In ecology, a niche is the match of a species to a specific environmental condition.
A Grinnellian niche is determined by the habitat in which a species lives and its accompanying behavioral adaptations. An Eltonian niche emphasizes that a species not only grows in and responds to an environment, it may also change the environment and its behavior as it grows. The Hutchinsonian niche uses mathematics and statistics to try to explain how species coexist within a given community.
The concept of ecological niche is central to ecological biogeography, which focuses on spatial patterns of ecological communities.
The majority of species exist in a standard ecological niche, sharing behaviors, adaptations, and functional traits similar to the other closely related species within the same broad taxonomic class, but there are exceptions. A premier example of a non-standard niche filling species is the flightless, ground-dwelling kiwi bird of New Zealand, which feeds on worms and other ground creatures, and lives its life in a mammal-like niche. Island biogeography can help explain island species and associated unfilled niches.
The Grinnellian niche concept embodies the idea that the niche of a species is determined by the habitat in which it lives and its accompanying behavioral adaptations. In other words, the niche is the sum of the habitat requirements and behaviors that allow a species to persist and produce offspring. For example, the behavior of the California thrasher is consistent with the chaparral habitat it lives in—it breeds and feeds in the underbrush and escapes from its predators by shuffling from underbrush to underbrush. Its 'niche' is defined by the felicitous complementing of the thrasher's behavior and physical traits (camouflaging color, short wings, strong legs) with this habitat.
Grinnellian niches can be defined by non-interactive (abiotic) variables and environmental conditions on broad scales. Variables of interest in this niche class include average temperature, precipitation, solar radiation, and terrain aspect which have become increasingly accessible across spatial scales. Most literature has focused on Grinnellian niche constructs, often from a climatic perspective, to explain distribution and abundance. Current predictions on species responses to climate change strongly rely on projecting altered environmental conditions on species distributions. However, it is increasingly acknowledged that climate change also influences species interactions and an Eltonian perspective may be advantageous in explaining these processes.
This perspective of niche allows for the existence of both ecological equivalents and empty niches. An ecological equivalent to an organism is an organism from a different taxonomic group exhibiting similar adaptations in a similar habitat, an example being the different Succulent plant found in American and African deserts, cactus and euphorbia, respectively.
As another example, the anole lizards of the Greater Antilles are a rare example of convergent evolution, adaptive radiation, and the existence of ecological equivalents: the anole lizards evolved in similar microhabitats independently of each other and resulted in the same across all four islands.
Elton classified niches according to foraging activities ("food habits"):
"Elton focused on the niche of a species as its functional role within the food chain and its impact upon the environment"
Conceptually, the Eltonian niche introduces the idea of a species' response to and effect on the environment. Unlike other niche concepts, it emphasizes that a species not only grows in and responds to an environment based on available resources, predators, and climatic conditions, but also changes the availability and behavior of those factors as it grows. In an extreme example, require certain resources in order to survive and reproduce, but also construct dams that alter water flow in the river where the beaver lives. Thus, the beaver affects the biotic and abiotic conditions of other species that live in and near the watershed.
Eltonian niches focus on biotic interactions and consumer–resource dynamics (biotic variables) on local scales. Because of the narrow extent of focus, data sets characterizing Eltonian niches typically are in the form of detailed Field research of specific individual phenomena, as the dynamics of this class of niche are difficult to measure at a broad geographic scale. However, the Eltonian niche may be useful in the explanation of a species' endurance of global change. Because adjustments in biotic interactions inevitably change abiotic factors, Eltonian niches can be useful in describing the overall response of a species to new environments.
The niche concept was popularized by the zoologist G. Evelyn Hutchinson in 1957. Hutchinson inquired into the question of why there are so many types of organisms in any one habitat. His work inspired many others to develop models to explain how many and how similar coexisting species could be within a given community, and led to the concepts of 'niche breadth' (the variety of resources or habitats used by a given species), 'niche partitioning' (resource differentiation by coexisting species), and 'niche overlap' (overlap of resource use by different species).
Statistics were introduced into the Hutchinson niche by Robert MacArthur and Richard Levins using the 'resource-utilization' niche employing histograms to describe the 'frequency of occurrence' as a function of a Hutchinson coordinate. So, for instance, a Gaussian might describe the frequency with which a species ate prey of a certain size, giving a more detailed niche description than simply specifying some median or average prey size. For such a bell-shaped distribution, the position, width and form of the niche correspond to the mean, standard deviation and the actual distribution itself. One advantage in using statistics is illustrated in the figure, where it is clear that for the narrower distributions (top) there is no competition for prey between the extreme left and extreme right species, while for the broader distribution (bottom), niche overlap indicates competition can occur between all species. The resource-utilization approach postulates that not only can competition occur, but that it does occur, and that overlap in resource utilization directly enables the estimation of the competition coefficients. This postulate, however, can be misguided, as it ignores the impacts that the resources of each category have on the organism and the impacts that the organism has on the resources of each category. For instance, the resource in the overlap region can be non-limiting, in which case there is no competition for this resource despite niche overlap.
