Keystone Species

Keystone species are those species whose importance to an ecosystems structure, composition, and function is disproportionately large relative to their abundance. These species can be of any life form, but they have in common an eect on their environment that is always greater than what can be expected based on their biomass . Well-studied examples include sea stars, beavers, bears, corals, elephants, and hummingbirds.

species whose importance to community and ecosystem structure, composition, and function is disproportionately large relative to its abundance is referred to as a keystone species. As the name implies, keystone species play key roles in ecosystems. They are distinguishable from dominant species, which also have large roles in ecosystems but solely by virtue of being abundant. Keystone species, even when rare, can drastically modify or create habitats and inuence the interaction between species in a community. An example of this can be beavers that create dams on rivers and streams, notably changing the previous habitat. Because keystone species are so important within communities, the removal of one often results in signicant loss of biodiversity. The concept of the keystone species, originally proposed by the US zoologist and University of Washington professor Robert T. Paine, was a transformative notion in biology. Keystone species can be any type of organism, including plants, animals, bacteria, or fungi. Ways to detect them vary, but an eective strategy to determine what is and what is not a keystone species is through removal experiments, in which a researcher excludes the suspected keystone species from some parts of a habitat and compares

areas with and without the species. This is how Paine conducted his groundbreaking 1966 experiment, in which he excluded the sea star (Pisaster ochraceous) from a stretch of shoreline in Makah Bay, Washington, in the United States. (The sea stars in the photograph above by Marjolijn Kaiser are in Oregon.) His comparison showed that the relatively uncommon sea star had a huge inuence on the tidal pool community. When the sea star was excluded from pools, the ecosystem lost almost half its resident diversity. Similar experiments involving other predators, such as bass, wolves, and jaguars, or herbivores, such as deer and elephants, have shown similar eects. One factor that can help de ne a keystone species is functional redundancy. In other words, if a species were to disappear from its community, are there other species that can ll its role? Some communities have more species redundancy than others, and therefore fewer keystone species (i.e., fewer species with fundamental roles in the ecosystems that cannot be replaced by other species). In a given community, the extinction of a keystone species will produce drastic changes. Th erefore, to maintain ecosystem functioning and services (like water purication and carbon sequestration), it may be critical to identify and protect those species.

Types of Keystone Species

There are many types of keystone species, and some of them have been thoroughly studied. Predators are typically dened as keystone species, because it takes only a few to regulate populations of other species in lower trophic levels. Many species that create or modify habitats, called ecosystem engineers, are also keystone species.

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AFRICAN ELEPHANTS AS A KEYSTONE SPECIES

The World Wildlife Fund (WWF) is one organization that works to protect the habitat of numerous species; one of its programs is the African Elephant Program, which aims to conserve forest and savanna elephant populations through projects and policies. The following is an excerpt from their website: African elephants once numbered in the millions across Africa, but by the mid-1980s their populations had been devastated by poaching. The status of the species now varies greatly across the continent. Some populations remain in danger due to poaching for meat and ivory, habitat loss, and conict with humans. Elephants are important because their future is tied to much of Africas rich biodiversity. Scientists consider African elephants to be keystone species as they help to maintain suitable habitats for many other species in savanna and forest ecosystems. Elephants directly inuence forest composition and density, and can alter the broader landscape. In tropical forests, elephants create clearings and gaps in the canopy that encourage tree regeneration. In the savannas, they can reduce bush cover to create an environment favorable to a mix of browsing and grazing animals. Many plant species also have evolved seeds that are dependent on passing through an elephants digestive tract before they can germinate; it is calculated that at least a third of tree species in west African forests rely on elephants in this way for distribution of their future generations.
Source: World Wildlife Fund (WWF). (2011). African elephant. Retrieved December 27, 2011, from http://www.worldwildlife.org/ species/nder/africanelephants/africanelephant.html

Th is is the case of the Canadian beaver and some species of African termites (in the genus Odontotermes), which build mounds that contain high levels of nutrients and thus can be colonized by many plant species. The presence of these nutritionally rich termite mounds can change an entire landscape. Large herbivores may also modify the habitat and the community through their feeding activity. One example of this is the African elephant (Loxodonta africana) in the savannas of southern Africa (discussed below). Also, many invasive species, which are exotic species that produce signicant changes in a native ecosystem, can be keystone species in the invaded ecosystem. The main types of keystone species and the effects if they become locally extinct are described below.

competitors. Th e e ects can cascade down to lower trophic levels; for example, the elimination of wolves leads to great increases in populations of deer, which in turn leads to destruction of certain plant species favored by deer.

