251:, a species of mussel, as a primary example. The ochre starfish is a generalist predator and feeds on chitons, limpets, snails, barnacles, echinoids, and even decapod crustacea. The favourite food for these starfish is the mussel which is a dominant competitor for the space on the rocks. The ochre starfish keeps the population numbers of the mussels in check along with the other preys allowing the other seaweeds, sponges, and anemones, that ochre starfish do not consume, to co-exist. When Paine removed the ochre starfish, the mussels quickly outgrew the other species crowding them out. At the start, the rock pools held 15 rock-clinging species. Three years later there were 8 such species; and ten years later the pools were largely occupied by a single species, mussels. The concept became popular in conservation, and was deployed in a range of contexts and mobilized to engender support for conservation, especially where human activities had damaged ecosystems, such as by removing keystone predators.
470:. Without predation, herbivores began to over-graze many woody browse species, affecting the area's plant populations. In addition, wolves often kept animals from grazing in riparian areas, which protected beavers from having their food sources encroached upon. The removal of wolves had a direct effect on beaver populations, as their habitat became grazing territory. Increased browsing on willows and conifers along Blacktail Creek due to a lack of predation caused channel incision because the beavers helped slow the water down, allowing soil to stay in place. Furthermore, predation keeps hydrological features such as creeks and streams in normal working order. When wolves were reintroduced, the beaver population and the whole riparian ecosystem recovered dramatically within a few years.
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systems. The term has been applied widely in different ecosystems and to predators, prey, and plants (primary producers), inevitably with differing ecological meanings. For instance, removing a predator may allow other animals to increase to the point where they wipe out other species; removing a prey species may cause predator populations to crash, or may allow predators to drive other prey species to extinction; and removing a plant species may result in the loss of animals that depend on it, like
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The community ecologist Bruce Menge states that the keystone concept has been stretched far beyond Paine's original concept. That stretching can be quantified: the researcher Ishana Shukla has listed 230 species identified as keystones in some 157 studies in the 50 years since Paine's paper. Menge's
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areas by cutting down older trees to use for their dams. This allows younger trees to take their place. Beaver dams alter the riparian area they are established in. Depending on topography, soils, and many factors, these dams change the riparian edges of streams and rivers into wetlands, meadows, or
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Although the concept of the keystone species has a value in describing particularly strong inter-species interactions, and for allowing easier communication between ecologists and conservation policy-makers, it has been criticized by L. S. Mills and colleagues for oversimplifying complex ecological
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so heavily that the kelp forests largely disappeared, along with all the species that depended on them. Reintroducing the sea otters has enabled the kelp ecosystem to be restored. For example, in
Southeast Alaska some 400 sea otters were released, and they have bred to form a population approaching
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from damage by sea urchins. When the sea otters of the North
American west coast were hunted commercially for their fur, their numbers fell to such low levels – fewer than 1000 in the north Pacific ocean – that they were unable to control the sea urchin population. The urchins, in turn, grazed the
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species. If prey numbers are low, keystone predators can be even less abundant and still be effective. Yet without the predators, the herbivorous prey would explode in numbers, wipe out the dominant plants, and dramatically alter the character of the ecosystem. The exact scenario changes in each
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and seed dispersers. Beavers too have been called keystone, not for eating other species but for modifying the environment in ways that affected other species. The term has thus been given quite different meanings in different cases. In Mills's view, Paine's work showed that a few species could
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The jaguar is an umbrella species, flagship species, and wilderness quality indicator. It promotes the goals of carnivore recovery, protecting and restoring connectivity through
Madrean woodland and riparian areas, and protecting and restoring riparian areas. ... A reserve system that protects
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Szpak, Paul; Orchard, Trevor J.; Salomon, Anne K.; Gröcke, Darren R. (2013). "Regional ecological variability and impact of the maritime fur trade on nearshore ecosystems in southern Haida Gwaii (British
Columbia, Canada): evidence from stable isotope analysis of rockfish (Sebastes spp.) bone
134:. While the keystone is under the least pressure of any of the stones in an arch, the arch still collapses without it. Similarly, an ecosystem may experience a dramatic shift if a keystone species is removed, even though that species was a small part of the ecosystem by measures of
455:. Introduction or removal of a keystone predator, or changes in its population density, can have drastic cascading effects on the equilibrium of many other populations in the ecosystem. For example, grazers of a grassland may prevent a single dominant species from taking over.
154:. Although the concept is valued as a descriptor for particularly strong inter-species interactions, and has allowed easier communication between ecologists and conservation policy-makers, it has been criticized for oversimplifying complex ecological systems.
314:
has been labeled a keystone species for its unparalleled nest size, colony size, and high rate of brood production. The diversity of its prey and the quantity necessary to sustain its high rate of growth have a direct impact on other species around it.
