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Keystone species
A key stone species is a species that has a disproportionately
large effect on its natural env ironment relativ e to its
abundance. Such species are described as play ing a critical role
in maintaining the structure of an ecological community ,
affecting many other organisms in an ecosy stem and helping to
determine the ty pes and numbers of v arious other species in
the community . A key stone species is a plant or animal that
play s a unique and crucial role in the way an ecosy stem
functions. Without key stone species, the ecosy stem would be
dramatically different or cease to exist altogether. Some
key stone species, such as the wolf, are also apex predators.

The role that a key stone species play s in its ecosy stem is
analogous to the role of a key stone in an arch. While the
key stone is under the least pressure of any of the stones in an
arch, the arch still collapses without it. Similarly , an ecosy stem
may experience a dramatic shift if a key stone species is
remov ed, ev en though that species was a small part of the
ecosy stem by measures of biomass or productiv ity . It became a The jaguar, a keystone, flagship, and umbrella
species, and an apex predator
popular concept in conserv ation biology , alongside flagship and
umbrella species. Although the concept is v alued as a
descriptor for particularly strong inter-species interactions, and it has allowed easier communication between
ecologists and conserv ation policy -makers, it has been criticized for ov ersimplify ing complex ecological sy stems.

Contents
History
Definitions
Predators
Sea otters and kelp forests
The wolf, Yellowstone's apex predator
Sea stars and other non-apex predators
Mutualists
Engineers
Limitations
See also
References
Further reading

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History
The concept of the key stone species was
introduced in 1969 by the zoologist Robert T.
Paine. [1 ][2 ] Paine dev eloped the concept to
explain his observ ations and experiments on the
relationships between marine inv ertebrates of the
intertidal zone (between the high and low tide
lines), including starfish and mussels. He remov ed
the starfish from an area, and documented the Ochre seastars (Pisaster California mussels (Mytilus
effects on the ecosy stem. [3 ] In his 1966 paper, ochraceus), a keystone californianus), the seastar's prey
predator
Food Web Complexity and Species Diversity, Paine
had described such a sy stem in Makah Bay in
Washington. [4 ] In his 1969 paper, Paine proposed the key stone species concept, using Pisaster ochraceus, a species
of starfish, and Mytilus californianus, a species of mussel, as a primary example. [1 ] The concept became popular in
conserv ation, and was deploy ed in a range of contexts and mobilized to engender support for conserv ation,
especially where human activ ities had damaged ecosy stems, such as by remov ing key stone predators. [5 ][6 ]

Definitions
A key stone species was defined by Paine as a species that has a disproportionately large effect on its env ironment
relativ e to its abundance. [7 ] It has been defined operationally by R. D. Dav ic in 2003 as "a strongly interacting
species whose top-down effect on species div ersity and competition is large relativ e to its biomass dominance within
a functional group."[8 ]

A classic key stone species is a predator that prev ents a particular herbiv orous species from eliminating dominant
plant species. If prey numbers are low, key stone predators can be ev en less abundant and still be effectiv e. Y et
without the predators, the herbiv orous prey would explode in numbers, wipe out the dominant plants, and
dramatically alter the character of the ecosy stem. The exact scenario changes in each example, but the central idea
remains that through a chain of interactions, a non-abundant species has an outsized impact on ecosy stem functions.
For example, the herbiv orous weev il Euhrychiopsis lecontei is thought to hav e key stone effects on aquatic plant
div ersity by foraging on nuisance Eurasian watermilfoil in North American waters. [9 ] Similarly , the wasp species
Agelaia vicina has been labeled a key stone species for its unparalleled nest size, colony size, and high rate of brood
production. The div ersity of its prey and the quantity necessary to sustain its high rate of growth hav e a direct
impact on other species around it. [7 ]

The key stone concept is defined by its ecological effects, and these in turn make it important for conserv ation. In
this it ov erlaps with sev eral other species conserv ation concepts such as flagship species, indicator species, and
umbrella species. For example, the jaguar is a charismatic big cat which meets all of these definitions:[1 0 ]

The jaguar is an umbrella species, flagship species, and wilderness quality indicator. It promotes the
goals of carniv ore recov ery , protecting and restoring connectiv ity through Madrean woodland and
riparian areas, and protecting and restoring riparian areas. ... A reserv e sy stem that protects jaguars is
an umbrella for many other species. ... the jaguar [is] a key stone in subtropical and tropical America ...