An organism free of interference from other species could use the full range of conditions (biotic and abiotic) and resources in which it could survive and reproduce which is called its fundamental niche. However, as a result of pressure from, and interactions with, other organisms (i.e. inter-specific competition) species are usually forced to occupy a niche that is narrower than this, and to which they are mostly highly adaptation; this is termed the realized niche. Hutchinson used the idea of competition for resources as the primary mechanism driving ecology, but overemphasis upon this focus has proved to be a handicap for the niche concept. In particular, overemphasis upon a species' dependence upon resources has led to too little emphasis upon the effects of organisms on their environment, for instance, colonization and invasions.
The term "adaptive zone" was coined by the paleontologist George Gaylord Simpson to explain how a population could jump from one niche to another that suited it, jump to an 'adaptive zone', made available by virtue of some modification, or possibly a change in the food chain, that made the adaptive zone available to it without a discontinuity in its way of life because the group was 'pre-adapted' to the new ecological opportunity.
Hutchinson's "niche" (a description of the ecological space occupied by a species) is subtly different from the "niche" as defined by Grinnell (an ecological role, that may or may not be actually filled by a species—see vacant niches).
A niche is a very specific segment of ecospace occupied by a single species. On the presumption that no two species are identical in all respects (called Hardin's 'axiom of inequality') and the competitive exclusion principle, some resource or adaptive dimension will provide a niche specific to each species. Species can however share a 'mode of life' or 'autecological strategy' which are broader definitions of ecospace. For example, Australian grasslands species, though different from those of the Great Plains grasslands, exhibit similar modes of life. Glossary for the Nature of Alberta
Once a niche is left vacant, other organisms can fill that position. For example, the niche that was left vacant by the extinction of the tarpan has been filled by other animals (in particular a small horse breed, the konik). Also, when plants and animals are introduced into a new environment, they have the potential to occupy or invade the niche or niches of native organisms, often outcompeting the indigenous species. Introduction of non-indigenous species to non-native by humans often results in biological pollution by the exotic or invasive species.
The mathematical representation of a species' fundamental niche in ecological space, and its subsequent projection back into geographic space, is the domain of niche modelling.On the logic of the relation between the niche and the corresponding geographic environment, see:
In contemporary niche theory, the "impact niche" is defined as the combination of effects that a given consumer has on both a). the resources that it uses, and b). the other consumers in the ecosystem. Therefore, the impact niche is equivalent to the Eltonian niche since both concepts are defined by the impact of a given species on its environment.
The range of environmental conditions where a species can successfully survive and reproduce (i.e. the Hutchinsonian definition of a realized niche) is also encompassed under contemporary niche theory, termed the "requirement niche". The requirement niche is bounded by both the availability of resources as well as the effects of coexisting consumers (e.g. competitors and predators).
These requirements are interesting and controversial because they require any two species to share a certain environment (have overlapping requirement niches) but fundamentally differ the ways that they use (or "impact") that environment. These requirements have repeatedly been violated by nonnative (i.e. introduced and Invasive species) species, which often coexist with new species in their nonnative ranges, but do not appear to be constricted these requirements. In other words, contemporary niche theory predicts that species will be unable to invade new environments outside of their requirement (i.e. realized) niche, yet many examples of this are well-documented. Additionally, contemporary niche theory predicts that species will be unable to establish in environments where other species already consume resources in the same ways as the incoming species, however examples of this are also numerous.
As an example of niche partitioning, several anole lizards in the Caribbean islands share common diets—mainly insects. They avoid competition by occupying different physical locations. Although these lizards might occupy different locations, some species can be found inhabiting the same range, with up to 15 in certain areas.Joshua Anderson. "Interspecific Competition, Competitive Exclusion, and Niche Differentiation"
In reality, this still leaves the question of how much differentiation is needed for coexistence. A vague answer to this question is that the more similar two species are, the more finely balanced the suitability of their environment must be in order to allow coexistence. There are limits to the amount of niche differentiation required for coexistence, and this can vary with the type of resource, the nature of the environment, and the amount of variation both within and between the species.
To answer questions about niche differentiation, it is necessary for ecologists to be able to detect, measure, and quantify the niches of different coexisting and competing species. This is often done through a combination of detailed ecology studies, controlled experiments (to determine the strength of competition), and mathematical models. To understand the mechanisms of niche differentiation and competition, much data must be gathered on how the two species interact, how they use their resources, and the type of ecosystem in which they exist, among other factors. In addition, several mathematical models exist to quantify niche breadth, competition, and coexistence (Bastolla et al. 2005). However, regardless of methods used, niches and competition can be distinctly difficult to measure quantitatively, and this makes detection and demonstration of niche differentiation difficult and complex.
Although "resource" generally refers to food, species can partition other non-consumable objects, such as parts of the habitat. For example, are thought to coexist because they nest in different parts of trees. Species can also partition habitat in a way that gives them access to different types of resources. As stated in the introduction, anole lizards appear to coexist because each uses different parts of the forests as perch locations. This likely gives them access to different species of insects as well.
