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• Ecology: Definition, Types, Importance & Examples

Ecological Niche: Definition, Types, Importance & Examples

Ecology is the study of the interactions between organisms and their environments, which comprise an ecosystem. The places organisms live in are called habitats.

An ecological niche , in contrast, is the ecological role an organism plays within its habitat.

Ecological Niche Definition

Several branches of ecology have adopted the concept of the ecological niche .

The ecological niche describes how a species interacts within an ecosystem. The niche of a species depends on both biotic and abiotic factors, which affect the ability of a species to survive and endure.

Biotic factors affecting a species' niche include food availability and predators. Abiotic factors affecting ecological niche include temperature, landscape characteristics, soil nutrients, light and other non-living factors.

An example of an ecological niche is that of the dung beetle. The dung beetle, as its name suggests, consumes dung both in larval and adult form. Dung beetles store dung balls in burrows, and females lay eggs within them.

This allows hatched larvae immediate access to food. The dung beetle in turn influences the surrounding environment by aerating soil and rereleasing beneficial nutrients. Therefore, the dung beetle performs a unique role in its environment.

The definition of a niche has changed since it was first introduced. A field biologist named Joseph Grinnell took the basic concept of the niche and further developed it, claiming that a niche distinguished between different species that occupied the same space. In other words, only one species could have a particular niche. He was influenced by species distribution.

Types of Ecological Niches

Ecologist Charles Elton’s definition of niche focused on the role of a species, such as its trophic role. His tenets emphasized more on community similarity and less on competition .

In 1957, Zoologist G. Evelyn Hutchinson provided a sort of compromise of these trains of thought. Hutchinson described two forms of niche. The fundamental niche focused on the conditions in which a species could exist with no ecological interactions. The realized niche , in contrast, considered the population’s existence in the presence of interactions, or competition.

The adoption of the ecological niche concept has allowed ecologists to understand the roles of species in ecosystems .

Importance of Ecological Niches

Ecologists use the concept of the ecological niche to help understand how communities relate to environmental conditions, fitness, trait evolution and predator-prey interactions in communities. This becomes ever more important as climate change affects community ecology .

Ecological niches allow species to exist in their environment. Under the right conditions, the species will thrive and play a unique role. Without the ecological niches, there would be less biodiversity, and the ecosystem would not be in balance.

Interspecies competition: Ecologists refer to coexistence when describing ecological niches. Two competing species cannot exist in one ecological niche. This is due to limited resources.

Competition affects the fitness of species, and can lead to evolutionary changes. An example of interspecies competition is an animal that forages for pollen or nectar from a specific plant species, competing with other such animals.

In the case of some species of ants, the insects will compete for nests and prey as well as water and food.

Competitive exclusion principle: Ecologists use the competitive exclusion principle to help understand how species coexist. The competitive exclusion principle dictates that two species cannot exist in the same ecological niche. This is due to competition for resources in a habitat.

Early champions of the competitive exclusion principle were Joseph Grinnell, T. I. Storer, Georgy Gause and Garrett Hardin in the early and mid 20th century.

Competition in a niche either leads each species to specialize in a different way, so as not to use the same resources, or leads one of the competing species to become extinct. This is another way of looking at natural selection. There are two theories used to address competitive exclusion.

In R* Theory , multiple species cannot exist with the same resources unless they differentiate their niches. When resource density is at its lowest, those species populations most limited by the resource will be competitively excluded.

In P* Theory , consumers can exist at high density due to having shared enemies.

Competition plays out even at the microbial level. For example, if Paramecium aurelia and Paramecium caudatum are grown together, they will compete for resources. P. aurelia will eventually overtake P. caudatum and cause it to go extinct.

Overlapping Niches/Resource Partitioning

Given the fact that organisms cannot exist in a bubble and must therefore naturally interact with other species, occasionally niches can overlap. To avoid competitive exclusion, similar species can change over time to use different resources.

In other cases, they can exist in the same area but use resources at different times. This scenario is called resource partitioning .

Resource partitioning: Partitioning means separating. Simply put, species can use their resources in ways that reduce depletion. This allows the species to coexist and even evolve.

An example of resource partitioning is that of lizards like anoles, which used different parts of their overlapping habitats in different ways. Some of the anoles might live on the forest floor; others might live high in the canopy or along the trunk and branches. Still other anoles might move away from plant environments and live in deserts or near oceans.

Another example would be dolphins and seals, which eat similar species of fish. However, their home ranges differ, allowing for a partitioning of resources.

Another example would be Darwin’s finches, which specialized their beak shapes over over time in their evolution. In this way, they were able to use their resources in different ways.

Examples of Ecological Niches

Several examples of ecological niches exist in various ecosystems.

For example, in the jack pine forest of Michigan, the Kirtland’s warbler occupies an area ideally suited for the bird. The birds prefer nesting on the ground between the trees, not in them, among small undergrowth.

But the jack pine tree must be only up to eight years old and around 5 feet tall. Once the tree ages or grows taller, the Kirtland’s warbler will not thrive. These highly specialized kinds of niches can be put at great risk due to human development.

Desert plants such as succulents adapted to arid ecological niches by storing water in their leaves and growing long roots. Unlike most plants, succulents open up their stomata only at night so as to reduce water loss from scorching daytime heat.

Thermophiles are organisms that thrive in extreme ecological niches such as thermal vents with high temperatures.

Channel Islands Ecosystem

In Southern California, mere miles away from one of the most populous areas of human settlement in the United States, the chain of islands known as the Channel Islands provides a fascinating ecosystem for studying ecological niches.

Nicknamed the “Galapagos of North America,” this delicate ecosystem plays host to numerous plants and animals. The islands vary in size and shape, and they provide unique habitats for various animals and plants.

Birds: Several birds call the Channel Islands home, and despite their overlap they have each managed to occupy special ecological niches on the islands. For example, the California brown pelican nests on Anacapa Island by the thousands. The island scrub jay is unique to the Channel Islands.

Fish: Over 2,000 fish species live in the waters around these islands. The kelp beds under the ocean provide habitat for both fish and mammals.

The Channel Islands have suffered from the introduction of invasive species by European settlers, as well as from pollutants such as DDT. Bald eagles disappeared, and taking their place, golden eagles made a home. However, bald eagles have been reintroduced to the islands. Peregrine falcons underwent a similar crisis and are making a comeback.

