DOI: 10.3724/SP.J.1003.2009.09141

Biodiversity Science (生物多样性) 2009/17:6 PP.605-612

Local and regional processes control species richness of plant communities: the species pool hypothesis

Exploring the mechanisms underlying community species richness is a key issue in ecology and conservation biology,and many hypotheses based on small-scale,local processes have traditionally been used as explanations.The species pool hypothesis developed by Zobel et al.suggests that the variation in community species richness is not only associated with contemporary environmental factors and ecological processes(e.g.competition and predation),but also limited by the regional species pool.The regional species pool is the set of species in a certain region that are capable of coexisting in a target community,which is shaped by historical(e.g.glaciation and geological age)and regional processes(e.g.speciation,immigration,dispersion,and extinction).The species pool hypothesis suggests that the larger the area of a habitat type and the greater its geological age,the greater the opportunity for speciation and hence the larger the number of available species adapted to that particular habitat,which will in turn lead to higher community diversity.The species pool is generally studied at two spatial scales:the regional and the actual scales.While the regional species pool is primarily determined by biogeographic processes,the actual species pool(species present inthe target community)is determined by both ecological processes(e.g.competition)and the regional pool.In this review,we introduce and discuss the concepts relating to,and evidence for the species pool hypothesis,together with methods for estimating the species pool.

Key words:competition,disturbance,species richness,species pool,habimt,community productivity,localprocess,regional process

ReleaseDate:2015-03-12 14:28:06

Begon M,Townsend CA,Harper JL(2006)Ecology:From Individuals to Ecosystems.Blackwell Publishing,Oxford.

Brudvig LA,Mabry CM(2008)Trait-based filtering of the regional species pool to guide understOry plant reintroductions in midwestem oak savannas,U.S.A.Restoration Ecology,16,290-304.

Chytry M,Danihelka J,Ermakov N,Haiek M,Hajkova P,Koei M,Kubesova S,Lustyk P,Otypkova Z,Popov D,Roleeek J,Reznickova M,Smarda P,Valachovic M(2007)Plant species richness in continental southern Siberia:effects of pH and climate in the context of the species pool hypothesis.Global Ecology and Biogeography,16,668-678.

Cornell HV,Lawton JH(1992)Species interactions,local and regional processes,and limits to the richness of ecological communities:a theoreticaI perspective.dournal of Animal Ecology,61,1-12.

Cornwell WK,Grubb PJ(2003)Regional and local pauerns in plant species richness with respect to resource availability.Oikos,100,417-428.

Dupr(e) C(2000)How to determine a regional species pool:a study in two Swedish regions.Oikos,89,128-136.

Dupr(e) C,Wessberg C,Diekmann M(2002)Species richness in deciduous forests:effects of species pools and environmental vailables.Journal of Vegetation Science,13,505-516.

Ellenberg H,Weber HE,Duell R,Wirth V,Wemer W,PaulisSen D(1991)Indicator Values of Plants in Central Europe.Goltze,Goettingen.

Eriksson O(1993)The species-pool hypothesis and plant community diversity.Oikos,68,371-374.

Ewald J(2003)The calcareous riddle:why are there so many calciphilous species in the central European flora?Folio Geobotanica,38,357-366.

Gaston KJ(2000)Global patterns in biodiversity.Nature,405,220-227.

Goldberg DE,Rajaniemi T,Gurevitch J,Stewart-Oaten A(1999)Empirical approaches to quantifying interaction intensity:competition and facilitation along productivity gradient.Ecology,80,1118-1131.

Grace JB(1999)The factors controlling species density in herbaceous plant communities:an assessment.Perspectives in Plant Ecology,Evolution and Systematics,2,1-28.

Grace JB(2001)Difficulties with estimating and interpreting species pools and the implications for understanding patterns of diversity.Folia Geobotanica,36,71-83.

Grime JP(1973a)Competitive exclusion in herbaceous vegetation.Nature,242,344-347.

Grime JP(1973b)Control of species diversity in herbaceous vegetation.Journal of Environmental Management,1,151-167.

Grime JP(1979)Plant Strategies and Vegetation Processes.John Wiley & Sons.London.

Grubb PJ(1987)Global trends in species-richness in terrestrial vegetation:a view from the northem hemisphere.In:Organization of Communities:Past and Present(eds Gee JHR,Giller PS),pp.99-118.Blackwell Scientific Publications,Oxford,UK.

Houseman GR,Gross KL(2006)Does ecological filtering across a productivity gradient explain variation in species pool-richness relationships?Oikos,115,148-154.

Huston MA(1979)A general hypothesis of species diversity.The American Naturalist,113,81-101.

Huston MA(1999)Local processes and regional patterns:appropriate scales for understanding variation in the diversity of plants and animals.Oikos,86,393-401.

Jetz W,Rahbek C(2002)Geographic range size and determinants of avian species richness.Science,297,1548-1551.