Prey
When a prey species is removed from an ecosystem, this leaves fewer prey available to feed the predators. If the remaining prey species are more sensitive to the increased predation pressure, they might become rare or extinct within the ecosystem. Further loss of prey species could eventually lead to collapse of the predator population.

Predator
Like wolves and sea stars, some predator species play unique roles in their ecosystem by regulating the populations of their prey. Th eir extirpation can aect the abundance and presence of other predators and lead to the elimination of both prey and

Plant
Many herbivores, pollinators, and seed dispersers specialize in and depend on specic plant species for food or shelter. The extinction of that plant could lead to a population crash of these other dependent animal species.

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228 THE BERKSHIRE ENCYCLOPEDIA OF SUSTAINABILITY: ECOSYSTEM MANAGEMENT AND SUSTAINABILITY

Link
Some species, such as bees and hummingbirds, play key roles in the maintenance of plant populations by providing pollination services that maintain gene ow and secure plant fecundity. Therefore, absence of these pollinators can aect all species that depend on them directly or indirectly.

sh are spawning in upstream rivers. The bears feed and deposit salmon carcasses further inland, where they decompose and fertilize the riparian areas with nutrients that otherwise may not be incorporated into the local terrestrial ecosystem. Brown bears thus act as nutrient vectors that aect an entire ecosystem.

Beavers
Beavers (Castor canadensis) are the classic example of an ecosystem engineer because they create dams in rivers. These dams significantly alter the ux of nutrients and therefore the growth and abundance of local plants and animals. Their tremendous eect can be observed in Tierra del Fuego, an area of South America (in Chile and Argentina) where they have been introduced. Beaver are not native to South America, and no other native species has the ability to create dams in rivers, so beavers are altering the local ecosystems, replacing the slow-growing Nothofagus trees for meadows. This change in the structure of the ecosystem provides evidence on the fundamental role that this species has in its native and exotic range.

Ecosystem Engineer
Species that create or modify habitats, such as beavers (Castor canadensis), can strongly affect ecosystem nutrient cycling. Shifts in available nutrients can directly and indirectly aect animal or plant species that use the same habitat.

Examples of Keystone Species

Even prior to Paines seminal work and terminology, biologists had studied and dened many species as unique and necessary components of a given ecosystem, despite their rarity or low numbers. Many species have been widely studied in their role as keystone species.

Sea Stars
Th is is the quintessential example of a keystone species since Paines experiment in 1966. Sea stars are a key predator of mussels. The absence of sea stars can drastically impact ecosystems, including changes in diversity and abundance of other species in the habitat, aecting dierent trophic levels. For example, in the absence of sea stars, diversity was reduced from fteen species to only eight.

Corals
The compact ivory bush coral (Oculina arbuscula) is considered a keystone species because it creates new habitat. This coral species is endemic to the inshore and nearshore bottomland habitats of North and South Carolina in the United States. It is the only coral species found in this region. It forms complex branching colonies that provide shelter to over three hundred species of invertebrates that are known to live and complete much of their life cycle around the corals branches.

Bears
Brown bears (Ursus arctos) as a predator constitute a keystone species by regulating the population of their prey species, but they also have a keystone species role regarding the cycling of nutrients, primarily nitrogen, by incorporating nutrients from rivers into riparian ecosystems. These bears capture Pacic salmon when the
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African Elephants
In the savannas of Africa, elephants (Loxodonta africana) are destructive herbivores that consume large quantities

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of woody plants and often uproot, break, and destroy the trees and shrubs on which they feed. The decreased cover and density of woody vegetation favors the growth and production of grasses, rapidly changing an area from woods to savanna. Many other herbivores that feed on the grasses benet from the activities of elephants.

Hummingbirds
Hummingbirds are functionally important in many ecosystems by providing pollination services to many plant species. They exemplify a link keystone species. These highly specialized birds pollinate plants that have adapted to be pollinated only by these bird species. They serve as mobile links between plant populations in dierent landscapes, facilitating pollen movement (and therefore gene ow) often over considerable distances. Pollination triggers seed production and therefore plant population survival. An example of hummingbirds fundamental role can be found in southern South America in forests of Patagonia (in Argentina and Chile). The hummingbird species Sephanoides sephanoides pollinates nearly 20 percent of the local woody ora. These plant species could go extinct or become very rare if this bird disappeared, since no other species is adapted to pollinate them.