Native mammals: Four native mammals reside in the Channel Islands: the island fox, harvest mouse, island deer mouse and spotted skunk. The fox and the deer mouse in turn have subspecies on separate islands; each island therefore hosts separate niches.

The island spotted skunk prefers habitat of different types depending on the island it lives on. On Santa Rosa Island, the skunk favors canyons, riparian areas and open woodlands. In contrast, on Santa Cruz Island, spotted skunks prefer open grassland mixed with chaparral. They play the role of predator on both islands.

The island spotted skunk and the island fox are competitors for resources on the islands. However, the spotted skunks are more carnivorous, and they are nocturnal. So in this manner, they are able to coexist in overlapping niches . This is another example of resource partitioning.

The island fox nearly went extinct. Recovery efforts have brought the species back.

Reptiles and Amphibians : The highly specialized niches extend to reptiles and amphibians. There is one salamander species, one frog species, two non-venomous snake species and four lizard species. And yet they are not found on every island. For example, only three islands play host to the island night lizard.

Bats also occupy niches on the islands of Santa Cruz and Santa Rosa, working as both pollinators and consumers of insects. Santa Cruz Island is a home for Townsend’s big-eared bats.

Today the islands are recovering. They now comprise Channel Islands National Park and the Channel Islands National Marine Sanctuary, and ecologists continue to monitor the many creatures that call the islands home.

Niche Construction Theory

Ecologists more recently have focused on niche construction theory , which describes how organisms modify their environments to make them better suited as niches. Examples of this include making burrows, building nests, creating shade, building beaver dams and other methods in which organisms alter their surroundings to suit their needs.

Niche construction arose from biologist John Odling-Smee. Odling-Smee argued that niche construction should be considered a process of evolution, a form of “ecological inheritance” passed on to descendents rather than a genetic inheritance.

There are four core principles behind niche construction theory:

• One involves non-random modification of the environment by a species, helping aid their evolution. 
• Second, the “ecological” inheritance alters evolution due to parents passing on the altering skills to their offspring.
• Third, new characteristics that are adopted become evolutionarily significant. The environments are affected systematically. 
• Fourth, what was considered adaptation is essentially the result of organisms making their environments more complementary via niche construction .

An example would be a seabird’s feces that lead to plant fertilization and a transition from scrubland to grassland. This is not an intentional adaptation, but it has brought implications for evolution. The seabird would therefore have significantly modified the environment.

Other modifications to the environment must affect the selection pressures on an organism. The selective feedback is unrelated to genes.

Examples of Niche Construction

More examples of niche construction include nesting and burrowing animals, yeast that modify themselves to attract more fruit flies and the modification of shells by hermit crabs. Even by moving around, organisms can affect the environment, in turn influencing gene flow in a population.

This is seen on a grand scale with humans, who have so altered the environment to suit their needs that it has led to worldwide consequences. This certainly can be evidenced by the transition from hunter-gatherers to agrarian cultures, which altered the landscape in order to raise food sources. In turn, humans altered animals for domestication.

Ecological niches offer rich potential knowledge for understanding how species interact with environmental variables. Ecologists can use this information to learn more about how to manage species and to conserve them, and how to plan for future development as well.

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About the Author

J. Dianne Dotson is a science writer with a degree in zoology/ecology and evolutionary biology. She spent nine years working in laboratory and clinical research. A lifelong writer, Dianne is also a content manager and science fiction and fantasy novelist. Dianne features science as well as writing topics on her website, jdiannedotson.com.

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• Published: 12 August 2009

The importance of niches for the maintenance of species diversity

• Jonathan M. Levine 1 &
• Janneke HilleRisLambers 2  

Nature volume  461 ,  pages 254–257 ( 2009 ) Cite this article

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Ecological communities characteristically contain a wide diversity of species with important functional, economic and aesthetic value. Ecologists have long questioned how this diversity is maintained 1 , 2 , 3 . Classic theory shows that stable coexistence requires competitors to differ in their niches 4 , 5 , 6 ; this has motivated numerous investigations of ecological differences presumed to maintain diversity 3 , 6 , 7 , 8 . That niche differences are key to coexistence, however, has recently been challenged by the neutral theory of biodiversity, which explains coexistence with the equivalence of competitors 9 . The ensuing controversy has motivated calls for a better understanding of the collective importance of niche differences for the diversity observed in ecological communities 10 , 11 . Here we integrate theory and experimentation to show that niche differences collectively stabilize the dynamics of experimental communities of serpentine annual plants. We used field-parameterized population models to develop a null expectation for community dynamics without the stabilizing effects of niche differences. The population growth rates predicted by this null model varied by several orders of magnitude between species, which is sufficient for rapid competitive exclusion. Moreover, after two generations of community change in the field, Shannon diversity was over 50 per cent greater in communities stabilized by niche differences relative to those exhibiting dynamics predicted by the null model. Finally, in an experiment manipulating species’ relative abundances, population growth rates increased when species became rare—the demographic signature of niche differences. Our work thus provides strong evidence that species differences have a critical role in stabilizing species diversity.

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Acknowledgements

This work was supported by US National Science Foundation grants 0743365 and 0743183, and a fellowship from the David and Lucile Packard Foundation. P. Adler, C. Briggs, B. Cardinale, P. Chesson, M. Levine, D. Murrell and the Levine and HilleRisLambers laboratories provided comments on the manuscript. C. Cowan, R. Harris and C. Peters conducted the field work.

Author Contributions J.M.L. and J.H. jointly conducted the project, analysed the data and prepared the manuscript.

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Correspondence to Jonathan M. Levine or Janneke HilleRisLambers .

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Levine, J., HilleRisLambers, J. The importance of niches for the maintenance of species diversity. Nature 461 , 254–257 (2009). https://doi.org/10.1038/nature08251

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AP®︎/College Biology

Course: ap®︎/college biology   >   unit 8.

• Interactions between populations
• Interactions in communities

Niches & competition

• Predator-prey cycles
• Tropical rainforest diversity
• Community structure
• Simpson's index of diversity
• Community ecology

Key points:

• In interspecies competition , two species use the same limited resource. Competition has a negative effect on both of the species (-/- interaction).
• A species' niche is basically its ecological role, which is defined by the set of conditions, resources, and interactions it needs (or can make use of).
• The competitive exclusion principle says that two species can't coexist if they occupy exactly the same niche (competing for identical resources).
• Two species whose niches overlap may evolve by natural selection to have more distinct niches, resulting in resource partitioning .

