Zoologist Robert T. Paine, who coined the term "keystone species," had an unorthodox way of doing his work. Instead of just observing the habitat of the Pisaster ochraceus sea star, Paine experimented by actually changing the habitat. Paine and his students from the University of Washington spent 25 years removing the sea stars from a tidal area on the coast of Tatoosh Island, Washington, in order to see what happened when they were gone. He was one of the first scientists in his field to experiment in nature in this manner.Nutrient VectorsKeystone species can sometimes be “nutrient vectors,” transferring nutrients from one habitat to another. Grizzly bears, for instance, prey on salmon. They can deposit salmon carcasses miles from rivers and streams. Salmon carcasses decompose and fertilize the soil with nutrients that may not be available from local terrestrial ecosystems.Keystone PreyKeystone prey are species that can maintain healthy populations despite being preyed upon. Wildebeests, prey for predators from lions to crocodiles of the African savanna, are an example of keystone prey.Keystone HostsPlants and other producers that provide food and shelter for keystone species are sometimes called keystone hosts. Kelp is a keystone host. Kelp forests provide stabilizing shelter for sea otters, and nutrient-rich food for their prey, such as fish and sea urchins.Keystone TrophicsKeystone species are often predators, but not always apex predators. Instead, they are usually secondary consumers. Sea stars, while voracious predators of mussels and barnacles, for example, are a prey species for sea anemones and fishes.A keystone species is an organism that helps define an entire ecosystem. Without its keystone species, the ecosystem would be dramatically different or cease to exist altogether.Keystone species have low functional redundancy. This means that if the species were to disappear from the ecosystem, no other species would be able to fill its ecological niche. The ecosystem would be forced to radically change, allowing new and possibly invasive species to populate the habitat.Any organism, from plants to fungi, may be a keystone species; they are not always the largest or most abundant species in an ecosystem. However, almost all examples of keystone species are animals that have a huge influence on food webs. The way these animals influence food webs varies from habitat to habitat.Carnivores, Herbivores, and MutualistsPredatorsA keystone species is often, but not always, a predator. Just a few predators can control the distribution and population of large numbers of prey species.The entire concept of keystone species was founded on research surrounding the influence of a marine predator on its environment. American zoology professor Robert T. Paine's research showed that removing a single species, the Pisaster ochraceus sea star, from a tidal plain on Tatoosh Island in the U.S. state of Washington, had a huge effect on the ecosystem. Pisaster ochraceus, commonly known as purple sea stars, are a major predator of mussels and barnacles on Tatoosh Island. With the sea stars gone, mussels took over the area and crowded out other species, including benthic algae that supported communities of sea snails, limpets, and bivalves. Lacking a keystone species, the tidal plain’s biodiversity was cut in half within a year.Another example of a predator acting as a keystone species is the presence of gray wolves in the Greater Yellowstone Ecosystem. The Greater Yellowstone Ecosystem (GYE) is an enormous and diverse temperate ecosystem stretching across the boundaries of the U.S. states of Montana, Wyoming, and Idaho. The GYE includes active geothermal basins, mountains, forests, meadows, and freshwater habitats.The elk, bison, rabbit, and bird species in the Greater Yellowstone Ecosystem are at least partly controlled by the presence of wolves. The feeding behavior of these prey species, as well as where they choose to make their nests and burrows, are largely a reaction to wolf activity. Scavenger species, such as vultures, are also controlled by the wolf activity.When the U.S. government designated land for Yellowstone National Park in the late 19th century, hundreds of wolves roamed the GYE, preying primarily on abundant herds of elk and bison. Fearing the wolves’ impact on those herds, as well as local livestock, governments at the local, state, and federal level worked to eradicate wolves from the GYE. The last remaining wolf pups in Yellowstone were killed in 1924.This started a top-down trophic cascade in the Greater Yellowstone Ecosystem. A trophic cascade describes changes in an ecosystem due to the addition or removal a predator. A top-down trophic cascade describes changes that result from the removal of an ecosystem’s top predator. (A bottom-up