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sea star that Paine had studied was a powerful keystone species in places exposed to strong wave action, but was far less important in sheltered places. Paine had indeed stated that in Alaska, without the relevant mussel species as prey, the predatory
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and helping to determine the types and numbers of various other species in the community. Without keystone species, the ecosystem would be dramatically different or cease to exist altogether. Some keystone species, such as the
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of numerous plant species. Therefore, the loss of this one species of tree would probably cause the honeyeater population to collapse, with profound implications for the entire ecosystem. Another example is
727:, shape their environment. The elephants destroy trees, making room for the grass species and creating habitat for various small animal species. Without these animals, much of the savanna would turn into
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and reversing soil compaction that can be a result of cattle grazing. Prairie dogs also trim the vegetation around their colonies, perhaps to remove any cover for predators. Grazing species such as
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The keystone concept is defined by its ecological effects, and these in turn make it important for conservation. In this it overlaps with several other species conservation concepts such as
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Keystone mutualists are organisms that participate in mutually beneficial interaction, the loss of which would have a profound impact upon the ecosystem as a whole. For example, in the
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that have no other natural predators. If the sea star is removed from the ecosystem, the mussel population explodes uncontrollably, driving out most other species. The recent onset of
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is a well known ecosystem engineer and keystone species. It transforms its territory from a stream to a pond or swamp. Beavers affect the environment first altering the edges of
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example, but the central idea remains that through a chain of interactions, a non-abundant species has an outsized impact on ecosystem functions. For example, the herbivorous
1958:
Coverdale, Tyler C.; Kartzinel, Tyler R.; Grabowski, Kathryn L.; Shriver, Robert K.; Hassan, Abdikadir A.; Goheen, Jacob R.; Palmer, Todd M.; Pringle, Robert M. (2016).
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sometimes have extremely strong interactions within a particular ecosystem, but that does not automatically imply that other ecosystems have a similar structure.
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are the only reef fish that consistently scrape and clean the coral on the reef. Without these animals, the Great
Barrier Reef would be under severe strain.
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1606:
Estes, J. A.; Tinker, M. T.; Williams, T. M.; Doak, D. F. (1998-10-16). "Killer whale predation on sea otters linking oceanic and nearshore ecosystems".
1304:
Estes, James E.; Smith, Norman S.; Palmisano, John F. (1978). "Sea otter predation and community organization in the
Western Aleutian Islands, Alaska".
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of communities by preventing a single species from becoming dominant. They can have a profound influence on the balance of organisms in a particular
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was "just another sea star". In other words, the extent to which a species could be described as a keystone depended on the ecological context.
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2155:
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Wright, J. P.; Jones, C. G.; Flecker, A. S. (2002). "An ecosystem engineer, the beaver, increases species richness at the landscape scale".
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relative to its abundance. It has been defined operationally by Davic in 2003 as "a strongly interacting species whose top-down effect on
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Hale, Sarah L.; Koprowski, John L. (February 2018). "Ecosystem-level effects of keystone species reintroduction: a literature review".
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riverine forests. These dams have been shown to be beneficial to a myriad of species including amphibians, salmon, and song birds.
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1088:
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1960:"Elephants in the understory: opposing direct and indirect effects of consumption and ecosystem engineering by megaherbivores"
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Landscape
Planning for Biodiversity Conservation in Agricultural Regions: A Case Study from the Wheatbelt of Western Australia
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611:, which spreads the seeds of many different trees. Some seeds will not grow unless they have been through a cassowary.
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Wilmers, Christopher C.; Crabtree, Robert L.; Smith, Douglas W.; Murphy, Kerry M.; Getz, Wayne M. (November 2003).
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1442:"Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park"
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2864:
2731:
2443:
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Barua, Maan (2011). "Mobilizing metaphors: the popular use of keystone, flagship and umbrella species concepts".
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jaguars is an umbrella for many other species. ... the jaguar a keystone in subtropical and tropical
America ...
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1150:"Linking Keystone Species and Functional Groups: A New Operational Definition of the Keystone Species Concept"
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around the United States has indirectly caused mussel populations to dominate in many intertidal habitats.
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227:. He removed the starfish from an area, and documented the effects on the ecosystem. In his 1966 paper,
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211:. Paine developed the concept to explain his observations and experiments on the relationships between
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Creed, R. P. Jr. (2000). "Is there a new keystone species in North
American lakes and rivers?".
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1064:"Some Animals Are More Equal than Others: Keystone Species and Trophic Cascades – HHMI (2016)"
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Conservation by proxy: indicator, umbrella, keystone, flagship, and other surrogate species
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is thought to have keystone effects on aquatic plant diversity by foraging on nuisance
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175:
20:
1760:
Walker, Brian (1995). "Conserving
Biological Diversity through Ecosystem Resilience".
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The role that a keystone species plays in its ecosystem is analogous to the role of a
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676:, and can also serve to change the composition of the soil in a region by increasing
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Large Mammal Restoration: Ecological And Sociological Challenges In The 21St Century
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1979:
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1827:"Associated Species : Prairie Dogs are a Keystone Species of the Great Plains"
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Paine, R. T. (1995). "A Conversation on Refining the Concept of Keystone Species".
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that are utilized by a wide variety of species. Australian studies have found that
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1905:"Elephants as agents of habitat creation for small vertebrates at the patch scale"
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2046:"How the overlooked peccary engineers the Amazon, an interview with Harald Beck"
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538:, acts as a keystone predator by its widely varied diet, helping to balance the
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2013:
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relative to its abundance. The concept was introduced in 1969 by the zoologist
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1991:
1936:
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1549:
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863:
Paine, R. T. (1969). "A Note on Trophic Complexity and Community Stability".
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is an ecosystem engineer. Prairie dog burrows provide the nesting areas for
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1999:
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have shown a proclivity for grazing on the same land used by prairie dogs.
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jungle ecosystem with its consumption of 87 different species of prey. The
520:
448:
268:
98:. Keystone species play a critical role in maintaining the structure of an
28:
1635:
1491:"Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems?"