Introduction

The niche concept, niche as an n -dimensional hypervolume, competitive exclusion principle, resource partitioning, attribution:.

• " Community ecology ," by Robert Bear and David Rintoul, CC BY 4.0 . Download the original article for free at http://cnx.org/contents/[email protected] .
• " Community ecology ," by OpenStax College, Concepts of Biology, CC BY 4.0 . Download the original article for free at http://cnx.org/contents/[email protected] .

Works cited:

• G. Tyler Miller and Scott E. Spoolman, "Each Species Plays a Unique Role in Its Ecosystem," in Essentials of Ecology , 5th ed. (Belmont: Cengage Learning, 2009), 91.
• G. Tyler Miller and Scott E. Spoolman, "Most Species Compete with One Another for Certain Resources," in Essentials of Ecology , 5th ed. (Belmont: Cengage Learning, 2009), 102.
• E. E. Williams, "Ecomorphs, Faunas, Island Size, and Diverse End Points in Island Radiations of Anolis," in Lizard Ecology: Studies of a Model Organism , ed. R. B. Huey et al. (Harvard University Press, 1983).

References:

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5.1: The Ecological Niche

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• Page ID 62278

• Laci M. Gerhart-Barley
• College of Biological Sciences - UC Davis

An important concept in ecology, which will be discussed in several contexts throughout the quarter is the ecological niche. A species’ ecological niche is the abiotic and biotic conditions the species requires in order to grow, reproduce, and survive. Every species, therefore, has a fundamental niche, which are the abiotic conditions the species can physiologically tolerate, as well as a realized niche, which is driven by species interactions. For example, consider a species of fish living in the ocean (Fig 5.1.1).

This species has certain physiological tolerances in temperature, salinity, etc, that determine where the species can exist. Each of the abiotic factors has an optimum range, where the species will be abundant. Outside of this optimum range (both above and below it), are zones of stress, where the species can survive, but experiences physiological stress and will be less common than in the optimum range. Beyond the zones of stress, are zones of intolerance, which represent conditions the species cannot tolerate and in which it will not be found. The tolerance ranges for all relevant abiotic conditions represent the species fundamental niche (Fig 5.1.2). Species interactions such as predation, parasitism, mutualisms, etc may result in this species not being found throughout its entire fundamental niche. For example, if this fish’s prey species do not occur in some areas of the fundamental niche, then this fish species may not occur there even though it could physiologically tolerate these conditions.

The example above illustrates how a species' realized niche might be reduced or contracted compared to the fundamental niche; however, species interactions can also expand the realized niche. For example, the resource acquisition section discussed root mutualisms that many plants form with mycorrhizal fungi and Rhizobium bacteria, which help the plant access soil nutrients. These mutualisms can expand the minimum soil nutrient availability in which the plant can survive, expanding its fundamental niche (Fig 5.1.3). We will consider the impact of species interactions on the niche in more detail in the species interactions sections.

Handbook of Evolutionary Thinking in the Sciences pp 547–586 Cite as

The Ecological Niche: History and Recent Controversies

• Arnaud Pocheville 5  
• First Online: 01 January 2014

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In this chapter, we first trace the history of the concept of ecological niche and see how its meanings varied with the search for a theory of ecology. The niche concept has its roots in the Darwinian view of ecosystems that are structured by the struggle for survival and, originally, the niche was perceived as an invariant place within the ecosystem, that would preexist the assembly of the ecosystem. The concept then slipped towards a sense in which the niche, no longer a pre-existing ecosystem structure, eventually became a variable that would in turn have to be explained by the competitive exclusion principle and the coevolution of species. This concept, while more operational from an empirical point of view than the previous one, suffered from an ill-founded definition. A recent refoundation by Chase & Leibold enabled to overcome some of the definitional difficulties.

We then present how, in contemporary ecology, the niche concept is recruited to explain biodiversity and species coexistence patterns. In parallel, neutralist models, by successfully explaining some ecological patterns without resorting to explanations in terms of niche, have questioned the explanatory virtues of the niche concept.

After this presentation, it seems that the fortunes and misfortunes of the niche concept can be seen as a reflection of the difficulties of ecology to give birth to a theory that would be both predictive and explanatory.

• Ecological niche
• Neutral theory
• Coexistence theories
• Competitive exclusion principle

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Julve ( 2005 ) provides a synthetic list of actors of seemingly ecological ideas since ancient times.

For both authors: (1) the ecological equivalents were the rationale for the concept, as an evidence that similar niches existed in different places, (2) the niche was seen as a place that existed independently of its occupant, (3) food was a major component of the niche but the niche was not restricted to food, as it also included the micro-habitat factors and the relationship to predators. However, Elton’s definition being more vague, several species could share the same niche (Griesemer 1992 : 235). In addition, Elton explicitly excluded macro-habitat factors, which was not the case for Grinnell. (See Schoener 1989 : 86–87 for a detailed discussion of the relationship of these two concepts.)

Griesemer ( 1992 : 235–236) notices that the two concepts are better distinguished with respect to the research programs in which they were inserted, rather than to differences between some of their respective definitions: Grinnell focused on the environment to explain speciation, while Elton focused on the structure of the communities.

Schoener ( 1989 : 85), acknowledging Gaffney (1973, here cited as 1975 ), notices in particular the precedence of Johnson ( 1910 ). Johnson used the word in a way similar to Grinnell’s concept: species must occupy different niches in a region, because of the importance of competition in the Darwinian theory. However, Johnson observed that the lady-beetles he studied did not seem to show a clear niche distinction – an observation, Schoener remarks, that was to be repeated many times on arthropods in later studies. Hutchinson ( 1978 : 156), who studied the books available to Grinnell from 1910 to 1914, did not find Johnson’s work in them (Schoener 1989 : 85).

Schoener also reports the work of another contemporary, Taylor ( 1916 ), who worked with Grinnell, and who also focused on ecological equivalents (Schoener 1989 : 84). Taylor however, Schoener notices, rather than imagining that the repetition of local adaptive radiations to similar niches between different locations would lead to convergences, suggested that the same group of organisms would fill the same niche in different geographical areas. Barriers to dispersal could thus prevent some niches to be filled.