How Keystone Species Aect Ecosystems

Many ecosystem eects are attributed to keystone species. For example, in Paines original work, he reported that diversity of the tidal pool community decreased dramatically when the keystone species (the sea star) was removed. The predator preferred to forage on the most abundant mussel (Mytilus californianus), and when the predator was removed, the population of this mussel exploded in numbers that prevented many other species from existing in the tidal pools. So this keystone predator increases community diversity by foraging on the most abundant species, which benets less abundant prey species. Other studies on predator species have found similar results. Many dominant species depend on mutualism for their survival. Therefore, such mutualistic species can play a fundamental role in ecosystem function, and their removal can change drastically its dynamics, such as in food webs containing hummingbirds. Similarly, keystone species that are ecosystem engineers can, by creating or altering habitats, directly aect other species that need these areas to obtain food or shelter.

mean any species that has a very large impact on the studied ecosystem, no matter its abundance or biomass. This casual use of the term has led to attacks on the concept because it may be too vague and therefore meaningless. The phrase has even been freely and loosely borrowed outside biology; for example, it has migrated into business and economics, where keystone is used to describe organizations that enhance the business ecosystem by incorporating technological innovations, simplifying the connection between network participants, and/or providing a stable environment. Their importance to business ecosystems is such that their removal would lead to the collapse of the entire ecosystem. The concept of keystones species helps determine priority species for conservation and habitats in need of protection. Identifying keystone species, however, is not simple, owing to the complexity of nature and its temporal and spatial variability. A species can be keystone under certain circumstances (e.g., a dry year) but redundant in others (e.g., wet or normal years). Th is complicates the use and detection of keystone species. Furthermore, there may be inherent problems with basing conservation on keystone species. The concept implies that some species are more important than others in maintaining a given ecosystem, which suggests that more resources should be devoted to protecting them rather than other, more redundant species. Th is can be a problem given the complexity of natural systems and the well-known fact that interaction strengths change with space and time. Also, it is important to consider that keystone species may just be a human construct based on our limited observational and experimental capacities (i.e., the dierence between redundant and keystone species may not exist in nature). Therefore, conservation plans made around them may not be ideal.

The Future
Keystone species are a central concept in biology. The term is widely used in theoretical, applied, and conservation biology and also serves as a heuristic tool in ecology to explain food webs and ecosystem functioning. Keystone species are recognized as a concept to help understand ecosystem diversity and functioning. A bibliographic analysis using the academic search tool Web of Science revealed that the term is not only widely used, but also that the number of publications using it is trending upward. Th is indicates that the study of keystone species remains an active part of the biological sciences and likely will continue, especially in areas related to invasive species, ecosystem engineers, and biodiversity conservation. Although it may be problematic and controversial to base conservation plans on keystone species,

Problems with Other Denitions

The term keystone species has many denitions; and, in some areas of science, the term is more casually used to
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230 THE BERKSHIRE ENCYCLOPEDIA OF SUSTAINABILITY: ECOSYSTEM MANAGEMENT AND SUSTAINABILITY understanding what they are, how to detect them, and their full inuence on ecosystems may be key for understanding, conserving, and protecting nature. Th is is especially important for sustainability of ecosystem processes and services that can be fundamental for human well-being and the functioning of ecosystems at the global scale. Martin A. NUEZ and Romina D. DIMARCO The University of Tennessee See also Biodiversity; Biodiversity Hotspots; Charismatic Megafauna, Complexity Theory; Edge Eects; Food Webs; Hunting; Indicator Species; Outbreak Species; Plant-Animal Interactions; Refugia; Regime Shifts; Species Reintroduction; Wilderness Areas

FURTHER READING
K areiva, Peter, & L evin, Simon A. (2003) The importance of species: Perspectives on expendability and triage. Princeton, NJ: Princeton University Press. Mills, L. Scott; Soule, Michael E.; & Doak, Daniel F. (1993). The keystone-species concept in ecology and conservation. Bioscience, 43, 219224. Paine, Robert T. (1966). Food web complexity and species diversity. American Naturalist, 100, 6575. Paine, Robert. T. (1969). A note on trophic complexity and community stability. American Naturalist, 103, 9192. Simberlo, Daniel. (1998). Flagships, umbrellas, and keystones: Is single-species management pass in the landscape era? Biological Conservation, 83, 247257. Power, Mary E., et al. (1996). Challenges in the quest for keystones. Bioscience, 46, 609620.

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