keystone species

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 has allowed easier communication between ecologists and conservation policy-makers, it has been criticized for oversimplifying complex ecological systems.

History

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 1966 paper, ''Food Web Complexity and Species Diversity'', Paine had described such a system in Makah Bay in Washington.
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. 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 to co-exist that ochre starfish do not consume. When Paine removed the ochre starfish the mussels quickly outgrew the other species crowding them out. 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.

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.

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:

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 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.

The wolf, Yellowstone's 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 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.

The elimination of the