trophic cascade describes changes that result from the removal of a producer or primary consumer.)Lacking an apex predator, elk populations in Yellowstone exploded. Elk herds competed for food resources, and plants such as grasses, sedges, and reeds did not have time or space to grow. Overgrazing influenced the populations of other species, such as fish, beaver, and songbirds. These animals rely on plants and their products—roots, flowers, wood, seeds—for survival.The physical geography of the Greater Yellowstone Ecosystem was also impacted by the loss of wolves and subsequent elk overgrazing. Stream banks eroded as wetland plants failed to anchor valuable soil and sediments. Lake and river temperatures increased as trees and shrubs failed to provide shaded areas.Starting in the 1990s, the U.S. government began reintroducing wolves to the Greater Yellowstone Ecosystem. The results have been noteworthy. Elk populations have shrunk, willow heights have increased, and beaver and songbird populations have recovered.HerbivoresHerbivores can also be keystone species. Their consumption of plants helps control the physical and biological aspects of an ecosystem.In African savannas such as the Serengeti plains in Tanzania, elephants are a keystone species. Elephants eat shrubs and small trees, such as acacia, that grow on the savanna. Even if an acacia tree grows to a height of a meter or more, elephants are able to knock it over and uproot it. This feeding behavior keeps the savanna a grassland and not a forest or woodland.With elephants to control the tree population, grasses thrive and sustain grazing animals such as antelopes, wildebeests, and zebras. Smaller animals such as mice and shrews are able to burrow in the warm, dry soil of a savanna. Predators such as lions and hyenas depend on the savanna for prey.Keystone MutualistsKeystone mutualists are two or more species that engage in mutually beneficial interactions. A change in one species would impact the other, and change the entire ecosystem. Keystone mutualists are often pollinators, such as bees. Pollinators often maintain gene flow and dispersal throughout widespread ecosystems.In the woody grasslands of Patagonia (at the southern tip of South America) a species of hummingbird and indigenous plants act together as keystone mutualists. Local trees, shrubs, and flowering plants have evolved to only be pollinated by Sephanoides sephanoides, a hummingbird known as the green-backed firecrown. Green-backed firecrowns pollinate 20% of local plant species. In turn, these plants provide the sugary nectar that makes up most of the hummingbird’s diet.Pockets of the existing Patagonian habitat would collapse without green-backed firecrowns, because their functional redundancy is nearly zero—no other pollinator has adapted to pollinate these plants.Other Organisms Crucial to EcosystemsIn addition to keystone species, there are other categories of organisms crucial to their ecosystems' survival.Umbrella SpeciesUmbrella species are often conflated with keystone species. Both terms describe a single species on which many other species depend. The key distinction between umbrella species and keystone species is that the value of an umbrella species is tied to its geographic species range.Umbrella species have large habitat needs, and the requirements of that habitat impact many other species living there. Most umbrella species are migratory, and their range may include different habitat types.The identification of an umbrella species can be an important aspect for conservation. The minimum species range of an umbrella species is often the basis for establishing the size of a protected area.The Siberian tiger, an endangered species, is an umbrella species with a range of more than 1,000 kilometers (620 miles) in Russia’s far east, with territory stretching into China and North Korea. The species range includes heavily forested ecosystems in both temperate and boreal (subarctic) biomes. Populations of deer, boar, and moose are under the snowy “umbrella” of the Siberian tiger range.Foundation SpeciesFoundation species play a major role in creating or maintaining a habitat.Corals are a key example of a foundation species across many islands in the South Pacific Ocean. These tiny animals grow as a colony of thousands and even millions of individual polyps. The rocky exoskeletons of these polyps create enormous structures around islands: coral reefs.Coral reefs are one of the most vibrant and biologically diverse ecosystems on the planet. Microscopic plankton, as well as crustaceans, mollusks, sponges, fish, and marine reptiles and mammals are all part of healthy coral reef ecosystems.Coral reef ecosystems also contribute to the human geography of a region. Pummeled by waves and ocean currents, coral