147:
first, Huffaker experienced difficulties similar to those of Gause in creating a stable predatorāprey interaction. By using oranges only, the prey species quickly became 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. These changes resulted in increased habitat patches and in turn provided more areas for the prey to seek temporary protection. When the prey would become 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. Although the term metapopulation had not yet been coined, the environmental factors of
474:, synthetic habitat landscapes have been fabricated on a chip by building a collection of bacterial mini-habitats with nano-scale channels providing them with nutrients for habitat renewal, and connecting them by corridors in different topological arrangements, generating a spatial mosaic of patches of opportunity distributed in time. This can be used for landscape experiments by studying the bacteria metapopulations on the chip, for example their
22:
459:
502:. 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.
394:
Huffaker's studies of spatial structure and species interactions are an example of early experimentation in metapopulation dynamics. Since the experiments of
Huffaker and Levins, models have been created which integrate stochastic factors. These models have shown that the combination of environmental
160:
Levins' original model applied to a metapopulation distributed over many patches of suitable habitat with significantly less interaction between patches than within a patch. Population dynamics within a patch were simplified to the point where only presence and absence were considered. Each patch in
151:
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 predatorāprey interactions
69:
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.
88:
Metapopulation theory was first developed for terrestrial ecosystems, and subsequently applied to the marine realm. 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
57:
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
25:
Metapopulations are important in fisheries. The local population (1.) serves as a source for hybridization with surrounding subspecies populations (1.a, 1.b, and 1.c).The populations are normally spatially separated and independent but spatial overlap between them during breeding times allows for
146:
In order to study predation and population oscillations, Huffaker used mite species, one being the predator and the other being the prey. 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. At
137:
accurately depicted the oscillations predicted by the Lotka-Volterra equation, with the peaks in prey abundance shifted slightly to the left of the peaks of the predator densities. Huffaker's experiments expanded on those of Gause by examining how both the factors of migration and spatial
93:, 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.
132:
over time based on the initial densities of predator and prey. Gause's early experiments to prove the predicted oscillations of this theory failed because the predatorāprey interactions were not influenced by immigration. However, once immigration was introduced, the
85:, emphasised the importance of connectivity between seemingly isolated populations. Although no single population may be able to guarantee the long-term survival of a given species, the combined effect of many populations may be able to do this.
395:
variability (stochasticity) and relatively small migration rates cause indefinite or unpredictable persistence. However, Huffaker's experiment almost guaranteed infinite persistence because of the controlled immigration variable.
128:, which was formulated in the mid-1920s, but no further application had been conducted. The Lotka-Volterra equation suggested that the relationship between predators and their prey would result in population
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423:
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
381:
1772:
1025:
1916:
755:
Keymer J.E; P.A. Marquet; J.X. Velasco-HernƔndez; S.A. Levin (November 2000). "Extinction
Thresholds and Metapopulation Persistence in Dynamic Landscapes".
1986:
867:
Petranka, J. W. (2007), "Evolution of complex life cycles of amphibians: bridging the gap between metapopulation dynamics and life history evolution",
1523:
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nature of extinction and colonisation. Also, in order to apply these models, the extinctions and colonisations of the patches must be asynchronous.
1996:
1724:
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purposes, metapopulation models must include (a) the finite nature of metapopulations (how many patches are suitable for habitat), and (b) the
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2001:
1558:
959:
Fahrig, L. 2003. Effects of
Habitat Fragmentation on Biodiversity. Annual Review of ecology, evolution, and systematics. 34:1, p. 487.
2189:
1606:
1942:
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954:
498:. Alternative ecological strategies have evolved. For example, some salamanders forgo metamorphosis and sexually mature as aquatic
2021:
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of insect pests in agricultural fields, but the idea has been most broadly applied to species in naturally or artificially
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2006:
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Levins, R. (1969), "Some demographic and genetic consequences of environmental heterogeneity for biological control",
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1991:
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Bascompte J.; SolƩ R. V. (1996), "Habitat
Fragmentation and Extinction Thresholds in spatially explicit models",
416:
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82:
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Huffaker, C.B. (1958), "Experimental
Studies on Predation: Dispersion factors and predatorāprey oscillations",
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events); the smaller the population, the more chances of inbreeding depression and prone to extinction.
