thumb|The [[jaguar: a keystone, flagship, and
umbrella species, and an apex predator ]]
thumb|The [[beaver: a keystone species, and habitat creator, responsible for ecological effects such as the creation of lakes, canals and wetlands irrigating large forests and creating ecosystems ]]
A keystone species is a species that has a disproportionately large effect on its natural environment relative to its abundance. The concept was 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 and lion, 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 has allowed easier communication between ecologists and conservation policy-makers, it has been criticized for oversimplifying complex ecological systems.
History
thumb|Keystone species will disrupt the environment if removed, And when kept they regulate the environment.
The concept of the keystone species was introduced in 1969 by zoologist Robert T. Paine. 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.
In his 1969 paper, Paine proposed the keystone species concept, using Pisaster ochraceus, a species of starfish generally known as ochre starfish, and Mytilus californianus, a species of mussel, as a primary example.
Definitions
A keystone species was defined by Paine as a species that has a disproportionately large effect on its environment relative to its abundance. 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."
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. 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.