2949:
2496:
2458:
2433:
2423:
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2335:
2315:
2017:
1904:
1390:"My Turn: Southeast Alaska's sea otters: The restoration of an ecosystem"
665:
653:
623:
599:
408:
287:
188:
239:. In his 1969 paper, Paine proposed the keystone species concept, using
207:
The concept of the keystone species was introduced in 1969 by zoologist
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1983:
755:
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724:
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539:
534:, whose numbers in Central and South America have been classified as
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107:
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283:
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103:
72:
1957:
1563:
Paine, R. T. (1966). "Food web complexity and species diversity".
973:
Paine, R. T. (1966). "Food Web Complexity and Species Diversity".
664:. Prairie dog tunnel systems also help channel rainwater into the
167:
51:
3612:
3419:
3289:
3284:
2911:
2859:
2519:
2164:
1524:"perspective: The keystone species concept: a critical appraisal"
720:
695:
673:
276:
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likelihood, which in turn promotes tree growth. The documentary
364:
1522:
Cottee-Jones, Henry Eden W; Whittaker, Robert J. (2012-09-28).
740:
708:
591:
531:
512:
493:
334:
is a charismatic big cat which meets all of these definitions:
331:
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245:, a species of starfish generally known as ochre starfish, and
224:
68:
56:
33:
1736:
291:
64:
1508:
10.1641/0006-3568(2004)054[0755:WATEOF]2.0.CO;2
543:
524:
417:
131:
116:
112:
1439:
936:"The Keystone-Species Concept in Ecology and Conservation"
559:, is periodically the sole source of nectar for important
1259:
1089:"Ecologists Struggle to Get a Grip on 'Keystone Species'"
759:
473:
1233:
Maehr, David; Noss, Reed F.; Larkin, Jeffery L. (2001).
424:
2085:"Single keystone species may be the key to reef health"
308:
in North American waters. Similarly, the wasp species
1664:
Wild Cats, Status Survey and Conservation Action Plan
1656:
1521:
1414:
703:
which has a transformative effect on the environment
397:, making it a keystone species for the kelp forests.
2022:
The sixth extinction: biodiversity and its survival
1802:. Nebraska Game and Park Commission. Archived from
1667:. IUCN/SSC Cat Specialist Group. pp. 118–122.
263:that has a disproportionately large effect on its
90:that has a disproportionately large effect on its
1824:
1681:
219:(between the high and low tide lines), including
3699:
1194:
1014:
1012:
934:Mills, L. S.; Soule, M. E.; Doak, D. F. (1993).
374:can damage kelp forests by chewing through kelp
1489:Ripple, William J.; Beschta, Robert L. (2004).
1417:Environmental Science: Earth as a living planet
1232:
2060:"Where Peccaries Wallow, Other Animals Follow"
933:
354:
16:Species with a large effect on its environment
2896:
2149:
1818:
1687:
1488:
1009:
433:Riparian willow recovery at Blacktail Creek,
259:A keystone species was defined by Paine as a
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1724:
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1228:
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1147:
1112:
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1080:
972:
862:
2012:
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1346:
1263:Archaeological and Anthropological Sciences
1108:
1106:
929:
927:
858:
856:
3117:Latitudinal gradients in species diversity
2903:
2889:
2156:
2142:
2006:
1845:
1482:
909:. University of Washington. Archived from
2020:(1999) . "11 The modern elephant story".
1599:
1539:
1506:
1465:
1364:
1297:
1253:
1223:
1188:
1173:
1077:
684:, which is another keystone species, the
229:Food Web Complexity and Species Diversity
63:creator, responsible for the creation of
3015:Predator–prey (Lotka–Volterra) equations
2654:Tritrophic interactions in plant defense
1340:
1141:
1103:
966:
924:
853:
723:, the larger herbivores, especially the
694:
618:
548:
428:
50:
27:
3047:Random generalized Lotka–Volterra model
1902:
1829:. Prairie Dog Coalition. Archived from
1730:
1657:Nowell, K.; Jackson, P., eds. (1996). "
231:, Paine had described such a system in
3700:
2855:Herbivore adaptations to plant defense
1759:
1735:. Biodiversity Technical Paper No. 2.
590:(acorn banksia) is the sole source of
584:, there is a period of each year when
474:Sea stars and other non-apex predators
279:dominance within a functional group."
75:irrigating large forests and creating
2884:
2137:
1387:
1087:Ogwen, Lesley Evans (24 April 2024).
1086:
1018:
778:
425:The wolf, Yellowstone's apex predator
2870:Predator avoidance in schooling fish
1788:
1419:. John Wiley & Sons. p. 2.
1062:HHMI, BioInteractive (29 May 2017).
1061:
796:
478:As described by Paine in 1966, some
447:Keystone predators may increase the
3320:Intermediate disturbance hypothesis
2068:. 27 September 2014. Archived from
1782:10.1046/j.1523-1739.1995.09040747.x
1501:(8). Oxford University Press: 755.
1135:10.1046/j.1523-1739.1995.09040962.x
784:own work has shown that the purple
13:
3073:Ecological effects of biodiversity
2105:
2098:How Wildebeest Saved the Serengeti
2083:Gruber, Karl (26 September 2014).
1739:Division of Wildlife and Ecology.
644:A term used alongside keystone is
598:, which play an important role in
290:species from eliminating dominant
14:
3734:
2409:Generalist and specialist species
1217:10.1034/j.1600-0706.2000.910222.x
437:, after reintroduction of wolves.
142:. It became a popular concept in
3132:Occupancy–abundance relationship
2119:. Washington, DC: Island Press.
1467:10.1046/j.1365-2656.2003.00766.x
1388:Brown, Tina M. (14 March 2013).