— David Maehr et al, 2001 [10]

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Predators

Sea otters and kelp forests

Sea otters protect kelp forests 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 lev els – fewer than
1000 in the north Pacific ocean – that they were
unable to control the sea urchin population. The Sea urchins like this The sea otter is an important
urchins in turn grazed the holdfasts of kelp so purple sea urchin can predator of sea urchins, making it
heav ily that the kelp forests largely disappeared, damage kelp forests by a keystone species for the kelp
along with all the species that depended on them. chewing through kelp forests.
holdfasts
Reintroducing the sea otters has enabled the kelp
ecosy stem to be restored. For example, in Southeast
Alaska some 400 sea otters were released, and they hav e bred to form a population approaching
25,000. [1 1 ][1 2 ][1 3 ][1 4 ]

The wolf, Yellowstone's apex predator

Key stone predators may increase the
biodiv ersity of communities by
prev enting a single species from
becoming dominant. They can hav e a
profound influence on the balance of
organisms in a particular ecosy stem.
Introduction or remov al of this Riparian willow recovery at Blacktail Creek, Yellowstone National Park, after
reintroduction of wolves
predator, or changes in its population
density , can hav e drastic cascading
effects on the equilibrium of many other populations in the ecosy stem. For example, grazers of a grassland may
prev ent a single dominant species from taking ov er. [1 5 ]

The elimination of the gray wolf from Y ellowstone National Park had profound impacts on the trophic py ramid.
Without predation, herbiv ores began to ov er-graze many woody browse species, affecting the area's plant
populations. In addition, wolv es often kept animals from grazing in riparian areas, which protected beav ers from
hav ing their food sources encroached upon. The remov al of wolv es had a direct effect on beav er populations, as
their habitat became territory for grazing. Increased browsing on willows and conifers along Blacktail Creek due to a
lack of predation caused channel incision because the beav ers helped slow the water down, allowing soil to stay in
place. Furthermore, predation keeps hy drological features such as creeks and streams in normal working order.
When wolv es were reintroduced, the beav er population and the whole riparian ecosy stem recov ered dramatically
within a few y ears. [1 6 ]

Sea stars and other non-apex predators

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As described by Paine in 1966, some sea stars (e.g., Pisaster ochraceus) may prey on sea urchins, mussels, and other
shellfish that hav e no other natural predators. If the sea star is remov ed from the ecosy stem, the mussel population
explodes uncontrollably , driv ing out most other species. [1 7 ]

These creatures need not be apex predators. Sea stars are prey for sharks, ray s, and sea anemones. Sea otters are
prey for orca. [1 8 ]

The jaguar, whose numbers in Central and South America hav e been classified as near threatened, acts as a key stone
predator by its widely v aried diet, helping to balance the mammalian jungle ecosy stem with its consumption of 87
different species of prey . [1 9 ] The lion is another key stone species. [2 0 ]

Mutualists
Key stone mutualists are organisms that participate in mutually beneficial
interaction and the loss of which would hav e a profound impact upon the
ecosy stem as a whole. For example, in the Av on Wheatbelt region of
Western Australia, there is a period of each y ear when Banksia prionotes
(acorn banksia) is the sole source of nectar for honey eaters, which play an
important role in pollination of numerous plant species. Therefore, the loss
of this one species of tree would probably cause the honey eater population
Acorn banksia, Bank sia prionotes, is
to collapse, with profound implications for the entire ecosy stem. Another the sole source of nectar for
example is frugiv ores such as the cassowary , which spreads the seeds of important pollinators, honeyeaters.
many different trees, and some will not grow unless they hav e been
through a cassowary . [2 1 ][2 2 ]

