomass with eutrophication.^[83]^[84]^[85]^[1]

Changes in benthic biota have far-reaching impacts on biogeochemical cycles in the coastal zone and beyond. In the illuminated zone, benthic microphytes and macrophytes mediate biogeochemical fluxes through primary production, nutrient storage and sediment stabilization and act as a habitat and food source for a variety of animals, as shown in the diagram on the left above. Benthic animals contribute to biogeochemical transformations and fluxes between water and sediments both directly through their metabolism and indirectly by physically reworking the sediments and their porewaters and stimulating bacterial processes. Grazing on pelagic organic matter and biodeposition of feces and pseudofeces by suspension-feeding fauna increases organic matter sedimentation rates.^[86]^[87] In addition, nutrients and carbon are retained in biomass and transformed from organic to inorganic forms through metabolic processes.^[88]^[85]^[89] Bioturbation, including sediment reworking and burrow ventilation activities (bioirrigation), redistributes particles and solutes within the sediment and enhances sediment-water fluxes of solutes.^[90]^[91] Bioturbation can also enhance resuspension of particles, a phenomenon termed "bioresuspension".^[92] Together, all these processes affect physical and chemical conditions at the sediment-water interface,^[93] and strongly influence organic matter degradation.^[94] When up-scaled to the ecosystem level, such modified conditions can significantly alter the functioning of coastal ecosystems and ultimately, the role of the coastal zone in filtering and transforming nutrients and carbon.^[1]

Artisan fisheries

Artisanal fisheries use simple fishing gears and small vessels.^[95] Their activities tend to be confined to coastal areas.

Top-down and bottom-up forces determine ecosystem function and dynamics. Fisheries as a top-down force can shorten and destabilize food webs, while effects driven by climate change can alter the bottom-up forces of primary productivity.^[95] Direct human impacts and the full suite of drivers of global change are the main cause of species extinctions in Anthropocene ecosystems,^[96]^[97], with detrimental consequences on ecosystem functioning and their services to human societies.^[98]^[99] The world fisheries crisis is among those consequences, which cuts across fishing strategies, oceanic regions, species, and includes countries that have little regulation and those that have implemented rights-based co-management strategies to reduce overharvesting.^[100]^[101]^[102]^[103]^[95]

Chile has been one of the countries implementing Territorial Use Rights (TURFs)^[104]^[105] over an unprecedented geographic scale to manage the diverse coastal benthic resources using a co-management strategy.^[106]^[107] These TURFS are used for artisanal fisheries. Over 60 coastal benthic species are actively harvested by these artisanal fisheries,^[108] with species that are extracted from intertidal and shallow subtidal habitats.^[109]^[110] The Chilean TURFs system brought significant improvements in sustainability of this complex socio-ecological system, helping to rebuild benthic fish stocks,^[108]^[106] improving fishers’ perception towards sustainability and increasing compliance9, as well as showing positive ancillary effects on conservation of biodiversity.^[111]^[112] However, the situation of most artisanal fisheries is still far from sustainable, and many fish stocks and coastal ecosystems show signs of over exploitation and ecosystem degradation, a consequence of the low levels of cooperation and low enforcement of TURF regulations, which leads to high levels of free-riding and illegal fishing.^[113]^[114]^[115] Thus, it is imperative to improve our understanding of the effects of these multi-species fisheries which simultaneously harvest species at all trophic levels, from kelp primary producers to top carnivores.^[110]^[116]^[95]

Network ecology

