Publications of Michael L. Rosenzweig


Species diversity

(co-authored publications list authors in order)

What accounts for the number of species that exist? The following works present a basic, continent-scale, mathematical theory of diversity as a process, a process resulting from the differential equations of speciation and extinction. They identify area (e.g., km2) as a principal variable and help to explain why the tropics are so rich. They predict several patterns of diversity with area, patterns that have proved accurate in the decades since the theory first emerged. They predict long-term equilibria (actually steady states) in diversity. These too have found empirical confirmation, turning up repeatedly in examinations of the fossil records of marine invertebrates, mammals, and land plants.

The contributions also reveal a new pattern of diversity: on a regional scale, animal diversity peaks in locales with intermediate energy flow. (Previously, ecology had assumed that diversity and productivity were inexorably and positively correlated.) They have contributed to the methodology of diversity and extinction rate estimation. And they have suggested a mode of speciation (competitive speciation) that joins ecology and evolution, and appears to be implicated in biotic revolutions, the sort that sees major taxa such as dinosaurs supplanted by other major taxa such as mammals. Much of this work is described and synthesized in the 1995 book. The book also attempts to sweep across all scales of space and time available to life, and put the many contributions made by other ecologists into the context of a general view of the development and maintenance of diversity.

2001 The four questions: What does the introduction of exotic species do to diversity? Evolutionary Ecology Research 3 (preprint edition URL:

2000 Wisheu, I.C., M.L.Rosenzweig, L. Olsvig-Whittaker, and A. Shmida: What makes nutrient-poor mediterranean heathlands so rich in plant diversity? Evolutionary Ecology Research 2: 935-955.

1999 MLR & Y. Ziv: The echo pattern of species diversity: pattern & process. Ecography 22: 614-628.

1999 W. Turner, W. Leitner & MLR: Ws2m; software for estimating diversity. URL:

1999 Heeding the warning in biodiversity's basic law. Science 284:276-277

1999 Species diversity, Chapter 9, p. 249-281 in McGlade, J. (ed.), Advanced Theoretical Ecology: principles and applications, Blackwell Science, Oxford, England.

1998 Articles on Species Diversity in Encyclopedia of Ecology and Environmental Management (P. Calow et al., Eds.) Blackwell Scientific Publications Ltd., Oxford, England.

1) Diversity, alpha, beta and gamma: three measures of species diversity in space (p.195 )

2) Diversity gradient: A correlation of diversity with another spatial or temporal variable (p.195-7)

3) Diversity, methods of measurement and analysis (p. 200)

1998 Preston's ergodic conjecture: the accumulation of species in space and time. Ch. 14 (pp.311-348) in McKinney, M.L. & J. Drake (eds.) Biodiversity Dynamics; turnover of populations, taxa and communities. Columbia Univ. Press, NY.

Looks at fossil data to construct the first species-time accumulation curves. These are analogues of species-area curves and suggest the existence of an ergodic property in diversity relationships.

1998 Davidowitz, G. & MLR, The latitudinal gradient of species diversity among North American grasshoppers within a single habitat: a test of the spatial heterogeneity hypothesis. J. Biogeog. 25:553-560.

Further evidence that the latitudinal gradient does not come from high habitat diversity in the tropics.

1997 Tempo and mode of speciation. Science 277:1622-1623.

1997 MLR & E.A. Sandlin: Species diversity and latitude: listening to area's signal. Oikos 80:172-176.

1997 Leitner, W.A. & MLR. Nested species-area curves and stochastic sampling: a new theory. Oikos 79:503-512.

Overturns the major theory of species-area relationships, i.e., the lognormal abundance distribution. Replaces previous theories with a closed form equation validated with simulations. Introduces the connection between species-area relationships, population densities and the size of geographical ranges.

1995 Species Diversity In Space and Time, Cambridge University Press, Cambridge, UK., 436 pp. (Revised ed., 1996) Now in third printing.

1994 MLR & C.W. Clark. Island extinction rates from regular censuses. Conserv. Biol. 8:491-494; reprinted pp. 112-115 in Ehrenfeld, D. (ed.), Readings from Conservation Biology: Genes Populations & Species, 1995, Blackwell Science, Inc. & Society for Conservation Biology, Cambridge, MA

1994 Clark, C.W. & MLR. Extinction and colonization processes: parameter estimates from sporadic surveys. Amer. Natur. 143:583-596.

1993 MLR & Z. Abramsky. How are diversity and productivity related? Pp. 52-65 in Ricklefs, R. & D. Schluter (eds.). Species diversity in ecological communities: historical and geographical perspectives. Univ. Chicago Press.

1992 Species diversity gradients: we know more and less than we thought. J. Mamm. 73:715-730.

1992 MLR & S. Vetault. Calculating speciation and extinction rates in fossil clades. Evolutionary Ecology 6:90-93.

1984 Z. Abramsky & MLR. Tilman's predicted productivity-diversity relationship shown by desert rodents. Nature 309:150-1.

1980 MLR & J. Taylor. Speciation and diversity in Ordovician invertebrates: filling niches quickly and carefully. Oikos 35:236-243.

1979 MLR & J.L. Duek. Species diversity and turnover in an Ordovician marine invertebrate assemblage, pp. 109-119 in Patil & Rosenzweig ibid.

1978 Competitive speciation. Biol. J. Linnaean Soc. 10:275-289.

1977 On interpreting the results of perturbation experiments performed by nature. Paleobiol. 3:322-324.

1977 Geographical speciation: on range size and the probability of isolate formation. Pp. 172-194 in D. Wollkind (ed.), Proc. Wash. State Univ. Conf., Biomath. and Biostatistics, Pullman, WA.

1975 On continental steady states of species diversity, pp.121-140 in The Ecology and Evolution of Communities, M. Cody & J. Diamond (eds.), Harvard Univ. Press.