Engineers
A term used alongside key stone is ecosy stem engineer. [5 ] In North
America, the prairie dog is an ecosy stem engineer. Prairie dog burrows
prov ide the nesting areas for mountain plov ers and burrowing owls. Prairie
dog tunnel sy stems also help channel rainwater into the water table to
prev ent runoff and erosion, and can also serv e to change the composition
of the soil in a region by increasing aeration and rev ersing soil compaction
that can be a result of cattle grazing. Prairie dogs also trim the v egetation
Prairie dog town. Drawing by Josiah around their colonies, perhaps to remov e any cov er for predators. [2 3 ]
Gregg, 1844
Grazing species such as plains bison, pronghorn, and mule deer hav e shown
a procliv ity for grazing on the same land used by prairie dogs. [2 4 ]

The beav er is a well known ecosy stem engineer and key stone species. It transforms its territory from a stream to a
pond or swamp. Beav ers affect the env ironment first altering the edges of riparian areas by cutting down older trees
to use for their dams. This allows y ounger trees to take their place. Beav er 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 riv ers into wetlands, meadows, or riv erine forests. These dams hav e been shown to be beneficial to a my riad of
species including amphibians, salmon, and song birds. [2 5 ]

In the African sav anna, the larger herbiv ores, especially the elephants, shape their env ironment. The elephants
destroy trees, making room for the grass species. Without these animals, much of the sav anna would turn into
woodland. [2 6 ]

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Australian studies hav e found that parrotfish on the Great Barrier Reef 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 sev ere
strain. [2 7 ]

Limitations
Although the concept of the key stone species has a v alue in describing
particularly strong inter-species interactions, and for allowing easier
communication between ecologists and conserv ation policy -makers, it has Beaver dam, an animal construction
which has a transformative effect on
been criticized by L. S. Mills and colleagues for ov ersimplify ing complex
the environment
ecological sy stems. The term has been applied widely in different
ecosy stems and to predators, prey , and plants (primary producers),
inev itably with differing ecological meanings. For instance, remov ing a predator may allow other animals to increase
to the point where they wipe out other species; remov ing a prey species may cause predator populations to crash, or
may allow predators to driv e other prey species to extinction; and remov ing a plant species may result in the loss of
animals that depend on it, like pollinators and seed dispersers. Beav ers too hav e been called key stone, not for eating
other species but for modify ing the env ironment in way s that affected other species. The term has thus been giv en
quite different meanings in different cases. In Mills's v iew, Paine's work showed that a few species could sometimes
hav e extremely strong interactions within a particular ecosy stem, but that does not automatically imply that other
ecosy stems hav e a similar structure. [3 ]

See also
Cultural keystone species
Foundation species
Indigenous
Introduced species

References
1. Paine, R. T. (1969). "A Note on Trophic Complexity and Community Stability". The American Naturalist. 103 (929): 91–
93. doi:10.1086/282586 (https://doi.org/10.1086%2F282586). JSTOR 2459472 (https://www.jstor.org/stable/2459472).
2. "Keystone Species Hypothesis" (https://web.archive.org/web/20110110060757/http://www.washington.edu/research/path
breakers/1969g.html). University of Washington. Archived from the original (http://www.washington.edu/research/pathbre
akers/1969g.html) on 2011-01-10. Retrieved 2011-02-03.
3. Mills, L. S.; Soule, M. E.; Doak, D. F. (1993). "The Keystone-Species Concept in Ecology and Conservation".
BioScience. 43 (4): 219–224. doi:10.2307/1312122 (https://doi.org/10.2307%2F1312122). JSTOR 1312122 (https://www.j
stor.org/stable/1312122).
4. Paine, R. T. (1966). "Food Web Complexity and Species Diversity". The American Naturalist. 100 (910): 65–75.
doi:10.1086/282400 (https://doi.org/10.1086%2F282400). JSTOR 2459379 (https://www.jstor.org/stable/2459379).
5. Barua, Maan (2011). "Mobilizing metaphors: the popular use of keystone, flagship and umbrella species concepts".
Biodiversity and Conservation. 20 (7): 1427–1440. doi:10.1007/s10531-011-0035-y (https://doi.org/10.1007%2Fs10531-01
1-0035-y).
6. HHMI, BioInteractive. "Some Animals Are More Equal than Others: Keystone Species and Trophic Cascades – HHMI
(2016)" (http://naturedocumentaries.org/14877/animals-equal-others-keystone-species-trophic-cascades-hhmi-2016/).
Retrieved 6 June 2017.