To compound things, removal of biomass from the ocean occurs simultaneously with multiple other stressors associated to climate change that compromise the capacity of these socio-ecological systems to respond to perturbations.^[117]^[118]^[119] Besides sea surface temperature, climate change also affects many other physical–chemical characteristics of marine coastal waters (stratification, acidification, ventilation)^[120]^[121] as well as the wind regimes that control surface water productivity along the productive coastal upwelling ecosystems.^[122]^[123]^[124]^[125]^[126] Changes in the productivity of the oceans are reflected in changes of plankton biomass. Plankton contributes approximately half of the global primary production, supports marine food webs, influences the biogeochemical process in the ocean, and strongly affects commercial fisheries.^[127]^[128]^[129] Indeed, an overall decrease in marine plankton productivity is expected over global scales.^[121]^[127]^[130] Long-term increases and decreases in plankton productivity have already occurred over the past two decades ^[131]^[132] along extensive regions of the Humboldt upwelling ecosystem off Chile, and are expected to propagate up the pelagic and benthic food webs. We therefore analyse the bottom-up impact of fluctuations in plankton productivity in combination with fisheries exploitation of these food webs, using the concepts and methods of network ecology.^[95]

Network ecology has advanced understanding of ecosystems by providing a powerful framework to analyse biological communities.^[133] Previous studies used this framework to assess food web robustness against species extinctions, defined as the fraction of initial species that remain present in the ecosystem after a primary extinction.^[134]^[135]^[136]^[137]^[138]^[139]^[140]^[141] These studies showed the importance for food web persistence of highly connected species (independent of trophic position),^[134]^[137]^[142] basal species,^[135] and highly connected species that, at the same time, trophically support other highly connected species.^[138] Most of these studies used a static approach, which stems from network theory and analyzes the impacts of structural changes on food webs represented by nodes (species) and links (interactions) that connect nodes, but ignores interaction strengths and population dynamics of interacting species.^[134] Other studies used a dynamic approach, which considers not only the structure and intensity of interactions in a food web, but also the changes in species biomasses through time and the indirect effects that these changes have on other species.^[135]^[136]^[136]^[143]^[144]^[145]^[95]

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@@ Line 139: / Line 139: @@
 ==Artisan fisheries==
 {{see also|Artisanal fisheries|Coastal fish|Fishing down the food web}}
-[[File:Intertidal food web highlighting nodes and links of artisanal fisheries and plankton.webp|thumb|upright=1.3| {{center|Intertidal food web highlighting nodes and links<br />of (A) artisanal fisheries and (B) plankton{{hsp}}<ref name="Ávila-Thieme2021">{{cite journal |doi = 10.1038/s41598-021-81392-4|title = Alteration of coastal productivity and artisanal fisheries interact to affect a marine food web|year = 2021|last1 = Ávila-Thieme|first1 = M. Isidora|last2 = Corcoran|first2 = Derek|last3 = Pérez-Matus|first3 = Alejandro|last4 = Wieters|first4 = Evie A.|last5 = Navarrete|first5 = Sergio A.|last6 = Marquet|first6 = Pablo A.|last7 = Valdovinos|first7 = Fernanda S.|journal = Scientific Reports|volume = 11|issue = 1|page = 1765|pmid = 33469119|pmc = 7815714|bibcode = 2021NatSR..11.1765A}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref>}}]]
 [[Artisanal fisheries]] use simple fishing gears and small vessels.<ref name="Ávila-Thieme2021" /> Their activities tend to be confined to coastal areas.
@@ Line 147: / Line 145: @@
 Chile has been one of the countries implementing Territorial Use Rights (TURFs)<ref>[https://fisherysolutionscenter.edf.org/catch-share-basics/turfs Sustainable fisheries: Territorial Use Rights for Fishing (TURF) Programs] ''Fishery Solutions Center'', Environmental Defense Fund. Retrieved: 2 November 2021. </ref><ref>Christy, F.T. (1982) "Territorial use rights in marine fisheries: definitions and conditions", ''Food and Agriculture Organisation'', Fishing Technical Paper 227. United Nations, Rome.</ref> over an unprecedented geographic scale to manage the diverse coastal benthic resources using a co-management strategy.<ref name=Gelcich2010>{{cite journal |doi = 10.1073/pnas.1012021107|title = Navigating transformations in governance of Chilean marine coastal resources|year = 2010|last1 = Gelcich|first1 = S.|last2 = Hughes|first2 = T. P.|last3 = Olsson|first3 = P.|last4 = Folke|first4 = C.|last5 = Defeo|first5 = O.|last6 = Fernandez|first6 = M.|last7 = Foale|first7 = S.|last8 = Gunderson|first8 = L. H.|last9 = Rodriguez-Sickert|first9 = C.|last10 = Scheffer|first10 = M.|last11 = Steneck|first11 = R. S.