382:
363:
282:A classic keystone species is a
187:
166:
3152:Relative abundance distribution
2865:Plant defense against herbivory
2732:Competitive exclusion principle
2444:Mesopredator release hypothesis
2091:
2076:
2052:
2038:
1951:
1896:
1650:
1556:
1515:
1433:
1415:Botkin, D.; Keller, E. (2003).
1408:
1392:. Juneau Empire. Archived from
1381:
2737:Consumer–resource interactions
1055:
899:
773:
254:
1:
3583:Biological data visualization
3410:Environmental niche modelling
3137:Population viability analysis
2024:. Phoenix. pp. 216–217.
1021:Biodiversity and Conservation
907:"Keystone Species Hypothesis"
847:
758:, the presence of sufficient
571:
546:is another keystone species.
464:Greater Yellowstone Ecosystem
3068:Density-dependent inhibition
1628:10.1126/science.282.5388.473
1239:. Island Press. p. 73.
762:in these grasslands reduces
614:
507:These creatures need not be
466:had profound impacts on the
393:is an important predator of
349:
7:
3537:Liebig's law of the minimum
3372:Resource selection function
2263:Metabolic theory of ecology
2065:National Geographic Society
1903:Pringle, Robert M. (2008).
1731:Lambeck, Robert J. (1999).
810:
355:Sea otters and kelp forests
286:that prevents a particular
10:
3739:
3437:Niche apportionment models
3157:Relative species abundance
2361:Primary nutritional groups
2258:List of feeding behaviours
1825:Rosmarino, Nicole (2007).
1349:"Understanding Sea Otters"
770:documents this in detail.
633:
523:. Sea otters are prey for
440:
157:
59:: a keystone species, and
18:
3686:
3618:Ecosystem based fisheries
3560:
3460:
3385:
3258:
3230:Interspecific competition
3195:
3122:Minimum viable population
3055:
2980:Maximum sustainable yield
2965:Intraspecific competition
2960:Effective population size
2923:
2840:Anti-predator adaptations
2825:
2704:
2631:
2588:
2510:
2477:
2374:
2351:Photosynthetic efficiency
2286:
2180:
1874:10.1007/s00442-002-0929-1
1528:Frontiers of Biogeography
1446:Journal of Animal Ecology
1283:10.1007/s12520-013-0122-y
1041:10.1007/s10531-011-0035-y
817:Cultural keystone species
511:. Sea stars are prey for
435:Yellowstone National Park
275:is large relative to its
3608:Ecological stoichiometry
3573:Alternative stable state
2087:. Australian Geographic.
1166:10.5751/ES-00502-0701r11
502:sea star wasting disease
19:Not to be confused with
3452:Ontogenetic niche shift
3315:Ideal free distribution
3225:Ecological facilitation
2975:Malthusian growth model
2945:Consumer-resource model
2802:Paradox of the plankton
2767:Energy systems language
2487:Chemoorganoheterotrophy
2454:Optimal foraging theory
2429:Heterotrophic nutrition
975:The American Naturalist
865:The American Naturalist
458:The elimination of the
344:David Maehr et al, 2001
102:, affecting many other
3598:Ecological forecasting
3542:Marginal value theorem
3340:Landscape epidemiology
3275:Cross-boundary subsidy
3210:Biological interaction
2560:Microbial intelligence
2248:Green world hypothesis
1800:Wildlife Species Guide
704:
631:
568:
438:
370:Sea urchins like this
347:
301:Euhrychiopsis lecontei
79:
48:
3603:Ecological humanities
3502:Ecological energetics
3447:Niche differentiation
3310:Habitat fragmentation
3078:Ecological extinction
3025:Small population size
2777:Feed conversion ratio
2757:Ecological succession
2689:San Francisco Estuary
2603:Ecological efficiency
2545:Microbial cooperation
1148:Davic, R. D. (2003).
739:produce and maintain
698:
634:Further information:
622:
552:
441:Further information:
432:
336:
306:Eurasian watermilfoil
248:Mytilus californianus
200:), the seastar's prey
197:Mytilus californianus
54:
31:
3713:Conservation biology
3628:Evolutionary ecology
3593:Ecological footprint
3588:Ecological economics
3512:Ecological threshold
3507:Ecological indicator
3377:Source–sink dynamics
3330:Land change modeling
3325:Insular biogeography
3177:Species distribution
2916:Modelling ecosystems
2575:Microbial metabolism
2414:Intraguild predation
2203:Biogeochemical cycle
2169:Modelling ecosystems
2048:. 20 September 2010.
1762:Conservation Biology
1347:Cohn, J. P. (1998).
1154:Conservation Ecology
1115:Conservation Biology
213:marine invertebrates
194:California mussels (
144:conservation biology
100:ecological community
3723:Biology terminology
3678:Theoretical ecology
3653:Natural environment
3517:Ecosystem diversity
3487:Ecological collapse
3477:Bateman's principle
3432:Limiting similarity
3345:Landscape limnology
3167:Species homogeneity
3005:Population modeling
3000:Population dynamics
2817:Trophic state index
1976:2016Ecol...97.3219C
1921:2008Ecol...89...26P
1866:2002Oecol.132...96W
1833:on 10 November 2013
1774:1995ConBi...9..747W
1702:2018ResEc..26..439H
1690:Restoration Ecology
1620:1998Sci...282..473E
1565:American Naturalist
1458:2003JAnEc..72..909W
1318:1978Ecol...59..822E
1275:2013ArAnS...5..159S
1209:2000Oikos..91..405C
1127:1995ConBi...9..962P
1033:2011BiCon..20.1427B
768:The Serengeti Rules
701:animal construction
330:. For example, the
92:natural environment
3689:Outline of ecology
3638:Industrial ecology
3633:Functional ecology
3497:Ecological deficit
3442:Niche construction
3405:Ecosystem engineer
3182:Species–area curve
3103:Introduced species
2918:: Other components
2850:Deimatic behaviour
2752:Ecological network
2684:North Pacific Gyre
2669:hydrothermal vents
2608:Ecological pyramid
2555:Microbial food web
2366:Primary production
2311:Foundation species
2072:on April 18, 2020.