exoskeletons can experience bioerosion. These eroded fragments of coral (along with bony fragments of organisms such as foraminifera, mollusks, and crustaceans) create a soft sand known as coral sand. Coral sand beaches are among the most popular tourist destinations in the world.Ecosystem EngineersLike foundation species, ecosystem engineers contribute to the physical geography of their habitat. Ecosystem engineers modify, create, and maintain habitats.Ecosystem engineers modify their habitats through their own biology or by physically changing biotic and abiotic factors in the environment.Autogenic engineers modify their environment by modifying their own biology. Corals and trees are autogenic engineers. As they grow, they are a living part of the environment, providing food and shelter to other organisms. (The hard exoskeletons left as corals die continue to define and modify the ecosystem.)Allogenic engineers physically change their environment from one state to another. Beavers are a classic example of allogenic engineers. Beavers help maintain woodland ecosystems by thinning out older trees and allowing young saplings to grow. Converting these trees into timber for dams radically alters woodland meadows and streams, changing them into wetland habitats.Invasive species are often ecosystem engineers. Lacking natural predators or abiotic factors to constrain them, these introduced species modify the existing environment in ways that inhibit the growth of the indigenous ecosystem.Kudzu, the so-called “vine that ate the South,” is an invasive species of plant that modified the environment of the southeastern United States. Kudzu regularly outcompetes native species for space and nutrients. As it crowds out native species, kudzu limits the pollinators, insects, and bird species that inhabit an area.Indicator SpeciesAn indicator species describes an organism that is very sensitive to environmental changes in its ecosystem. Indicator species are almost immediately affected by changes to the ecosystem and can give early warning that a habitat is suffering.Changes associated with external influences such as water pollution, air pollution, or climate change first appear in indicator species. For this reason, indicator species are sometimes known as “sentinel species.”In the “nation’s estuary” of the Chesapeake Bay, oysters are an indicator species. Oysters and other bivalves are filter feeders, meaning they filter water as they strain it for food particles. Oysters filter nutrients, sediments, and pollutants that enter the bay through natural or anthropogenic sources. Oyster beds help protect fisheries, coastal habitats, and even benthic ecosystems. The health of oyster populations in the Chesapeake, therefore, is used to indicate the health of the entire ecosystem.Flagship SpeciesA flagship species acts as a symbol for an environmental habitat, movement, campaign, or issue. They can be mascots for entire ecosystems.The identification of a flagship species relies heavily on the social, cultural, and economic value of a species. They are often “charismatic megafauna,”—large animals with popular appeal due to their appearance or cultural significance. Flagship species may or may not be keystone or indicator species.Flagship species can sometimes be symbols of general ideas about conservation, not representatives of specific ecosystems. However, specific issues are often associated with a specific animal. The movement to end seal hunting in the Arctic found its flagship species in the juvenile harp seal. Polar bears are the unchallenged flagship species associated with climate change.The giant panda is perhaps the most familiar flagship species. Pandas are the global symbol of endangered species and the value of captive breeding.
Term Part of Speech Definition Encyclopedic Entry abiotic Adjective
characterized by the absence of life or living organisms
in large amounts.
tree or shrub that is often thorny.
air pollution Noun
harmful chemicals in the atmosphere.
Encyclopedic Entry: air pollution algae Plural Noun
(singular: alga) diverse group of aquatic organisms, the largest of which are seaweeds.
allogenic engineer Noun
organism that changes biotic and/or abiotic resources from one physical state to another.
anthropogenic source Noun
caused by people.
apex predator Noun
species at the top of the food chain, with no predators of its own. Also called an alpha predator or top predator.
region at Earth's extreme north, encompassed by the Arctic Circle.
Encyclopedic Entry: Arctic autogenic engineer Noun
organism that modifies its environment by modifying its own biology.
a slope of land adjoining a body of water, or a large elevated area of the sea floor.
body of water partially surrounded by land, usually with a wide mouth to a larger body of water.