|last12 = Castilla|first12 = J. C.|journal = Proceedings of the National Academy of Sciences|volume = 107|issue = 39|pages = 16794–16799|pmid = 20837530|pmc = 2947917|doi-access = free}}</ref><ref>{{cite journal |doi = 10.5343/bms.2015.1082|title = Fishers' perceptions on the Chilean coastal TURF system after two decades: Problems, benefits, and emerging needs|year = 2017|last1 = Gelcich|first1 = S.|last2 = Cinner|first2 = J.|last3 = Donlan|first3 = CJ|last4 = Tapia-Lewin|first4 = S.|last5 = Godoy|first5 = N.|last6 = Castilla|first6 = JC|journal = Bulletin of Marine Science|volume = 93|pages = 53–67}}</ref> These TURFS are used for artisanal fisheries. Over 60 coastal benthic species are actively harvested by these artisanal fisheries,<ref name=Castilla2007>{{cite book |doi = 10.1002/9780470996072.ch2|chapter = Successes, Lessons, and Projections from Experience in Marine Benthic Invertebrate Artisanal Fisheries in Chile|title = Fisheries Management|year = 2007|last1 = Castilla|first1 = Juan Carlos|last2 = Gelcich|first2 = Stefan|last3 = Defeo|first3 = Omar|pages = 23–42|isbn = 9780470996072}}</ref> with species that are extracted from intertidal and shallow subtidal habitats.<ref>{{cite journal |doi = 10.1890/13-1424.1|title = Network structure beyond food webs: Mapping non‐trophic and trophic interactions on Chilean rocky shores|year = 2015|last1 = Kéfi|first1 = Sonia|last2 = Berlow|first2 = Eric L.|last3 = Wieters|first3 = Evie A.|last4 = Joppa|first4 = Lucas N.|last5 = Wood|first5 = Spencer A.|last6 = Brose|first6 = Ulrich|last7 = Navarrete|first7 = Sergio A.|journal = Ecology|volume = 96|issue = 1|pages = 291–303|pmid = 26236914}}</ref><ref name="Pérez-Matus2017">{{cite journal |doi = 10.3354/meps12057|title = Temperate rocky subtidal reef community reveals human impacts across the entire food web|year = 2017|last1 = Pérez-Matus|first1 = A.|last2 = Ospina-Alvarez|first2 = A.|last3 = Camus|first3 = PA|last4 = Carrasco|first4 = SA|last5 = Fernandez|first5 = M.|last6 = Gelcich|first6 = S.|last7 = Godoy|first7 = N.|last8 = Ojeda|first8 = FP|last9 = Pardo|first9 = LM|last10 = Rozbaczylo|first10 = N.|last11 = Subida|first11 = MD|last12 = Thiel|first12 = M.|last13 = Wieters|first13 = EA|last14 = Navarrete|first14 = SA|journal = Marine Ecology Progress Series|volume = 567|pages = 1–16|bibcode = 2017MEPS..567....1P}}</ref> The Chilean TURFs system brought significant improvements in sustainability of this complex socio-ecological system, helping to rebuild benthic fish stocks,<ref name=Castilla2007 /><ref name=Gelcich2010 /> improving fishers’ perception towards sustainability and increasing compliance9, as well as showing positive ancillary effects on conservation of biodiversity.<ref>{{cite journal |doi = 10.1002/ecs2.1808|title = Exploring the effects of fishing pressure and upwelling intensity over subtidal kelp forest communities in Central Chile|year = 2017|last1 = Pérez-Matus|first1 = Alejandro|last2 = Carrasco|first2 = Sergio A.|last3 = Gelcich|first3 = Stefan|last4 = Fernandez|first4 = Miriam|last5 = Wieters|first5 = Evie A.|journal = Ecosphere|volume = 8|issue = 5|pages = e01808}}</ref><ref>{{cite journal |doi = 10.1111/j.1523-1739.2012.01928.x|title = Territorial User Rights for Fisheries as Ancillary Instruments for Marine Coastal Conservation in Chile|year = 2012|last1 = Gelcich|first1 = Stefan|last2 = Fernández|first2 = Miriam|last3 = Godoy|first3 = Natalio|last4 = Canepa|first4 = Antonio|last5 = Prado|first5 = Luis|last6 = Castilla|first6 = Juan Carlos|journal = Conservation Biology|volume = 26|issue = 6|pages = 1005–1015|pmid = 22971114}}</ref> However, the situation of most artisanal fisheries is still far from sustainable, and many fish stocks and coastal ecosystems show signs of over exploitation and ecosystem degradation, a consequence of the low levels of cooperation and low enforcement of TURF regulations, which leads to high levels of free-riding and illegal fishing.<ref>{{cite journal |doi = 10.1111/cobi.13048|title = Illegal fishing and territorial user rights in Chile|year = 2018|last1 = Oyanedel|first1 = Rodrigo|last2 = Keim|first2 = Andres|last3 = Castilla|first3 = Juan Carlos|last4 = Gelcich|first4 = Stefan|journal = Conservation Biology|volume = 32|issue = 3|pages = 619–627|pmid = 29114934|s2cid = 3703022}}</ref><ref>{{cite journal |doi = 10.1038/s41598-020-69311-5|title = Estimating illegal fishing from enforcement officers|year = 2020|last1 = Donlan|first1 = C. Josh|last2 = Wilcox|first2 = Chris|last3 = Luque|first3 = Gloria M.