Research has determined that plants can recognize each other's root systems and differentiate between a clone, a plant grown from the same mother plants seeds, and other species. Based on the root secretions, also called exudates, plants can make this determination. The communication between plants starts with the secretions from plant roots into the rhizosphere. If another plant that is kin is entering this area the plant will take up exudates. The exudate, being several different compounds, will enter the plants root cell and attach to a receptor for that chemical halting growth of the root meristem in that direction, if the interaction is kin.
Simonsen discusses how plants accomplish root communication with the addition of beneficial rhizobia and fungal networks and the potential for different genotypes of the kin plants, such as the legume M. Lupulina, and specific strains of nitrogen fixing bacteria and rhizomes can alter relationships between kin and non-kin competition. This means there could be specific subsets of genotypes in kin plants that selects well with specific strains that could outcompete other kin. What might seem like an instance in kin competition could just be different genotypes of organisms at play in the soil that increase the symbiotic efficiency.
The other paradigm assumes that niche space is to a large degree vacant, i.e., that there are many vacant niches. It is based on many empirical studies Rohde, K. 1980. Warum sind ökologische Nischen begrenzt? Zwischenartlicher Antagonismus oder innerartlicher Zusammenhalt? Naturwissenschaftliche Rundschau, 33, 98–102.Rohde, K. 2005a. Eine neue Ökologie. Aktuelle Probleme der evolutionären Ökologie. Naturwissenschaftliche Rundschau, 58, 420–426.K. Rohde: Nonequilibrium Ecology, Cambridge University Press, Cambridge, 2005b, 223 pp. auf http://www.cambridge.org/9780521674553 and theoretical investigations especially of Kauffman 1993.Kauffman, S.A. 1993. The origins of order. Self-organization and selection in evolution. Oxford University Press, New York Oxford. Causes of vacant niches may be evolutionary contingencies or brief or long-lasting environmental disturbances.
Both paradigms agree that species are never "universal" in the sense that they occupy all possible niches; they are always specialized, although the degree of specialization varies. For example, there is no universal parasite which infects all host species and microhabitats within or on them. However, the degree of host specificity varies strongly. Thus, Toxoplasma (Protista) infects numerous vertebrates including humans, Enterobius vermicularis infects only humans.
The following mechanisms for niche restriction and segregation have been proposed:
Niche restriction:
Niche segregation:
For a detailed discussion, especially of competition and reinforcement of reproductive barriers, see
This example illustrates that the evidence for niche differentiation is by no means universal. Niche differentiation is also not the only means by which coexistence is possible between two competing species. However, niche differentiation is a critically important ecological idea which explains species coexistence, thus promoting the high biodiversity often seen in many of the world's biomes.
Research using mathematical modelling is indeed demonstrating that predation can indeed stabilize lumps of very similar species. Willow warbler and chiffchaff and other very similar warblers can serve as an example. The idea is that it is also a good strategy to be very similar to a successful species or have enough dissimilarity. Other examples of nearly identical species clusters occupying the same niche were water beetles, prairie birds and algae. The basic idea is that there can be clusters of very similar species all applying the same successful strategy and between them open spaces. Here the species cluster takes the place of a single species in the classical ecological models.
An early study on ecological niches conducted by Joseph H. Connell analyzed the environmental factors that limit the range of a barnacle ( Chthamalus stellatus) on Scotland's Isle of Cumbrae. In his experiments, Connell described the dominant features of C. stellatus niches and provided explanation for their distribution on intertidal zone of the rocky coast of the Isle. Connell described the upper portion of C. stellatus's range is limited by the barnacle's ability to resist dehydration during periods of low tide. The lower portion of the range was limited by interspecific interactions, namely competition with a cohabiting barnacle species and predation by a snail. By removing the competing B. balanoides, Connell showed that C. stellatus was able to extend the lower edge of its realized niche in the absence of competitive exclusion. These experiments demonstrate how biotic and abiotic factors limit the distribution of an organism.
Eltonian niche
Hutchinsonian niche
Contemporary niche theory
Coexistence under contemporary niche theory
Niche differentiation
/ref> The competitive exclusion principle states that if two species with identical niches (ecological roles) compete, then one will inevitably drive the other to extinction. This rule also states that two species cannot occupy the same exact niche in a habitat and coexist together, at least in a stable manner.Khan Academy. "Niches & Competition"
/ref> When two species differentiate their niches, they tend to compete less strongly, and are thus more likely to coexist. Species can differentiate their niches in many ways, such as by consuming different foods, or using different areas of the environment.
/ref> For example, some live on the ground while others are arboreal. Species who live in different areas compete less for food and other resources, which minimizes competition between species. However, species who live in similar areas typically compete with each other.
Detection and quantification
Development
Current competition (The Ghost of Competition Present)
Via past extinctions (The Ghost of Competition Past)
Evolving differences
Types
Resource partitioning
Predator partitioning
Conditional differentiation
Competition-predation trade-off
Segregation versus restriction
Relative significance of the mechanisms
Coexistence without niche differentiation: exceptions to the rule
Niche and Geographic Range
Parameters
See also
Further reading
External links
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