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Definition
The definition given here is somewhat qualitative in nature because there is not yet an accepted, rigorous definition.[3] An issue is how to measure both abundance and impact and at what point to draw the line. Abundance in this context can be measured by biomass or productivity, among other metrics. Impact is even harder to define. It has been suggested that such keystone predators can be identified in ecosystems by their biomass dominance within ecological functional groups, even though they may be relatively rare in relation to the ecosystem as a whole.[4]
The term is often misused in places where foundation species would be more appropriate. A keystone species is not simply one whose disappearance would alter the ecosystem. Removing any abundant species from any ecosystem will drastically alter that ecosystem by definition. What makes the concept of keystone species attractive to conservationists is that a species with a relatively tiny physical footprint can be critical to the health of its ecosystem.[3]
Examples
Without a consensus on its exact definition, we are left to illustrate the concept of keystone species with a list of examples.
A classic keystone species is a small predator that prevents a particular herbivorous species from eliminating dominant plant species. Since the prey numbers are low, the keystone predator numbers can be even lower 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 out-sized impact on ecosystem functions. One example is the weevil and its suggested keystone effects on aquatic plant species diversity by prey activities on nuisance Euransian Watermilfoil.[5]
Predators
Some starfish may perform this function by preying 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, while the urchin population annihilates coral reefs. In his classic 1966 paper, Dr. Robert Paine described such a system in Mukkaw Bay in Washington State.[6] This led to his 1969 paper where he proposed the keystone species concept.[7]
Similarly, sea otters in kelp forests keep sea urchins in check. Kelp roots are merely anchors, and not the vast nutrient gathering networks of land plants. Thus the urchins only need to eat the roots of the kelp, a tiny fraction of the plant's biomass, to remove it from the ecosystem.[8] [9]
These creatures need not be apex predators. Sea stars are prey for sharks, rays, and sea anemones. Sea otters are prey for orca.[10]
Engineers
In North America, the grizzly bear is a keystone species - not as a predator but as ecosystem engineers. They transfer nutrients from the oceanic ecosystem to the forest ecosystem. The first stage of the transfer is performed by salmon, rich in nitrogen and potassium, who swim up rivers, sometimes for hundreds of miles. The bears then capture the salmon and carry them onto dry land, dispersing nutrient-rich feces and partially-eaten carcasses. It has been estimated that the bears leave up to half of the salmon they harvest on the forest floor.citation needed
Another ecosystem engineering keystone species is the beaver, which transforms its territory from a stream to a pond or swamp.[11]
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.[12]
References
- ^ Paine, R.T. (1995). "A Conversation on Refining the Concept of Keystone Species". Conservation Biology 9 (4): 962–964. doi:.
- ^ Mills, L.S.; Soule, M.E.; Doak, D.F. (1993). "The Keystone-Species Concept in Ecology and Conservation". BioScience 43 (4): 219–224. doi:. Retrieved on 2007-10-04.
- ^ a b Power, M.E.; Tilman, D.; Estes, J.A.; Menge, B.A.; Bond, W.J.; Mills, L.S.; Daily, G.; Castilla, J.C.; Lubchenco, J.; Paine, R.T. (1996). "Challenges in the Quest for Keystones". BioScience 46 (8): 609–620. doi:. Retrieved on 2007-10-04.
- ^ Davic, R.D. (2003). "Linking Keystone Species and Functional Groups: A New Operational Definition of the Keystone Species Concept". Conservation Ecology 7 (1): r11. Retrieved on 2007-10-04.
- ^ Creed Jr, R.P. (2000). "Is there a new keystone species in North American lakes and rivers?". OIKOS 91 (2): 405. doi:.
- ^ Paine, R.T. (1966). "Food Web Complexity and Species Diversity". The American Naturalist 100 (910): 65–75. doi:. Retrieved on 2007-10-04.
- ^ Paine, R.T. (1969). "A Note on Trophic Complexity and Community Stability". The American Naturalist 103 (929): 91–93. doi:. Retrieved on 2007-10-04.
- ^ Estes, James E.; Norman S. Smith, John F. Palmisano (1978). "Sea otter predation and community organization in the Western Aleutian Islands, Alaska". Ecology 59 (4): 822–833. doi:. Retrieved on 2007-10-04.
- ^ Cohn, J.P. (1998). "Understanding Sea Otters". BioScience 48 (3): 151–155. doi:. Retrieved on 2007-10-04.
- ^ 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. doi:. PMID 9774274. Retrieved on 2007-10-04.
- ^ 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. doi:. Retrieved on 2007-10-04.
- ^ Leakey, Richard; Roger Lewin [1995] (1999). "11 The modern elephant story", The sixth extinction: biodiversity and its survival. London: Phoenix, pp. 216-217. ISBN 1-85799-473-6.
See also
External links
