Priority effect

{{Short description|Ecological effect of a species arriving first at a site}} {{multiple issues| {{Tone|date=April 2010}} {{Synthesis|date=April 2014}} }}

In ecology, a '''priority effect''' refers to the impact that a particular species can have on community development as a result of its prior arrival at a site.<ref name="tansley1935"/><ref name="clements1916"/><ref name="clements1936"/> There are two basic types of priority effects: inhibitory and facilitative. An inhibitory priority effect occurs when a species that arrives first at a site negatively affects a species that arrives later by reducing the availability of space or resources. In contrast, a facilitative priority effect occurs when a species that arrives first at a site alters abiotic or biotic conditions in ways that positively affect a species that arrives later.<ref name="clements1936"/><ref name="watt1947"/> Inhibitory priority effects have been documented more frequently than facilitative priority effects.{{Citation needed|date=April 2024}} Studies indicate that both abiotic (e.g., resource availability) and biotic (e.g., predation) factors can affect the strength of priority effects.{{Citation needed|date=April 2024}} . Priority effects are a central and pervasive element of ecological community development that have significant implications for natural systems and ecological restoration efforts.<ref name="clements1936"/><ref name="gleason1926"/>{{Citation needed|date=April 2024}}

==Theoretical foundation==

===Community succession theory=== Early in the 20th century, Frederic Clements and other plant ecologists suggested that ecological communities develop in a linear, directional manner towards a final, stable endpoint: the climax community.<ref name="clements1936"/> Clements indicated that a site's climax community would reflect local climate. He conceptualized the climax community as a "superorganism" that followed a defined developmental sequence.<ref name="clements1916"/>

Early ecological succession theory maintained that the directional shifts from one stage of succession to the next were induced by the plants themselves.<ref name="tansley1935"/> In this sense, succession theory implicitly recognized priority effects; the prior arrival of certain species had important impacts on future community composition. At the same time, the climax concept implied that species shifts were predetermined. This implies that a given species would always appear at the same point during the development of the climax community and have a predictable impact on community development.

This static view of priority effects remained essentially unchanged by the concept of patch dynamics, introduced by Alex Watt in 1947.<ref name="watt1947"/> Watt conceived of plant communities as dynamic "mechanisms" that followed predetermined succession cycles. He viewed succession as a process driven by facilitation, in which each species made local conditions more suitable for another species.

===Individualistic approach=== In 1926, Henry Gleason presented an alternative hypothesis in which plants were conceptualized as individuals rather than components of a superorganism.<ref name="gleason1926"/> This hypothesis suggested that the distribution of various species across the landscape reflected species-specific dispersal limitations and environmental requirements rather than predetermined associations among species. Gleason contested the idea of a predetermined climax community, recognizing that different colonizing species could produce alternative trajectories of community development. For example, initially identical ponds colonized by different species could develop through succession into very different communities.

The Initial Floristic Composition model was put forward by Frank Egler to describe community development in abandoned agricultural fields.<ref name="egler1954"/> According to this model, the set of species present in a field immediately after abandonment had strong influences on community development and final community composition.<ref name="young2001"/>

===Alternative stable states=== In the 1970s, it was suggested that natural communities could be characterized by multiple or alternative stable states.<ref name="lewontin1969"/><ref name="holling1973"/><ref name="may1977"/> Multiple stable state models suggested that the same environment could support several combinations of species.<ref name="gleason1926"/><ref name="egler1954"/> Theorists argued that historical context could play a central role in determining which stable state would be present at any given time. Robert May explained, "If there is a unique stable state, historical accidents are unimportant; if there are many alternative locally stable states, historical accidents can be of overriding significance."<ref name="may1977"/>

===Community assembly theory=== Assembly theory explains community development processes in the context of multiple stable states: it asks why a particular type of community developed when other stable community types are possible. In contrast to succession theory, assembly theory was developed largely by animal ecologists and explicitly incorporated historical context.<ref name="young2001"/>

In 1975, Jared Diamond<ref name="diamond1975"/> developed quantitative "assembly rules" to predict avian community composition on an archipelago. This approach emphasizes historical contingency and multiple stable states. Although the idea of deterministic community assembly initially drew criticism,<ref name="connor1979"/> the approach continued to gain support.<ref name="may1977"/><ref name="hughes1989"/> In 1991, Drake used an assembly model to demonstrate that different community types result from different sequences of species invasions.<ref name="drake1991"/> In this model, early invaders have major impacts on the invasion success of species that arrive later. Other modelling studies suggested that priority effects may be especially important when invasion frequency is low enough to allow species to become established before replacement,<ref name="lockwood1997"/> or when other factors that could drive assembly (e.g., competition, abiotic stress) are relatively unimportant.<ref name="weiher1995"/>

In a 1999 review, Belyea and Lancaster described three basic determinants of community assembly: dispersal constraints, environmental constraints, and internal dynamics.<ref name="belyea1999"/> They identified priority effects as a manifestation of the interaction between dispersal constraints and internal dynamics.

