Keystone species

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Not to be confused with foundation species.

The jaguar, a keystone, flagship, and umbrella species, and an apex

predator

A keystone species is a species which has a

disproportionately large effect on its natural environment
relative to its abundance, a concept introduced in 1969 by the
zoologist Robert T. Paine. Keystone species play a critical role
in maintaining the structure of an ecological community,
affecting many other organisms in an ecosystem 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 wolf, are also apex
predators.

The role that a keystone species plays in its ecosystem is

analogous to the role of a keystone in an arch. 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 biomass or productivity. It became
a popular concept in conservation biology, alongside flagship
and umbrella species. Although the concept is valued as a
descriptor for particularly strong inter-species interactions,
and it has allowed easier communication between ecologists
and conservation policy-makers, it has been criticized for
oversimplifying complex ecological systems.

Contents

1
History

2
Definitions

3
Predators
3.1
Sea otters and kelp forests

3.2
The wolf, Yellowstone's apex predator

3.3
Sea stars and other non-apex predators

4
Mutualists

5
Engineers

6
Limitations
 7
See also

8
References

9
Further reading

History[edit]

Ochre seastars (Pisaster ochraceus), a keystone predator

California mussels (Mytilus californianus), the seastar's prey

The concept of the keystone species was introduced in 1969

by the zoologist Robert T. Paine.[1][2] Paine developed the
concept to explain his observations and experiments on the
relationships between marine invertebrates of the intertidal
zone (between the high and low tide lines), including starfish
and mussels. He removed the starfish from an area, and
documented the effects on the ecosystem.[3] In his 1966
paper, Food Web Complexity and Species Diversity, Paine
had described such a system in Makah Bay in Washington.[4]
In his 1969 paper, Paine proposed the keystone 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 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.[5][6]

Definitions[edit]
A keystone species was defined by Paine as a species that
has a disproportionately large effect on its environment
relative to its abundance.[7] It has been defined operationally
by Davic in 2003 as "a strongly interacting species whose
top-down effect on species diversity and competition is large
relative to its biomass dominance within a functional group."[8]

A classic keystone species is a predator that prevents a

particular herbivorous species from eliminating dominant plant
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 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
weevil Euhrychiopsis lecontei is thought to have keystone
effects on aquatic plant diversity by foraging on nuisance
Eurasian watermilfoil in North American waters. [9] Similarly,
the wasp species Agelaia vicina 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.[7]

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 flagship species, indicator species, and umbrella
species. For example, the jaguar is a charismatic big cat
which meets all of these definitions:[10]

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
jaguars is an umbrella for many other species. ... the jaguar
[is] a keystone in subtropical and tropical America ...

— David Maehr et al, 2001[10]

Predators[edit]

Sea otters and kelp forests[edit]

Sea urchins like this purple sea urchin can damage kelp forests by
chewing through kelp holdfasts
 The sea otter is an important predator of sea urchins, making it a
keystone species for the 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 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 holdfasts of kelp 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
25,000.[11][12][13][14]

The wolf, Yellowstone's apex

predator[edit]

Riparian willow recovery at Blacktail Creek, Yellowstone National Park,

after reintroduction of wolves

Further information: Apex predator

Keystone predators may increase the biodiversity of
communities by preventing a single species from becoming
dominant. They can have a profound influence on the balance
of organisms in a particular ecosystem. Introduction or
removal of this 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.[15]

The elimination of the gray wolf from the Greater Yellowstone

Ecosystem had profound impacts on the trophic pyramid.
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
territory for grazing. 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.[16]

Sea stars and other non-apex

predators[edit]
As described by Paine in 1966, some sea stars (e.g., Pisaster
ochraceus) may prey on sea urchins, mussels, and other
shellfish 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.[17] The recent
onset of sea star wasting disease around the United States
has indirectly caused mussel populations to dominate in many
intertidal habitats.

These creatures need not be apex predators. Sea stars are

prey for sharks, rays, and sea anemones. Sea otters are prey
for orca.[18]

The jaguar, whose numbers in Central and South America

have been classified as near threatened, acts as a keystone
predator by its widely varied diet, helping to balance the
mammalian jungle ecosystem with its consumption of 87
different species of prey.[19] The lion is another keystone

species.[20]

Acorn banksia, Banksia prionotes, is the sole source of nectar for

important pollinators, honeyeaters.

Mutualists[edit]
Keystone mutualists are organisms that participate in mutually
beneficial interaction and the loss of which would have a
profound impact upon the ecosystem as a whole. For
example, in the Avon Wheatbelt region of Western Australia,
there is a period of each year when Banksia prionotes (acorn
banksia) is the sole source of nectar for honeyeaters, which
play an important role in pollination 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 frugivores such as the cassowary, which spreads
the seeds of many different trees, and some will not grow
unless they have been through a cassowary.[21][22]

Engineers[edit]

Prairie dog town. Drawing by Josiah Gregg, 1844

Further information: Ecosystem engineer and Niche

construction

A term used alongside keystone is ecosystem engineer.[5] In

North America, the prairie dog is an ecosystem engineer.
Prairie dog burrows provide the nesting areas for mountain
plovers and burrowing owls. Prairie dog tunnel systems also
help channel rainwater into the water table to prevent runoff
and erosion, and can also serve to change the composition of
the soil in a region by increasing aeration 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.[23] Grazing species such as
plains bison, pronghorn, and mule deer have shown a
proclivity for grazing on the same land used by prairie
dogs.[24]

Beaver dam, an animal construction which has a transformative effect on

the environment

The beaver 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 riparian 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 riverine forests. These dams have
been shown to be beneficial to a myriad of species including
amphibians, salmon, and song birds.[25]