1541:10.21425/F54312533
837:Introduced species
827:Foundation species
779:Depends on context
749:Great Barrier Reef
733:Amazon river basin
705:
646:ecosystem engineer
640:Niche construction
636:Ecosystem engineer
632:
569:
485:Pisaster ochraceus
439:
242:Pisaster ochraceus
176:Pisaster ochraceus
80:
49:
21:foundation species
3708:Community ecology
3695:
3694:
3578:Balance of nature
3335:Landscape ecology
3220:Community ecology
3162:Species diversity
3098:Indicator species
3093:Gradient analysis
2970:Logistic function
2878:
2877:
2835:Animal coloration
2812:Trophic mutualism
2550:Microbial ecology
2341:Photoheterotrophs
2326:Myco-heterotrophy
2238:Ecosystem ecology
2223:Carrying capacity
2188:Abiotic component
1970:(11): 3219–3230.
1929:10.1890/07-0776.1
1806:on 19 August 2009
1746:978-0-642-21423-2
1710:10.1111/rec.12684
1674:978-2-8317-0045-8
1614:(5388): 473–476.
1426:978-0-471-38914-9
1246:978-1-55963-817-3
1091:. Quanta Magazine
822:Ecosystem service
797:Multiple meanings
626:town. Drawing by
587:Banksia prionotes
582:Western Australia
556:Banksia prionotes
372:purple sea urchin
324:indicator species
269:species diversity
3730:
3395:Ecological niche
3367:selection theory
3187:Umbrella species
3172:Species richness
3108:Invasive species
3088:Flagship species
2995:Population cycle
2990:Overexploitation
2955:Ecological yield
2905:
2898:
2891:
2882:
2881:
2787:Mesotrophic soil
2727:Climax community
2659:Marine food webs
2598:Biomagnification
2399:Chemoorganotroph
2253:Keystone species
2213:Biotic component
2158:
2151:
2144:
2135:
2134:
2130:
2100:
2095:
2089:
2088:
2080:
2074:
2073:
2056:
2050:
2049:
2042:
2036:
2035:
2010:
2004:
2003:
1984:10.1002/ecy.1557
1955:
1949:
1948:
1900:
1894:
1893:
1849:
1843:
1842:
1840:
1838:
1822:
1816:
1815:
1813:
1811:
1792:
1786:
1785:
1757:
1751:
1750:
1728:
1722:
1721:
1685:
1679:
1678:
1654:
1648:
1647:
1603:
1597:
1596:
1560:
1554:
1553:
1543:
1519:
1513:
1512:
1510:
1486:
1480:
1479:
1469:
1437:
1431:
1430:
1412:
1406:
1405:
1403:
1401:
1396:on 23 April 2018
1385:
1379:
1378:
1368:
1344:
1338:
1337:
1301:
1295:
1294:
1257:
1251:
1250:
1230:
1221:
1220:
1192:
1186:
1185:
1183:
1182:
1177:
1145:
1139:
1138:
1110:
1101:
1100:
1098:
1096:
1084:
1075:
1074:
1072:
1070:
1059:
1053:
1052:
1027:(7): 1427–1440.
1016:
1007:
1006:
970:
964:
963:
931:
922:
921:
919:
918:
903:
897:
896:
860:
842:Sentinel species
658:mountain plovers
386:
367:
345:
328:umbrella species
320:flagship species
191:
173:Ochre seastars (
170:
152:umbrella species
84:keystone species
42:umbrella species
3738:
3737:
3733:
3732:
3731:
3729:
3728:
3727:
3698:
3697:
3696:
3691:
3682:
3668:Systems ecology
3556:
3527:Extinction debt
3492:Ecological debt
3482:Bioluminescence
3463:
3456:
3425:marine habitats
3400:Ecological trap
3381:
3261:
3254:
3197:
3191:
3147:Rapoport's rule
3142:Priority effect
3083:Endemic species
3051:
3010:Population size
2926:
2919:
2909:
2879:
2874:
2827:
2821:
2807:Trophic cascade
2717:Bioaccumulation
2700:
2627:
2584:
2506:
2473:
2370:
2282:
2243:Ecosystem model
2176:
2162:
2127:
2111:
2108:
2106:Further reading
2103:
2096:
2092:
2081:
2077:
2058:
2057:
2053:
2044:
2043:
2039:
2032:
2014:Leakey, Richard
2011:
2007:
1956:
1952:
1901:
1897:
1850:
1846:
1836:
1834:
1823:
1819:
1809:
1807:
1794:
1793:
1789:
1758:
1754:
1747:
1729:
1725:
1686:
1682:
1675:
1655:
1651:
1604:
1600:
1561:
1557:
1520:
1516:
1487:
1483:
1438:
1434:
1427:
1413:
1409:
1399:
1397:
1386:
1382:
1366:10.2307/1313259
1345:
1341:
1326:10.2307/1938786
1302:
1298:
1258:
1254:
1247:
1231:
1224:
1193:
1189:
1180:
1178:
1146:
1142:
1111:
1104:
1094:
1092:
1085:
1078:
1068:
1066:
1060:
1056:
1017:
1010:
971:
967:
952:10.2307/1312122
932:
925:
916:
914:
905:
904:
900:
861:
854:
850:
813:
799:
781:
776:
719:In the African
699:Beaver dam, an
642:
617:
574:
553:Acorn banksia,
536:near threatened
476:
468:trophic pyramid
445:
427:
402:
401:
400:
399:
398:
387:
379:
378:
368:
357:
352:
346:
343:
257:
217:intertidal zone
209:Robert T. Paine
205:
204:
203:
202:
201:
192:
184:
183:
171:
160:
96:Robert T. Paine
24:
17:
12:
11:
5:
3736:
3726:
3725:
3720:
3715:
3710:
3693:
3692:
3687:
3684:
3683:
3681:
3680:
3675:
3670:
3665:
3660:
3655:
3650:
3648:Microecosystem
3645:
3640:
3635:
3630:
3625:
3620:
3615:
3610:
3605:
3600:
3595:
3590:
3585:
3580:
3575:
3570:
3564:
3562:
3558:
3557:
3555:
3554:
3549:
3547:Thorson's rule
3544:
3539:
3534:
3529:
3524:
3519:
3514:
3509:
3504:
3499:
3494:
3489:
3484:
3479:
3474:
3472:Assembly rules
3468:
3466:
3458:
3457:
3455:
3454:
3449:
3444:
3439:
3434:
3429:
3428:
3427:
3417:
3412:
3407:
3402:
3397:
3391:
3389:
3383:
3382:
3380:
3379:
3374:
3369:
3357:
3355:Patch dynamics
3352:
3350:Metapopulation
3347:
3342:
3337:
3332:
3327:
3322:
3317:
3312:
3307:
3302:
3297:
3292:
3287:
3282:
3277:
3272:
3266:
3264:
3256:
3255:
3253:
3252:
3247:
3245:Storage effect
3242:
3237:
3232:
3227:
3222:
3217:
3212:
3207:
3201:
3199:
3193:
3192:
3190:
3189:
3184:
3179:
3174:
3169:
3164:
3159:
3154:
3149:
3144:
3139:
3134:
3129:
3127:Neutral theory
3124:
3119:
3114:
3112:Native species
3105:
3100:
3095:
3090:
3085:
3080:
3075:
3070:
3065:
3059:
3057:
3053:
3052:
3050:
3049:
3044:
3043:
3042:
3037:
3027:
3022:
3017:
3012:
3007:
3002:
2997:
2992:
2987:
2985:Overpopulation
2982:
2977:
2972:
2967:
2962:
2957:
2952:
2947:
2942:
2937:
2931:
2929:
2921:
2920:
2908:
2907:
2900:
2893:
2885:
2876:
2875:
2873:
2872:
2867:
2862:
2857:
2852:
2847:
2842:
2837:
2831:
2829:
2823:
2822:
2820:
2819:
2814:
2809:
2804:
2799:
2794:
2792:Nutrient cycle
2789:
2784:
2782:Feeding frenzy
2779:
2774:
2769:
2764:
2762:Energy quality
2759:
2754:
2749:
2744:
2739:
2734:
2729:
2724:
2722:Cascade effect
2719:
2714:
2708:
2706:
2702:
2701:
2699:
2698:
2697:
2696:
2691:
2686:
2681:
2676:
2671:
2666:
2656:
2651:
2646:
2641:
2635:
2633:
2629:
2628:
2626:
2625:
2620:
2615:
2610:
2605:
2600:
2594:
2592:
2586:
2585:
2583:
2582:
2577:
2572:
2567:
2565:Microbial loop
2562:
2557:
2552:
2547:
2542:
2537:
2532:
2530:Lithoautotroph
2527:
2522:
2516:
2514:
2512:Microorganisms
2508:
2507:
2505:
2504:
2499:
2494:
2489:
2483:
2481:
2475:
2474:
2472:
2471:
2469:Prey switching
2466:
2461:
2456:
2451:
2446:
2441:
2436:
2431:
2426:
2421:
2416:
2411:
2406:
2401:
2396:
2391:
2386:
2380:
2378:
2372:
2371:
2369:
2368:
2363:
2358:
2353:
2348:
2346:Photosynthesis
2343:
2338:
2333:
2328:
2323:
2318:
2313:
2308:
2303:
2301:Chemosynthesis
2298:
2292:
2290:
2284:
2283:
2281:
2280:
2275:
2270:
2265:
2260:
2255:
2250:
2245:
2240:
2235:
2230:
2225:
2220:
2215:
2210:
2205:
2200:
2195:
2193:Abiotic stress
2190:
2184:
2182:
2178:
2177:
2161:
2160:
2153:
2146:
2138:
2132:
2131:
2125:
2107:
2104:
2102:
2101:
2090:
2075:
2051:
2037:
2030:
2005:
1950:
1895:
1844:
1817:
1796:"Prairie Dogs"
1787:
1768:(4): 747–752.
1752:
1745:
1723:
1696:(3): 439–445.
1680:
1673:
1649:
1598:
1577:10.1086/282400
1571:(910): 65–75.
1555:
1514:
1481:
1452:(6): 909–916.
