262:, 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.
481:. 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
726:
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
812:
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
431:
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
422:
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
305:
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
817:
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
349:
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
1271:
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
145:. 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
466:. 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.
165:. 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.
325:
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
121:
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
613:
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
738:, 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
2070:
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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
329:
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
587:
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
511:
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
1400:
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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
306:
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
1969:
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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2080:
3127:
1617:
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".
1315:
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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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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1971:"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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Wilmers, Christopher C.; Crabtree, Robert L.; Smith, Douglas W.; Murphy, Kerry M.; Getz, Wayne M. (November 2003).
1453:"Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park"
3628:
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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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1161:"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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238:. He removed the starfish from an area, and documented the effects on the ecosystem. In his 1966 paper,
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222:. 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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1075:"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
199:
1771:
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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687:, and can also serve to change the composition of the soil in a region by increasing
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1247:
Large Mammal Restoration: Ecological And Sociological Challenges In The 21St Century
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1900:
1838:"Associated Species : Prairie Dogs are a Keystone Species of the Great Plains"
1654:
1124:
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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459:
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1916:"Elephants as agents of habitat creation for small vertebrates at the patch scale"
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2444:
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2346:
2326:
2057:"How the overlooked peccary engineers the Amazon, an interview with Harald Beck"
2028:
676:
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549:, acts as a keystone predator by its widely varied diet, helping to balance the
419:
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relative to its abundance. The concept was introduced in 1969 by the zoologist
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17:
2002:
1947:
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874:
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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401:
118:
2010:
1955:
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have shown a proclivity for grazing on the same land used by prairie dogs.
553:
jungle ecosystem with its consumption of 87 different species of prey. The
375:
109:. Keystone species play a critical role in maintaining the structure of an
1646:
1502:"Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems?"
3633:
3562:
3093:
2600:
2512:
2459:
2414:
2123:
1915:
1401:"My Turn: Southeast Alaska's sea otters: The restoration of an ecosystem"
774:
739:
723:
688:
294:
231:
191:
114:
83:
250:. In his 1969 paper, Paine proposed the keystone species concept, using
218:
The concept of the keystone species was introduced in 1969 by zoologist
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1939:
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1994:
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545:, whose numbers in Central and South America have been classified as
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79:
67:
44:
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1336:
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997:
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1747:
1574:
Paine, R. T. (1966). "Food web complexity and species diversity".
984:
Paine, R. T. (1966). "Food Web Complexity and Species Diversity".
705:
675:. Prairie dog tunnel systems also help channel rainwater into the
629:
559:
439:
302:
75:
61:
38:
1535:"perspective: The keystone species concept: a critical appraisal"
777:
likelihood, which in turn promotes tree growth. The documentary
554:
535:
428:
142:
127:
123:
2895:
2148:
1533:
Cottee-Jones, Henry Eden W; Whittaker, Robert J. (2012-09-28).
345:
is a charismatic big cat which meets all of these definitions:
256:, a species of starfish generally known as ochre starfish, and
770:
1519:
10.1641/0006-3568(2004)054[0755:WATEOF]2.0.CO;2
947:"The Keystone-Species Concept in Ecology and Conservation"
570:, is periodically the sole source of nectar for important
1100:"Ecologists Struggle to Get a Grip on 'Keystone Species'"
1244:
Maehr, David; Noss, Reed F.; Larkin, Jeffery L. (2001).
2096:"Single keystone species may be the key to reef health"
319:
in North American waters. Similarly, the wasp species
1675:
Wild Cats, Status Survey and Conservation Action Plan
714:
which has a transformative effect on the environment
408:, making it a keystone species for the kelp forests.
3571:
3471:
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2836:
2715:
2642:
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2521:
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2385:
2297:
2191:
2033:
The sixth extinction: biodiversity and its survival
1813:. Nebraska Game and Park Commission. Archived from
1678:. IUCN/SSC Cat Specialist Group. pp. 118–122.
274:that has a disproportionately large effect on its
101:that has a disproportionately large effect on its
230:(between the high and low tide lines), including
1025:
1023:
945:Mills, L. S.; Soule, M. E.; Doak, D. F. (1993).
385:can damage kelp forests by chewing through kelp
1500:Ripple, William J.; Beschta, Robert L. (2004).
