A keystone species is a species that 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.
The concept of the keystone species was introduced in 1969 by 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 generally known as ochre starfish, and Mytilus californianus , a species of mussel, as a primary example. [1] 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, that ochre starfish do not consume, to co-exist. 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. [5] [6]
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]
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]
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. [15]
The elimination of the gray wolf from the Greater Yellowstone Ecosystem had profound impacts on the trophic pyramid. [16] 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 grazing territory. 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. [17]
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. [18] If the sea star is removed from the ecosystem, the mussel population explodes uncontrollably, driving out most other species. [19] 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. [20]
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. [21] The lion is another keystone species. [22]
Keystone mutualists are organisms that participate in mutually beneficial interaction, 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. Some seeds will not grow unless they have been through a cassowary. [23] [24]
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. [25] Grazing species such as plains bison, which is another keystone species, the pronghorn, and the mule deer have shown a proclivity for grazing on the same land used by prairie dogs. [26]
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. [27]
In the African savanna, the larger herbivores, especially the elephants, shape their environment. The elephants destroy trees, making room for the grass species and creating habitat for various small animal species. [28] [29] Without these animals, much of the savanna would turn into woodland. [30] In the Amazon river basin, peccaries produce and maintain wallows that are utilized by a wide variety of species. [31] [32] 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. [33]
In the Serengeti, the presence of sufficient gnus in these grasslands reduces wildfire likelihood, which in turn promotes tree growth. The documentary The Serengeti Rules documents this in detail. [34]
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]
Ecology is the natural science of the relationships among living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps with the closely related sciences of biogeography, evolutionary biology, genetics, ethology, and natural history.
Theoretical ecology is the scientific discipline devoted to the study of ecological systems using theoretical methods such as simple conceptual models, mathematical models, computational simulations, and advanced data analysis. Effective models improve understanding of the natural world by revealing how the dynamics of species populations are often based on fundamental biological conditions and processes. Further, the field aims to unify a diverse range of empirical observations by assuming that common, mechanistic processes generate observable phenomena across species and ecological environments. Based on biologically realistic assumptions, theoretical ecologists are able to uncover novel, non-intuitive insights about natural processes. Theoretical results are often verified by empirical and observational studies, revealing the power of theoretical methods in both predicting and understanding the noisy, diverse biological world.
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation and parasitoidism. It is distinct from scavenging on dead prey, though many predators also scavenge; it overlaps with herbivory, as seed predators and destructive frugivores are predators.
Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea. Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish are also known as asteroids due to being in the class Asteroidea. About 1,900 species of starfish live on the seabed in all the world's oceans, from warm, tropical zones to frigid, polar regions. They are found from the intertidal zone down to abyssal depths, at 6,000 m (20,000 ft) below the surface.
An ecosystem engineer is any species that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on species richness and landscape-level heterogeneity of an area. As a result, ecosystem engineers are important for maintaining the health and stability of the environment they are living in. Since all organisms impact the environment they live in one way or another, it has been proposed that the term "ecosystem engineers" be used only for keystone species whose behavior very strongly affects other organisms.
An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own.
Intertidal ecology is the study of intertidal ecosystems, where organisms live between the low and high tide lines. At low tide, the intertidal is exposed whereas at high tide, the intertidal is underwater. Intertidal ecologists therefore study the interactions between intertidal organisms and their environment, as well as between different species of intertidal organisms within a particular intertidal community. The most important environmental and species interactions may vary based on the type of intertidal community being studied, the broadest of classifications being based on substrates—rocky shore and soft bottom communities.
Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce the abundance, or alter the behavior of their prey, thereby releasing the next lower trophic level from predation.
The California mussel is a large edible mussel, a marine bivalve mollusk in the family Mytilidae.
In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological community, or life assemblage. The term community has a variety of uses. In its simplest form it refers to groups of organisms in a specific place or time, for example, "the fish community of Lake Ontario before industrialization".
The mesopredator release hypothesis is an ecological theory used to describe the interrelated population dynamics between apex predators and mesopredators within an ecosystem, such that a collapsing population of the former results in dramatically increased populations of the latter. This hypothesis describes the phenomenon of trophic cascade in specific terrestrial communities.
Ecological extinction is "the reduction of a species to such low abundance that, although it is still present in the community, it no longer interacts significantly with other species".
Rewilding is a form of ecological restoration aimed at increasing biodiversity and restoring natural processes. It differs from other forms of ecological restoration in that rewilding aspires to reduce human influence on ecosystems. It is also distinct from other forms of restoration in that, while it places emphasis on recovering geographically specific sets of ecological interactions and functions that would have maintained ecosystems prior to human influence, rewilding is open to novel or emerging ecosystems which encompass new species and new interactions.
Pisaster ochraceus, generally known as the purple sea star, ochre sea star, or ochre starfish, is a common seastar found among the waters of the Pacific Ocean. Identified as a keystone species, P. ochraceus is considered an important indicator for the health of the intertidal zone.
Robert Treat "Bob" Paine III was an American ecologist who spent most of his career at the University of Washington. Paine coined the keystone species concept to explain the relationship between Pisaster ochraceus, a species of starfish, and Mytilus californianus, a species of mussel.
Intraguild predation, or IGP, is the killing and sometimes eating of a potential competitor of a different species. This interaction represents a combination of predation and competition, because both species rely on the same prey resources and also benefit from preying upon one another. Intraguild predation is common in nature and can be asymmetrical, in which one species feeds upon the other, or symmetrical, in which both species prey upon each other. Because the dominant intraguild predator gains the dual benefits of feeding and eliminating a potential competitor, IGP interactions can have considerable effects on the structure of ecological communities.
Stichaster australis, the reef starfish, is a species of starfish found in the shallow waters of the rocky intertidal of New Zealand. Typically, the animal is endemic to the west coast shores of the North and South Islands, where wave action is increased. They do not usually inhabit ecosystems that have reduced wave action and calm conditions as they prefer a higher-energy environment. These marine invertebrates range in color from pink to purple, but can also be orange. They typically have eleven arms, but sometimes they may have either ten or twelve. As full-grown adults, they are 8 to 10 cm in diameter.
A refuge is a concept in ecology, in which an organism obtains protection from predation by hiding in an area where it is inaccessible or cannot easily be found. Due to population dynamics, when refuges are available, populations of both predators and prey are significantly higher, and significantly more species can be supported in an area.
Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers is larger than the biomass of primary producers. This happens because the ocean's primary producers are tiny phytoplankton which grow and reproduce rapidly, so a small mass can have a fast rate of primary production. In contrast, many significant terrestrial primary producers, such as mature forests, grow and reproduce slowly, so a much larger mass is needed to achieve the same rate of primary production.
The ecology of fear is a conceptual framework describing the psychological impact that predator-induced stress experienced by animals has on populations and ecosystems. Within ecology, the impact of predators has been traditionally viewed as limited to the animals that they directly kill, while the ecology of fear advances evidence that predators may have a far more substantial impact on the individuals that they predate, reducing fecundity, survival and population sizes. To avoid being killed, animals that are preyed upon will employ anti-predator defenses which aid survival but may carry substantial costs.
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