In the African savanna, the larger herbivores, especially the

elephants, shape their environment. The elephants destroy
trees, making room for the grass species. Without these
animals, much of the savanna would turn into woodland.[26] In
the Amazon river basin, peccaries produce and maintain
wallows that are utilized by a wide variety of species.[27][28]
Australian studies have 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 severe strain.[29]

In the Serengeti, the presence of sufficient gnus in these

grasslands promote tree growth, which in turn reduces wildfire
likelihood. The documentary The Serengeti Rules documents
this in detail.[30]

Limitations[edit]
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
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 pollinators 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
sometimes have extremely strong interactions within a
particular ecosystem, but that does not automatically imply
that other ecosystems have a similar structure.[3]
See also[edit]
● Cultural keystone species
● Ecosystem service
● Foundation species
● Indigenous
● Introduced species

References[edit]
● ^
● Jump up to:
ab
● Paine, R. T. (1969). "A Note on Trophic Complexity and
Community Stability". The American Naturalist. 103 (929):
91–93. doi:10.1086/282586. JSTOR 2459472. S2CID
83780992.
● ^ "Keystone Species Hypothesis". University of Washington.
Archived from the originalon 2011-01-10. Retrieved
2011-02-03.
● ^
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ab
● 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. JSTOR
1312122.
● ^ Paine, R. T. (1966). "Food Web Complexity and Species
Diversity". The American Naturalist. 100 (910): 65–75.
doi:10.1086/282400. JSTOR 2459379. S2CID 85265656.
● ^
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ab
● 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. S2CID 11030284.
● ^ HHMI, BioInteractive. "Some Animals Are More Equal than
Others: Keystone Species and Trophic Cascades – HHMI
(2016)". Retrieved 6 June 2017.
● ^
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ab
● 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.
● ^ Davic, R. D. (2003). "Linking Keystone Species and
Functional Groups: A New Operational Definition of the
Keystone Species Concept". Conservation Ecology. Retrieved
2011-02-03.
● ^ 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.
● ^
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 ab
● Maehr, David; Noss, Reed F.; Larkin, Jeffery L. (2001).
Large Mammal Restoration: Ecological And Sociological
Challenges In The 21St Century. Island Press. p. 73. ISBN
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● ^ Szpak, Paul; Orchard, Trevor J.; Salomon, Anne K.; Gröcke,
Darren R. (2013). "Regional ecological variability and impact of
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● ^ Estes, James E.; Smith, Norman S.; Palmisano, John F.
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● ^ Cohn, J. P. (1998). "Understanding Sea Otters". BioScience.
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● ^ Brown, Tina M. (14 March 2013). "My Turn: Southeast
Alaska's sea otters: The restoration of an ecosystem". Juneau
Empire. Archived from the original on 23 April 2018. Retrieved
23 April 2018.
● ^ Botkin, D.; Keller, E. (2003). Environmental Science: Earth
as a living planet. John Wiley & Sons. p. 2. ISBN
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● ^ Ripple, William J.; Beschta, Robert L. (2004). "Wolves and
the Ecology of Fear: Can Predation Risk Structure
Ecosystems?". BioScience. Oxford University Press. 54 (8):
755.
doi:10.1641/0006-3568(2004)054[0755:WATEOF]2.0.CO;2.
● ^ Paine, R. T. (1966). "Food web complexity and species
diversity". American Naturalist. 100 (910): 65–75.
doi:10.1086/282400. JSTOR 2459379. S2CID 85265656.
● ^ 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.
doi:10.1126/science.282.5388.473. PMID 9774274.
● ^ Nowell, K.; Jackson, P., eds. (1996). Panthera onca. Wild
Cats, Status Survey and Conservation Action Plan. IUCN/SSC
Cat Specialist Group. pp. 118–122. ISBN 978-2-8317-0045-8.
● ^ Hale, Sarah L.; Koprowski, John L. (February 2018).
"Ecosystem-level effects of keystone species reintroduction: a
literature review". Restoration Ecology. 26 (3): 439–445.
doi:10.1111/rec.12684.
● ^ 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.
● ^ Walker, Brian (1995). "Conserving Biological Diversity
through Ecosystem Resilience". Conservation Biology. 9 (4):
747–752. doi:10.1046/j.1523-1739.1995.09040747.x.
● ^ "Prairie Dogs". Wildlife Species Guide. Nebraska Game and
Park Commission. Archived from the original on 19 August
2009. Retrieved 10 November 2013.
 ● ^ Rosmarino, Nicole (2007). "Associated Species : Prairie
Dogs are a Keystone Species of the Great Plains". Prairie Dog
Coalition. Archived from the original on 10 November 2013.
Retrieved 10 November 2013.
● ^ Wright, J. P.; Jones, C. G.; Flecker, A. S. (2002). "An
ecosystem engineer, the beaver, increases species richness at
the landscape scale". Oecologia. 132 (1): 96–101.
Bibcode:2002Oecol.132...96W.
doi:10.1007/s00442-002-0929-1. PMID 28547281. S2CID
5940275.
● ^ 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.
● ^
https://news.mongabay.com/2010/09/how-the-overlooked-pecc
ary-engineers-the-amazon-an-interview-with-harald-beck/
● ^
https://www.nationalgeographic.com/news/2014/9/140927-pec
cary-wallow-amazon-rainforest-camera-trap-biodiversity-scienc
e/
● ^ Gruber, Karl (26 September 2014). "Single keystone species
may be the key to reef health". Australian Geographic.
● ^ How Wildebeest Saved the Serengeti

Further reading[edit]
● Caro, Tim (2010). Conservation by proxy: indicator,
umbrella, keystone, flagship, and other surrogate
species. Washington, DC: Island Press. ISBN
9781597261920.
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