In their historical introduction, Chase and Leibold ( 2003 : 7–8) give a quick and edifying portrait of such studies in vegetal ecology: “For example, Tansley ( 1917 ) performed experiments that showed how plant species competed and coexisted, in a sense vying for shared niche space. Tansley also explicitly contrasted the conditions in which a species could theoretically exist with the actual conditions in which it did exist: ideas generally attributed to Hutchinson ( 1957 ) in his discussion of “fundamental” and “realized” niches (…). Salisbury ( 1929 ) furthered this distinction and suggested that the similarity in species requirements was strongly related to the intensity of their competition – much the same concept as appears in the more widely appreciated work of Gause (1936)” (referred here as Gause 1934 ).

“… if the species lay claim to the very same “niche”, and are more or less equivalent as concerns the utilization of the medium, then the coefficient α [in Lotka-Volterra’s equations] will approach unity” (Gause 1934 : chap. III).

“It appears that the properties of the corresponding [Lotka-Volterra] equation of the struggle for existence are such that if one species has any advantage over the other it will inevitably drive it out completely (Chapter III). It must be noted here that it is very difficult to verify these conclusions under natural conditions. (…) There being but a single niche in the conditions of the experiment it is very easy to investigate the course of the displacement of one species by another.” (Gause 1934 : chap. V)

By contrast, in France, L’Héritier and Teissier ( 1935 ), who carried out experiments on the coexistence of two species of Drosophila, came (in agreement with some experimental results of Gause 1934 ) to the conclusion that “two species sharing the same resource in an environment and using it in an apparently identical way may survive side by side in a state of approximate balance.” (see Gayon and Veuille 2001 : 88). On the status of the competitive exclusion principle, seen as an a priori , and therefore irrefutable, principle, see Hardin ( 1960 : 1293).

The first formulation of the niche concept by Hutchinson is to be found in a footnote, in a paper in limnology (Hutchinson 1944 : 20). Schoener ( 1989 : 91) reports a very similar formulation (in french) in a book by Kostitzin ( 1935 : 43): “Imagine a multi-dimensional symbolic space representing the vital factors: p = pressure, T = temperature, I = illumination, etc.. In this space every living creature at a given time occupies a point, a species may be represented by a set of points.”. Hutchinson ( 1978 : 158, quoted in Schoener 1989 : 91) acknowledged having been informed of Kostitzin’s work in the 1940s, without, however, remembering it when formulating his definition in 1944.

Note that predation will also be set aside in the development of the neutral theory.

The environmental and populational niches are however incommensurable if one holds the view that species make some ecological factors relevant that could not be suspected to be so before observing the species (that is, if species and niches are co-constituted, see e.g. similar views in Drake et al. 2007 ; Longo et al. 2012 ).

The operational difficulties of Hutchinson’s concept come from the (binary) formalism of the set theory he used. They are already partly mentioned by Hutchinson ( 1957 : 417) and discussed in length by Schoener ( 1989 : 93).

All points of the fundamental niche represent the possibility of indefinite existence while all points outside the fundamental niche represent non-indefinite viability. Now, for the ecologist, the performance of a species cannot be reduced to a binary variable. (I thank François Munoz for an insightful comment on this point.) Despite this simplification, a major difficulty is to empirically determine the environmental states that allow the population to survive, because the viability of a population is difficult to assess – especially in the field. Similarly, it is physically impossible to measure the survival of a population at one point of the environmental values, and less precise measurements are likely to ignore the extent of the impact of competing species on the realized niche. Hutchinson ( 1978 : 159, quoted in Schoener 1989 : 93) proposed to use the average values instead, but this would lack both biological relevance (the same average can represent very different biological realities) and relevance for the limiting similarity (the niche width and overlap would not be represented).

Another difficulty concerns the nature of the environmental variables considered: strictly speaking, it is the occurrence of a factor (for example, the frequency of the seeds of a certain size) that is one axis of the niche, and not the measurement of this factor (seed size) (see Hutchinson 1957 : 421, fig. 1 shown above: the axes are respectively “temperature” and “size of food”). This is because organisms compete, if any competition, for places in the biotope space, not for places in the niche space. This gets particularly clear if one considers possible biotopes where the places corresponding to the intersection of the two fundamental niches would be non-limiting. As Schoener ( 1989 : 94) puts it: “Hutchinson’s formulation of niche overlap acts as if competing species are placed together in arenas having single values of such niche dimensions as food size or temperature. (…) But real arenas where populations interact are characterized by distributions of values over axes of resource availability, not by single values.”. A similar problem exists with the concept of utilization niche, as it also uses the measurement of a factor and not the measurement of its occurrence (see below).

Besides, niche theory was considered as inappropriate or of limited use by some botanists, who insisted on the fact that all autotrophic plants “need light, carbon dioxide, water and the same mineral nutrients” (Grubb 1977 : 107) and that a substantial partitioning of these resources seems impossible (but see Sect. 3.4.3). Among them, Grubb pleaded for an extended definition of the niche, including notably the regeneration niche – that is, the way plants colonize the gaps arising in the environment (Grubb 1977 : 119). Fagerström and Agren ( 1979 ) have used models to show how different regeneration properties (i.e. temporal average and variance, and phenology, of diaspore production) could enable coexistence.

See also the treatment by Looijen ( 1998 : Chap. 11 , esp. pp. 184–185).

See the review by Schoener ( 1989 : 97), and the discussions by e.g. Schoener ( 1974 ), Neill ( 1974 ), May ( 1975 ), and references therein.

See Schoener ( 1989 : 97), Chase and Leibold ( 2003 : 13), and references therein.

These and other models are briefly reviewed in Schoener ( 1989 : 98–99).

The word comes from Chase and Leibold ( 2003 : 11).

Pielou ( 1975 : e.g. 80, 1977 ) seems to have been a pioneer (Keddy 1998 : 753) who has been overlooked, which might be brought into perspective with Simberloff’s style, which was “perceived as arrogant and combative” (Lewin 1983 : 639).

See e.g. Gotelli and Graves ( 1996 : chap. 1 ), Looijen ( 1998 : chap. 13 ), Chase and Leibold ( 2003 : 13), and references therein.

On adaptation, See Grandcolas, and Downes, this volume (Ed. note)

Stress: a factor having a negative impact on the organism and on which the organism has no impact ( sensu Chase and Leibold 2003 : 26, table 2).