|last4 = Gelcich|first4 = Stefan|journal = Scientific Reports|volume = 10|issue = 1|page = 12478|pmid = 32719385|pmc = 7385102|bibcode = 2020NatSR..1012478D}}</ref><ref>{{cite journal |doi = 10.1371/journal.pone.0180012|doi-access = free|title = Exploitation rates of two benthic resources across management regimes in central Chile: Evidence of illegal fishing in artisanal fisheries operating in open access areas|year = 2017|last1 = Andreu-Cazenave|first1 = Miguel|last2 = Subida|first2 = Maria Dulce|last3 = Fernandez|first3 = Miriam|journal = PLOS ONE|volume = 12|issue = 6|pages = e0180012|pmid = 28666013|bibcode = 2017PLoSO..1280012A}}</ref> Thus, it is imperative to improve our understanding of the effects of these multi-species fisheries which simultaneously harvest species at all trophic levels, from kelp primary producers to top carnivores.<ref name="Pérez-Matus2017" /><ref>{{cite journal |doi = 10.1016/s0169-5347(99)01602-x|title = Coastal marine communities: Trends and perspectives from human-exclusion experiments|year = 1999|last1 = Castilla|first1 = Juan Carlos|journal = Trends in Ecology & Evolution|volume = 14|issue = 7|pages = 280–283|pmid = 10370266}}</ref><ref name="Ávila-Thieme2021" />
+==Network ecology==
+{{see also|Network ecology}}
+[[File:Intertidal food web highlighting nodes and links of artisanal fisheries and plankton.webp|thumb|upright=1.3| {{center|Intertidal food web highlighting nodes and links<br />of (A) artisanal fisheries and (B) plankton{{hsp}}<ref name="Ávila-Thieme2021">{{cite journal |doi = 10.1038/s41598-021-81392-4|title = Alteration of coastal productivity and artisanal fisheries interact to affect a marine food web|year = 2021|last1 = Ávila-Thieme|first1 = M. Isidora|last2 = Corcoran|first2 = Derek|last3 = Pérez-Matus|first3 = Alejandro|last4 = Wieters|first4 = Evie A.|last5 = Navarrete|first5 = Sergio A.|last6 = Marquet|first6 = Pablo A.|last7 = Valdovinos|first7 = Fernanda S.|journal = Scientific Reports|volume = 11|issue = 1|page = 1765|pmid = 33469119|pmc = 7815714|bibcode = 2021NatSR..11.1765A}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref>}}]]
+To compound things, removal of biomass from the ocean occurs simultaneously with multiple other stressors associated to climate change that compromise the capacity of these socio-ecological systems to respond to perturbations.<ref>{{cite journal |doi = 10.1242/jeb.037473|title = The physiology of climate change: How potentials for acclimatization and genetic adaptation will determine 'winners' and 'losers'|year = 2010|last1 = Somero|first1 = G. N.|journal = Journal of Experimental Biology|volume = 213|issue = 6|pages = 912–920|pmid = 20190116|s2cid = 16838841}}</ref><ref>{{cite journal |doi = 10.1126/science.1189930|title = The Impact of Climate Change on the World's Marine Ecosystems|year = 2010|last1 = Hoegh-Guldberg|first1 = Ove|last2 = Bruno|first2 = John F.|journal = Science|volume = 328|issue = 5985|pages = 1523–1528|pmid = 20558709|bibcode = 2010Sci...328.1523H|s2cid = 206526700}}</ref><ref>{{cite journal |doi = 10.1098/rstb.2012.0232|title = Climate change in size-structured ecosystems|year = 2012|last1 = Brose|first1 = Ulrich|last2 = Dunne|first2 = Jennifer A.|last3 = Montoya|first3 = Jose M.|last4 = Petchey|first4 = Owen L.|last5 = Schneider|first5 = Florian D.|last6 = Jacob|first6 = Ute|journal = Philosophical Transactions of the Royal Society B: Biological Sciences|volume = 367|issue = 1605|pages = 2903–2912|pmid = 23007078|pmc = 3479741}}</ref> Besides sea surface temperature, climate change also affects many other physical–chemical characteristics of marine coastal waters (stratification, acidification, ventilation)<ref>{{cite journal |doi = 10.1146/annurev-marine-041911-111611|title = Climate Change Impacts on Marine Ecosystems|year = 2012|last1 = Doney|first1 = Scott C.|last2 = Ruckelshaus|first2 = Mary|last3 = Emmett Duffy|first3 = J.|last4 = Barry|first4 = James P.|last5 = Chan|first5 = Francis|last6 = English|first6 = Chad A.|last7 = Galindo|first7 = Heather M.|last8 = Grebmeier|first8 = Jacqueline M.|last9 = Hollowed|first9 = Anne B.|last10 = Knowlton|first10 = Nancy|last11 = Polovina|first11 = Jeffrey|last12 = Rabalais|first12 = Nancy N.|last13 = Sydeman|first13 = William J.|last14 = Talley|first14 = Lynne D.