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They may also emigrate to a small population and rescue that population from extinction (called the
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521:
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350:
288:
90:
704:
Kareiva, P. (1987), "Habitat
Fragmentation and the Stability of PredatorāPrey Interactions",
386:
At equilibrium, therefore, some fraction of the species's habitat will always be unoccupied.
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2401:
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2256:
2174:
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1977:
1957:
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2149:
2129:
2011:
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1676:
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1112:
1038:
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effects take place in these configurations predicting more drastic extinction thresholds.
78:
may occur because declining populations leave niche opportunities open to the "rescuers".
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825:
717:
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1981:
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43:
745:
Janssen, A. et al. 1997. Metapopulation
Dynamics of a Persisting PredatorāPrey system.
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Metapopulation models have been used to explain life-history evolution, such as the
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2834:
2755:
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2139:
2084:
1967:
1932:
1809:
1308:
794:
733:
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121:
97:
21:
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The development of metapopulation theory, in conjunction with the development of
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2884:
2874:
2820:
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1819:
1548:
1366:
1328:
1303:
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531:
63:
411:. Metapopulations are particularly useful when discussing species in disturbed
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1714:
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Legendre, P.; Fortin, M.J. (1989), "Spatial pattern and ecological analysis",
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59:
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199:, each unoccupied patch can become occupied with a colonization probability
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113:
27:
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1952:
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Keymer J.E.; P. Galajda; C. Muldoon R. & R. Austin (November 2006).
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39:
54:. In Levins' own words, it consists of "a population of populations".
725:
938:
42:
which interact at some level. The term metapopulation was coined by
976:
768:
168:
be the fraction of patches occupied at a given time. During a time
38:
consists of a group of spatially separated populations of the same
457:
20:
930:
141:
2571:
1754:
1007:
427:. Simple modifications to this model can also incorporate for
963:
Levin S.A. (1974), "Dispersion and
Population Interactions",
203:. Accordingly, the time rate of change of occupied patches,
142:
Huffaker's experiments on predatorāprey interactions (1958)
454:
Microhabitat patches (MHPs) and bacterial metapopulations
16:
Group of separated yet interacting ecological populations
2567:
810:"Bacterial metapopulations in nanofabricated landscapes"
615:
Foundations of
Ecology: Classic papers with commentaries
183:
of the patches are unoccupied. Assuming a constant rate
407:, whereas the fundamental metapopulation processes are
104:
was an important contributor to metapopulation theory.
403:
One major drawback of the Levins model is that it is
353:
304:
216:
172:, each occupied patch can become unoccupied with an
2605:
2430:
2330:
2255:
2128:
2065:
1925:
1793:
1695:
1574:
1501:
1458:
1380:
1347:
1244:
1156:
1050:
375:
330:
287:This equation is mathematically equivalent to the
276:
138:heterogeneity lead to predatorāprey oscillations.
571:Bulletin of the Entomological Society of America
62:(fluctuations in population size due to random
627:
625:
623:
597:Kritzer, J. P. & Sale, P. F. (eds) (2006)
277:{\displaystyle {\frac {dN}{dt}}=cN(1-N)-eN.\,}
2583:
1766:
1019:
8:
564:
562:
617:. The University of Chicago Press, Chicago.
593:
591:
152:and in turn influence population dynamics.
2590:
2576:
2568:
1987:Latitudinal gradients in species diversity
1773:
1759:
1751:
1026:
1012:
1004:
613:Real, Leslie A. and Brown, James H. 1991.
843:
833:
776:
673:
645:
399:Stochastic patch occupancy models (SPOMs)
372:
352:
327:
317:
303:
273:
217:
215:
1885:Predatorāprey (LotkaāVolterra) equations
1524:Tritrophic interactions in plant defense
609:
607:
1917:Random generalized LotkaāVolterra model
558:
1725:Herbivore adaptations to plant defense
419:, i.e., how likely they are to become
161:his model is either populated or not.
