Beta diversity and the scale‐dependence of the productivity‐diversity relationship: a test in the Californian serpentine flora

Summary 1 The relationship of productivity to species diversity is usually positive at regional scales, but is often neutral, unimodal or negative at local spatial scales. Recent studies have pointed out that beta diversity, or among-locality and within-region variation in species composition, must therefore tend to increase with productivity. 2 We tested for a positive relationship of productivity to beta diversity in herbaceous plants at 105 widely distributed sites on serpentine soil in California. We also asked whether any such pattern could be explained, as previously proposed, by increased environmental heterogeneity at higher levels of productivity. 3 We found that one measure of beta diversity (the species dissimilarity between paired 500 m2 plots on adjacent north and south slopes) was positively related to productivity (as measured by the normalized difference vegetation index, a remotely sensed index). However, this effect was not strong enough to transform the neutral relationship of productivity with alpha (1 m2) diversity to a positive relationship of productivity with gamma (1000 m2) diversity. 4 The positive effect of productivity on beta diversity was not related to increasing heterogeneity in coarse measures of vegetation structure (percentage cover of shade, litter, rocks, moss, bare soil, animal disturbance). We speculate that the effect may instead have a ‘top-down’ explanation: higher beta diversity may be caused by the positive influence of productivity on the size of the regional species pool. 5 Our study illustrates the principle of a ‘scale transition’, in which an ecological pattern is qualitatively different at different spatial scales. Careful attention to scale-dependence may help narrow the search for mechanisms for such long-studied ecological puzzles as the latitudinal diversity gradient.

[1]  J. C. Hickman,et al.  The Jepson Manual: Higher Plants of California , 1993 .

[2]  S. Ware,et al.  California Serpentines: Flora, Vegetation, Geology, Soils and Management Problems. , 1985 .

[3]  R. Whittaker Vegetation of the Siskiyou Mountains, Oregon and California , 1960 .

[4]  Peter J. Richerson,et al.  Patterns of Plant Species Diversity in California: Relation to Weather and Topography , 1980, The American Naturalist.

[5]  J. Viers,et al.  CalJep: A Spatial Distribution Database of CalFlora and Jepson Plant Species , 2006 .

[6]  J. Oksanen Is the humped relationship between species richness and biomass an artefact due to plot size , 1996 .

[7]  James B. Grace,et al.  The factors controlling species density in herbaceous plant communities: an assessment , 1999 .

[8]  Jonathan M. Chase,et al.  Spatial scale dictates the productivity–biodiversity relationship , 2002, Nature.

[9]  James C. Hickman,et al.  The Jepson Manual: Higher Plants of California , 1993 .

[10]  Eric E. Porter,et al.  Does Herbivore Diversity Depend on Plant Diversity? The Case of California Butterflies , 2002, The American Naturalist.

[11]  J. Weitz,et al.  Scale-dependence of resource-biodiversity relationships. , 2003, Journal of theoretical biology.

[12]  Kevin J. Gaston,et al.  Measuring beta diversity for presence–absence data , 2003 .

[13]  M. Loreau Are communities saturated? On the relationship between α, β and γ diversity , 2000 .

[14]  H. Safford,et al.  Does the Age of Exposure of Serpentine Explain Variation in Endemic Plant Diversity in California? , 2004 .

[15]  David M. Stoms,et al.  Anthropogenic impacts upon plant species richness and net primary productivity in California , 2004 .

[16]  Michael A. Huston,et al.  Local processes and regional patterns : appropriate scales for understanding variation in the diversity of plants and animals , 1999 .

[17]  Marcel Rejmánek,et al.  Species pools and the "hump-back" model of plant species diversity : an empirical analysis at a relevant spatial scale , 2001 .

[18]  David J. Currie,et al.  A Globally Consistent Richness‐Climate Relationship for Angiosperms , 2003, The American Naturalist.

[19]  M. Willig,et al.  The Relationship Between Productivity and Species Richness , 1999 .

[20]  R. Brooks,et al.  Serpentine and Its Vegetation: A Multidisciplinary Approach , 1987 .

[21]  James B. Grace,et al.  REGIONAL AND LOCAL SPECIES RICHNESS IN AN INSULAR ENVIRONMENT: SERPENTINE PLANTS IN CALIFORNIA , 2006 .

[22]  D. Tilman Resource competition and community structure. , 1983, Monographs in population biology.

[23]  Robert K. Colwell,et al.  Estimating terrestrial biodiversity through extrapolation. , 1994, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[24]  J. P. Grime,et al.  Competitive Exclusion in Herbaceous Vegetation , 1973, Nature.

[25]  Richard Field,et al.  ENERGY, WATER, AND BROAD‐SCALE GEOGRAPHIC PATTERNS OF SPECIES RICHNESS , 2003 .

[26]  T. Fukami,et al.  Productivity–biodiversity relationships depend on the history of community assembly , 2003, Nature.

[27]  J. P. Grime,et al.  Plant Strategies, Vegetation Processes, and Ecosystem Properties , 2006 .

[28]  John H. Lawton,et al.  Beta diversity on geographic gradients in Britain , 1992 .

[29]  J. Viers,et al.  Climatic and spatial patterns of diversity in the serpentine plants of California , 2000 .

[30]  D. Janzen Why Mountain Passes are Higher in the Tropics , 1967, The American Naturalist.

[31]  Jonathan M. Chase,et al.  Connectivity, scale‐dependence, and the productivity–diversity relationship , 2004 .

[32]  H. Safford,et al.  Fire effects on plant diversity in serpentine vs. sandstone chaparral , 2004 .

[33]  A. Magurran Ecological Diversity and Its Measurement , 1988, Springer Netherlands.

[34]  M. Willig,et al.  Patterns of species density and productivity at different spatial scales in herbaceous plant communities , 2000 .

[35]  Helmut Hillebrand,et al.  On the Generality of the Latitudinal Diversity Gradient , 2004, The American Naturalist.

[36]  J. Lovett-Doust,et al.  Plant strategies, vegetation processes, and ecosystem properties , 2002 .

[37]  J. Viers,et al.  SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA: A DATABASE OF SERPENTINE AFFINITY , 2005 .

[38]  Katherine L. Gross,et al.  WHAT IS THE OBSERVED RELATIONSHIP BETWEEN SPECIES RICHNESS AND PRODUCTIVITY , 2001 .

[39]  J. Cihlar,et al.  Relation between the normalized difference vegetation index and ecological variables , 1991 .

[40]  W. Carson,et al.  The significance of assemblage‐level thinning for species richness , 1999 .

[41]  S. Williams,et al.  Spatial scale, species diversity, and habitat structure: small mammals in Australian tropical rain forest , 2002 .

[42]  T. Hughes,et al.  Coral communities are regionally enriched along an oceanic biodiversity gradient , 2004, Nature.

[43]  Don A. Driscoll,et al.  Metacommunities: Spatial Dynamics and Ecological Communities , 2006 .