==Empirical evidence== Although early research focused on animals and aquatic systems, more recent{{when|date=April 2014}} studies have begun to examine terrestrial and plant-based priority effects.

===Marine=== Most of the earliest empirical evidence for priority effects came from studies on aquatic animals. Sutherland (1974) found that final community composition varied depending on the initial order of larval recruitment in a community of small marine organisms (sponges, tunicates, hydroids, and other species).<ref name="sutherland1974"/> Shulman (1983) found strong priority effects among coral reef fish.<ref name="shulman1983"/> The study found that prior establishment by a territorial damselfish reduced establishment rates of other fish. The authors also identified cross-trophic priority effects; prior establishment by a predator fish reduced establishment rates of prey fishes.

In the late 1980s, several studies examined priority effects in marine microcosms. Robinson and Dickerson (1987) found that priority effects were important in some cases, but suggested, "Being the first to invade a habitat does not guarantee success; there must be sufficient time for the early colonist to increase its population size for it to pre-empt further colonization."<ref name="robinson1987"/> Robinson and Edgemon (1988) later developed 54 communities of phytoplankton species by varying invasion order, rate, and timing. They found that although invasion order (priority effects) could explain a small fraction of the resulting variation in community composition, most of the variation was explained by changes in invasion rate and invasion timing.<ref name="robinson1988"/> These studies indicate that priority effects may not be the only or the most important historical factor affecting the trajectory of community development.

In a striking example of cross-trophic priority effects, Hart (1992) found that priority effects explain the maintenance of two alternate stable states in stream ecosystems. While a macroalga is dominant in some patches, sessile grazers maintain a "lawn" of small microalgae in others. If the sessile grazers colonize a patch first, they exclude the macroalga, and vice versa.<ref name="hart1992"/>

===Amphibian=== In two of the most commonly cited empirical studies on priority effects, Alford and Wilbur documented inhibitory and facilitative priority effects among toad larvae in experimental ponds.<ref name="alford1985"/><ref name="wilbur1985"/> They found that hatchlings of a toad species (''Bufo americanus'') exhibited higher growth and survivorship when introduced to a pond before those of a frog species (''Rana sphenocephala''). The frog larvae, however, did best when introduced after the toad larvae. Thus, prior establishment by the toad species facilitated the frog species, while prior establishment by the frog species inhibited the toad species. Studies on tree frogs have also documented both types of priority effects.<ref name="morin1987"/><ref name="warner1991"/> Morin (1987) also observed that priority effects became less important in the presence of a predatory salamander. He hypothesized that predation mediated priority effects by reducing competition between frog species.<ref name="morin1987"/> Studies on larval insects and frogs in water-filled tree holes and stumps found that abiotic factors such as space, resource availability, and toxin levels can also be important in mediating priority effects.<ref name="fincke1999"/><ref name="sunahara2002"/>

===Terrestrial=== Terrestrial studies on priority effects are rare, with most studies focusing on arthropods or grassland plant species. In a lab experiment, Shorrocks and Bingley (1994) showed that prior arrival increased survivorship for two species of fruit flies; each fly species had inhibitory impacts on the other.<ref name="shorrocks1994"/> A 1996 field study on desert spiders by Ehmann and MacMahon showed that the presence of species from one spider guild reduced establishment of spiders from a different guild.<ref name="ehmann1996"/> Palmer (2003) demonstrated that priority effects allowed a competitively subordinate ant species to avoid exclusion by a competitively dominant species.<ref name="palmer2003"/> If the competitively subordinate ants were able to colonize first, they altered their host tree’s morphology in ways that made it less suitable for other ant species. This study was especially important because it was able to identify a mechanism driving observed priority effects.

A study on two species of introduced grasses in Hawaiian woodlands found that the species with inferior competitive abilities may be able to persist through priority effects.<ref name="dantonio2001"/> At least three studies have come to similar conclusions about the coexistence of native and exotic grasses in California grassland ecosystems.<ref name="seabloom2003"/><ref name="corbin2004"/><ref name="lulow2006"/> If given time to establish, native species can successfully inhibit the establishment of exotics. The authors of the various studies attributed the prevalence of exotic grasses in California to the low seed production and relatively poor dispersal ability of native species.

