A multi-scale analysis of fragmentation effects on remnant plant species richness in Phoenix, Arizona
Corresponding Author
Arthur Stiles
School of Life Sciences, Arizona State University, Tempe, AZ 85287, USA
Correspondence: Arthur Stiles, School of Life Sciences, Arizona State University, Tempe, AZ 85287, USA.E-mail: [email protected]Search for more papers by this authorSamuel M. Scheiner
Division of Environmental Biology, National Science Foundation, Arlington, VA 22230, USA
Search for more papers by this authorCorresponding Author
Arthur Stiles
School of Life Sciences, Arizona State University, Tempe, AZ 85287, USA
Correspondence: Arthur Stiles, School of Life Sciences, Arizona State University, Tempe, AZ 85287, USA.E-mail: [email protected]Search for more papers by this authorSamuel M. Scheiner
Division of Environmental Biology, National Science Foundation, Arlington, VA 22230, USA
Search for more papers by this authorAbstract
Aim Understanding complex ecological phenomena, such as the determinants of species richness, is best achieved by investigating their properties at different spatial scales. Factors significantly affecting the number of species occurring at one scale may not impact on richness at other scales. While this scale dependence has become increasingly recognized, there still remains a need to elucidate exactly how richness is structured across scales, and which mechanisms are influential for determining this important community property. This study explores how woody plant species richness varies in a fragmented system at multiple scales, and which factors are primarily responsible for these patterns.
Location The study area is located in the Sonoran Desert within the bounds of metropolitan Phoenix, Arizona, which is the locus of the Central Arizona–Phoenix Long-Term Ecological Research (CAP-LTER) site.
Methods Estimates of local and fragment plant species richness were generated from field data collected from 22 sites. Independent variables describing fragment sites were also calculated, including area, habitat heterogeneity, density of individuals, mean elevation, and extent of isolation. Structural equation modelling, multiple regression, and analysis of covariance were used to assess the contribution of independent variables to richness at the fragment and local scales.
Results Fragment species richness was significantly influenced by area, though not isolation, habitat heterogeneity, mean elevation, or density of individuals. Local richness was not significantly related to fragment area, but was positively related to fragment richness, plant density, and elevation.
Main conclusions The fragment species–area effect resulted from larger remnants supporting higher numbers of individuals at comparable densities, increasing richness through either passive sampling of progressively less common species and/or lower extinction rates among larger populations. Without using multi-temporal data it is not possible to disentangle these mechanisms. We found that patterns evident at one scale are not necessarily apparent at other scales, as elevation and density of individuals significantly affected richness at the local scale but not at the fragment scale. These results lend support to the concept that mechanisms influencing the species richness of natural communities may be operable only within certain domains and that relevant scales should be specified.
Supporting Information
Appendix S1 Map of remnant sites examined in this study.
Appendix S2 Data for the dependent variables.
Appendix S3 Data for the independent variables.
Appendix S4 Plots showing the relationship between local-scale species richness and fragment area.
As a service to our authors and readers, this journal provides supporting information supplied by the authors. Such materials are peer-reviewed and may be re-organized for online delivery, but are not copy-edited or typeset. Technical support issues arising from supporting information (other than missing files) should be addressed to the authors.
Filename | Description |
---|---|
JBI_2333_sm_AppendixS1-S4.doc932.5 KB | Supporting info item |
Please note: The publisher is not responsible for the content or functionality of any supporting information supplied by the authors. Any queries (other than missing content) should be directed to the corresponding author for the article.
References
- Anderson, T.M., Metzger, K.L. & McNaughton, S.J. (2007) Multi-scale analysis of plant species richness in Serengeti grasslands. Journal of Biogeography, 34, 313–323.
- Arbuckle, J.L. (2003) Amos 5. Small Waters Corp., Chicago, IL.
- Arrhenius, O. (1921) Species and area. Journal of Ecology, 9, 95–99.
-
Barbour, M.G. (1973) Larrea plant communities on bajada and moisture gradients in the United States and Argentina.
Vegetatio, 28, 335–352.
10.1007/BF02386497 Google Scholar
- Bowers, J.E. (2005) Effects of drought on shrub survival and longevity in the northern Sonoran Desert. Bulletin of the Torrey Botanical Society, 132, 421–431.
- Bowers, J.E. & Turner, R.M. (2002) The influence of climatic variability on local population dynamics of Cercidium microphyllum (foothill paloverde). Oecologia, 130, 105–113.
- Bowers, J.E., Turner, R.M. & Burgess, T.L. (2004) Temporal and spatial patterns in emergence and early survival of perennial plants in the Sonoran Desert. Plant Ecology, 172, 107–119.
- Bowers, M.A. & Lowe, C.H. (1986) Plant-form gradients on Sonoran Desert bajadas. Oikos, 46, 284–291.