1428:Environmental Science: Earth as a living planet
347:
2071:"Where Peccaries Wallow, Other Animals Follow"
27:Species with a large effect on its environment
2907:
2160:
444:Riparian willow recovery at Blacktail Creek,
270:A keystone species was defined by Paine as a
8:
1239:
1237:
1093:
1091:
1274:Archaeological and Anthropological Sciences
1119:
1117:
940:
938:
869:
867:
3128:Latitudinal gradients in species diversity
2914:
2900:
2892:
2167:
2153:
2145:
920:. University of Washington. Archived from
2031:(1999) . "11 The modern elephant story".
1550:
1517:
1476:
1375:
1184:
695:, which is another keystone species, the
240:Food Web Complexity and Species Diversity
74:creator, responsible for the creation of
3026:Predator–prey (Lotka–Volterra) equations
2665:Tritrophic interactions in plant defense
734:, the larger herbivores, especially the
3058:Random generalized Lotka–Volterra model
1840:. Prairie Dog Coalition. Archived from
1668:Nowell, K.; Jackson, P., eds. (1996). "
863:
242:, Paine had described such a system in
2866:Herbivore adaptations to plant defense
1746:. Biodiversity Technical Paper No. 2.
601:(acorn banksia) is the sole source of
595:, there is a period of each year when
485:Sea stars and other non-apex predators
290:dominance within a functional group."
86:irrigating large forests and creating
1098:Ogwen, Lesley Evans (24 April 2024).
436:The wolf, Yellowstone's apex predator
7:
2881:Predator avoidance in schooling fish
1430:. John Wiley & Sons. p. 2.
1073:HHMI, BioInteractive (29 May 2017).
489:As described by Paine in 1966, some
458:Keystone predators may increase the
3331:Intermediate disturbance hypothesis
2079:. 27 September 2014. Archived from
1793:10.1046/j.1523-1739.1995.09040747.x
1512:(8). Oxford University Press: 755.
1146:10.1046/j.1523-1739.1995.09040962.x
795:own work has shown that the purple
3084:Ecological effects of biodiversity
2109:How Wildebeest Saved the Serengeti
2094:Gruber, Karl (26 September 2014).
1750:Division of Wildlife and Ecology.
655:A term used alongside keystone is
609:, which play an important role in
301:species from eliminating dominant
25:
2420:Generalist and specialist species
1228:10.1034/j.1600-0706.2000.910222.x
448:, after reintroduction of wolves.
153:. It became a popular concept in
3143:Occupancy–abundance relationship
2130:. Washington, DC: Island Press.
1478:10.1046/j.1365-2656.2003.00766.x
1399:Brown, Tina M. (14 March 2013).
393:
374:
293:A classic keystone species is a
198:
177:
3163:Relative abundance distribution
2876:Plant defense against herbivory
2743:Competitive exclusion principle
2455:Mesopredator release hypothesis
1426:Botkin, D.; Keller, E. (2003).
1403:. Juneau Empire. Archived from
2748:Consumer–resource interactions
1:
3594:Biological data visualization
3421:Environmental niche modelling
3148:Population viability analysis
2035:. Phoenix. pp. 216–217.
1032:Biodiversity and Conservation
918:"Keystone Species Hypothesis"
769:, the presence of sufficient
557:is another keystone species.
475:Greater Yellowstone Ecosystem
435:
3079:Density-dependent inhibition
1639:10.1126/science.282.5388.473
1250:. Island Press. p. 73.
773:in these grasslands reduces
518:These creatures need not be
477:had profound impacts on the
404:is an important predator of
3548:Liebig's law of the minimum
3383:Resource selection function
2274:Metabolic theory of ecology
2076:National Geographic Society
1914:Pringle, Robert M. (2008).
1742:Lambeck, Robert J. (1999).
366:Sea otters and kelp forests
297:that prevents a particular
3750:
3448:Niche apportionment models
3168:Relative species abundance
2372:Primary nutritional groups
2269:List of feeding behaviours
1836:Rosmarino, Nicole (2007).
1360:"Understanding Sea Otters"
781:documents this in detail.
644:
534:. Sea otters are prey for
451:
70:: a keystone species, and
29:
3697:
3629:Ecosystem based fisheries
3241:Interspecific competition
3133:Minimum viable population
2991:Maximum sustainable yield
2976:Intraspecific competition
2971:Effective population size
2851:Anti-predator adaptations
2362:Photosynthetic efficiency
1885:10.1007/s00442-002-0929-1
1539:Frontiers of Biogeography
1457:Journal of Animal Ecology
1294:10.1007/s12520-013-0122-y
1052:10.1007/s10531-011-0035-y
828:Cultural keystone species
522:. Sea stars are prey for
446:Yellowstone National Park
286:is large relative to its
3619:Ecological stoichiometry
3584:Alternative stable state
2098:. Australian Geographic.