See Chase and Leibold ( 2003 : 13–14) and references therein.

See also e.g. MacArthur and Levins ( 1964 : 1208), MacArthur ( 1972 : e.g. 37–40) and other predecessors cited in Chase and Leibold ( 2003 : 16).

To be precise, Leibold ( 1995 ) and Chase and Leibold ( 2003 : 15–61) refer to the union of the requirements of the organism and its impacts: “[the niche is] the joint description of the environmental conditions that allow a species to satisfy its minimum requirements so that the birth rate of a local population is equal to or greater than its death rate along with the set of per capita effects of that species on these environmental conditions” (p. 15). The generalization of the definition to the organism responses seems natural (see e.g . Meszéna et al. 2006 ).

See Chase and Leibold ( 2003 : chap. 2 , esp. table 2, p. 26).

See e.g. Chase and Leibold ( 2003 : fig. 2.4 p. 27, p. 44).

Pocheville ( 2010 : chap. 2 , esp. pp. 75–77) provides a more thorough critique of the symmetry between niche construction and natural selection. This point will be further deepened in a forthcomming paper, aimed at showing in which cases niche construction theory produces radical theoretical novelty.

Watt and Hogan ( 2000 : 1427) give the following definition: “Although [the question of what a stem cell is] remains contentious after 30 years of debate (…) the prevailing view is that stem cells are cells with the capacity for unlimited or prolonged self-renewal that can produce at least one type of highly differentiated descendant. Usually, between the stem cell and its terminally differentiated progeny there is an intermediate population of committed progenitors with limited proliferative capacity and restricted differentiation potential, sometimes known as transit amplifying cells.” Laplane ( 2013 ) provides a thorough discussion of the stem cell concept.

Though the stem cell niche concept has been later claimed to come by analogy with the ecological niche concept (e.g. Powell 2005 : 268, see also Papayannopoulou and Scaddeb 2008 ), it does not seem to have been imported from the ecological literature by Schofield. I thank Lucie Laplane for drawing my attention to this point.

It has been shown that tumoral cells can mobilize normal bone marrow cells, have them migrate to particular regions and change the local environment so that it attracts and supports the development of a metastasis (Steeg 2005 ).

Work on cell niche sometimes explicitly refers to the concept of ecological niche (e.g. Powell 2005 : 269). Work on the “niche construction” by the cells, however, does not seem to have been inspired by Odling-Smee’s and colleagues’ program (e.g. Bershad et al. 2008 ).

We briefly discussed this point in Pocheville ( 2010 : chap. III).

See Delord, Chap. 25 , this volume. (Ed. note)

See Meszéna et al. ( 2006 ) for an examination of the structural stability (robustness of coexistence against changes of parameters) of models of stable coexistence.

Here, we use the word “mechanism” in the – very broad – sense used in ecology: practically any form of generation of a pattern can be considered as a mechanism (e.g. Strong et al. 1984 : 5&220, Bell 2000 : 606, Hubbell 2001 : 114, Leigh 2007 : 2087; see the brief discussions in Turner et al. 2001 : 53 and McGill et al. 2007 : 1001). For example, the intensity of competition in a Lotka-Volterra model can be seen, in our view, as a mechanism for the exclusion of two species, while the consumption of the same resource by two species in a Tilman model can be seen as a mechanism, among other possible mechanisms, for the intensity of competition (Tilman 1982 : 6, 1987 : 769; Chesson 2000 : 345). In this sense we say that a Tilman model is “more mechanistic” than a Lotka-Volterra model (e.g. Chase and Leibold 2003 : 13), qualified as “more phenomenological” (see Mikkelson 2005 : 561).

We draw reader’s attention to the fact that here, fitness is not averaged over time but over all environmental states, e.g. the different values of resource availability (Chesson 2000 : 346–7353) or the relative frequency of species (Adler et al. 2007 : 96: fig. 1, 97: fig. 2). Last, we also speak of an average fitness in the sense of the per capita growth rate, averaged among individuals within a population.

Negative frequency-dependence : most frequent populations are disadvantaged. Negative density-dependence: for each population, the per capita growth rate decreases as density increases. While negative frequency-dependence can emerge from negative density-dependence (e.g., when each species has a specific niche which can support a given maximum density), density-dependence is not sufficient to generate frequency-dependence: each species must, in addition, reduce its own growth more than those of others (Chesson 2000 : 348; Adler et al. 2007 : 97). (Note that density-dependence is not necessary for frequency-dependence to occur: for instance rock-paper-scissors games can arise without any obvious link to underlying limiting conditions (e.g. Sinervo and Lively 1996 ).)

In neutral theory, fitness equality is defined at the individual level (regardless of the species), which implies equality at the population level (the reverse is not true).

We would like to draw once again the reader’s attention to the fact that these stabilizing feedbacks are not sufficient in themselves to ensure the stability of coexistence. Put in the graphical terms of Fig. 26.2 given here, niche partitioning will be expressed as a correlation between zero net growth isoclines and impact vectors, and equalizing mechanisms as a proximity of the intercepts of the zero net growth isoclines (Chase and Leibold 2003 : 43).

On predation and parasitism see also Chesson ( 2000 : 356–357) and references therein.

Robustness here is meant in the sense of structural stability (model robustness to parameters changes) (Meszéna et al. 2006 : 69–70). On the concept of model robustness see Levins ( 1966 : 423–427) and for instance, the critique by Orzack and Sober ( 1993 : 538), and the account by Lesne ( 2012 : 1–3).

A population is subject to an Allee effect when “the overall individual fitness, or one of its components, is positively related to population size or density” (Courchamp et al. 2008 : 4, see also p. 10: box 1.1). This effect can be explained by difficulties in finding breeding partners, or by the need for a group to reach a critical mass to be able to exploit a resource or deal with predation (Courchamp et al. 2008 : chap. 2 ).

We draw reader’s attention to the fact that this stabilizing mechanism is different from the niche partitioning with respect to predation exposed above.

To be precise, in this case we would speak of niche restriction rather than niche partitioning (e.g. Rohde 2005 : 51–52).

See Rohde ( 2005 : chap. 5 , quoted here from p. 82) and other works in the 1970s by the same author (e.g. Rohde 1979 ).

See e.g. the discussion by Looijen ( 1998 : chap. XIII).

“I believe that community ecology will have to rethink completely the classical niche-assembly paradigm from first principles.” (Hubbell 2001 : 320).

