Scale and species richness: towards a general, hierarchical theory of species diversity

Journal of Biogeography - Tập 28 Số 4 - Trang 453-470 - 2001
Robert J. Whittaker1, Katherine J. Willis1, Richard Field1
1School of Geography and the Environment, University of Oxford, Mansfield Road, Oxford, UK

Tóm tắt

AimCurrent weaknesses of diversity theory include: a failure to distinguish different biogeographical response variables under the general heading of diversity; and a general failure of ecological theory to deal adequately with geographical scale. Our aim is to articulate the case for a top‐down approach to theory building, in which scale is addressed explicitly and in which different response variables are clearly distinguished.LocationThe article draws upon both theoretical contributions and empirical analyses from all latitudes, focusing on terrestrial ecosystems and with some bias towards (woody) plants.MethodsWe review current diversity theory and terminology in relation to scale of applicability. As a starting point in developing a general theory, we take the issue of geographical gradients in species richness as a main theme and evaluate the extent to which commonly cited theories are likely to operate at scales from the macro down to the local.ResultsA degree of confusion surrounds the use of the terms alpha, beta and gamma diversity, and the terms local, landscape and macro‐scale are preferred here as a more intuitive framework. The distinction between inventory and differentiation diversity is highlighted as important as, in terms of scale of analysis, are the concepts of focus and extent. The importance of holding area constant in analysis is stressed, as is the notion that different environmental factors exhibit measurable heterogeneity at different scales. Evaluation of several of the most common diversity theories put forward for the grand clines in species richness, indicates that they can be collapsed to dynamic hypotheses based on climate or historical explanations. The importance of the many ecological/biological mechanisms that have been proposed is evident mainly at local scales of analysis, whilst at the macro‐scale they are dependent largely upon climatic controls for their operation. Local communities have often been found not to be saturated, i.e. to be non‐equilibrial. This is argued, perhaps counter‐intuitively, to be entirely compatible with the persistence through time of macro‐scale patterns of richness that are climatically determined. The review also incorporates recent developments in macroecology, Rapoport’s rule, trade‐offs, and the importance of isolation, landscape impedance and geometric constraints on richness (the mid‐domain effect) in generating richness patterns; highlighting those phenomena that are contributory to the first‐order climatic pattern, and those, such as the geometric constraints, that may confound or obscure these patterns.Main conclusionsA general theory of diversity must necessarily cover many disparate phenomena, at various scales of analysis, and cannot therefore be expressed in a simple formula, but individual elements of this general theory may be. In particular, it appears possible to capture in a dynamic climate‐based model and ‘capacity rule’, the form of the grand cline in richness of woody plants at the macro‐scale. This provides a starting point for a top‐down, global‐to‐local, macro‐to‐micro scale approach to modelling richness variations in a variety of taxa. Patterns in differentiation/endemicity, on the other hand, require more immediate attention to historical events, and to features of geography such as isolation. Thus, whilst we argue that there are basic physical principles and laws underlying certain diversity phenomena (e.g. macro‐scale richness gradients), a pluralistic body of theory is required that incorporates dynamic and historical explanation, and which bridges equilibrial and nonequilibrial concepts and ideas.

Từ khóa


Tài liệu tham khảo

10.1038/339699a0

Allen T.F.H.&Starr T.B.(1982)Hierarchy: perspectives for ecological complexity. Chicago University Press Chicago.

10.1111/j.1469-7998.2000.tb00605.x

10.1016/0169-5347(87)90005-X

10.1111/j.1744-7429.2000.tb00474.x

10.2307/3236782

Bennett K.D.(1997)Evolution and ecology: the pace of life. Cambridge University Press Cambridge.

10.2307/2844972

10.1006/bijl.1998.0285

10.2307/3236373

10.1046/j.1466-822x.2001.00213.x

10.1093/icb/21.4.877

Brown J.H.(1988)Species diversity.Analytical biogeography(eds A.A. Myers and P.S. Giller) pp. 57–89. Chapman and Hall London.

Brown J.H. (1995)Macroecology. Chicago University Press Chicago.

10.2307/3546991

10.1007/s004420000536

Brown J.H.&Lomolino M.V.(1998)Biogeography 2nd edn. Sinauer Sunderland MA.

10.1046/j.1365-2699.2000.00186.x

10.1146/annurev.ecolsys.27.1.597

Brown J.H.&West G.B.(eds) (2000)Scaling in biology. Oxford University Press Oxford.

10.2307/2845600

10.1046/j.1523-1739.1995.09030559.x

10.1007/BF00115314

Cody M.L.(1975)Towards a theory of continental species diversities: bird distributions over Mediterranean habitat gradients.Ecology and Evolution of Communities(eds M.L. Cody and J.M. Diamond) pp. 214–257. Harvard University Press Cambridge MA.

Colinvaux P.A.(1993)Ecology 2. Wiley New York.

10.1016/S0169-5347(99)01767-X

10.1126/science.288.5470.1414

10.1126/science.199.4335.1302

10.1126/science.246.4930.675

10.1126/science.291.5505.864

10.1086/285144

10.1038/329326a0

10.1111/j.1466-8238.1998.00269.x

10.1046/j.1466-822X.1999.00126.x

10.1007/BF01237657

10.1038/44819

Ernest S.K.M.&Brown J.H.(2001)Homeostasis and compensation: the role of species and resources in ecosystem stability.Ecology(in press).

10.1111/j.1466-8238.1998.00278.x

10.2307/3546780

10.1046/j.1466-822X.1998.00294.x

10.1086/285915

Gaston K.J.(ed.) (1996)Biodiversity: a biology of numbers and difference. Blackwell Science Oxford.