==Emerging concepts==

===Long-term implications: convergence and divergence=== Although many studies have documented priority effects, the persistence of these effects over time often remains unclear. Young (2001) indicated that both convergence (in which "communities proceed towards a pre-disturbance state regardless of historical conditions") and divergence (in which historical factors continue to affect the long-term trajectory of community development) are present in nature.<ref name="young2001"/> Among studies of priority effects, both trends seem to have been observed.<ref name="lulow2004"/><ref name="hart1992"/> Fukami (2005) argued that a community could be both convergent and divergent at different levels of community organization. The authors studied experimentally assembled plant communities and found that while the identities of individual species remained unique across different community replicates, species traits generally became more similar.<ref name="fukami2005"/>

===Trophic ecology=== Some studies indicate that priority effects can occur across guilds<ref name="ehmann1996"/> or trophic levels.<ref name="hart1992"/> Such priority effects could have dramatic impacts on community composition and food web structure. Even intra-guild priority effects could have important consequences at multiple trophic levels if the affected species are associated with unique predator or prey species. Consider, for example, a plant species that is eaten by a host-specific herbivore. Priority effects that influence the ability of the plant species to establish would indirectly affect the establishment success of the associated herbivore. Theoretical models have described cyclical assembly dynamics in which species associated with different suites of predators can repeatedly replace one another.<ref name="leibold2004"/><ref name="steiner2004"/>

===Intra-specific aggregation=== In situations where two species are introduced at the same time, spatial aggregation of a species' propagules could cause priority effects by initially reducing interspecific competition.<ref name="inouye1999"/> Aggregation during recruitment and establishment could allow inferior competitors to coexist with or even displace competitive dominants over the long-term. Several modelling efforts have begun to examine the implications of spatial priority effects for species coexistence.<ref name="shorrocks1994"/><ref name="chesson2000"/><ref name="hartley2002"/><ref name="molofsky2002"/>

===Mechanisms and new organisms=== A few studies have begun to explore the mechanisms driving observed priority effects.<ref name="palmer2003"/> Moreover, although past studies focused on a small subset of species, recent papers indicate that priority effects may be important for a wide range of organisms, including fungi,<ref name="kennedy2005"/><ref name="rohlfs2005"/> birds,<ref name="gamarra2005"/> lizards,<ref name="mcloskey1998"/> and salamanders.<ref name="eitam2005"/>

===Ecological restoration=== Priority effects have important implications for ecological restoration. In many systems, information about priority effects can help practitioners identify cost-effective strategies for improving the survival and persistence of certain species, especially species of inferior competitive ability.<ref name="lulow2004"/><ref name="young2005"/><ref name="suding2004"/> For example, in a study on the restoration of native Californian grasses and forbs, Lulow (2004) found that forbs could not establish in plots where bunchgrasses had been previously planted. When bunchgrasses were added to plots where forbs had already been growing for a year, forbs were able to coexist with grasses for at least 3–4 years.<ref name="lulow2004"/>