7:
1740:Predator avoidance in schooling fish
331:{\displaystyle K=1-{\frac {e}{c}}\,}
2190:Intermediate disturbance hypothesis
1943:Ecological effects of biodiversity
14:
1279:Generalist and specialist species
462:E. coli metapopulation on a chip.
390:Stochasticity and metapopulations
2002:Occupancyāabundance relationship
999:Helsinki-science: Metapopulation
195:occupied patches, during a time
2022:Relative abundance distribution
1735:Plant defense against herbivory
1602:Competitive exclusion principle
1314:Mesopredator release hypothesis
949:Oxford University Press. 1999.
46:in 1969 to describe a model of
1607:Consumerāresource interactions
417:viability of their populations
258:
246:
179:. Additionally, 1 ā
1:
2453:Biological data visualization
2280:Environmental niche modelling
2007:Population viability analysis
537:Population viability analysis
1938:Density-dependent inhibition
191:generation from each of the
2407:Liebig's law of the minimum
2242:Resource selection function
1133:Metabolic theory of ecology
601:, Academic Press, New York.
291:, with a carrying capacity
124:in the 1930s, based on the
112:The first experiments with
58:consequence of demographic
2947:
2307:Niche apportionment models
2027:Relative species abundance
1231:Primary nutritional groups
1128:List of feeding behaviours
108:Predation and oscillations
2556:
2488:Ecosystem based fisheries
2100:Interspecific competition
1992:Minimum viable population
1850:Maximum sustainable yield
1835:Intraspecific competition
1830:Effective population size
1710:Anti-predator adaptations
1221:Photosynthetic efficiency
911:Journal of Animal Ecology
889:10.1007/s10682-006-9149-1
2478:Ecological stoichiometry
2443:Alternative stable state
527:LotkaāVolterra equations
376:{\displaystyle r=c-e.\,}
30:between the populations.
2322:Ontogenetic niche shift
2185:Ideal free distribution
2095:Ecological facilitation
1845:Malthusian growth model
1815:Consumer-resource model
1672:Paradox of the plankton
1637:Energy systems language
1357:Chemoorganoheterotrophy
1324:Optimal foraging theory
1299:Heterotrophic nutrition
965:The American Naturalist
835:10.1073/pnas.0607971103
757:The American Naturalist
647:10.3733/hilg.v27n14p343
492:amphibian metamorphosis
126:LotkaāVolterra equation
2468:Ecological forecasting
2412:Marginal value theorem
2210:Landscape epidemiology
2145:Cross-boundary subsidy
2080:Biological interaction
1430:Microbial intelligence
1118:Green world hypothesis
947:Metapopulation Ecology
599:Marine metapopulations
482:Life history evolution
463:
377:
332:
278:
174:extinction probability
102:University of Helsinki
31:
2473:Ecological humanities
2372:Ecological energetics
2317:Niche differentiation
2180:Habitat fragmentation
1948:Ecological extinction
1895:Small population size
1647:Feed conversion ratio
1627:Ecological succession
1559:San Francisco Estuary
1473:Ecological efficiency
1415:Microbial cooperation
583:10.1093/besa/15.3.237
547:Spatial heterogeneity
512:Competition (biology)
461:
437:habitat fragmentation
433:percolation threshold
378:
333:
279:
149:spatial heterogeneity
118:spatial heterogeneity
24:
2498:Evolutionary ecology
2463:Ecological footprint
2458:Ecological economics
2382:Ecological threshold
2377:Ecological indicator
2247:Sourceāsink dynamics
2200:Land change modeling
2195:Insular biogeography
2047:Species distribution