For simplicity, we use in this section the term “niche theory” in a broad sense (equivalent to the niche-assembly perspective in Hubbell’s terms, 2001 : 8), to mean the corpus of models that are based on the niche concept – and not, in the strict sense, the research program of MacArthur & Levins evoked in Sect. 1.4 .

To be precise, we already find the idea of neutral variation in Darwin (e.g. 1859 : 46): “These facts [an inordinate amount of variation in some genera] seem to be very perplexing, for they seem to show that this kind of variability is independent of the conditions of life. I am inclined to suspect that we see in these polymorphic genera variations in points of structure which are of no service or disservice to the species, and which consequently have not been seized on and rendered definite by natural selection (…)”

However, Hutchinson still considered the competitive exclusion principle as a starting point (Hutchinson 1961 : 143), envisageing to explain unexpectedly high levels of diversity in functional terms, among others: non-equilibrium competitive dynamics (Hutchinson 1941 , cited and deepened in Hutchinson 1961 : 138), the mosaic nature of the environment (Hutchinson 1959 : 154), and the supposed stability of more complex trophic relationships (Hutchinson 1959 : 150).

Schoener ( 1983b ) cited in Loreau and Mouquet ( 1999 : 427), Chase and Leibold ( 2003 : 177–178).

Drift: variation in frequency (here, allelic frequency) due to a random sampling effect in the population: the offspring population of alleles represents a (finite) sample of the parental population. In virtue of the law of large numbers, the larger the sample, the more representative it is.

On neutrality in population genetics, see Leigh ( 2007 : 2076), and references therein.

See Chave ( 2004 : 244) for a discussion on the emergence of neutral models in ecology. Alonso et al. ( 2006 : 452: table 1) provide a useful comparison of the main parameters used in the two neutral theories.

Migration had already been studied in population genetics, but never had a central status as in Hubbell’s theory (Alonso et al. 2006 : 452).

See also Hubbell ( 1997 ).

See Hubbell ( 2001 : esp. chap. 1 , 5 , 6 ) and the presentations by Chave ( 2004 : esp. p. 245.: fig. 2) and Leigh ( 2007 ). Beeravolu et al. ( 2009 ) provide a remarkable review of neutral models. McGill et al. ( 2006 : table 1) provide a usefull comparison of existing neutral models.

It is in particular the case when two species are exactly similar (for instance, if they have exactly the same genes and allelic frequencies as for the functional aspects) and are only inter-sterile: there would be intraspecific, but not interspecific, competition. Hubbell ( 2006 ) proposed (without, however, stating it explicitly) such a mechanism to explain the evolution of neutrality at the interspecific level. (A similar result would probably be obtained assuming no limitation on (epi)mutations at the intraspecific level.) Chave ( 2004 : 249) quicky discusses how restrictive the assumption of individual equivalence is.

Bell ( 2000 : 613) proposed a different – and compatible – definition: “Even the notion of ecological equivalence is rather vague; I shall take it to refer to a set of species for each member of which no interaction with another member is positive. If community structure is determined to some extent by competition, then at least one interaction for each member is negative; the neutral model is the limiting case in which all interactions are negative and equal.”

Neutral theory considers fitness equivalence at the individual level (e.g. Hubbell 2001 : 6), which implies fitness equivalence at the population level.

On the use of these terms, see e.g. Hubbell ( 2001 : 6, 2005 : 166, 2006 ), and the discussion in Clark ( 2009 : 9). For instance Hubbell’s following statement shows a slippage between demographic and functional equivalence: “These life history trade-offs equalize the per capita relative fitness of species in the community, which set the stage for ecological drift.” (Hubbell 2001 : 346, briefly discussed in Alonso et al. 2006 : 455, similar statements can be found elsewhere in the literature, see e.g. Kraft et al. 2008 : 582: note 11). Notice, however, that a full ecological drift would in addition require the absence of any stabilizing mechanisms (an absence that seems to be implicitly hypothesized by Hubbell 2001 : 327–328). The word trade-off itself is ambiguous, as trade-offs can theoretically produce both equalizing and/or stabilizing effects (Chesson 2000 : 346–347), be they trophic (e.g. Clark et al. 2003 ) or life-history trade-offs (e.g. Clark et al. 2004 ). Chase and Leibold ( 2003 ), as for them, seem to use trade-offs (here in niche use) as explanantes of stabilization in their whole book: “That is, Hubbell’s hypothetical species show no niche differences or trade-offs.” (p. 42, note the contrast with Hubbell’s quote above). Clark ( 2009 : 9) shows, using Lotka-Volterra equations, how species can have identical parameters (demographic equivalence) while displaying stable coexistence, in particular if there are trade-offs that entail that each species negatively impacts itself more than it impacts the other (functional differences). (Functional equivalence would in this case be represented by an equivalence of the intra- and inter-specific competition terms for each, and all, species. Notice that, still, it would not imply that species be ecologically equivalent, as Lotka-Volterra parameters can be ecologically multiply realized (see Clark 2009 : fig. 1).)

That is, complete overlap of responses and impacts to environmental factors in Chase’s and Leibold’s ( 2003 : 23) account. Note that with this concept, two species having exactly the same niche behave neutrally, and the only “competitive exclusion” occuring is mere drift.

See Chave ( 2008 : 18–20) for a short comparison of niche vs dispersal assembly frameworks. See Beeravolu et al. ( 2009 : 2605–7) for a review of the different kinds of spatial neutral models.

See esp. Hubbell ( 2001 : chap. 5 ) and the quick and didactic presentation by Alonso et al. ( 2006 : 453: box 2).

See Bell ( 2001 : 2417), Bell et al. ( 2001 : 121–128), Bell ( 2005 ).

See Gotteli and Graves ( 1996 : chap. I), Bell ( 2001 : 2416), Bell ( 2005 : 1757–1758) and references therein.

As we have seen, Darwin ( 1859 : 46, quoted above) already aknowledged the possibility of neutral differences in phenotypes; he however supposed that the abundances of species in an ecosystem could not be explained by chance, but by the struggle between kinds: “When we look at the plants and bushes clothing an entangled bank, we are tempted to attribute their proportional numbers and kinds to what we call chance. But how false a view is this! Every one has heard that when an American forest is cut down, a very different vegetation springs up; but it has been observed that the trees now growing on the ancient Indian mounds, in the Southern United States, display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. What a struggle between the several kinds of trees must here have gone on during long centuries, each annually scattering its seeds by the thousand; (…)” (Darwin 1859 : 74–75).