==References== {{reflist|1=2|refs= <ref name="alford1985">{{cite journal | last1=Alford | first1=Ross A. | last2=Wilbur | first2=Henry M. | title=Priority Effects in Experimental Pond Communities: Competition between Bufo and Rana | journal=Ecology | date=1985-08-01 | doi=10.2307/1939161 | issn=0012-9658 | volume=66 | issue=4 | pages=1097–1105 | jstor=1939161 | bibcode=1985Ecol...66.1097A }}</ref> <ref name="belyea1999">{{cite journal | last1=Belyea | first1=Lisa R. | last2=Lancaster | first2=Jill | title=Assembly Rules within a Contingent Ecology | journal=Oikos |date=September 1999| jstor=3546646 | doi=10.2307/3546646 | issn=0030-1299 | volume=86 | issue=3 | pages=402 | bibcode=1999Oikos..86..402B }}</ref> <ref name="chesson2000">{{cite journal | last=Chesson | first=Peter | title=Mechanisms of Maintenance of Species Diversity | journal=Annual Review of Ecology and Systematics | year=2000 | doi=10.1146/annurev.ecolsys.31.1.343 | volume=31 | issue=1 | pages=343–366| bibcode=2000AnRES..31..343C | s2cid=403954}}</ref> <ref name="clements1916">{{cite book | last=Clements | first=Frederic Edward | title=Plant Succession: An Analysis of the Development of Vegetation | url=https://archive.org/details/in.ernet.dli.2015.351403 | year=1916 | publisher=Carnegie Institution of Washington}}</ref> <ref name="clements1936">{{cite journal | last=Clements | first=Frederic E. | title=Nature and Structure of the Climax | journal=The Journal of Ecology |date=February 1936| url=http://www.bgu.ac.il/desert_agriculture/Vegecology/Papers/Clements36.pdf | doi=10.2307/2256278 | issn=0022-0477 | volume=24 | issue=1 | pages=252–284 | jstor=2256278 | bibcode=1936JEcol..24..252C }}</ref> <ref name="connor1979">{{cite journal | last1=Connor | first1=Edward F. | last2=Simberloff | first2=Daniel | title=The Assembly of Species Communities: Chance or Competition? | journal=Ecology |date=December 1979| jstor=1936961 | doi=10.2307/1936961 | issn=0012-9658 | volume=60 | issue=6 | pages=1132 | bibcode=1979Ecol...60.1132C }}</ref> <ref name="corbin2004">{{cite journal | last1=Corbin | first1=Jeffrey D. | last2=D'Antonio | first2=Carla M. | title=Competition Between Native Perennial and Exotic Annual Grasses: Implications for a Historical Invasion | journal=Ecology | date=2004-05-01 | url=http://www.elkhornsloughctp.org/uploads/files/1108491037Corbin04.pdf | doi=10.1890/02-0744 | issn=0012-9658 | volume=85 | issue=5 | pages=1273–1283 | bibcode=2004Ecol...85.1273C }}</ref> <ref name="dantonio2001">{{cite journal | last1=D'Antonio | first1=Carla M. | last2=Hughes | first2=R. Flint | last3=Vitousek | first3=Peter M. | title=Factors Influencing Dynamics of two Invasive C4 Grasses in Seasonally Dry Hawaiian Woodlands | journal=Ecology | date=2001-01-01 | doi=10.1890/0012-9658(2001)082[0089:FIDOTI]2.0.CO;2 | issn=0012-9658 | volume=82 | issue=1 | pages=89–104}}</ref> <ref name="diamond1975">{{cite book | author=Diamond, Jared M. | editor-last=Cody | editor-first=Martin L. | editor2-last=Diamond | editor2-first=Jared M. | title=Ecology and Evolution of Communities | date=1975-01-01 | publisher=Harvard University Press | isbn=9780674224445 | chapter=Assembly of species communities | pages=[https://archive.org/details/ecologyevolution00gres/page/342 342–444] | chapter-url-access=registration | chapter-url=https://archive.org/details/ecologyevolution00gres/page/342}}</ref> <ref name="drake1991">{{cite journal | last=Drake | first=James A | title=Community-assembly mechanics and the structure of an experimental species ensemble | journal=American Naturalist | year=1991 | issn=0003-0147 | pages=1–26 | doi = 10.1086/285143 | jstor=2462154 | volume=137| issue=1 | bibcode=1991ANat..137....1D | s2cid=53478943}}</ref> <ref name="egler1954">{{cite journal | last=Egler | first=Frank E. | title=Vegetation science concepts I. Initial floristic composition, a factor in old-field vegetation development with 2 figs. | journal=Vegetatio | date=1954-11-01 | issn=1573-5052 | volume=4 | issue=6 | pages=412–417 | doi=10.1007/BF00275587| s2cid=44573990}}</ref> <ref name="ehmann1996">{{cite journal | last1=Ehmann | first1=William J | last2=MacMahon | first2=James A | title=Initial tests for priority effects