https://en.wikipedia.org/wiki/Keystone_species 5/7
3/28/2019 Keystone species - Wikipedia

7. Paine, R. T. (1995). "A Conversation on Refining the Concept of Keystone Species". Conservation Biology. 9 (4): 962–
964. doi:10.1046/j.1523-1739.1995.09040962.x (https://doi.org/10.1046%2Fj.1523-1739.1995.09040962.x).
8. Davic, Robert D. (2003). "Linking Keystone Species and Functional Groups: A New Operational Definition of the
Keystone Species Concept" (http://www.consecol.org/vol7/iss1/resp11/). Conservation Ecology. Retrieved 2011-02-03.
9. Creed, R. P., Jr. (2000). "Is there a new keystone species in North American lakes and rivers?". OIKOS. 91 (2): 405.
doi:10.1034/j.1600-0706.2000.910222.x (https://doi.org/10.1034%2Fj.1600-0706.2000.910222.x).
10. Maehr, David; Noss, Reed F.; Larkin, Jeffery L. (2001). Large Mammal Restoration: Ecological And Sociological
Challenges In The 21St Century (https://books.google.com/books?id=x3n_pBQFxVQC&pg=PA73). Island Press. p. 73.
ISBN 978-1-55963-817-3.
11. 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 collagen". Archaeological and Anthropological Sciences. 5
(2): 159–182. doi:10.1007/s12520-013-0122-y (https://doi.org/10.1007%2Fs12520-013-0122-y).
12. Estes, James E.; Smith, Norman S.; Palmisano, John F. (1978). "Sea otter predation and community organization in the
Western Aleutian Islands, Alaska". Ecology. Ecology. 59 (4): 822–833. doi:10.2307/1938786 (https://doi.org/10.2307%2
F1938786). JSTOR 1938786 (https://www.jstor.org/stable/1938786).
13. Cohn, J. P. (1998). "Understanding Sea Otters". BioScience. BioScience. 48 (3): 151–155. doi:10.2307/1313259 (https://
doi.org/10.2307%2F1313259). JSTOR 1313259 (https://www.jstor.org/stable/1313259).
14. Brown, Tina M. (14 March 2013). "My Turn: Southeast Alaska's sea otters: The restoration of an ecosystem" (http://june
auempire.com/opinion/2013-03-14/my-turn-southeast-alaskas-sea-otters-restoration-ecosystem). Juneau Empire.
Retrieved 23 April 2018.
15. Botkin, D.; Keller, E. (2003). Environmental Science: Earth as a living planet. John Wiley & Sons. p. 2. ISBN 978-0-471-
38914-9.
16. Ripple, William J.; Beschta, Robert L. (2004). "Wolves and the Ecology of Fear: Can Predation Risk Structure
Ecosystems?" (https://academic.oup.com/bioscience/article/54/8/755/238242). BioScience. Oxford University Press. 54
(8): 755. doi:10.1641/0006-3568(2004)054[0755:WATEOF]2.0.CO;2 (https://doi.org/10.1641%2F0006-3568%282004%29
054%5B0755%3AWATEOF%5D2.0.CO%3B2).
17. Paine, R. T. (1966). "Food web complexity and species diversity". American Naturalist. 100: 65–75. JSTOR 2459379 (htt
ps://www.jstor.org/stable/2459379).
18. 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". Science. 282 (5388): 473–476. Bibcode:1998Sci...282..473E (http://adsabs.harvar
d.edu/abs/1998Sci...282..473E). doi:10.1126/science.282.5388.473 (https://doi.org/10.1126%2Fscience.282.5388.473).
PMID 9774274 (https://www.ncbi.nlm.nih.gov/pubmed/9774274).
19. Nowell, K.; Jackson, P., eds. (1996). Panthera onca (https://portals.iucn.org/library/node/6998). Wild Cats, Status
Survey and Conservation Action Plan. IUCN/SSC Cat Specialist Group. pp. 118–122. ISBN 978-2-8317-0045-8.
20. Hale, Sarah L.; Koprowski, John L. (February 2018). "Ecosystem-level effects of keystone species reintroduction: a
literature review". Restoration Ecology. doi:10.1111/rec.12684 (https://doi.org/10.1111%2Frec.12684).
21. Lambeck, Robert J. (1999). "Landscape Planning for Biodiversity Conservation in Agricultural Regions: A Case Study
from the Wheatbelt of Western Australia". Biodiversity Technical Paper No. 2. CSIRO Division of Wildlife and Ecology.
ISBN 978-0-642-21423-2.
22. Walker, Brian (1995). "Conserving Biological Diversity through Ecosystem Resilience". Conservation Biology. 9 (4): 747–
752. doi:10.1046/j.1523-1739.1995.09040747.x (https://doi.org/10.1046%2Fj.1523-1739.1995.09040747.x).
23. "Prairie Dogs" (https://web.archive.org/web/20090819031734/http://www.ngpc.state.ne.us/wildlife/prairiedogs.asp).
Wildlife Species Guide. Nebraska Game and Park Commission. Archived from the original (http://www.ngpc.state.ne.us/
wildlife/prairiedogs.asp) on 19 August 2009. Retrieved 10 November 2013.
24. Rosmarino, Nicole (2007). "Associated Species : Prairie Dogs are a Keystone Species of the Great Plains" (https://web.
archive.org/web/20131110163932/http://www.prairiedogcoalition.org/pd-associated-species.php). Prairie Dog Coalition.
Archived from the original (http://www.prairiedogcoalition.org/pd-associated-species.php) on 10 November 2013.
Retrieved 10 November 2013.