Encyclopedic Entry: bay benthic Adjective
having to do with the bottom of a deep body of water.
all the different kinds of living organisms within a given area.
Encyclopedic Entry: biodiversity bioerosion Noun
the process in which a living organism wears away at rock or another hard substance.
study of living things.
area of the planet which can be classified according to the plant and animal life in it.
Encyclopedic Entry: biome biotic Adjective
having to do with living or once-living organisms.
type of animal (mollusk) with two shells hinged together, such as a clam or mussel.
boreal forest Noun
land covered by evergreen trees in cool, northern latitudes. Also called taiga.
bottom-up trophic cascade Noun
ecological phenomenon in which a producer or primary consumer is removed from the environment.
small hole or tunnel used for shelter.
captive breeding Noun
reproduction of rare species controlled by humans in a closed environment, such as a zoo.
to stop or end.
charismatic megafauna Noun
large animals with popular appeal due to their appearance or cultural significance.
climate change Noun
gradual changes in all the interconnected weather elements on our planet.
Encyclopedic Entry: climate change collapse Verb
to fall apart completely.
group of one species of organism living close together.
to merge or fuse into one entity.
management of a natural resource to prevent exploitation, destruction, or neglect.
Encyclopedic Entry: conservation coral reef Noun
rocky ocean features made up of millions of coral skeletons.
type of animal (an arthropod) with a hard shell and segmented body that usually lives in the water.
learned behavior of people, including their languages, belief systems, social structures, institutions, and material goods.
structure built across a river or other waterway to control the flow of water.
to name or single out.
foods eaten by a specific group of people or other organisms.
Encyclopedic Entry: diet dispersal Noun
spread of something to a new area.
the way something is spread out over an area.
Encyclopedic Entry: distribution economic Adjective
having to do with money.
community and interactions of living and nonliving things in an area.
Encyclopedic Entry: ecosystem ecosystem engineer Noun
organism that creates, modifies, maintains or destroys a habitat.
endangered species Noun
organism threatened with extinction.
Encyclopedic Entry: endangered species environment Noun
conditions that surround and influence an organism or community.
to destroy or remove.
mouth of a river where the river's current meets the sea's tide.
Encyclopedic Entry: estuary exoskeleton Noun
the hard external shell or covering of some animals.
feeding behavior Noun
methods by which an organism obtains food and eats.
filter feeder Noun
aquatic animal that strains nutrients from water.
industry or occupation of harvesting fish, either in the wild or through aquaculture.
flagship species Noun
organism that serves as a symbol for an environmental habitat, movement, campaign, or issue.
food web Noun
all related food chains in an ecosystem. Also called a food cycle.
Encyclopedic Entry: food web foraminifera Plural Noun
(singular: foraminifer.) Type of microscopic organism (protist) that forms a shell and lives in marine or salty conditions.
ecosystem filled with trees and underbrush.
foundation species Noun
species that creates or maintains an ecosystem.
functional redundancy Noun
characteristic of species whose contributions to an ecosystem are matched by other species.
gene flow Noun
movement and exchange of genes between interbreeding populations.
geothermal energy Noun
heat energy generated within the Earth.
system or order of a nation, state, or other political unit.
ecosystem with large, flat areas of grasses.
grazing animal Noun
animal that feeds on grasses, trees, and shrubs.
environment where an organism lives throughout the year or for shorter periods of time.
Encyclopedic Entry: habitat herbivore Noun
organism that eats mainly plants and other producers.
Encyclopedic Entry: herbivore herd Noun
group of animals.
human geography Noun
the study of the way human communities and systems interact with their environment.
indicator species Noun
organism that is very sensitive to changes in its environment. Also called a sentinel species.
characteristic to or of a specific place.
to slow or prevent.
introduced species Noun
a species that does not naturally occur in an area. Also called alien, exotic, or non-native species.
invasive species Noun
type of plant or animal that is not indigenous to a particular area and causes economic or environmental harm.