|journal = Annual Review of Marine Science|volume = 4|pages = 11–37|pmid = 22457967|bibcode = 2012ARMS....4...11D}}</ref><ref name=Kwiatkowski2019>{{cite journal |doi = 10.1111/gcb.14468|title = Consistent trophic amplification of marine biomass declines under climate change|year = 2019|last1 = Kwiatkowski|first1 = Lester|last2 = Aumont|first2 = Olivier|last3 = Bopp|first3 = Laurent|journal = Global Change Biology|volume = 25|issue = 1|pages = 218–229|pmid = 30295401|bibcode = 2019GCBio..25..218K|s2cid = 52930402}}</ref> as well as the wind regimes that control surface water productivity along the productive coastal upwelling ecosystems.<ref>{{cite journal |doi = 10.1126/science.247.4939.198|title = Global Climate Change and Intensification of Coastal Ocean Upwelling|year = 1990|last1 = Bakun|first1 = Andrew|journal = Science|volume = 247|issue = 4939|pages = 198–201|pmid = 17813287|bibcode = 1990Sci...247..198B|s2cid = 32516158}}</ref><ref>{{cite journal |doi = 10.1111/j.1365-2486.2009.02094.x|title = Greenhouse gas, upwelling-favorable winds, and the future of coastal ocean upwelling ecosystems|year = 2010|last1 = Bakun|first1 = Andrew|last2 = Field|first2 = David B.|last3 = Redondo-Rodriguez|first3 = ANA|last4 = Weeks|first4 = Scarla J.|journal = Global Change Biology|volume = 16|issue = 4|pages = 1213–1228|bibcode = 2010GCBio..16.1213B}}</ref><ref>Thiel, M., Castilla, J.C., Fernández, M. and Navarrete, S., (2007). "The Humboldt current system of northern and central Chile". ''Oceanography and Marine Biology'': An Annual Review, '''45''': 195-344</ref><ref>{{cite journal |doi = 10.3390/rs5115550|doi-access = free|title = Time-Space Variability of Chlorophyll-a and Associated Physical Variables within the Region off Central-Southern Chile|year = 2013|last1 = Morales|first1 = Carmen|last2 = Hormazabal|first2 = Samuel|last3 = Andrade|first3 = Isabel|last4 = Correa-Ramirez|first4 = Marco|journal = Remote Sensing|volume = 5|issue = 11|pages = 5550–5571|bibcode = 2013RemS....5.5550M}}</ref><ref>{{cite journal |doi = 10.1029/2011JG001731|title = Potential changes in larval dispersal and alongshore connectivity on the central Chilean coast due to an altered wind climate|year = 2011|last1 = Aiken|first1 = C. M.|last2 = Navarrete|first2 = S. A.|last3 = Pelegrí|first3 = J. L.|journal = Journal of Geophysical Research|volume = 116|issue = G4|bibcode = 2011JGRG..116.4026A}}</ref> Changes in the productivity of the oceans are reflected in changes of plankton biomass. Plankton contributes approximately half of the global primary production, supports marine food webs, influences the biogeochemical process in the ocean, and strongly affects commercial fisheries.<ref name=Blanchard2012>{{cite journal |doi = 10.1098/rstb.2012.0231|title = Potential consequences of climate change for primary production and fish production in large marine ecosystems|year = 2012|last1 = Blanchard|first1 = Julia L.|last2 = Jennings|first2 = Simon|last3 = Holmes|first3 = Robert|last4 = Harle|first4 = James|last5 = Merino|first5 = Gorka|last6 = Allen|first6 = J. Icarus|last7 = Holt|first7 = Jason|last8 = Dulvy|first8 = Nicholas K.|last9 = Barange|first9 = Manuel|journal = Philosophical Transactions of the Royal Society B: Biological Sciences|volume = 367|issue = 1605|pages = 2979–2989|pmid = 23007086|pmc = 3479740}}</ref><ref>{{cite journal |doi = 10.3389/fmars.2018.00179|doi-access = free|title = Temporal Variability in Net Primary Production in an Upwelling Area off Central Chile (36°S)|year = 2018|last1 = Testa|first1 = Giovanni|last2 = Masotti|first2 = Italo|last3 = Farías|first3 = Laura|journal = Frontiers in Marine Science|volume = 5}}</ref><ref>{{cite journal |doi = 10.3389/fmars.2019.00321|doi-access = free|title = A Global Plankton Diversity Monitoring Program|year = 2019|last1 = Batten|first1 = Sonia D.|last2 = Abu-Alhaija|first2 = Rana|last3 = Chiba|first3 = Sanae|last4 = Edwards|first4 = Martin|last5 = Graham|first5 = George|last6 = Jyothibabu|first6 = R.|last7 = Kitchener|first7 = John A.|last8 = Koubbi|first8 = Philippe|last9 = McQuatters-Gollop|first9 = Abigail|last10 = Muxagata|first10 = Erik|last11 = Ostle|first11 = Clare|last12 = Richardson|first12 = Anthony J.|last13 = Robinson|first13 = Karen V.|last14 = Takahashi|first14 = Kunio T.|last15 = Verheye|first15 = Hans M.|last16 = Wilson|first16 = Willie|journal = Frontiers in Marine Science|volume = 6}}</ref> Indeed, an overall decrease in marine plankton productivity is expected over global scales.