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-A '''metapopulation''' consists of a group of spatially separated [[population]]s of the same [[species]] which interact at some level. The term metapopulation was coined by [[Richard Levins]] in 1970 to describe a model of population dynamics of insect pests in agricultural fields, but the idea has been most broadly applied to species in naturally or artificially [[habitat fragmentation|fragmented habitats]]. In Levins' own words, it consists of "a population of populations".<ref name=Levins1969>{{citation  | last = Levins | first =  R.  | year = 1969  | title = Some demographic and genetic consequences of environmental heterogeneity for biological control  | journal = Bulletin of the Entomological Society of America  | volume = 15  | pages = 237–240 }}</ref>
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-A metapopulation is generally considered to consist of several distinct populations together with areas of suitable habitat which are currently unoccupied. In classical metapopulation theory, each population cycles in relative independence of the other populations and eventually goes extinct as a consequence of demographic [[stochastic]]ity (fluctuations in population size due to random [[demographic]] events); the smaller the population, the more prone it is to extinction.
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-Although individual populations have finite life-spans, the metapopulation as a whole is often stable because immigrants from one population (which may, for example, be experiencing a population boom) are likely to re-colonize habitat which has been left open by the extinction of another population. They may also emigrate to a small population and rescue that population from extinction (called the ''rescue effect''). Such a rescue effect may occur because declining populations leave niche opportunities open to the "rescuers".
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-Metapopulation theory was first developed for terrestrial ecosystems, and subsequently applied to the marine realm.<ref name="Kritzer06">Kritzer, JP & Sale, PF (eds) (2006) Marine metapopulations, Academic Press, New York.</ref> In fisheries science, the term "sub-population" is equivalent to the metapopulation science term "local population". Most marine examples are provided by relatively sedentary species occupying discrete patches of habitat, with both local recruitment and recruitment from other local populations in the larger metapopulation. Kritzer & Sale have argued against strict application of the metapopulation definitional criteria that extinction risks to local populations must be non-negligible.<ref name="Kritzer06"/>{{rp|32}}
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-== Huffaker's experiments on predator-prey interactions (1958) ==
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-In order to study predation and population oscillations, Huffaker used mite species, one being the predator and the other being the prey.<ref name=Huffaker1958>{{citation  | last = Huffaker | first =  C.B.  | year = 1958  | title = Experimental Studies on Predation: Dispersion factors and predator-prey oscillations  | journal = Hilgardia  | volume = 27  | issue = 343  | pages = 83 }}</ref> He set up a controlled experiment using oranges, which the prey fed on, as the spatially structured habitat in which the predator and prey would interact.<ref name=Legendre1989>{{citation  | last1 = Legendre | first1 = P.  | last2 = Fortin | first2 = M.J.  | year = 1989  | title = Spatial pattern and ecological analysis  | journal = Plant Ecology  | volume = 80  | issue = 2  | pages = 107  | doi = 10.1007/BF00048036 }}</ref> At first, Huffaker experienced difficulties similar to those of Gause in creating a stable predator-prey interaction. By using oranges only, the prey species quickly went extinct followed consequently with predator extinction. However, he discovered that by modifying the spatial structure of the habitat, he could manipulate the population dynamics and allow the overall survival rate for both species to increase. He did this by altering the distance between the prey and oranges (their food), establishing barriers to predator movement, and creating corridors for the prey to disperse.<ref name="Real91"/> These changes resulted in increased habitat patches and in turn provided more areas for the prey to seek temporary protection. When the prey would go extinct locally at one habitat patch, they were able to reestablish by migrating to new patches before being attacked by predators. This habitat spatial structure of patches allowed for coexistence between the predator and prey species and promoted a stable population oscillation model.<ref name=Kareiva1987>{{citation  | last = Kareiva | first =  P.  | year = 1987  | title = Habitat Fragmentation and the Stability of Predator-Prey Interactions  | journal = Nature  | volume = 326  | issue = 6111  | pages = 388  | doi = 10.1038/326388a0 |bibcode = 1987Natur.326..388K }}</ref> Although the term metapopulation had not yet been coined, the environmental factors of [[spatial heterogeneity]] and habitat patchiness would later describe the conditions of a metapopulation relating to how groups of spatially separated populations of species interact with one another. Huffaker's experiment is significant because it showed how metapopulations can directly affect the predator-prey interactions and in turn influence population dynamics.<ref>Janssen, A. et al. 1997. Metapopulation Dynamics of a Persisting Predator-Prey system.</ref>
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-and growth rate ''r''
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-At equilibrium, therefore, some fraction of the species's habitat will always be unoccupied.
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-== Stochasticity and metapopulations ==
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-Huffaker's<ref name="Huffaker1958" /> studies of spatial structure and species interactions are an example of early experimentation in metapopulation dynamics. Since the experiments of Huffaker<ref name="Huffaker1958"/> and Levins,<ref name="Levins1969" /> models have been created which integrate stochastic factors. These models have proven that the combination of environmental variability (stochascity) and relatively small migration rates cause indefinite or unpredictable persistence. However, Huffaker's experiment almost guaranteed infinite persistence because of the controlled immigration variable.