https://en.wikipedia.org/wiki/Keystone_species 6/7
3/28/2019 Keystone species - Wikipedia

25. Wright, J. P.; Jones, C. G.; Flecker, A. S. (2002). "An ecosystem engineer, the beaver, increases species richness at
the landscape scale" (http://www.springerlink.com/index/0637GF0979LRU90J.pdf) (PDF). Oecologia. 132 (1): 96–101.
Bibcode:2002Oecol.132...96W (http://adsabs.harvard.edu/abs/2002Oecol.132...96W). doi:10.1007/s00442-002-0929-1 (h
ttps://doi.org/10.1007%2Fs00442-002-0929-1). PMID 28547281 (https://www.ncbi.nlm.nih.gov/pubmed/28547281).
26. Leakey, Richard; Lewin, Roger (1999) [1995]. "11 The modern elephant story". The sixth extinction: biodiversity and its
survival. Phoenix. pp. 216–217. ISBN 1-85799-473-6.
27. Gruber, Karl (26 September 2014). "Single keystone species may be the key to reef health" (http://www.australiangeogra
phic.com.au/news/2014/09/single-keystone-species-may-be-key-to-reef-health/). Australian Geographic.

Further reading
Caro, Tim (2010). Conservation by proxy: indicator, umbrella, k eystone, flagship, and other surrogate species.
Washington, DC: Island Press. ISBN 9781597261920.

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