1786:Modelling ecosystems
1445:Microbial metabolism
1284:Intraguild predation
1073:Biogeochemical cycle
1039:Modelling ecosystems
869:Evolutionary Ecology
517:Conservation biology
488:ecological stability
476:evolutionary ecology
444:conservation biology
351:
302:
214:
83:sourceāsink dynamics
2606:Domains and methods
2548:Theoretical ecology
2523:Natural environment
2387:Ecosystem diversity
2357:Ecological collapse
2347:Bateman's principle
2302:Limiting similarity
2215:Landscape limnology
2037:Species homogeneity
1875:Population modeling
1870:Population dynamics
1687:Trophic state index
923:1996JAnEc..65..465B
881:2007EvEco..21..751P
826:2006PNAS..10317290K
718:1987Natur.326..388K
89:occupying discrete
52:fragmented habitats
48:population dynamics
2921:Population ecology
2652:Meta-communication
2559:Outline of ecology
2508:Industrial ecology
2503:Functional ecology
2367:Ecological deficit
2312:Niche construction
2275:Ecosystem engineer
2052:Speciesāarea curve
1973:Introduced species
1788:: Other components
1720:Deimatic behaviour
1622:Ecological network
1554:North Pacific Gyre
1539:hydrothermal vents
1478:Ecological pyramid
1425:Microbial food web
1236:Primary production
1181:Foundation species
684:10.1007/BF00048036
464:
373:
328:
274:
120:were conducted by
96:Finnish biologist
91:patches of habitat
32:
2926:Landscape ecology
2908:
2907:
2786:Meta-organization
2781:Meta-optimization
2565:
2564:
2448:Balance of nature
2205:Landscape ecology
2090:Community ecology
2032:Species diversity
1968:Indicator species
1963:Gradient analysis
1840:Logistic function
1748:
1747:
1705:Animal coloration
1682:Trophic mutualism
1420:Microbial ecology
1211:Photoheterotrophs
1196:Myco-heterotrophy
1108:Ecosystem ecology
1093:Carrying capacity
1058:Abiotic component
820:(46): 17290ā295.
712:(6111): 388ā390,
522:Landscape ecology
472:landscape ecology
325:
235:
135:population cycles
2938:
2801:Metaepistemology
2618:Metabibliography
2592:
2585:
2578:
2569:
2265:Ecological niche
2237:selection theory
2057:Umbrella species
2042:Species richness
1978:Invasive species
1958:Flagship species
1865:Population cycle
1860:Overexploitation
1825:Ecological yield
1775:
1768:
1761:
1752:
1657:Mesotrophic soil
1597:Climax community
1529:Marine food webs
1468:Biomagnification
1269:Chemoorganotroph
1123:Keystone species
1083:Biotic component
1028:
1021:
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726:10.1038/326388a0
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640:(343): 343ā383,
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341:and growth rate
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156:The Levins model
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2870:Meta-regulation
2845:Metaprogramming
2826:Metametaphysics
2746:Metamathematics
2601:
2596:
2566:
2561:
2552:
2538:Systems ecology
2426:
2397:Extinction debt
2362:Ecological debt
2352:Bioluminescence
2333:
2326:
2295:marine habitats
2270:Ecological trap
2251:
2131:
2124:
2067:
2061:
2017:Rapoport's rule
2012:Priority effect
1953:Endemic species
1921:
1880:Population size
1796:
1789:
1779:
1749:
1744:
1697:
1691:
1677:Trophic cascade
1587:Bioaccumulation
1570:
1497:
1454:
1376:
1343:
1240:
1152:
1113:Ecosystem model
1046:
1032:
995:
962:
908:
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904:
866:
865:
861:
807:
806:
802:
763:(5): 478ā4945.