See Bell et al. ( 2006 ) and Leigh ( 2007 : 2081), for reviews.

Overyielding: positive correlation between the productivity and the diversity of a community.

Hubbell ( 2006 : 1395) argues that he found no evidence for overyielding in the tropical forest on Barro Colorado Island.

See Beeravolu et al. ( 2009 : 2607).

Munoz et al. ( 2007 ) have proposed an approach that relaxes the speciation modalities and do not imply any estimation of the speciation parameter. The estimation of the speciation parameter seems generally highly unreliable, contrary to the estimation of the migration parameter, that seems more robust (on parameter estimation, see also Beeravolu et al. 2009 ). I thank François Munoz for an insightful comment on this point.

Unless explicitly stated, this part draws on the remarkable review by Bell et al. ( 2006 ).

E.g. Watterson ( 1974 ), Caswell ( 1976 ), Hubbell ( 1979 , 1997 , 2001 : 11&17, chap. 5 ), Volkov et al. ( 2003 ).

E.g. Bramson et al. ( 1996 , 1998 ), Hubbell ( 2001 : chap. 6 ), but see Leigh ( 2007 : 2080).

See e.g. Bell ( 2001 , 2005 ), Bell et al. ( 2006 ).

See e.g. Hubbell ( 2001 : 320–321), or this interview of Hubbell by Baker ( 2002 ): “Look, I think the biggest question to come out of the neutral theory is: “Why does it work so well?” I’m as puzzled as the next person. But one idea is these trade-offs.” (Notice that here Hubbell still seeks to explain neutrality in functional terms, while a possibly more neutral explanation would be that environmental variations in space and time are such that the environment is not selective, as for instance with fractal perturbations; a case briefly discussed in Pocheville 2010 : 85–86).

See Puyeo et al. ( 2007 : 1017), McGill et al. ( 2007 : esp. 1001) and references therein; see also Chave ( 2004 : 247–248).

The controversy about SADs draws back to Fisher et al. ( 1943 ) and Preston ( 1948 ). According to Fisher et al. ( 1943 ) the expected number N of species having n individuals in a sample can be described by a log-serie: N  = α n /n , where α (a parameter now known as Fisher’s α) is a measure of species diversity. According to Preston ( 1948 ), the log-serie lacked the bell-shape he observed in his data on bird abundances, a phenomenon he attributed to the presence of trully rare species that are hardly detectable in small samples (a concept now known as Preston’s veil line). Preston ( 1948 ) remarked that, by contrast, a log-normal distribution fitted his data. See Hubbell ( 2001 : 31–37) and McGill et al. ( 2007 : 998–999,1004–1005) for short historical introductions, emphasizing respectively the theoretical and empirical sides.

The maximum entropy technique consists in describing the microscopic degrees of freedom of a system (e.g. the species abundances) by the probability distribution that maximizes the Shannon entropy, under a set of macroscopic constraints (such as bounded mean abundance). On entropy maximization in ecology, see also Banavar and Maritan ( 2007 ), Banavar et al. ( 2010 ), Dewar and Porté ( 2008 ) and the controversy between Shipley et al. ( 2006 ) and Shipley ( 2009 ), and Haegeman and Loreau ( 2008 , 2009 ). Haegeman and Loreau ( 2008 ) provide a nice and critical introduction to the technique.

See Hubbell ( 2001 : 125–126, 150, chap. 6 , 280).

E.g. Hengeveld and Haeck ( 1981 , cited in Brown 1995 : 24, 1982 ), Brown ( 1995 : 32, et al. 1996 )

Note that qualitative patterns (e.g. Bell et al. 2001 : 133) could be an insufficient method to detect selective processes.

See Bell et al. ( 2001 : 129,132), Bell ( 2003 ), Bell ( 2005 ), Bell et al. ( 2006 : 1380–1381, 1383–1384)

See McGill et al. ( 2006 : 1414). Such a question is already mentionned by MacArthur ( 1972 : 21), and is repeated, in a less general form, in Chesson and Huntly ( 1997 : 520), quoted in Hubbell ( 2001 : 9–10). Leigh ( 2007 : 2080) raises, in passing, a similar question. Hubbell ( 1997 : S9) interprets the niche assembly perspective of ecologists (vs the dispersal assembly perspective of biogeographers) as a mark of the different processes occuring on the respective scales of these disciplines. See also the three-levels spatially implicit neutral model of Munoz et al. ( 2008 : 117)

A major difficulty of this research program is to separate the effects of the environmental variability (on fitness) from the effects of physical/biological distances (on dispersal), for there is a covariation between environment similarity and distance in natural landscapes: environmental variability tends to increase with the geographic distance, and the biologically perceived distance tends to increase with environmental variability (due to barriers to dispersal for instance – such barriers need not be, of course, purely “neutral”, i.e . , equivalent for all species). Fort short discussions of this issue, see Bell (2006: 1382), Chave ( 2008 : 21–23). Borcard et al. ( 1992 , see also Legendre and Legendre 2012 ) proposed a method to statistically partition environment from distance, implemented in Gilbert and Lechowicz ( 2004 : 7653) who found “strong evidence of niche-structuring but almost no support for neutral predictions” ( 2004 : 7651). Jeliazkov ( 2013 : Chap. III) performed an implementation in a similar vein, finding that the environment explained a major part of the community variation only when it was joined to a spatial component. On dispersal as a non-neutral phenomenon see Clark ( 2009 : 12).

In other terms, while the composition of a neutral community does not show any equilibrium nor resilience, it is not the case for the caracteristics of this composition (species number, relative frequencies, etc.).

A similar counterargument has been opposed by Hubbell ( 2001 : 330–331) to the conclusions reached by Terborgh et al. ( 1996 ) on floodplain forests and Pandolfi ( 1996 ) on a paleo-reconstruction of coral reefs. Leigh ( 2007 : 2082) points to the fact that Hubbell’s ( 2001 : 331) and Volkov et al.’s ( 2004 ) arguments rely on “the fictitious concept of a panmictic source pool”, a fiction that contrasts with a – desirable – approach studying the long-range correlations produced by local dispersal alone (as hypothesized by Bell et al. 2006 : 1382). As another step in the controversy, Dornelas et al. ( 2006 ) have shown that Indo-Pacific coral communities exhibit far more variable, and lower on average, community similarities than expected by neutrality.