1177:10.5751/ES-00502-0701r11
513:sea star wasting disease
30:Not to be confused with
3463:Ontogenetic niche shift
3326:Ideal free distribution
3236:Ecological facilitation
2986:Malthusian growth model
2956:Consumer-resource model
2813:Paradox of the plankton
2778:Energy systems language
2498:Chemoorganoheterotrophy
2465:Optimal foraging theory
2440:Heterotrophic nutrition
986:The American Naturalist
876:The American Naturalist
469:The elimination of the
355:David Maehr et al, 2001
113:, affecting many other
3609:Ecological forecasting
3553:Marginal value theorem
3351:Landscape epidemiology
3286:Cross-boundary subsidy
3221:Biological interaction
2571:Microbial intelligence
2259:Green world hypothesis
1811:Wildlife Species Guide
715:
642:
579:
449:
381:Sea urchins like this
358:
312:Euhrychiopsis lecontei
90:
59:
3614:Ecological humanities
3513:Ecological energetics
3458:Niche differentiation
3321:Habitat fragmentation
3089:Ecological extinction
3036:Small population size
2788:Feed conversion ratio
2768:Ecological succession
2700:San Francisco Estuary
2614:Ecological efficiency
2556:Microbial cooperation
1159:Davic, R. D. (2003).
750:produce and maintain
709:
645:Further information:
633:
563:
452:Further information:
443:
317:Eurasian watermilfoil
259:Mytilus californianus
211:), the seastar's prey
208:Mytilus californianus
65:
42:
3724:Conservation biology
3639:Evolutionary ecology
3604:Ecological footprint
3599:Ecological economics
3523:Ecological threshold
3518:Ecological indicator
3388:Source–sink dynamics
3341:Land change modeling
3336:Insular biogeography
3188:Species distribution
2927:Modelling ecosystems
2586:Microbial metabolism
2425:Intraguild predation
2214:Biogeochemical cycle
2180:Modelling ecosystems
2059:. 20 September 2010.
1773:Conservation Biology
1358:Cohn, J. P. (1998).
1165:Conservation Ecology
1126:Conservation Biology
224:marine invertebrates
205:California mussels (
155:conservation biology
111:ecological community
3734:Biology terminology
3689:Theoretical ecology
3664:Natural environment
3528:Ecosystem diversity
3498:Ecological collapse
3488:Bateman's principle
3443:Limiting similarity
3356:Landscape limnology
3178:Species homogeneity
3016:Population modeling
3011:Population dynamics
2828:Trophic state index
1987:2016Ecol...97.3219C
1932:2008Ecol...89...26P
1877:2002Oecol.132...96W
1844:on 10 November 2013
1785:1995ConBi...9..747W
1713:2018ResEc..26..439H
1701:Restoration Ecology
1631:1998Sci...282..473E
1576:American Naturalist
1469:2003JAnEc..72..909W
1329:1978Ecol...59..822E
1286:2013ArAnS...5..159S
1220:2000Oikos..91..405C
1138:1995ConBi...9..962P
1044:2011BiCon..20.1427B
779:The Serengeti Rules
712:animal construction
341:. For example, the
103:natural environment
3700:Outline of ecology
3649:Industrial ecology
3644:Functional ecology
3508:Ecological deficit
3453:Niche construction
3416:Ecosystem engineer
3193:Species–area curve
3114:Introduced species
2929:: Other components
2861:Deimatic behaviour
2763:Ecological network
2695:North Pacific Gyre
2680:hydrothermal vents
2619:Ecological pyramid
2566:Microbial food web
2377:Primary production
2322:Foundation species
2083:on April 18, 2020.