10.1038/35012228

Gaston K.J.&Blackburn T.M.(2000)Pattern and process in macroecology. Blackwell Science Oxford.

10.1046/j.1365-2745.1999.00333.x

10.1046/j.1466-822X.2001.00249.x

10.1034/j.1600-0706.2000.890301.x

10.1038/35006630

10.1046/j.1365-2699.2000.00163.x

10.1007/BF00328360

10.1046/j.1365-2656.2000.00377.x

Hulme M., 1996, A 1961–90 climatology for Africa south of the Equator and a comparison of potential evapotranspiration estimates, South African Journal of Science, 92, 334

10.2307/3546645

Innes J.L.(1998)Measuring environmental change.Ecological scale: theory and applications(eds D.L. Peterson and V.T. Parker) pp. 429–457. Columbia University Press New York.

10.1111/j.1654-1103.2005.tb02371.x

10.2307/2997393

10.1046/j.1365-2699.1999.00315.x

10.1046/j.1466-822X.2001.00234.x

10.1046/j.1365-2486.2000.00332.x

10.2307/2997373

10.2307/3545479

10.1111/j.1600-0587.2000.tb00265.x

10.1046/j.1365-2656.2000.00418.x

10.2307/2845687

10.1046/j.1365-2699.1998.2540735.x

10.1086/303369

10.1046/j.1365-2699.2000.00185.x

10.1046/j.1365-2699.2000.00377.x

10.1046/j.1466-822x.2001.00229.x

10.1046/j.1466-822x.2001.00221.x

10.1046/j.1365-2699.2001.00550.x

10.1046/j.1461-0248.2000.00127.x

10.1890/0012-9658(1999)080[2483:AHAOSD]2.0.CO;2

Maurer B.A.(1999)Untangling ecological complexity: the macroscopic perspective. University of Chicago Press Chicago.

MacArthur R.H.&Wilson E.O.(1967)The theory of island biogeography. Princeton University Press Princeton.

10.1038/35012234

10.2307/2997586

10.1038/35020160

10.1046/j.1365-2699.1999.00305.x

10.1017/s0266467499000942

10.2307/2845670

10.1046/j.1365-2699.1998.252166.x

10.1034/j.1600-0706.2000.890319.x

10.1111/j.1600-0587.1998.tb00441.x

10.2307/2261364

10.1038/35005072

10.1086/285704

10.1046/j.1466-822X.2001.00233.x

10.1093/icb/34.1.134

Peterson D.L.&Parker V.T.(eds) (1998)Ecological scale: theory and applications. Columbia University Press New York.

10.1038/35012221

10.1038/35025052

10.1086/283645

10.2307/3546454

10.2307/3545569

10.2307/3546928

Rosenzweig M.C.(1995)Species diversity in space and time. Cambridge University Press Cambridge.

Rosenzweig M.C., 1997, Species diversity and latitude: listening to area’s signal, Oikos, 80, 172, 10.2307/3546528

Schumm S.A.(1991)To interpret the earth: ten ways to be wrong. CUP Cambridge.

10.1023/A:1009755320731

10.2307/3237310

10.2307/2845026

10.2307/2844907

10.1038/21877

10.1086/285067

10.1086/284913

Stevens G.C.&Enquist B.J.(1998)Macroecological limits to the distribution and abundance ofPinus.Ecology and Biogeography of Pinus(ed. D.M. Richardson) pp. 183–190. Cambridge University Press Cambridge.

10.1111/j.1744-7429.2000.tb00468.x

10.1890/0012-9658(1999)080[2474:CRSRBR]2.0.CO;2

10.1038/35012217

Tokeshi M.(1999)Species coexistence: ecological and evolutionary perspectives. Blackwell Science Oxford.

10.4319/lo.1997.42.5.0938

10.2307/1366721

10.1038/335539a0

Turner J.R.G. Lennon J.J. Greenwood J.D.(1996)Does climate cause the global biodiversity gradient?Aspects of the genesis and maintenance of biological diversity(eds M. Hochberg J. Claubert and R. Barbault) pp. 199–220. Oxford University Press Oxford.

10.1046/j.1365-2745.1999.00412.x

10.1038/23251

10.1111/j.1558-5646.1969.tb03503.x

10.2307/1943563

Whittaker R.H.(1975)Communities and ecosystems. 2nd edn. Macmillan New York.

Whittaker R.H.(1977)Evolution of species diversity in land communities.Evolutionary biology Vol 10(eds M.K. Hecht W.C. Steere and B. Wallace) pp. 250–268. Plenum Press New York.

Whittaker R.J.(1998)Island biogeography: ecology evolution and conservation. Oxford University Press Oxford.

10.1038/44732

10.1046/j.1365-2699.2000.00200.x

10.1034/j.1600-0706.2000.890222.x

10.2307/2388920

10.1023/A:1018335007666

10.1098/rstb.1999.0527

10.1126/science.287.5457.1406

Wilson J.B., 1990, Mechanisms of species coexistence: twelve explanations for Hutchinson’s ‘paradox of the plankton’: evidence from New Zealand plant communities, New Zealand Journal of Ecology, 13, 17

Wilson J.B., 1994, The intermediate disturbance hypothesis of species coexistence is based on patch dynamics, New Zealand Journal of Ecology, 18, 176

10.2307/3544109

Thông tin
Thông tin xuất bản

Journal of Biogeography

Tập 28 Số 4

453-470

Thông tin tác giả