The debate is quickly summarized in Bell et al. ( 2006 : 1379) and Leigh ( 2007 : 2081–2082).

E.g. Hubbell ( 2001 : 220), McGill et al. ( 2005 : 16706), Bell et al. ( 2006 : 1379), Gewin ( 2006 : 1309), Daleo et al. ( 2009 : 547). These terms are not new, as in the 1970s Lewontin for instance could write: “Genetic variation is removed from populations by both random and deterministic forces” (Lewontin 1974 : 192).

See Malaterre & Merlin, Chap. 17 , this volume (Ed. note)

We mean here by “directionnal” a direction in the composition dynamics (of alleles or species frequencies for instance) or in spatial patterns of distributions. Drift, by contrast, can be considered as a noise: it “explains” to what extent we cannot know the direction. (This, of course, does not hold for parameters that are explananda of neutral theory, such as the number of alleles/species, mentionned in the preceding section.)

This notion of epistemic randomness is, to our knowledge, the most common notion of randomness in ecology (e.g. Clark 2009 : 10: “First, there is no evidence for stochasticity in nature at observable scales. Stochasticity is an attribute of models”). To be precise, random terms could also be considered to reflect deterministically random phenomena, as in classical physics, or intrinsically random phenomena, as in quantum physics. Other concepts of randomness could be developed for ecology. The distinction between direction/dispersion proposed here holds for epitemic randomness.

This explanandum is significant in, for instance, the review by Lavergne et al. ( 2010 ).

Huneman ( 2012 ) questions in the same vein the conception of causation (counterfactual vs statistical) required to make sense of natural selection (by contrast with drift) in evolutionary biology.

Neutral theory has not always been perceived as a null hypothesis. Bell ( 2001 : 2418) distinguishes two versions of the theory: “The weak version recognizes that the NCM [neutral community model] is capable of generating patterns that resemble those arising from survey data, without acknowledging that it correctly identifies the underlying mechanism responsible for generating these patterns. The role of the NCM is then restricted to providing the appropriate null hypothesis when evaluating patterns of abundance and diversity. (…) The strong version is that the NCM is so successful precisely because it has correctly identified the principal mechanism underlying patterns of abundance and diversity. This has much more revolutionary consequences, because it involves accepting that neutral theory will provide a new conceptual foundation for community ecology and therefore for its applied arm, conservation biology.”

A similar argument would hold for implicitly spatial models (involving limited dispersal without necessarily defining a distance between communities): dispersal, even symmetric, is not “null” regarding the niche.

The a priori principle belongs to the same familiy than the strong adaptationist principle, that can be formulated as, for example: “every trait is an adaptation to a selection pressure, even if this pressure is not shown”, or: “it is the fittest who survives, even if fitness is not shown”. (On adaptationism, see Orzack and Sober 2001 , in particular the chapter by Godfrey-Smith.)

I am indebted to Philippe Huneman for having drawn my attention to this point.

The author wish to thank Frédéric Bouchard, Antoine Collin, Régis Ferrière, Jean Gayon, Philippe Huneman, Maël Montévil, Michel Morange, François Munoz, Aurélien Pocheville and Marc Silberstein, whose suggestions enabled to greatly improve previous versions of the manuscript. Sylvie Beaud and Robert Pocheville were of considerable help for the translation of the french version. This work consists in a partial update of a previous work in French (Pocheville 2009 ), realized while the author was benefiting from a funding from the Frontiers in Life Sciences PhD Program and from the Liliane Bettencourt Doctoral Program. This update was realized while the author was benefiting from a Postdoctoral Fellowship from the Center for Philosophy of Science, University of Pittsburgh.

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Arnaud Pocheville

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Pocheville, A. (2015). The Ecological Niche: History and Recent Controversies. In: Heams, T., Huneman, P., Lecointre, G., Silberstein, M. (eds) Handbook of Evolutionary Thinking in the Sciences. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9014-7_26

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ENCYCLOPEDIC ENTRY

A species’ niche is all of the environmental factors and interspecies relationships that influence the species.

Biology, Ecology

Kirtland's Warbler

A species' niche describes how it fits within its environment. The Kirtland's Warbler (Setophaga kirtlandii) has quite a specific niche, only nesting in young jack pine trees (Pinus banksiana).

Photograph by Jeff Rzepka/500px

In ecology , the term “ niche ” describes the role an organism plays in a community. A species’ niche encompasses both the physical and environmental conditions it requires (like temperature or terrain) and the interactions it has with other species (like predation or competition ). For example, the rare Kirtland’s warbler ( Setophaga kirtlandii ), a small songbird of North America, has a very limited niche. It nests only among young jack pine trees ( Pinus banksiana ), which require periodic wildfires for their seeds to germinate . In this environment, one of the species interactions it must contend with is nest parasitism by the brown-headed cowbird ( Molothrus ater ). Cowbirds lay their eggs in nests built by other bird species and these host birds then incubate and raise the cowbird’s young, often at the expense of their own babies. In general, species that have narrow or limited niches are considered to be specialist species . Koalas ( Phascolarctos cinereus ), which feed only on leaves from eucalyptus trees in Australia, are an example of a specialist species. Species with broader niches, like coyotes ( Canis latrans ) or raccoons ( Procyon lotor ), are considered generalists. No two species can have the exact same niche , otherwise they would be in direct competition for resources with one another. If this occurs, then one species will outcompete the other. If the losing species then does not adapt , it would lead to its extinction . Joseph Grinnell, an American ecologist, was the first person to develop the idea of an ecological niche . His definition, which he wrote about in scientific papers starting in 1917, focused on the environmental factors that determined where a species could survive rather than interactions between species. Around the same time, an English ecologist named Charles Elton was developing his own ideas about niches . In his definition, a species’ niche was determined by its interactions with other species—namely its relationships with food and predators . Almost 40 years later, in the late 1950s, English ecologist G. Evelyn Hutchinson blended these two versions into a broader definition. This definition considers all abiotic and biotic factors that influence a species in a quantifiable way. This definition is still used by scientists today. It is important to keep learning about species’ niches today, because it can help us understand how organisms will respond to environmental changes caused by humans.

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