1432:
1425:
1407:
1380:
1359:(3): 151–155.
1339:
1312:(4): 822–833.
1296:
1269:(2): 159–182.
1252:
1245:
1222:
1187:
1140:
1121:(4): 962–964.
1102:
1076:
1054:
1008:
987:10.1086/282400
981:(910): 65–75.
965:
946:(4): 219–224.
923:
898:
877:10.1086/282586
871:(929): 91–93.
851:
849:
846:
845:
844:
839:
834:
829:
824:
819:
812:
809:
798:
795:
780:
777:
775:
772:
662:burrowing owls
616:
613:
607:, such as the
578:Avon Wheatbelt
573:
570:
509:apex predators
488:) may prey on
475:
472:
426:
423:
388:
381:
380:
369:
362:
361:
360:
359:
358:
356:
353:
351:
348:
341:
311:Agelaia vicina
256:
253:
193:
186:
185:
179:), a keystone
172:
165:
164:
163:
162:
161:
159:
156:
121:apex predators
36:: a keystone,
15:
9:
6:
4:
3:
2:
3735:
3724:
3721:
3719:
3716:
3714:
3711:
3709:
3706:
3705:
3703:
3690:
3685:
3679:
3676:
3674:
3673:Urban ecology
3671:
3669:
3666:
3664:
3661:
3659:
3656:
3654:
3651:
3649:
3646:
3644:
3641:
3639:
3636:
3634:
3631:
3629:
3626:
3624:
3621:
3619:
3616:
3614:
3611:
3609:
3606:
3604:
3601:
3599:
3596:
3594:
3591:
3589:
3586:
3584:
3581:
3579:
3576:
3574:
3571:
3569:
3566:
3565:
3563:
3559:
3553:
3550:
3548:
3545:
3543:
3540:
3538:
3535:
3533:
3532:Kleiber's law
3530:
3528:
3525:
3523:
3520:
3518:
3515:
3513:
3510:
3508:
3505:
3503:
3500:
3498:
3495:
3493:
3490:
3488:
3485:
3483:
3480:
3478:
3475:
3473:
3470:
3469:
3467:
3465:
3459:
3453:
3450:
3448:
3445:
3443:
3440:
3438:
3435:
3433:
3430:
3426:
3423:
3422:
3421:
3418:
3416:
3413:
3411:
3408:
3406:
3403:
3401:
3398:
3396:
3393:
3392:
3390:
3388:
3384:
3378:
3375:
3373:
3370:
3368:
3366:
3362:
3358:
3356:
3353:
3351:
3348:
3346:
3343:
3341:
3338:
3336:
3333:
3331:
3328:
3326:
3323:
3321:
3318:
3316:
3313:
3311:
3308:
3306:
3305:Foster's rule
3303:
3301:
3298:
3296:
3293:
3291:
3288:
3286:
3283:
3281:
3278:
3276:
3273:
3271:
3268:
3267:
3265:
3263:
3257:
3251:
3248:
3246:
3243:
3241:
3238:
3236:
3233:
3231:
3228:
3226:
3223:
3221:
3218:
3216:
3213:
3211:
3208:
3206:
3203:
3202:
3200:
3194:
3188:
3185:
3183:
3180:
3178:
3175:
3173:
3170:
3168:
3165:
3163:
3160:
3158:
3155:
3153:
3150:
3148:
3145:
3143:
3140:
3138:
3135:
3133:
3130:
3128:
3125:
3123:
3120:
3118:
3115:
3113:
3109:
3106:
3104:
3101:
3099:
3096:
3094:
3091:
3089:
3086:
3084:
3081:
3079:
3076:
3074:
3071:
3069:
3066:
3064:
3061:
3060:
3058:
3054:
3048:
3045:
3041:
3038:
3036:
3033:
3032:
3031:
3028:
3026:
3023:
3021:
3018:
3016:
3013:
3011:
3008:
3006:
3003:
3001:
2998:
2996:
2993:
2991:
2988:
2986:
2983:
2981:
2978:
2976:
2973:
2971:
2968:
2966:
2963:
2961:
2958:
2956:
2953:
2951:
2948:
2946:
2943:
2941:
2938:
2936:
2933:
2932:
2930:
2928:
2922:
2917:
2913:
2906:
2901:
2899:
2894:
2892:
2887:
2886:
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2623:Trophic level
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2570:Microbial mat
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2492:Decomposition
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1860:(1): 96–101.
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443:Apex predator
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46:apex predator
43:
39:
35:
30:
26:
22:
3658:Regime shift
3643:Macroecology
3364:
3360:
3300:Edge effects
3270:Biogeography
3215:Commensalism
3063:Biodiversity
2940:Allee effect
2679:kelp forests
2632:Example webs
2497:Detritivores
2336:Organotrophs
2316:Kinetotrophs
2268:Productivity
2252:
2116:
2093:
2078:
2070:the original
2063:
2054:
2040:
2021:
2018:Lewin, Roger
2008:
1967:
1963:
1953:
1915:(1): 26–33.