1552:10.21425/F54312533
848:Introduced species
838:Foundation species
790:Depends on context
760:Great Barrier Reef
744:Amazon river basin
716:
657:ecosystem engineer
651:Niche construction
647:Ecosystem engineer
643:
580:
496:Pisaster ochraceus
450:
253:Pisaster ochraceus
187:Pisaster ochraceus
91:
60:
32:foundation species
3719:Community ecology
3706:
3705:
3589:Balance of nature
3346:Landscape ecology
3231:Community ecology
3173:Species diversity
3109:Indicator species
3104:Gradient analysis
2981:Logistic function
2889:
2888:
2846:Animal coloration
2823:Trophic mutualism
2561:Microbial ecology
2352:Photoheterotrophs
2337:Myco-heterotrophy
2249:Ecosystem ecology
2234:Carrying capacity
2199:Abiotic component
1981:(11): 3219–3230.
1940:10.1890/07-0776.1
1817:on 19 August 2009
1757:978-0-642-21423-2
1721:10.1111/rec.12684
1685:978-2-8317-0045-8
1625:(5388): 473–476.
1437:978-0-471-38914-9
1257:978-1-55963-817-3
1102:. Quanta Magazine
833:Ecosystem service
808:Multiple meanings
637:town. Drawing by
598:Banksia prionotes
593:Western Australia
567:Banksia prionotes
383:purple sea urchin
335:indicator species
280:species diversity
16:(Redirected from
3741:
3406:Ecological niche
3378:selection theory
3198:Umbrella species
3183:Species richness
3119:Invasive species
3099:Flagship species
3006:Population cycle
3001:Overexploitation
2966:Ecological yield
2916:
2909:
2902:
2893:
2798:Mesotrophic soil
2738:Climax community
2670:Marine food webs
2609:Biomagnification
2410:Chemoorganotroph
2264:Keystone species
2224:Biotic component
2169:
2162:
2155:
2146:
2141:
2111:
2106:
2100:
2099:
2091:
2085:
2084:
2067:
2061:
2060:
2053:
2047:
2046:
2021:
2015:
2014:
1995:10.1002/ecy.1557
1966:
1960:
1959:
1911:
1905:
1904:
1860:
1854:
1853:
1851:
1849:
1833:
1827:
1826:
1824:
1822:
1803:
1797:
1796:
1768:
1762:
1761:
1739:
1733:
1732:
1696:
1690:
1689:
1665:
1659:
1658:
1614:
1608:
1607:
1571:
1565:
1564:
1554:
1530:
1524:
1523:
1521:
1497:
1491:
1490:
1480:
1448:
1442:
1441:
1423:
1417:
1416:
1414:
1412:
1407:on 23 April 2018
1396:
1390:
1389:
1379:
1355:
1349:
1348:
1312:
1306:
1305:
1268:
1262:
1261:
1241:
1232:
1231:
1203:
1197:
1196:
1194:
1193:
1188:
1156:
1150:
1149:
1121:
1112:
1111:
1109:
1107:
1095:
1086:
1085:
1083:
1081:
1070:
1064:
1063:
1038:(7): 1427–1440.
1027:
1018:
1017:
981:
975:
974:
942:
933:
932:
930:
929:
914:
908:
907:
871:
853:Sentinel species
669:mountain plovers
397:
378:
356:
339:umbrella species
331:flagship species
202:
184:Ochre seastars (
181:
163:umbrella species
95:keystone species
53:umbrella species
21:
3749:
3748:
3744:
3743:
3742:
3740:
3739:
3738:
3709:
3708:
3707:
3702:
3693:
3679:Systems ecology
3567:
3538:Extinction debt
3503:Ecological debt
3493:Bioluminescence
3474:
3467:
3436:marine habitats
3411:Ecological trap
3392:
3272:
3265:
3208:
3202:
3158:Rapoport's rule
3153:Priority effect
3094:Endemic species
3062:
3021:Population size
2937:
2930:
2920:
2890:
2885:
2838:
2832:
2818:Trophic cascade
2728:Bioaccumulation
2711:
2638:
2595:
2517:
2484:
2381:
2293:
2254:Ecosystem model
2187:
2173:
2138:
2122:
2119:
2117:Further reading
2114:
2107:
2103:
2093:
2092:
2088:
2069:
2068:
2064:
2055:
2054:
2050:
2043:
2025:Leakey, Richard
2023:
2022:
2018:
1968:
1967:
1963:
1913:
1912:
1908:
1862:
1861:
1857:
1847:
1845:
1835:
1834:
1830:
1820:
1818:
1805:
1804:
1800:
1770:
1769:
1765:
1758:
1741:
1740:
1736:
1698:
1697:
1693:
1686:
1667:
1666:
1662:
1616:
1615:
1611:
1573:
1572:
1568:
1532:
1531:
1527:
1499:
1498:
1494:
1450:
1449:
1445:
1438:
1425:
1424:
1420:
1410:
1408:
1398:
1397:
1393:
1377:10.2307/1313259
1357:
1356:
1352:
1337:10.2307/1938786
1314:
1313:
1309:
1270:
1269:
1265:
1258:
1243:
1242:
1235:
1205:
1204:
1200:
1191:
1189:
1158:
1157:
1153:
1123:
1122:
1115:
1105:
1103:
1097:
1096:
1089:
1079:
1077:
1072:
1071:
1067:
1029:
1028:
1021:
983:
982:
978:
963:10.2307/1312122
944:
943:
936:
927:
925:
916:
915:
911:
873:
872:
865:
861:
824:
810:
792:
787:
730:In the African
710:Beaver dam, an
653:
628:
585:
564:Acorn banksia,
547:near threatened
487:
479:trophic pyramid
456:
438:
413:
412:
411:
410:
409:
398:
390:
389:
379:
368:
363:
357:
354:
268:
228:intertidal zone
220:Robert T. Paine
216:
215:
214:
213:
212:
203:
195:
194:
182:
171:
107:Robert T. Paine
35:
28:
23:
22:
15:
12:
11:
5:
3747:
3745:
3737:
3736:
3731:
3726:
3721:
3711:
3710:
3704:
3703:
3698:
3695:
3694:
3692:
3691:
3686:
3681:
3676:
3671:
3666:
3661:
3659:Microecosystem
3656:
3651:
3646:
3641:
3636:
3631:
3626:
3621:
3616:
3611:
3606:
3601:
3596:
3591:
3586:
3581:
3575:
3573:
3569:
3568:
3566:
3565:
3560:
3558:Thorson's rule
3555:
3550:
3545:
3540:
3535:
3530:
3525:
3520:
3515:
3510:
3505:
3500:
3495:
3490:
3485:
3483:Assembly rules
3479:
3477:
3469:
3468:
3466:
3465:
3460:
3455:
3450:
3445:
3440:
3439:
3438:
3428:
3423:
3418:
3413:
3408:
3402:
3400:
3394:
3393:
3391:
3390:
3385:
3380:
3368:
3366:Patch dynamics
3363:
3361:Metapopulation
3358:
3353:
3348:
3343:
3338:
3333:
3328:
3323:
3318:
3313:
3308:
3303:
3298:
3293:
3288:
3283:
3277:
3275:
3267:
3266:
3264:
3263:
3258:
3256:Storage effect
3253:
3248:
3243:
3238:
3233:
3228:
3223:
3218:
3212:
3210:
3204:
3203:
3201:
3200:
3195:
3190:
3185:
3180:
3175:
3170:
3165:
3160:
3155:
3150:
3145:
3140:
3138:Neutral theory
3135:
3130:
3125:
3123:Native species
3116:
3111:
3106:
3101:
3096:
3091:
3086:
3081:
3076:
3070:
3068:
3064:
3063:
3061:
3060:
3055:
3054:
3053:
3048:
3038:
3033:
3028:
3023:
3018:
3013:
3008:
3003:
2998:
2996:Overpopulation
2993:
2988:
2983:
2978:
2973:
2968:
2963:
2958:
2953:
2948:
2942:
2940:
2932:
2931:
2921:
2919:
2918:
2911:
2904:
2896:
2887:
2886:
2884:
2883:
2878:
2873:
2868:
2863:
2858:
2853:
2848:
2842:
2840:
2834:
2833:
2831:
2830:
2825:
2820:
2815:
2810:
2805:
2803:Nutrient cycle
2800:
2795:
2793:Feeding frenzy
2790:
2785:
2780:
2775:
2773:Energy quality
2770:
2765:
2760:
2755:
2750:
2745:
2740:
2735:
2733:Cascade effect
2730:
2725:
2719:
2717:
2713:
2712:
2710:
2709:
2708:
2707:
2702:
2697:
2692:
2687:
2682:
2677:
2667:
2662:
2657:
2652:
2646:
2644:
2640:
2639:
2637:
2636:
2631:
2626:
2621:
2616:
2611:
2605:
2603:
2597:
2596:
2594:
2593:
2588:
2583:
2578:
2576:Microbial loop
2573:
2568:
2563:
2558:
2553:
2548:
2543:
2541:Lithoautotroph
2538:
2533:
2527:
2525:
2523:Microorganisms
2519:
2518:
2516:
2515:
2510:
2505:
2500:
2494:
2492:
2486:
2485:
2483:
2482:
2480:Prey switching
2477:
2472:
2467:
2462:
2457:
2452:
2447:
2442:
2437:
2432:
2427:
2422:
2417:
2412:
2407:
2402:
2397:
2391:
2389:
2383:
2382:
2380:
2379:
2374:
2369:
2364:
2359:
2357:Photosynthesis
2354:
2349:
2344:
2339:
2334:
2329:
2324:
2319:
2314:
2312:Chemosynthesis
2309:
2303:
2301:
2295:
2294:
2292:
2291:
2286:
2281:
2276:
2271:
2266:
2261:
2256:
2251:
2246:
2241:
2236:
2231:
2226:
2221:
2216:
2211:
2206:
2204:Abiotic stress
2201:
2195:
2193:
2189:
2188:
2174:
2172:
2171:
2164:
2157:
2149:
2143:
2142:
2136:
2118:
2115:
2113:
2112:
2101:
2086:
2062:
2048:
2041:
2016:
1961:
1906:
1855:
1828:
1807:"Prairie Dogs"
1798:
1779:(4): 747–752.