+ </ul>
-== Stochastic patch occupancy models (SPOMs) ==
-One major drawback of the Levins model is that it is [[deterministic]], whereas the fundamental metapopulation processes are [[stochastic]]. Metapopulations are particularly useful when discussing species in disturbed [[habitats]], and the [[minimum viable population|viability of their populations]], i.e., how likely they are to become [[extinct]] in a given time interval. The Levins model cannot address this issue. A simple way to extend the Levins' model to incorporate space and stochastic considerations is by using the [[contact process (mathematics)]]. Simple modifications to this model can also incorporate for [[patch dynamics]]. At a given [[percolation threshold]], [[habitat fragmentation]] effects take place in these configuration predicting more drastic extinction thresholds.
-<ref>{{cite  journal|author= Keymer J.E, P.A. Marquet, J.X. Velasco‐Hernández, S.A. Levin|year=2000 |month=November |title=Extinction Thresholds and Metapopulation Persistence in Dynamic Landscapes|journal=The American Naturalist |volume=156 |pages= 478–4945 | doi=10.1086/303407}}</ref>
-For [[conservation biology]] purposes, metapopulation models must include (a) the finite nature of metapopulations (how many patches are suitable for habitat), and (b) the [[probability theory|probabilistic]] nature of extinction and colonisation. Also, note that in order to apply these models, the extinctions and colonisations of the patches must be asynchronous.
-==Microhabitat patches (MHPs) and bacterial metapopulations==
-[[Image:ecoli metapopulation.jpg|thumb|E. coli metapopulation on-chip.]]
-By combining [[nanotechnology]] with [[landscape ecology]], a habitat landscape can be nanofabricated on-chip by building a collection of nanofabricated bacterial habitats, and connecting them by corridors in different topological arrangements and with nano-scale channels providing them with the local [[ecosystem service]] of habitat renewal. These landscapes of MHPs can be used as physical implementations of an [[adaptive landscape]]:<ref>{{cite  journal|author= Keymer J.E., P. Galajda, C. Muldoon R., and R. Austin|year=2006 |month=November |title=Bacterial metapopulations in nanofabricated landscapes|journal=PNAS |volume=103 |issue=46 |pages= 17290–295 |pmc= 1635019|doi=10.1073/pnas.0607971103 |bibcode=2006PNAS..10317290K |pmid=17090676}}</ref> by generating a spatial mosaic of [[patches]] of opportunity distributed in space and time. The patchy nature of these fluidic landscapes allows for the study of adapting bacterial cells in a metapopulation system operating on-chip within a synthetic ecosystem. The metapopulation biology and [[evolutionary ecology]] of these bacterial systems, in these synthetic ecosystems, can be addressed using experimental [[biophysics]].
-==Life history evolution==
-Metapopulation models have been used to explain life-history evolution, such as the [[ecological stability]] of [[Metamorphosis#Amphibian metamorphosis|amphibian metamorphosis]] in small [[vernal pools|vernal ponds]]. Alternative ecological strategies have evolved. For example, some salamanders forgo metamorphosis and sexually mature as aquatic [[Neoteny|neotenes]]. The seasonal duration of wetlands and the migratory range of the species determines which ponds are connected and if they form a metapopulation. The duration of the life history stages of amphibians relative to the duration of the vernal pool before it dries up regulates the ecological development of metapopulations connecting aquatic patches to terrestrial patches.<ref name="Petranka07">{{Citation |last = Petranka |first = J. W. |title = Evolution of complex life cycles of amphibians: bridging the gap between metapopulation dynamics and life history evolution |journal = Evolutionary Ecology |volume = 21 |issue = 6 |pages=751–764 |year = 2007 |doi = 10.1007/s10682-006-9149-1 |postscript = .}}</ref>
-== See also ==
-* [[Competition (biology)]]
-* [[Conservation biology]]
-* [[Landscape ecology]]
-* [[Lotka-Volterra equation]]
-* [[Oscillation]]
-* [[Population viability analysis]]
-* [[Predation]]
-* [[Spatial heterogeneity]]
-== References ==
-{{Reflist}}
-* {{Citation | author = Bascompte J., Solé R. V. | year = 1996 | title = Habitat Fragmentation and Extinction Thresholds in spatially explicit models | url = | journal = Journal of Animal Ecology | volume = 65 | issue = 4| pages = 465–473 | doi = 10.2307/5781 | postscript = . }}
-* Hanski, I. ''Metapopulation Ecology'' Oxford University Press. 1999. ISBN 0-19-854065-5
-* Fahrig, L. 2003. Effects of Habitat Fragmentation on Biodiversity. Annual Review of ecology, evolution, and systematics. 34:1, p.&nbsp;487.
-* {{Citation | author = Levin S.A. | year = 1974 | title = Dispersion and Population Interactions | url = | journal = The American Naturalist | volume = 108 | issue = 960| page = 207 | postscript = . }}
-== External links ==
-*[http://www.helsinki.fi/science/metapop/ Helsinki-science: Metapopulation]
-{{-}}
-{{modelling ecosystems|expanded=other}}
-[[Category:Population ecology]]
-[[Category:Landscape ecology]]
-[[Category:Demography]]
-[[Category:Population]]