1912:
1908:
1898:
1857:
1853:
1847:
1835:. Retrieved
1831:the original
1820:
1808:. Retrieved
1804:the original
1799:
1790:
1765:
1761:
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1398:. Retrieved
1394:the original
1383:
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1179:. Retrieved
1157:
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1093:. Retrieved
1067:. Retrieved
1057:
1024:
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915:. Retrieved
911:the original
901:
868:
864:
800:
790:
785:
782:
753:
718:
706:
682:plains bison
643:
628:Josiah Gregg
585:
575:
554:
529:
521:sea anemones
506:
496:, and other
483:
477:
457:
449:biodiversity
446:
409:kelp forests
403:
337:
317:
309:
299:
281:
258:
246:
240:
228:
206:
195:
174:
146:, alongside
140:productivity
125:
83:
81:
25:
3295:Disturbance
3198:interaction
3020:Recruitment
2950:Depensation
2742:Copiotrophs
2613:Energy flow
2535:Lithotrophy
2479:Decomposers
2459:Planktivore
2434:Insectivore
2424:Heterotroph
2389:Bacterivore
2356:Phototrophs
2306:Chemotrophs
2278:Restoration
2228:Competition
1837:10 November
1810:10 November
1261:collagen".
804:pollinators
774:Limitations
668:to prevent
666:water table
654:prairie dog
624:Prairie dog
600:pollination
596:honeyeaters
565:honeyeaters
561:pollinators
490:sea urchins
395:sea urchins
288:herbivorous
273:competition
265:environment
255:Definitions
119:, are also
3702:Categories
3663:Sexecology
3240:Parasitism
3205:Antibiosis
3040:Resistance
3035:Resilience
2925:Population
2845:Camouflage
2797:Oligotroph
2712:Ascendency
2674:intertidal
2664:cold seeps
2618:Food chain
2419:Herbivores
2394:Carnivores
2321:Mixotrophs
2296:Autotrophs
2175:components
1495:BioScience
1353:BioScience
1203:(2): 405.
1181:2011-02-03
1175:10535/2966
940:BioScience
917:2011-02-03
848:References
832:Indigenous
745:parrotfish
688:, and the
605:frugivores
580:region of
572:Mutualists
405:Sea otters
237:Washington
77:ecosystems
3568:Allometry
3522:Emergence
3250:Symbiosis
3235:Mutualism
3030:Stability
2935:Abundance
2747:Dominance
2705:Processes
2694:tide pool
2590:Food webs
2464:Predation
2449:Omnivores
2376:Consumers
2331:Mycotroph
2288:Producers
2233:Ecosystem
2198:Behaviour
2113:Caro, Tim
1992:0012-9658
1937:0012-9658
1854:Oecologia
1550:1948-6596
1476:0021-8790
756:Serengeti
737:peccaries
731:. In the
725:elephants
690:mule deer
686:pronghorn
615:Engineers
609:cassowary
540:mammalian
498:shellfish
480:sea stars
462:from the
460:gray wolf
453:ecosystem
414:holdfasts
391:sea otter
376:holdfasts
350:Predators
233:Makah Bay
108:ecosystem
104:organisms
44:, and an
3623:Endolith
3552:Xerosere
3464:networks
3280:Ecocline
2826:Defense,
2502:Detritus
2404:Foraging
2273:Resource
2115:(2010).
2000:27870025
1945:18376543
1882:28547281
1718:53686934
1593:85265656
1400:23 April
1291:84866250
1049:11030284
1003:85265656
893:83780992
811:See also
791:Pisaster
786:Pisaster
764:wildfire
729:woodland
713:riparian
678:aeration
421:25,000.
407:protect
342:—
284:predator
221:starfish
181:predator
148:flagship
128:keystone
73:wetlands
38:flagship
3718:Habitat
3613:Ecopath
3420:Habitat
3290:Ecotype
3285:Ecotone
3262:ecology
3260:Spatial
3196:Species
3056:Species
2927:ecology
2912:Ecology
2860:Mimicry
2828:counter
2772:f-ratio
2520:Archaea
2208:Biomass
2181:General
2173:Trophic
2165:Ecology
1972:Bibcode
1964:Ecology
1917:Bibcode
1909:Ecology
1890:5940275
1862:Bibcode
1770:Bibcode
1698:Bibcode
1644:8925215
1636:9774274
1616:Bibcode
1608:Science
1585:2459379
1454:Bibcode
1375:1313259
1334:1938786
1314:Bibcode
1306:Ecology
1271:Bibcode
1205:Bibcode
1123:Bibcode
1029:Bibcode
995:2459379
960:1312122
885:2459472
754:In the
747:on the
741:wallows
721:savanna
674:erosion
494:mussels
482:(e.g.,
277:biomass
261:species
225:mussels
215:of the
158:History
136:biomass
88:species
61:habitat
40:, and
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670:runoff
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630:, 1844
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532:jaguar
519:, and
513:sharks
332:jaguar
326:, and
297:weevil
130:in an
106:in an
69:canals
57:beaver
34:jaguar
3561:Other
3462:Other
3415:Guild
3387:Niche
2639:Lakes
1886:S2CID
1737:CSIRO
1714:S2CID
1640:S2CID
1589:S2CID
1581:JSTOR
1534:(3).
1371:JSTOR
1330:JSTOR
1287:S2CID
1197:OIKOS
1095:1 May
1045:S2CID
999:S2CID
991:JSTOR
956:JSTOR
889:S2CID
881:JSTOR
648:. In
292:plant
86:is a
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2121:ISBN
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1996:PMID
1988:ISSN
1941:PMID
1933:ISSN
1878:PMID
1839:2013
1812:2013
1741:ISBN
1669:ISBN
1632:PMID
1546:ISSN
1472:ISSN
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1402:2018
1241:ISBN
1097:2024
1071:2017
760:gnus
707:The
672:and
660:and
638:and
594:for
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517:rays
418:kelp
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271:and
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150:and
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71:and
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