1763:
1756:
1734:
1707:(3): 439–445.
1691:
1684:
1660:
1609:
1588:10.1086/282400
1582:(910): 65–75.
1566:
1525:
1492:
1463:(6): 909–916.
1443:
1436:
1418:
1391:
1370:(3): 151–155.
1350:
1323:(4): 822–833.
1307:
1280:(2): 159–182.
1263:
1256:
1233:
1198:
1151:
1132:(4): 962–964.
1113:
1087:
1065:
1019:
998:10.1086/282400
992:(910): 65–75.
976:
957:(4): 219–224.
934:
909:
888:10.1086/282586
882:(929): 91–93.
862:
860:
857:
856:
855:
850:
845:
840:
835:
830:
823:
820:
809:
806:
791:
788:
786:
783:
673:burrowing owls
627:
624:
618:, such as the
589:Avon Wheatbelt
584:
581:
520:apex predators
499:) may prey on
486:
483:
437:
434:
399:
392:
391:
380:
373:
372:
371:
370:
369:
367:
364:
362:
359:
352:
322:Agelaia vicina
267:
264:
204:
197:
196:
190:), a keystone
183:
176:
175:
174:
173:
172:
170:
167:
132:apex predators
47:: a keystone,
26:
24:
18:Keystone genus
14:
13:
10:
9:
6:
4:
3:
2:
3746:
3735:
3732:
3730:
3727:
3725:
3722:
3720:
3717:
3716:
3714:
3701:
3696:
3690:
3687:
3685:
3684:Urban ecology
3682:
3680:
3677:
3675:
3672:
3670:
3667:
3665:
3662:
3660:
3657:
3655:
3652:
3650:
3647:
3645:
3642:
3640:
3637:
3635:
3632:
3630:
3627:
3625:
3622:
3620:
3617:
3615:
3612:
3610:
3607:
3605:
3602:
3600:
3597:
3595:
3592:
3590:
3587:
3585:
3582:
3580:
3577:
3576:
3574:
3570:
3564:
3561:
3559:
3556:
3554:
3551:
3549:
3546:
3544:
3543:Kleiber's law
3541:
3539:
3536:
3534:
3531:
3529:
3526:
3524:
3521:
3519:
3516:
3514:
3511:
3509:
3506:
3504:
3501:
3499:
3496:
3494:
3491:
3489:
3486:
3484:
3481:
3480:
3478:
3476:
3470:
3464:
3461:
3459:
3456:
3454:
3451:
3449:
3446:
3444:
3441:
3437:
3434:
3433:
3432:
3429:
3427:
3424:
3422:
3419:
3417:
3414:
3412:
3409:
3407:
3404:
3403:
3401:
3399:
3395:
3389:
3386:
3384:
3381:
3379:
3377:
3373:
3369:
3367:
3364:
3362:
3359:
3357:
3354:
3352:
3349:
3347:
3344:
3342:
3339:
3337:
3334:
3332:
3329:
3327:
3324:
3322:
3319:
3317:
3316:Foster's rule
3314:
3312:
3309:
3307:
3304:
3302:
3299:
3297:
3294:
3292:
3289:
3287:
3284:
3282:
3279:
3278:
3276:
3274:
3268:
3262:
3259:
3257:
3254:
3252:
3249:
3247:
3244:
3242:
3239:
3237:
3234:
3232:
3229:
3227:
3224:
3222:
3219:
3217:
3214:
3213:
3211:
3205:
3199:
3196:
3194:
3191:
3189:
3186:
3184:
3181:
3179:
3176:
3174:
3171:
3169:
3166:
3164:
3161:
3159:
3156:
3154:
3151:
3149:
3146:
3144:
3141:
3139:
3136:
3134:
3131:
3129:
3126:
3124:
3120:
3117:
3115:
3112:
3110:
3107:
3105:
3102:
3100:
3097:
3095:
3092:
3090:
3087:
3085:
3082:
3080:
3077:
3075:
3072:
3071:
3069:
3065:
3059:
3056:
3052:
3049:
3047:
3044:
3043:
3042:
3039:
3037:
3034:
3032:
3029:
3027:
3024:
3022:
3019:
3017:
3014:
3012:
3009:
3007:
3004:
3002:
2999:
2997:
2994:
2992:
2989:
2987:
2984:
2982:
2979:
2977:
2974:
2972:
2969:
2967:
2964:
2962:
2959:
2957:
2954:
2952:
2949:
2947:
2944:
2943:
2941:
2939:
2933:
2928:
2924:
2917:
2912:
2910:
2905:
2903:
2898:
2897:
2894:
2882:
2879:
2877:
2874:
2872:
2869:
2867:
2864:
2862:
2859:
2857:
2854:
2852:
2849:
2847:
2844:
2843:
2841:
2835:
2829:
2826:
2824:
2821:
2819:
2816:
2814:
2811:
2809:
2806:
2804:
2801:
2799:
2796:
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2634:Trophic level
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2591:Phage ecology
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2581:Microbial mat
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2536:Bacteriophage
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2532:
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2503:Decomposition
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2450:Mesopredators
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661:North America
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57:apex predator
54:
50:
46:
41:
37:
33:
19:
3669:Regime shift
3654:Macroecology
3375:
3371:
3311:Edge effects
3281:Biogeography
3226:Commensalism
3074:Biodiversity
2951:Allee effect
2690:kelp forests
2643:Example webs
2508:Detritivores
2347:Organotrophs
2327:Kinetotrophs
2279:Productivity
2263:
2127:
2104:
2089:
2081:the original
2074:
2065:
2051:
2032:
2029:Lewin, Roger
2019:
1978:
1974:
1964:
1926:(1): 26–33.
1923:
1919:
1909:
1868:
1864:
1858:
1846:. Retrieved
1842:the original
1831:
1819:. Retrieved
1815:the original
1810:
1801:
1776:
1772:
1766:
1743:
1737:
1704:
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1421:
1409:. Retrieved
1405:the original
1394:
1367:
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1277:
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1266:
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1207:
1201:
1190:. Retrieved
1168:
1164:
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1104:. Retrieved
1078:. Retrieved
1068:
1035:
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989:
985:
979:
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950:
926:. Retrieved
922:the original
912:
879:
875:
811:
801:
796:
793:
764:
729:
717:
693:plains bison
654:
639:Josiah Gregg
596:
586:
565:
540:
532:sea anemones
517:
507:, and other
494:
488:
468:
460:biodiversity
457:
420:kelp forests
414:
348:
328:
320:
310:
292:
269:
257:
251:
239:
217:
206:
185:
157:, alongside
151:productivity
136:
94:
92:
36:
3306:Disturbance
3209:interaction
3031:Recruitment
2961:Depensation
2753:Copiotrophs
2624:Energy flow
2546:Lithotrophy
2490:Decomposers
2470:Planktivore
2445:Insectivore
2435:Heterotroph
2400:Bacterivore
2367:Phototrophs
2317:Chemotrophs
2289:Restoration
2239:Competition
1848:10 November
1821:10 November
1272:collagen".
815:pollinators
785:Limitations
679:to prevent
677:water table
665:prairie dog
635:Prairie dog
611:pollination
607:honeyeaters
576:honeyeaters
572:pollinators
501:sea urchins
406:sea urchins
299:herbivorous
284:competition
276:environment
266:Definitions
130:, are also
3713:Categories
3674:Sexecology
3251:Parasitism
3216:Antibiosis
3051:Resistance
3046:Resilience
2936:Population
2856:Camouflage
2808:Oligotroph
2723:Ascendency
2685:intertidal
2675:cold seeps
2629:Food chain
2430:Herbivores
2405:Carnivores
2332:Mixotrophs
2307:Autotrophs
2186:components
1506:BioScience
1364:BioScience
1214:(2): 405.