<ref name=Kwiatkowski2019 /><ref name=Blanchard2012 /><ref>{{cite journal |doi = 10.1111/gcb.12562|title = Biomass changes and trophic amplification of plankton in a warmer ocean|year = 2014|last1 = Chust|first1 = Guillem|last2 = Allen|first2 = J. Icarus|last3 = Bopp|first3 = Laurent|last4 = Schrum|first4 = Corinna|last5 = Holt|first5 = Jason|last6 = Tsiaras|first6 = Kostas|last7 = Zavatarelli|first7 = Marco|last8 = Chifflet|first8 = Marina|last9 = Cannaby|first9 = Heather|last10 = Dadou|first10 = Isabelle|last11 = Daewel|first11 = Ute|last12 = Wakelin|first12 = Sarah L.|last13 = Machu|first13 = Eric|last14 = Pushpadas|first14 = Dhanya|last15 = Butenschon|first15 = Momme|last16 = Artioli|first16 = Yuri|last17 = Petihakis|first17 = George|last18 = Smith|first18 = Chris|last19 = Garçon|first19 = Veronique|last20 = Goubanova|first20 = Katerina|last21 = Le Vu|first21 = Briac|last22 = Fach|first22 = Bettina A.|last23 = Salihoglu|first23 = Baris|last24 = Clementi|first24 = Emanuela|last25 = Irigoien|first25 = Xabier|journal = Global Change Biology|volume = 20|issue = 7|pages = 2124–2139|pmid = 24604761|bibcode = 2014GCBio..20.2124C|hdl = 11511/31726}}</ref> Long-term increases and decreases in plankton productivity have already occurred over the past two decades{{hsp}}<ref>{{cite journal |doi = 10.1016/j.gloplacha.2020.103259|title = Spatial shifts in productivity of the coastal ocean over the past two decades induced by migration of the Pacific Anticyclone and Bakun's effect in the Humboldt Upwelling Ecosystem|year = 2020|last1 = Weidberg|first1 = Nicolas|last2 = Ospina-Alvarez|first2 = Andres|last3 = Bonicelli|first3 = Jessica|last4 = Barahona|first4 = Mario|last5 = Aiken|first5 = Christopher M.|last6 = Broitman|first6 = Bernardo R.|last7 = Navarrete|first7 = Sergio A.|journal = Global and Planetary Change|volume = 193|page = 103259|arxiv = 2104.11698|bibcode = 2020GPC...19303259W|s2cid = 224872806}}</ref><ref>{{cite journal |doi = 10.1525/elementa.314|title = Insight into anthropogenic forcing on coastal upwelling off south-central Chile|year = 2018|last1 = Aguirre|first1 = Catalina|last2 = García-Loyola|first2 = Sebastián|last3 = Testa|first3 = Giovanni|last4 = Silva|first4 = Diego|last5 = Farías|first5 = Laura|journal = Elementa: Science of the Anthropocene|volume = 6}}</ref> along extensive regions of the Humboldt upwelling ecosystem off Chile, and are expected to propagate up the pelagic and benthic food webs. We therefore analyse the bottom-up impact of fluctuations in plankton productivity in combination with fisheries exploitation of these food webs, using the concepts and methods of network ecology.<ref name="Ávila-Thieme2021" />
+[[Network ecology]] has advanced understanding of ecosystems by providing a powerful framework to analyse biological communities.<ref>{{cite book | last=Pascual | first=Mercedes | title=Ecological networks : linking structure to dynamics in food webs | url=https://books.google.com/books?id=YpQRDAAAQBAJ |publisher=Oxford University Press | publication-place=Oxford New York | year=2006 | isbn=978-0-19-518816-5 | oclc=727944896}}</ref> Previous studies used this framework to assess food web robustness against species extinctions, defined as the fraction of initial species that remain present in the ecosystem after a primary extinction.<ref name=Dunne2002>{{cite journal |doi = 10.1046/j.1461-0248.2002.00354.x|title = Network structure and biodiversity loss in food webs: Robustness increases with connectance|year = 2002|last1 = Dunne|first1 = Jennifer A.|last2 = Williams|first2 = Richard J.|last3 = Martinez|first3 = Neo D.|journal = Ecology Letters|volume = 5|issue = 4|pages = 558–567}}</ref><ref name=Curtsdotter2011>{{cite journal |doi = 10.1016/j.baae.2011.09.008|title = Robustness to secondary extinctions: Comparing trait-based sequential deletions in static and dynamic food webs|year = 2011|last1 = Curtsdotter|first1 = Alva|last2 = Binzer|first2 = Amrei|last3 = Brose|first3 = Ulrich|last4 = De Castro|first4 = Francisco|last5 = Ebenman|first5 = Bo|last6 = Eklöf|first6 = Anna|last7 = Riede|first7 = Jens O.|last8 = Thierry|first8 = Aaron|last9 = Rall|first9 = Björn C.|journal = Basic and Applied Ecology|volume = 12|issue = 7|pages = 571–580|url = http://urn.kb.se/resolve?urn=urn:nbn:se:liu:diva-73611}}</ref><ref name="Ramos-Jiliberto2012">{{cite journal |doi = 10.1111/j.1365-2656.2012.01960.x|title = Topological plasticity increases robustness of mutualistic networks|year = 2012|last1 = Ramos-Jiliberto|first1 = Rodrigo|last2 = Valdovinos|first2 = Fernanda S.|last3 = Moisset De Espanés|first3 = Pablo|last4 = Flores|first4 = José D.