Encyclopedic Entry: invasive species keystone mutualists Noun
organisms that participate in mutually beneficial interactions, the loss of which would have a profound impact on the ecosystem.
keystone species Noun
organism that has a major influence on the way its ecosystem works.
Encyclopedic Entry: keystone species livestock noun, plural noun
animals raised for sale and profit.
animal with hair that gives birth to live offspring. Female mammals produce milk to feed their offspring.
having to do with the ocean.
animal, person, or thing representing a group of people or organization.
organisms that travel from one place to another at predictable times of the year.
to change or alter.
large phylum of invertebrate animal, all possessing a mantle with a significant cavity used for breathing and excretion, a radula (except for bivalves), and the structure of the nervous system.
native species Noun
species that occur naturally in an area or habitat. Also called indigenous species.
sweet plant material that attracts pollinators.
protected area built by birds to hatch their eggs and raise their young.
role and space of a species within an ecosystem.
substance an organism needs for energy, growth, and life.
Encyclopedic Entry: nutrient overgrazing Noun
process of too many animals feeding on one area of pasture or grassland.
large plateau in southern South America, stretching from the Andes Mountains to the Atlantic Ocean.
physical geography Noun
study of the natural features and processes of the Earth.
flat, smooth area at a low elevation.
Encyclopedic Entry: plain plankton Plural Noun
(singular: plankton) microscopic aquatic organisms.
organism that produces its own food through photosynthesis and whose cells have walls.
to transfer pollen from one part of a flower (the anther) to another (the stigma).
animal, object, or force such as wind that transfers pollen from one plant to another, allowing seeds to develop.
chemical or other substance that harms a natural resource.
a type of animal with a fixed base, a tubelike body, and tentacles for catching prey.
total number of people or organisms in a particular area.
animal that hunts other animals for food.
animal that is hunted and eaten by other animals.
primary consumer Noun
organism that eats plants or other autotrophs.
organism on the food chain that can produce its own energy and nutrients. Also called an autotroph.
highest-ranking teacher at a college or university.
to strike violently and repeatedly.
a ridge of rocks, coral, or sand rising from the ocean floor all the way to or near the ocean's surface.
Encyclopedic Entry: reef resource Noun
available supply of materials, goods, or services. Resources can be natural or human.
to wander or travel over a wide area without a specific destination.
type of tropical grassland with scattered trees.
organism that eats dead or rotting biomass, such as animal flesh or plant material.
Encyclopedic Entry: scavenger sediment Noun
solid material transported and deposited by water, ice, and wind.
Encyclopedic Entry: sediment shelter Noun
structure that protects people or other organisms from weather and other dangers.
type of plant, smaller than a tree but having woody branches.
having to do with a community or other group of organized people.
top layer of the Earth's surface where plants can grow.
species range Noun
native, geographic area in which an organism can be found. Range also refers to the geographic distribution of a particular species.
Encyclopedic Entry: species range subarctic Noun
region just south of the Arctic Circle.
something used to represent something else.
land an animal, human, or government protects from intruders.
to develop and be successful.
tidal plain Noun
large, flat area where mud and sediment are deposited by ocean tides. Also called tidal flat or mudflat.
wood in an unfinished form, either trees or logs.
top-down trophic cascade Noun
ecological phenomenon in which a top predator is removed from the environment.
person who travels for pleasure.
trophic cascade Noun
ecological phenomenon triggered by the addition or removal of predators from an environment.
umbrella species Noun
large, usually migratory species on which other species in an ecosystem depend.
to tear or remove a tree or other plant by the roots.
water pollution Noun
introduction of harmful materials into a body of water.
area of land covered by shallow water or saturated by water.
Encyclopedic Entry: wetland woodland Noun
land covered with trees, usually less dense than a forest.
Encyclopedic Entry: woodland zoology Noun
the study of animals.