Karison RH,Cornell HV,Hughes TP(2004)Coral communities are regionally enriched along an oceanic biodiversity gradient.Nature,429,867-870.

Laanisto L,Urbas P,Partel M(2008)Why does the tmimodal species richness-productivity relationship not apply to woody species:a lack of clonality or a legacy of tropical evolutionary history?Global Ecology and Biogeography,17,320-326.

Latham RE,Ricklefs RE(1993)Global pattems of tree species richness in moist forests:energY diversity theory does not account for variation in species richness.Oikos,67,325-333.

Oksanen J(1996)Is the humped relationship between species richness and biomass an artefact due to plot size?Journal of Ecology,84,293-295.

Parmentier I,Malhi Y,Senterre B,Whittaker RJ,A.T.DN,Alonso A,Balinga MPB,Bakayoko A,Bongers F,Chatelain C,Comiskey JA,Cortay R,Kamdem M-ND,Doucet J-L,Gautier L,Hawthome WD,Issembe YA,Kouame FN,Kouka LA,Leal ME,Lejoly J,Lewis SL,Nusbaumer L,Parren MPE,Peh KSH,Phillips OL,Sheil D,Sonke B,Sosef MSM,Sunderland TCH,Stropp J,Ter Steege H,Swaine MD,Tchouto MGP(2007)The odd man out?Might climate explain the lower tree α-diversity of African rain forests relative to Amazonian rain forests?Journal of Ecology,95,1058-1071.

P(a)tel M(2002)Local plant diversity paaems and evolutionary history at the regional scale.Ecology,83,2361-2366.

P(a)rtel M,Helm A,Ingerpuu N,Reier (U),Tuvi E-L(2004)Conservation of Northem European plant diversity:the correspondence with soil pH.Biological Conservation,120,525-531.

P(a)rtel M,Laanisto L,Zobel M(2007)Contrasting plant productivity-diversity relationships across latitude:the role of evolutionary history.Ecology,88,1091-1097.

P(a)rtel M,Zobel M,Liira J,Zobel K(2000)Species richness limitations in productive and oligotrophic plant communities.Oikos,90,191-193.

P(a)rtel M,Zobel M,Zobel K,van der Maarel E(1996)The species pool and its relation to species richness:evidence from Estonian plant communities.Oikos,75,111-117.

Peet RK,Fridley JD,Gramling JM(2003)Variation in species richness and species pool size across a pH gradient in forests of the southern Blue Ridge Mountains.Folio Geobotanica,38,391-401.

Rajaniemi TK,Goldberg DE,Turkington R,Dyer AR(2006)Quantitative partitioning of regional and local processes shaping regional diversity patterns.Ecology Letters,9,121-128.

Ricklefs RE(1987)Community diversity:relative roles of local and regional processes.Science,235,167-171.

Ricklefs RE(2004)A comprehensive framework for global patterns in biodiversity.Ecology Letters,7,1-15.

Ricklefs RE(2006)Evolutionary diversification and the origin of the diversity-environment relationship.Ecology,87,S3-S13.

Schuster B,Diekmann M(2003)Changes in species density along the soil pH gradient:evidence from German plant communities.Folia Geobotanica,38,367-379.

Taylor DR,Aarssen LW,Loehle C(1990)On the relationship between r/K selection and environmental carrying capacity:a new habitat templet for plant life history strategies.Oikos,58,239-250.

Tihnan D(1982)Resource Competition and Community Structure.Princeton University Press,Princeton.

Tilman D(1994)Competition and biodiversity in spatially structured habitats.Ecology,75,2-16.

Wamelink GWW,Joosten V,van Dobben HF,Berendse F(2002)Validity of Ellenberg indicator values judged from physico-chemical field measurements.Journal of Vegetation Science,13,269-278.

Weiher E,Forbes S,Schauwecker T,Grace JB(2004)Multivariate control of plant species richness and community biomass in blackland prairie.Oikos,106,151-157.

Wilson JB,Anderson BJ(2001)Species-pool relations:like a wooden light bulb?Folia Geobotanica,36,35-44.

Zinko U,Dynesius M,Nilsson C,Seibert J(2006)The role of soil pH in linking groundwater flow and plant species density in boreal forest landscapes.Ecography,29,515-524.

Zobel K(2001)On the species-pool hypothesis and on the quasi-neutral concept of plant community diversity.Folia Geobotanica,36,3-8.

Zobel M(1992)Plant species coexistence:the role of historical,evolutionary and ecological factors.Oikos,65,314-320.

Zobel M(1997)The relative role of species pools in determining plant species richness:an alternative explanation of species coexistence?Trends in Ecology and Evolution,12,266-269.

Zobel M,van der Maarel E,Dupr(e) C(1998)Species pool:the concept,its determination and significance for community restoration.Applied Vegetation Science,1,55-66.