|journal = Journal of Animal Ecology|volume = 81|issue = 4|pages = 896–904|pmid = 22313043}}</ref><ref name=Valdovinos2012>{{cite journal |doi = 10.1111/j.1600-0706.2012.20830.x|title = Adaptive foraging allows the maintenance of biodiversity of pollination networks|year = 2013|last1 = Valdovinos|first1 = Fernanda S.|last2 = Moisset De Espanés|first2 = Pablo|last3 = Flores|first3 = José D.|last4 = Ramos-Jiliberto|first4 = Rodrigo|journal = Oikos|volume = 122|issue = 6|pages = 907–917}}</ref><ref name=Allesina2009>{{cite journal |doi = 10.1371/journal.pcbi.1000494|title = Googling Food Webs: Can an Eigenvector Measure Species' Importance for Coextinctions?|year = 2009|last1 = Allesina|first1 = Stefano|last2 = Pascual|first2 = Mercedes|journal = PLOS Computational Biology|volume = 5|issue = 9|pages = e1000494|pmid = 19730676|pmc = 2725316|bibcode = 2009PLSCB...5E0494A}}</ref><ref name=deSantana2013>{{cite journal |doi = 10.3354/meps10073|title = Topological properties of polar food webs|year = 2013|last1 = De Santana|first1 = CN|last2 = Rozenfeld|first2 = AF|last3 = Marquet|first3 = PA|last4 = Duarte|first4 = CM|journal = Marine Ecology Progress Series|volume = 474|pages = 15–26|bibcode = 2013MEPS..474...15D|hdl = 11336/7024}}</ref><ref name="Eklöf2014">{{cite journal |doi = 10.1111/2041-210X.12062|title = Secondary extinctions in food webs: A Bayesian network approach|year = 2013|last1 = Eklöf|first1 = Anna|last2 = Tang|first2 = Si|last3 = Allesina|first3 = Stefano|journal = Methods in Ecology and Evolution|volume = 4|issue = 8|pages = 760–770}}</ref><ref name=Staniczenko2010>{{cite journal |doi = 10.1111/j.1461-0248.2010.01485.x|title = Structural dynamics and robustness of food webs|year = 2010|last1 = Staniczenko|first1 = Phillip P. A.|last2 = Lewis|first2 = Owen T.|last3 = Jones|first3 = Nick S.|last4 = Reed-Tsochas|first4 = Felix|journal = Ecology Letters|volume = 13|issue = 7|pages = 891–899|pmid = 20482578}}</ref> These studies showed the importance for food web persistence of highly connected species (independent of trophic position),<ref name=Dunne2002 /><ref name=Valdovinos2012 /><ref>{{cite journal |doi = 10.1038/nature02515|title = Food-web interactions govern the resistance of communities after non-random extinctions|year = 2004|last1 = Ives|first1 = Anthony R.|last2 = Cardinale|first2 = Bradley J.|journal = Nature|volume = 429|issue = 6988|pages = 174–177|pmid = 15141210|bibcode = 2004Natur.429..174I|s2cid = 4351240}}</ref> basal species,<ref name=Curtsdotter2011 /> and highly connected species that, at the same time, trophically support other highly connected species.<ref name=Allesina2009 /> Most of these studies used a static approach, which stems from network theory and analyzes the impacts of structural changes on food webs represented by nodes (species) and links (interactions) that connect nodes, but ignores interaction strengths and population dynamics of interacting species.<ref name=Dunne2002 /> Other studies used a dynamic approach, which considers not only the structure and intensity of interactions in a food web, but also the changes in species biomasses through time and the indirect effects that these changes have on other species.<ref name=Curtsdotter2011 /><ref name="Ramos-Jiliberto2012" /><ref name="Ramos-Jiliberto2012" /><ref>{{cite journal |doi = 10.1137/17M1153431|title = An Open-System Approach to Complex Biological Networks|year = 2019|last1 = Rebolledo|first1 = Rolando|last2 = Navarrete|first2 = Sergio A.|last3 = Kéfi|first3 = Sonia|last4 = Rojas|first4 = Sergio|last5 = Marquet|first5 = Pablo A.|journal = SIAM Journal on Applied Mathematics|volume = 79|issue = 2|pages = 619–640|s2cid = 146077530}}</ref><ref>{{cite journal |doi = 10.1038/35012234|title = The diversity–stability debate|year = 2000|last1 = McCann|first1 = Kevin Shear|journal = Nature|volume = 405|issue = 6783|pages = 228–233|pmid = 10821283|s2cid = 4319289}}</ref><ref>{{cite journal |doi = 10.1126/sciadv.aaz4891|title = Integrating economic dynamics into ecological networks: The case of fishery sustainability|year = 2020|last1 = Glaum|first1 = Paul|last2 = Cocco|first2 = Valentin|last3 = Valdovinos|first3 = Fernanda S.|journal = Science Advances|volume = 6|issue = 45|pmid = 33148659|pmc = 7673689|bibcode = 2020SciA....6.4891G}}</ref><ref name="Ávila-Thieme2021" />
 ==References==

v t e Ecology: Modelling ecosystems: Trophic components
General	Abiotic component Abiotic stress Behaviour Biogeochemical cycle Biomass Biotic component Biotic stress Carrying capacity Competition Ecosystem Ecosystem ecology Ecosystem model Keystone species List of feeding behaviours Metabolic theory of ecology Productivity Resource Restoration