754:
753:
749:
744:
740:
703:
702:
698:
675:10.1.1.330.8940
659:
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631:
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621:
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425:contact process
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5:
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2897:
2892:
2887:
2882:
2877:
2872:
2867:
2865:Meta-reference
2862:
2857:
2855:Metapsychology
2852:
2850:Metapsychiatry
2847:
2842:
2840:Metapopulation
2837:
2832:
2831:
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2829:
2828:
2818:
2813:
2808:
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2796:Metaphilosophy
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2788:
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2773:
2768:
2766:Metamotivation
2763:
2758:
2753:
2748:
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2729:Metapragmatics
2721:
2719:Meta-knowledge
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2603:
2602:
2597:
2595:
2594:
2587:
2580:
2572:
2563:
2562:
2557:
2554:
2553:
2551:
2550:
2545:
2540:
2535:
2530:
2525:
2520:
2518:Microecosystem
2515:
2510:
2505:
2500:
2495:
2490:
2485:
2480:
2475:
2470:
2465:
2460:
2455:
2450:
2445:
2440:
2434:
2432:
2428:
2427:
2425:
2424:
2419:
2417:Thorson's rule
2414:
2409:
2404:
2399:
2394:
2389:
2384:
2379:
2374:
2369:
2364:
2359:
2354:
2349:
2344:
2342:Assembly rules
2338:
2336:
2328:
2327:
2325:
2324:
2319:
2314:
2309:
2304:
2299:
2298:
2297:
2287:
2282:
2277:
2272:
2267:
2261:
2259:
2253:
2252:
2250:
2249:
2244:
2239:
2227:
2225:Patch dynamics
2222:
2220:Metapopulation
2217:
2212:
2207:
2202:
2197:
2192:
2187:
2182:
2177:
2172:
2167:
2162:
2157:
2152:
2147:
2142:
2136:
2134:
2126:
2125:
2123:
2122:
2117:
2115:Storage effect
2112:
2107:
2102:
2097:
2092:
2087:
2082:
2077:
2071:
2069:
2063:
2062:
2060:
2059:
2054:
2049:
2044:
2039:
2034:
2029:
2024:
2019:
2014:
2009:
2004:
1999:
1997:Neutral theory
1994:
1989:
1984:
1982:Native species
1975:
1970:
1965:
1960:
1955:
1950:
1945:
1940:
1935:
1929:
1927:
1923:
1922:
1920:
1919:
1914:
1913:
1912:
1907:
1897:
1892:
1887:
1882:
1877:
1872:
1867:
1862:
1857:
1855:Overpopulation
1852:
1847:
1842:
1837:
1832:
1827:
1822:
1817:
1812:
1807:
1801:
1799:
1791:
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1763:
1755:
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1743:
1742:
1737:
1732:
1727:
1722:
1717:
1712:
1707:
1701:
1699:
1693:
1692:
1690:
1689:
1684:
1679:
1674:
1669:
1664:
1662:Nutrient cycle
1659:
1654:
1652:Feeding frenzy
1649:
1644:
1639:
1634:
1632:Energy quality
1629:
1624:
1619:
1614:
1609:
1604:
1599:
1594:
1592:Cascade effect
1589:
1584:
1578:
1576:
1572:
1571:
1569:
1568:
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1566:
1561:
1556:
1551:
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1475:
1470:
1464:
1462:
1456:
1455:
1453:
1452:
1447:
1442:
1437:
1435:Microbial loop
1432:
1427:
1422:
1417:
1412:
1407:
1402:
1400:Lithoautotroph
1397:
1392:
1386:
1384:
1382:Microorganisms
1378:
1377:
1375:
1374:
1369:
1364:
1359:
1353:
1351:
1345:
1344:
1342:
1341:
1339:Prey switching
1336:
1331:
1326:
1321:
1316:
1311:
1306:
1301:
1296:
1291:
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1281:
1276:
1271:
1266:
1261:
1256:
1250:
1248:
1242:
1241:
1239:
1238:
1233:
1228:
1223:
1218:
1216:Photosynthesis
1213:
1208:
1203:
1198:
1193:
1188:
1183:
1178:
1173:
1171:Chemosynthesis
1168:
1162:
1160:
1154:
1153:
1151:
1150:
1145:
1140:
1135:
1130:
1125:
1120:
1115:
1110:
1105:
1100:
1095:
1090:
1085:
1080:
1075:
1070:
1065:
1063:Abiotic stress
1060:
1054:
1052:
1048:
1047:
1033:
1031:
1030:
1023:
1016:
1008:
1002:
1001:
994:
993:External links