1192:2011-02-03
1186:10535/2966
951:BioScience
928:2011-02-03
859:References
843:Indigenous
756:parrotfish
699:, and the
616:frugivores
591:region of
583:Mutualists
416:Sea otters
248:Washington
88:ecosystems
3579:Allometry
3533:Emergence
3261:Symbiosis
3246:Mutualism
3041:Stability
2946:Abundance
2758:Dominance
2716:Processes
2705:tide pool
2601:Food webs
2475:Predation
2460:Omnivores
2387:Consumers
2342:Mycotroph
2299:Producers
2244:Ecosystem
2209:Behaviour
2124:Caro, Tim
2003:0012-9658
1948:0012-9658
1865:Oecologia
1561:1948-6596
1487:0021-8790
767:Serengeti
748:peccaries
742:. In the
736:elephants
701:mule deer
697:pronghorn
626:Engineers
620:cassowary
551:mammalian
509:shellfish
491:sea stars
473:from the
471:gray wolf
464:ecosystem
425:holdfasts
402:sea otter
387:holdfasts
361:Predators
244:Makah Bay
119:ecosystem
115:organisms
55:, and an
3634:Endolith
3563:Xerosere
3475:networks
3291:Ecocline
2837:Defense,
2513:Detritus
2415:Foraging
2284:Resource
2126:(2010).
2011:27870025
1956:18376543
1893:28547281
1729:53686934
1604:85265656
1411:23 April
1302:84866250
1060:11030284
1014:85265656
904:83780992
822:See also
802:Pisaster
797:Pisaster
775:wildfire
740:woodland
724:riparian
689:aeration
432:25,000.
418:protect
353:—
295:predator
232:starfish
192:predator
159:flagship
139:keystone
84:wetlands
49:flagship
3729:Habitat
3624:Ecopath
3431:Habitat
3301:Ecotype
3296:Ecotone
3273:ecology
3271:Spatial
3207:Species
3067:Species
2938:ecology
2923:Ecology
2871:Mimicry
2839:counter
2783:f-ratio
2531:Archaea
2219:Biomass
2192:General
2184:Trophic
2176:Ecology
1983:Bibcode
1975:Ecology
1928:Bibcode
1920:Ecology
1901:5940275
1873:Bibcode
1781:Bibcode
1709:Bibcode
1655:8925215
1647:9774274
1627:Bibcode
1619:Science
1596:2459379
1465:Bibcode
1386:1313259
1345:1938786
1325:Bibcode
1317:Ecology
1282:Bibcode
1216:Bibcode
1134:Bibcode
1040:Bibcode
1006:2459379
971:1312122
896:2459472
765:In the
758:on the
752:wallows
732:savanna
685:erosion
505:mussels
493:(e.g.,
288:biomass
272:species
236:mussels
226:of the
169:History
147:biomass
99:species
72:habitat
51:, and
2655:Rivers
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720:beaver
681:runoff
663:, the
641:, 1844
603:nectar
543:jaguar
530:, and
524:sharks
343:jaguar
337:, and
308:weevil
141:in an
117:in an
80:canals
68:beaver
45:jaguar
3572:Other
3473:Other
3426:Guild
3398:Niche
2650:Lakes
1897:S2CID
1748:CSIRO
1725:S2CID
1651:S2CID
1600:S2CID
1592:JSTOR
1545:(3).
1382:JSTOR
1341:JSTOR
1298:S2CID
1208:OIKOS
1106:1 May
1056:S2CID
1010:S2CID
1002:JSTOR
967:JSTOR
900:S2CID
892:JSTOR
659:. In
303:plant
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76:lakes
2660:Soil
2132:ISBN
2037:ISBN
2007:PMID
1999:ISSN
1952:PMID
1944:ISSN
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1850:2013
1823:2013
1752:ISBN
1680:ISBN
1643:PMID
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1432:ISBN
1413:2018
1252:ISBN
1108:2024
1082:2017
771:gnus
718:The
683:and
671:and
649:and
605:for
555:lion
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536:orca
528:rays
429:kelp
400:The
282:and
234:and
161:and
143:arch
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