Producers	Autotrophs Chemosynthesis Chemotrophs Foundation species Kinetotrophs Mixotrophs Myco-heterotrophy Mycotroph Organotrophs Photoheterotrophs Photosynthesis Photosynthetic efficiency Phototrophs Primary nutritional groups Primary production
Consumers	Apex predator Bacterivore Carnivores Chemoorganotroph Foraging Generalist and specialist species Intraguild predation Herbivores Heterotroph Heterotrophic nutrition Insectivore Mesopredators Mesopredator release hypothesis Omnivores Optimal foraging theory Planktivore Predation Prey switching
Decomposers	Chemoorganoheterotrophy Decomposition Detritivores Detritus
Microorganisms	Archaea Bacteriophage Lithoautotroph Lithotrophy Marine Microbial cooperation Microbial ecology Microbial food web Microbial intelligence Microbial loop Microbial mat Microbial metabolism Phage ecology
Food webs	Biomagnification Ecological efficiency Ecological pyramid Energy flow Food chain Trophic level
Example webs	Lakes Rivers Soil Tritrophic interactions in plant defense Marine food webs cold seeps hydrothermal vents intertidal kelp forests North Pacific Gyre San Francisco Estuary tide pool
Processes	Ascendency Bioaccumulation Cascade effect Climax community Competitive exclusion principle Consumer–resource interactions Copiotrophs Dominance Ecological network Ecological succession Energy quality Energy systems language f-ratio Feed conversion ratio Feeding frenzy Mesotrophic soil Nutrient cycle Oligotroph Paradox of the plankton Trophic cascade Trophic mutualism Trophic state index
Defense, counter	Animal coloration Anti-predator adaptations Camouflage Deimatic behaviour Herbivore adaptations to plant defense Mimicry Plant defense against herbivory Predator avoidance in schooling fish

v t e Ecology: Modelling ecosystems: Other components
Population ecology	Abundance Allee effect Consumer-resource model Depensation Ecological yield Effective population size Intraspecific competition Logistic function Malthusian growth model Maximum sustainable yield Overpopulation Overexploitation Population cycle Population dynamics Population modeling Population size Predator–prey (Lotka–Volterra) equations Recruitment Small population size Stability Resilience Resistance Random generalized Lotka–Volterra model
Species	Biodiversity Density-dependent inhibition Ecological effects of biodiversity Ecological extinction Endemic species Flagship species Gradient analysis Indicator species Introduced species Invasive species / Native species Latitudinal gradients in species diversity Minimum viable population Neutral theory Occupancy–abundance relationship Population viability analysis Priority effect Rapoport's rule Relative abundance distribution Relative species abundance Species diversity Species homogeneity Species richness Species distribution Species–area curve Umbrella species
Species interaction	Antibiosis Biological interaction Commensalism Community ecology Ecological facilitation Interspecific competition Mutualism Parasitism Storage effect Symbiosis
Spatial ecology	Biogeography Cross-boundary subsidy Ecocline Ecotone Ecotype Disturbance Edge effects Foster's rule Habitat fragmentation Ideal free distribution Intermediate disturbance hypothesis Insular biogeography Land change modeling Landscape ecology Landscape epidemiology Landscape limnology Metapopulation Patch dynamics r/K selection theory Resource selection function Source–sink dynamics
Niche	Ecological niche Ecological trap Ecosystem engineer Environmental niche modelling Guild Habitat marine habitats Limiting similarity Niche apportionment models Niche construction Niche differentiation Ontogenetic niche shift
Other networks	Assembly rules Bateman's principle Bioluminescence Ecological collapse Ecological debt Ecological deficit Ecological energetics Ecological indicator Ecological threshold Ecosystem diversity Emergence Extinction debt Kleiber's law Liebig's law of the minimum Marginal value theorem Thorson's rule Xerosere
Other	Allometry Alternative stable state Balance of nature Biological data visualization Ecological economics Ecological footprint Ecological forecasting Ecological humanities Ecological stoichiometry Ecopath Ecosystem based fisheries Endolith Evolutionary ecology Functional ecology Industrial ecology Macroecology Microecosystem Natural environment Regime shift Sexecology Systems ecology Urban ecology Theoretical ecology
Outline of ecology