991:
990:
989:
977:10.1086/282900
960:
957:
943:
917:(4): 465ā473,
903:
902:
875:(6): 751ā764,
859:
800:
769:10.1086/303407
747:
738:
696:
652:
619:
603:
587:
577:(3): 237ā240,
557:
556:
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551:
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539:
534:
529:
524:
519:
514:
507:
504:
483:
480:
468:nanotechnology
455:
452:
429:patch dynamics
400:
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371:
368:
365:
362:
359:
356:
339:
338:
324:
321:
316:
313:
310:
307:
289:logistic model
285:
284:
272:
269:
266:
263:
260:
257:
254:
251:
248:
245:
242:
239:
233:
230:
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157:
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143:
140:
109:
106:
44:Richard Levins
36:metapopulation
15:
13:
10:
9:
6:
4:
3:
2:
2943:
2932:
2929:
2927:
2924:
2922:
2919:
2918:
2916:
2901:
2898:
2896:
2893:
2891:
2888:
2886:
2883:
2881:
2880:Metasociology
2878:
2876:
2873:
2871:
2868:
2866:
2863:
2861:
2858:
2856:
2853:
2851:
2848:
2846:
2843:
2841:
2838:
2836:
2833:
2827:
2824:
2823:
2822:
2819:
2817:
2816:Meta-ontology
2814:
2812:
2809:
2807:
2804:
2802:
2799:
2798:
2797:
2794:
2792:
2791:Metaphenomics
2789:
2787:
2784:
2782:
2779:
2777:
2774:
2772:
2771:Metanarrative
2769:
2767:
2764:
2762:
2761:Metamodernism
2759:
2757:
2754:
2752:
2749:
2747:
2744:
2742:
2741:Metamaterials
2739:
2735:
2734:Metasemantics
2732:
2730:
2727:
2726:
2725:
2724:Meta-language
2722:
2720:
2717:
2715:
2712:
2710:
2707:
2705:
2704:Metaheuristic
2702:
2700:
2697:
2695:
2692:
2690:
2687:
2683:
2680:
2679:
2678:
2675:
2673:
2672:Metadiscourse
2670:
2668:
2665:
2663:
2660:
2658:
2657:Metacomputing
2655:
2653:
2650:
2646:
2643:
2641:
2640:Meta-learning
2638:
2637:
2636:
2635:Metacognition
2633:
2629:
2626:
2625:
2624:
2621:
2619:
2616:
2614:
2613:Meta-analysis
2611:
2610:
2608:
2604:
2600:
2593:
2588:
2586:
2581:
2579:
2574:
2573:
2570:
2560:
2555:
2549:
2546:
2544:
2543:Urban ecology
2541:
2539:
2536:
2534:
2531:
2529:
2526:
2524:
2521:
2519:
2516:
2514:
2511:
2509:
2506:
2504:
2501:
2499:
2496:
2494:
2491:
2489:
2486:
2484:
2481:
2479:
2476:
2474:
2471:
2469:
2466:
2464:
2461:
2459:
2456:
2454:
2451:
2449:
2446:
2444:
2441:
2439:
2436:
2435:
2433:
2429:
2423:
2420:
2418:
2415:
2413:
2410:
2408:
2405:
2403:
2402:Kleiber's law
2400:
2398:
2395:
2393:
2390:
2388:
2385:
2383:
2380:
2378:
2375:
2373:
2370:
2368:
2365:
2363:
2360:
2358:
2355:
2353:
2350:
2348:
2345:
2343:
2340:
2339:
2337:
2335:
2329:
2323:
2320:
2318:
2315:
2313:
2310:
2308:
2305:
2303:
2300:
2296:
2293:
2292:
2291:
2288:
2286:
2283:
2281:
2278:
2276:
2273:
2271:
2268:
2266:
2263:
2262:
2260:
2258:
2254:
2248:
2245:
2243:
2240:
2238:
2236:
2232:
2228:
2226:
2223:
2221:
2218:
2216:
2213:
2211:
2208:
2206:
2203:
2201:
2198:
2196:
2193:
2191:
2188:
2186:
2183:
2181:
2178:
2176:
2175:Foster's rule
2173:
2171:
2168:
2166:
2163:
2161:
2158:
2156:
2153:
2151:
2148:
2146:
2143:
2141:
2138:
2137:
2135:
2133:
2127:
2121:
2118:
2116:
2113:
2111:
2108:
2106:
2103:
2101:
2098:
2096:
2093:
2091:
2088:
2086:
2083:
2081:
2078:
2076:
2073:
2072:
2070:
2064:
2058:
2055:
2053:
2050:
2048:
2045:
2043:
2040:
2038:
2035:
2033:
2030:
2028:
2025:
2023:
2020:
2018:
2015:
2013:
2010:
2008:
2005:
2003:
2000:
1998:
1995:
1993:
1990:
1988:
1985:
1983:
1979:
1976:
1974:
1971:
1969:
1966:
1964:
1961:
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1956:
1954:
1951:
1949:
1946:
1944:
1941:
1939:
1936:
1934:
1931:
1930:
1928:
1924:
1918:
1915:
1911:
1908:
1906:
1903:
1902:
1901:
1898:
1896:
1893:
1891:
1888:
1886:
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1881:
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1868:
1866:
1863:
1861:
1858:
1856:
1853:
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1841:
1838:
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1833:
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1800:
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1625:
1623:
1620:
1618:
1615:
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1603:
1600:
1598:
1595:
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1552:
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1517:
1515:
1512:
1510:
1507:
1506:
1504:
1500:
1494:
1493:Trophic level
1491:
1489:
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1481:
1479:
1476:
1474:
1471:
1469:
1466:
1465:
1463:
1461:
1457:
1451:
1450:Phage ecology
1448:
1446:
1443:
1441:
1440:Microbial mat
1438:
1436:
1433:
1431:
1428:
1426:
1423:
1421:
1418:
1416:
1413:
1411:
1408:
1406:
1403:
1401:
1398:
1396:
1395:Bacteriophage
1393:
1391:
1388:
1387:
1385:
1383:
1379:
1373:
1370:
1368:
1365:
1363:
1362:Decomposition
1360:
1358:
1355:
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1352:
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1346:
1340:
1337:
1335:
1332:
1330:
1327:
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1309:Mesopredators
1307:
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1280:
1277:
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1267:
1265:
1262:
1260:
1257:
1255:
1254:Apex predator
1252:
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1234:
1232:
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1227:
1224:
1222:
1219:
1217:
1214:
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1202:
1199:
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1192:
1189:
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1149:
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1131:
1129:
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1119:
1116:
1114:
1111:
1109:
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1104:
1101:
1099:
1096:
1094:
1091:
1089:
1088:Biotic stress
1086:
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1079:
1076:
1074:
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1022:
1017:
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1009:
1006:
1000:
997:
996:
992:
986:
982:
978:
974:
970:
966:
961:
958:
956:
955:0-19-854065-5
952:
948:
944:
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936:
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928:
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770:
766:
762:
758:
751:
748:
742:
739:
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731:
727:
723:
719:
715:
711:
707:
700:
697:
693:
689:
685:
681:
676:
671:
667:
663:
662:Plant Ecology
656:
653:
648:
643:
639:
635:
628:
626:
624:
620:
616:
610:
608:
604:
600:
594:
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543:
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538:
535:
533:
530:
528:
525:
523:
520:
518:
515:
513:
510:
509:
505:
503:
501:
497:
493:
489:
481:
479:
477:
473:
469:
460:
453:
451:
449:
448:probabilistic
445:
440:
438:
434:
431:. At a given
430:
426:
422:
418:
414:
410:
406:
405:deterministic
398:
396:
389:
387:
369:
366:
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347:
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344:
322:
319:
314:
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308:
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298:
297:
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270:
267:
264:
261:
255:
252:
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210:
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139:
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131:
127:
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119:
115:
107:
105:
103:
99:
94:
92:
86:
84:
79:
77:
76:rescue effect
73:
72:rescue effect
67:
65:
61:
60:stochasticity
55:
53:
49:
45:
41:
37:
29:
23:
19:
2839:
2835:Metapolitics
2756:Metamodeling
2699:Metagenomics
2677:Meta-emotion
2628:Semantic Web
2528:Regime shift
2513:Macroecology
2234:
2230:
2219:
2170:Edge effects
2140:Biogeography
2085:Commensalism
1933:Biodiversity
1810:Allee effect
1549:kelp forests
1502:Example webs
1367:Detritivores
1206:Organotrophs
1186:Kinetotrophs
1138:Productivity
971:(960): 207,
968:
964:
946:
931:10.2307/5781
914:
910:
872:
868:
862:
817:
813:
803:
778:10533/172124
760:
756:
750:
741:
709:
705:
699:
665:
661:
655:
637:
633:
614:
598:
574:
570:
496:vernal ponds
485:
465:
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402:
393:
385:
342:
340:
292:
286:
204:
200:
196:
192:
184:
180:
176:
169:
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163:
159:
145:
130:oscillations
111:
98:Ilkka Hanski
95:
87:
80:
71:
68:
56:
35:
33:
18:
2890:Metatheorem
2885:Meta-system
2875:Metascience
2821:Metaphysics
2776:Meta-object
2709:Metahistory
2689:Metafiction
2165:Disturbance
2068:interaction
1890:Recruitment
1820:Depensation
1612:Copiotrophs
1483:Energy flow
1405:Lithotrophy
1349:Decomposers
1329:Planktivore
1304:Insectivore
1294:Heterotroph
1259:Bacterivore
1226:Phototrophs
1176:Chemotrophs
1148:Restoration
1098:Competition
945:Hanski, I.
532:Oscillation
122:G. F. Gause
64:demographic
2931:Population
2915:Categories
2895:Metatheory
2860:Metapuzzle
2806:Metaethics
2694:Metagaming
2667:Metadesign
2645:Metamemory
2533:Sexecology
2110:Parasitism
2075:Antibiosis
1910:Resistance
1905:Resilience
1795:Population
1715:Camouflage
1667:Oligotroph
1582:Ascendency
1544:intertidal
1534:cold seeps
1488:Food chain
1289:Herbivores
1264:Carnivores
1191:Mixotrophs
1166:Autotrophs
1045:components
668:(2): 107,
553:References
466:Combining
415:, and the
409:stochastic
74:). Such a
2900:Metaverse
2811:Metalogic
2751:Metamedia
2714:Metahumor
2623:Metaclass
2438:Allometry
2392:Emergence
2120:Symbiosis
2105:Mutualism
1900:Stability
1805:Abundance
1617:Dominance
1575:Processes
1564:tide pool
1460:Food webs
1334:Predation
1319:Omnivores
1246:Consumers
1201:Mycotroph
1158:Producers
1103:Ecosystem
1068:Behaviour
670:CiteSeerX
634:Hilgardia
542:Predation
494:in small
364:−
315:−
295:given by
262:−
253:−
189:propagule
114:predation
28:gene flow
2682:Metamood
2662:Metadata
2493:Endolith
2422:Xerosere
2334:networks
2150:Ecocline
1696:Defense,
1372:Detritus
1274:Foraging
1143:Resource
985:83630608
897:38832436
854:17090676
787:29587508
692:17101938
506:See also
500:neotenes
413:habitats
2483:Ecopath
2290:Habitat
2160:Ecotype
2155:Ecotone
2132:ecology
2130:Spatial
2066:Species
1926:Species
1797:ecology
1782:Ecology
1730:Mimicry
1698:counter
1642:f-ratio
1390:Archaea
1078:Biomass
1051:General
1043:Trophic
1035:Ecology
919:Bibcode
877:Bibcode
845:1635019
822:Bibcode
795:4385886
734:4335135
714:Bibcode
421:extinct
100:of the
40:species
1514:Rivers
1410:Marine
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1509:Lakes
981:S2CID
935:JSTOR
893:S2CID
791:S2CID
730:S2CID
688:S2CID
470:with
207:, is
205:dN/dt
1519:Soil
951:ISBN
939:5781
850:PMID
814:PNAS
783:PMID
442:For
201:cNdt
164:Let
116:and
973:doi
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927:doi
885:doi
840:PMC
830:doi
818:103
773:hdl
765:doi
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722:doi
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642:doi
579:doi
490:of
187:of
177:edt
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241:c
238:=
232:t
229:d
224:N
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193:N
185:c
181:N
166:N
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