among spiders that co-occur on sagebrush shrubs | journal=Journal of Arachnology | year=1996 | issn=0161-8202 | volume=24 | issue=3 | pages=173–185}}</ref> <ref name="eitam2005">{{cite journal | last1=Eitam | first1=Avi | last2=Blaustein | first2=Leon | last3=Mangel | first3=Marc | title=Density and intercohort priority effects on larval Salamandra salamandra in temporary pools | journal=Oecologia | date=2005-11-01 | citeseerx = 10.1.1.118.8240 | doi=10.1007/s00442-005-0185-2 | pmid=16133198 | issn=0029-8549 | volume=146 | issue=1 | pages=36–42| bibcode=2005Oecol.146...36E | s2cid=14645093}}</ref> <ref name="fincke1999">{{cite journal | last=Fincke | first=OlA. M. | title=Organization of predator assemblages in Neotropical tree holes: effects of abiotic factors and priority | journal=Ecological Entomology | date=1999-02-01 | url=https://www.researchgate.net/publication/227768223 | doi=10.1046/j.1365-2311.1999.00166.x | issn=1365-2311 | volume=24 | issue=1 | pages=13–23| bibcode=1999EcoEn..24...13F | s2cid=84330573}}</ref> <ref name="fukami2005">{{cite journal | last1=Fukami | first1=Tadashi | last2=Martijn Bezemer | first2=T. | last3=Mortimer | first3=Simon R. | last4=van der Putten | first4=Wim H. | title=Species divergence and trait convergence in experimental plant community assembly | journal=Ecology Letters | date=2005-12-01 | url=http://xa.yimg.com/kq/groups/22969879/1652750489/name/Fukamy+ey+al.+2005.pdf | doi=10.1111/j.1461-0248.2005.00829.x | issn=1461-0248 | volume=8 | issue=12 | pages=1283–1290 | bibcode=2005EcolL...8.1283F }}</ref> <ref name="gamarra2005">{{cite journal | last1=Gamarra | first1=Javier G. P. | last2=Montoya | first2=José M. | last3=Alonso | first3=David | last4=Solé | first4=Ricard V. | title=Competition and introduction regime shape exotic bird communities in Hawaii | journal=Biological Invasions | date=2005-03-01 | url=https://www.researchgate.net/publication/226526326 | doi=10.1007/s10530-004-0876-3 | issn=1573-1464 | volume=7 | issue=2 | pages=297–307| bibcode=2005BiInv...7..297G | s2cid=23347191}}</ref> <ref name="gleason1926">{{cite journal | last=Gleason | first=H. A. | title=The Individualistic Concept of the Plant Association | journal=Bulletin of the Torrey Botanical Club | date=January 1926 | url=http://www.planta.cn/forum/files_planta/the_individualistic_concept_of_the_plant_association_857.pdf | doi=10.2307/2479933 | issn=0040-9618 | volume=53 | issue=1 | pages=7–26 | jstor=2479933 | access-date=2014-04-28 | archive-url=https://web.archive.org/web/20140429050302/http://www.planta.cn/forum/files_planta/the_individualistic_concept_of_the_plant_association_857.pdf | archive-date=2014-04-29 | url-status=dead}}</ref> <ref name="hart1992">{{cite journal | last=Hart | first=David D. | title=Community organization in streams: the importance of species interactions, physical factors, and chance | journal=Oecologia | date=1992-08-01 | url=http://rydberg.biology.colostate.edu/bz471/readings/Hart.pdf | doi=10.1007/BF00317787 | pmid=28313460 | issn=1432-1939 | volume=91 | issue=2 | pages=220–228| bibcode=1992Oecol..91..220H | s2cid=206774636}}</ref> <ref name="hartley2002">{{cite journal | last1=Hartley | first1=Stephen | last2=Shorrocks | first2=Bryan | title=A general framework for the aggregation model of coexistence | journal=Journal of Animal Ecology | date=2002-07-01 | doi=10.1046/j.1365-2656.2002.00628.x | issn=1365-2656 | volume=71 | issue=4 | pages=651–662| doi-access=free | bibcode=2002JAnEc..71..651H }}</ref> <ref name="holling1973">{{cite journal | last=Holling | first=Crawford S | title=Resilience and stability of ecological systems | journal=Annual Review of Ecology and Systematics | volume=4 | year=1973 | issue=1 | issn=0066-4162 | pages=1–23 | jstor=2096802|doi=10.1146/annurev.es.04.110173.000245| bibcode=1973AnRES...4....1H | s2cid=53309505 | url=http://pure.iiasa.ac.at/26/1/RP-73-003.pdf}}</ref> <ref name="hughes1989">{{cite journal | last=Hughes | first=Terence P. | title=Community Structure and Diversity of Coral Reefs: The Role of History | journal=Ecology |date=February 1989| url=http://izt.ciens.ucv.ve/ecologia/Archivos/ECOLOGIA_DE%20_POBLACIONES_Hasta%202004/ECOL_POBLAC_Hasta%202004_%28H-N%29/Hughes%201989.pdf | doi=10.2307/1938434 | issn=0012-9658 | volume=70 | issue=1 | pages=275–279| jstor=1938434 | bibcode=1989Ecol...70..275H }}</ref> <ref name="inouye1999">{{cite journal | last=Inouye | first=Brian D. | title=Integrating nested spatial scales: implications for the coexistence of competitors on a patchy resource | journal=Journal of Animal Ecology | date=1999-01-01 | doi=10.1046/j.1365-2656.1999.00272.x | issn=1365-2656 | volume=68 | issue=1 | pages=150–162| doi-access=free | bibcode=1999JAnEc..68..150I }}</ref> <ref name="kennedy2005">{{cite journal | last1=Kennedy | first1=Peter G. | last2=Bruns | first2=Thomas D. | title=Priority effects determine the outcome of ectomycorrhizal competition between two Rhizopogon species colonizing Pinus muricata seedlings | journal=New Phytologist | date=2005-05-01 | doi=10.1111/j.1469-8137.2005.01355.x | issn=1469-8137 | volume=166 | issue=2 | pages=631–638 | pmid=15819925| doi-access=free | bibcode=2005NewPh.166..631K }}</ref> <ref name="leibold2004">{{cite journal | last1=Leibold | first1=M. A. | last2=Holyoak | first2=M. | last3=Mouquet | first3=N. | last4=Amarasekare | first4=P. | last5=Chase | first5=J. M. | last6=Hoopes | first6=M. F. | last7=Holt | first7=R. D. | last8=Shurin | first8=J. B. | last9=Law | first9=R. | last10=Tilman | first10=D. | last11=Loreau | first11=M. | last12=Gonzalez | first12=A. | title=The metacommunity concept: a framework for multi-scale community ecology | journal=Ecology Letters | date=2004-07-01 | doi=10.1111/j.1461-0248.2004.00608.x | issn=1461-0248 | volume=7 | issue=7 | pages=601–613| doi-access= | bibcode=2004EcolL...7..601L }}</ref> <ref name="lewontin1969">{{cite journal | last=Lewontin | first=R C | title=The meaning of stability | journal=Brookhaven Symposia in Biology | year=1969 | pmid=5372787 | issn=0068-2799 | volume=22 | pages=13–24}}</ref> <ref name="lockwood1997">{{cite journal | last1=Lockwood | first1=Julie L. | last2=Powell | first2=Robert D. | last3=Nott | first3=M. Philip | last4=Pimm | first4=Stuart L. | title=Assembling Ecological Communities in Time and Space | journal=Oikos |date=December 1997| url=https://www.researchgate.net/publication/229068825 | doi=10.2307/3546628 | issn=0030-1299 | volume=80 | issue=3 | pages=549 | jstor=3546628 | bibcode=1997Oikos..80..549L }}</ref> <ref name="lulow2004">{{cite thesis | last=Lulow | first=Megan Elizabeth | title=Restoration of California's Inland Grasslands: The Role of Priority Effects and Management Strategies in Establishing Native Communities and the Ability of Native Grasses to Resist Invasion by Non-native Grasses | year=2004 | publisher=University of California, Davis | type=Ph.D.}}</ref> <ref name="lulow2006">{{cite journal | last=Lulow | first=Megan E. | title=Invasion by Non-Native Annual Grasses: The Importance of Species Biomass, Composition, and Time Among California Native Grasses of the Central Valley | journal=Restoration Ecology | date=2006-12-01 | url=http://tpyoung.ucdavis.edu/publications/LulowRestEcol06.pdf | doi=10.1111/j.1526-100X.2006.00173.x | issn=1526-100X | volume=14 | issue=4 | pages=616–626| bibcode=2006ResEc..14..616L | s2cid=12460270}}</ref> <ref name="may1977">{{cite journal | last=May | first=Robert M | title=Thresholds and breakpoints in ecosystems with a multiplicity of stable states | journal=Nature | year=1977 | volume=269 | issue=5628 | pages=471–477 | doi=10.1038/269471a0| bibcode=1977Natur.269..471M | s2cid=4283750}}</ref> <ref name="mcloskey1998">{{cite journal | last1=M'Closkey | first1=Robert T. | last2=Hecnar | first2=Stephen J. | last3=Chalcraft | first3=David R. | last4=Cotter | first4=Jill E. | title=Size distributions and sex ratios of colonizing lizards | journal=Oecologia | date=1998-10-01 | url=https://www.researchgate.net/publication/225388393 | doi=10.1007/s004420050615 | pmid=28307519 | issn=1432-1939 | volume=116 | issue=4 | pages=501–509| bibcode=1998Oecol.116..501M | s2cid=23042707}}</ref> <ref name="molofsky2002">{{cite journal | last1=Molofsky | first1=Jane | last2=Bever | first2=James D. | title=A novel theory to explain species diversity in landscapes: positive frequency dependence and habitat suitability | journal=Proceedings of the Royal Society B: Biological Sciences | date=2002-12-07 | pmid=12495479 | doi=10.1098/rspb.2002.2164 | issn=1471-2954 | volume=269 | issue=1508 | pages=2389–2393 | pmc=1691177 | bibcode=2002PBioS.269.2389M }}</ref> <ref name="morin1987">{{cite journal | last=Morin | first=Peter Jay | title=Predation, Breeding Asynchrony, and the Outcome of Competition Among Treefrog Tadpoles | journal=Ecology | date=1987-06-01 | doi=10.2307/1938473 | issn=0012-9658 | volume=68 | issue=3 | pages=675–683| jstor=1938473 | bibcode=1987Ecol...68..675M }}</ref> <ref name="palmer2003">{{cite journal | last1=Palmer | first1=Todd M. | last2=Stanton | first2=Maureen L. | last3=Young | first3=Truman P. | title=Competition and Coexistence: Exploring Mechanisms That Restrict and Maintain Diversity within Mutualist Guilds | journal=The American Naturalist |date=October 2003| doi=10.1086/378682 | issn=1537-5323 | volume=162 | issue=s4 | pages=S63–S79 | jstor=10.1086/378682 | pmid=14583858| bibcode=2003ANat..162S..63P | s2cid=28047182}}</ref> <ref name="robinson1987">{{cite journal | last1=Robinson | first1=James F. | last2=Dickerson | first2=Jaime E. | title=Does Invasion Sequence Affect Community Structure? | journal=Ecology | date=1987-06-01 | doi=10.2307/1938464 | issn=0012-9658 | volume=68 | issue=3 | pages=587–595| jstor=1938464 | bibcode=1987Ecol...68..587R }}</ref> <ref name="robinson1988">{{cite journal | last1=Robinson | first1=James V. | last2=Edgemon | first2=Michael A. | title=An Experimental Evaluation of the Effect of Invasion History on Community Structure | journal=Ecology |date=October 1988| doi=10.2307/1941638 | issn=0012-9658 | volume=69 | issue=5 | pages=1410–1417| jstor=1941638}}</ref> <ref name="rohlfs2005">{{cite journal | last=Rohlfs | first=Marko | title=Density-dependent insect-mold interactions: effects on fungal growth and spore production | journal=Mycologia | date=2005-09-01 | url=http://www.mycologia.org/content/97/5/996 | pmid=16596951 | doi=10.3852/mycologia.97.5.996 | issn=1557-2536 | volume=97 | issue=5 | pages=996–1001| url-access=subscription }}</ref> <ref name="seabloom2003">{{cite journal | last1=Seabloom | first1=Eric W. | last2=Harpole | first2=W. Stanley | last3=Reichman | first3=O. J. | last4=Tilman | first4=David | title=Invasion, competitive dominance, and resource use by exotic and native California grassland species | journal=Proceedings of the National Academy of Sciences | date=2003-11-11 | pmid=14595028 | doi=10.1073/pnas.1835728100 | issn=1091-6490 | volume=100 | issue=23 | pages=13384–13389 | pmc=263823| bibcode=2003PNAS..10013384S | doi-access=free}}</ref> <ref name="shorrocks1994">{{cite journal | last1=Shorrocks | first1=B. | last2=Bingley | first2=M. | title=Priority effects and species coexistence: experiments with fungal-breeding Drosophila | journal=Journal of Animal Ecology | year=1994 | issn=0021-8790 | volume=63 | issue=4 | pages=799–806 | doi=10.2307/5257| jstor=5257 | bibcode=1994JAnEc..63..799S }}</ref> <ref name="shulman1983">{{cite journal | last1=Shulman | first1=Myra J. | last2=Ogden | first2=John C. | last3=Ebersole | first3=John P. | last4=McFarland | first4=William N. | last5=Miller | first5=Steven L. | last6=Wolf | first6=Nancy G. | title=Priority Effects in the Recruitment of Juvenile Coral Reef Fishes | journal=Ecology | date=1983-12-01 | doi=10.2307/1937505 | issn=0012-9658 | volume=64 | issue=6 | pages=1508–1513| jstor=1937505 | bibcode=1983Ecol...64.1508S }}</ref> <ref name="steiner2004">{{cite journal | last1=Steiner | first1=Christopher F. | last2=Leibold | first2=Mathew A. | title=Cyclic assembly trajectories and scale-dependent productivity–diversity relationships | journal=Ecology | date=2004-01-01 | url=http://bio.wayne.edu/profhtml/ChrisSteiner/Pubs/LeiboldAssembly2004.pdf | doi=10.1890/03-3010 | issn=0012-9658 | volume=85 | issue=1 | pages=107–113 | bibcode=2004Ecol...85..107S }}</ref> <ref name="suding2004">{{cite journal | last1=Suding | first1=Katharine N. | last2=Gross | first2=Katherine L. | last3=Houseman | first3=Gregory R. | title=Alternative states and positive feedbacks in restoration ecology | journal=Trends in Ecology & Evolution | date=2004-01-01 | url=http://bio.fsu.edu/miller/docs/Suding_et%20al._2004.pdf | pmid=16701225 | doi=10.1016/j.tree.2003.10.005 | issn=0169-5347 | volume=19 | issue=1 | pages=46–53 | access-date=2014-04-28 | archive-url=https://web.archive.org/web/20140429080206/http://bio.fsu.edu/miller/docs/Suding_et%20al._2004.pdf | archive-date=2014-04-29 | url-status=dead}}</ref> <ref name="sunahara2002">{{cite journal | last1=Sunahara | first1=Toshihiko | last2=Mogi | first2=Motoyoshi | title=Priority effects of bamboo-stump mosquito larvae: influences of water exchange and leaf litter input | journal=Ecological Entomology | date=2002-06-01 | doi=10.1046/j.1365-2311.2002.00417.x | issn=1365-2311 | volume=27 | issue=3 | pages=346–354| bibcode=2002EcoEn..27..346S | s2cid=85188993}}</ref> <ref name="sutherland1974">{{cite journal | last=Sutherland | first=John P | title=Multiple stable points in natural communities | journal=American Naturalist | year=1974 | issn=0003-0147 | volume=108 | issue=964 | pages=859–873 | jstor=2459615 | doi=10.1086/282961| bibcode=1974ANat..108..859S | s2cid=85014132}}</ref> <ref name="tansley1935">{{cite journal | last=Tansley | first=A. G. | title=The Use and Abuse of Vegetational Concepts and Terms | journal=Ecology |date=July 1935| url=http://karljaspers.org/files/tansley.pdf | doi=10.2307/1930070 | issn=0012-9658 | volume=16 | issue=3 | pages=284–307 | jstor=1930070 | bibcode=1935Ecol...16..284T | url-status=dead | archive-url=https://web.archive.org/web/20110726213308/http://karljaspers.org/files/tansley.pdf | archive-date=2011-07-26}}</ref> <ref name="warner1991">{{cite journal | last1=Warner | first1=Susan C. | last2=Dunson | first2=William A. | last3=Travis | first3=Joseph | title=Interaction of pH, density, and priority effects on the survivorship and growth of two species of hylid tadpoles | journal=Oecologia | date=1991-11-01 | doi=10.1007/BF00317575 | pmid=28313793 | issn=1432-1939 | volume=88 | issue=3 | pages=331–339| bibcode=1991Oecol..88..331W | s2cid=19276988}}</ref> <ref name="watt1947">{{cite journal | last=Watt | first=Alex S. | title=Pattern and Process in the Plant Community | journal=The Journal of Ecology |date=December 1947| url=http://faculty.washington.edu/edford/Watt_1947.pdf | doi=10.2307/2256497 | issn=0022-0477 | volume=35 | issue=1/2 | pages=1–22 | jstor=2256497 | bibcode=1947JEcol..35....1W }}</ref> <ref name="weiher1995">{{cite journal | author=Weiher, E. | author2 = Keddy, P. A. | year=1995 | title=Assembly rules, null models, and trait dispersion: new questions front old patterns | journal=Oikos | volume=74 | issue = 1 | pages=159–164 | doi=10.2307/3545686| jstor = 3545686 | bibcode = 1995Oikos..74..159W }}</ref> <ref name="wilbur1985">{{cite journal | last1=Wilbur | first1=Henry M. | last2=Alford | first2=Ross A. | title=Priority Effects in Experimental Pond Communities: Responses of Hyla to Bufo and Rana | journal=Ecology |date=August 1985| doi=10.2307/1939162 | issn=0012-9658 | volume=66 | issue=4 | pages=1106–1114| jstor=1939162 | bibcode=1985Ecol...66.1106W }}</ref> <ref name="young2001">{{cite journal | last1=Young | first1=Truman P. | last2=Chase | first2=Jonathan M. | last3=Huddleston | first3=Russell T. | title=Community Succession and Assembly Comparing, Contrasting and Combining Paradigms in the Context of | journal=Ecological Restoration | date=2001-03-20 | url=http://tpyoung.ucdavis.edu/publications/2001SuccAssEcolRest.pdf | doi=10.3368/er.19.1.5 | issn=1543-4079 | volume=19 | issue=1 | pages=5–18| s2cid=16235361}}</ref> <ref name="young2005">{{cite journal | last1=Young | first1=T. P. | last2=Petersen | first2=D. A. | last3=Clary | first3=J. J. | title=The ecology of restoration: historical links, emerging issues and unexplored realms | journal=Ecology Letters | date=2005-06-01 | url=http://globalrestorationnetwork.org/uploads/files/LiteratureAttachments/320_the-ecology-of-restoration---historical-links-emerging-issues-and-unexplored-realms.pdf | doi=10.1111/j.1461-0248.2005.00764.x | issn=1461-0248 | volume=8 | issue=6 | pages=662–673| doi-access=free | bibcode=2005EcolL...8..662Y }}</ref> }}

{{modelling ecosystems|expanded=other}}

{{Improve categories|date=October 2023}}

Category:Ecology