- Brown, J.H., Fox, B.J. & Kelt, D.A. (2000) Assembly rules: desert rodent communities are structured at scales from local to continental. The American Naturalist, 156, 314–321.
- Caley, M.J. & Schluter, D. (1997) The relationship between local and regional diversity. Ecology, 78, 70–80.
- de Candolle, A. (1855) Géographie botanique raisonnée: ou l’exposition des faits principaux et des lois concernant la distribution géographique des plates de l’epoque actuelle. Maisson, Paris.
- Cave, G.H. & Patten, D.T. (1984) Short-term vegetation responses to fire in the upper Sonoran Desert. Journal of Range Management, 37, 491–496.
- Chalcraft, D.R., Williams, J.W., Smith, M.D. & Willig, M.R. (2004) Scale dependence in the species-richness–productivity relationship: the role of species turnover. Ecology, 85, 2701–2708.
- Chase, J.M. & Leibold, M.A. (2002) Spatial scale dictates the productivity–diversity relationship. Nature, 416, 427–430.
- Chesson, P., Donahue, M.J., Melbourne, B.A. & Sears, A.L. (2005) Scale transition theory for understanding mechanisms in metacommunities. Metacommunities: spatial dynamics and ecological communities (ed. by M. Holyoak, M.A. Leibold and R.D. Holt), pp. 279–306. University of Chicago Press, Chicago.
- Coleman, B. (1981) On random placement and species–area relationships. Mathematical Biosciences, 54, 191–215.
- Coleman, B.D., Mares, M.A., Willig, M.R. & Hsieh, Y.H. (1982) Randomness, area, and species richness. Ecology, 64, 1121–1133.
- Colwell, R.K. (1999) EstimateS: Statistical estimation of species richness and shared species from samples. Version 5.0.1. User’s guide and application. Available at: http://viceroy.eeb.uconn.edu/estimates.
- Crawley, M.J. & Harral, J.E. (2001) Scale dependence in plant biodiversity. Science, 291, 864–868.
- Drayton, B. & Primack, R.B. (1996) Plant species lost in an isolated conservation area in metropolitan Boston from 1894 to 1993. Conservation Biology, 10, 30–39.
- Drinnan, I.N. (2005) The search for fragmentation thresholds in a southern Sydney suburb. Biological Conservation, 124, 339–349.
- Dzwonko, Z. & Loster, S. (1988) Species richness of small woodlands on the western Carpathian foothills. Vegetatio, 76, 15–27.
- ESRI (2002a) ArcView 3.3. Environmental Systems Research Institute, Redlands, CA.
- ESRI (2002b) ArcGIS 8.3. Environmental Systems Research Institute, Redlands, CA.
-
Forman, R.T.T. (1995) Land mosaics: the ecology of landscapes and regions. Cambridge University Press, Cambridge.
10.1017/9781107050327 Google Scholar
-
Gleason, H.A. (1922) On the relation between species and area.
Ecology, 3, 158–162.
10.2307/1929150 Google Scholar
- Gleason, H.A. (1925) Species and area. Ecology, 6, 66–74.
- Goldberg, D.E. & Turner, R.M. (1986) Vegetation change and plant demography in permanent plots in the Sonoran Desert. Ecology, 67, 695–712.
-
Grace, J.B. (2006) Structural equation modeling and natural systems. Cambridge University Press, Cambridge.
10.1017/CBO9780511617799 Google Scholar
- Gross, K.L., Willig, M.R., Gough, L., Inouye, R. & Cox, S.B. (2000) Patterns of species density and productivity at different spatial scales in herbaceous plant communities. Oikos, 89, 417–427.
- Gustafson, E.J. & Parker, G.R. (1992) Relationships between landcover proportion and indices of landscape spatial pattern. Landscape Ecology, 7, 101–110.
- Halvorson, W.L. & Patten, D.T. (1974) Seasonal water potential changes in Sonoran Desert shrubs in relation to topography. Ecology, 55, 173–177.
- Harrison, S., Safford, H.D., Grace, J.B., Viers, J.H. & Davies, K.F. (2006) Regional and local species richness in an insular environment: serpentine plants in California. Ecological Monographs, 76, 41–56.
- He, Z., Zhao, W., Chang, X., Chang, X. & Fang, J. (2006) Scale dependence in desert plant diversity. Biodiversity and Conservation, 15, 3055–3064.
- Hope, D., Gries, C., Zhu, W., Fagan, W.F., Redman, C.L., Grimm, N.B., Nelson, A.L., Martin, C. & Kinzig, A. (2003) Socioeconomics drive urban plant diversity. Proceedings of the National Academy of Sciences USA, 100, 8788–8792.
- Lack, D. (1976) Island birds. Blackwell, Oxford.
- Lane, D.R., Coffin, D.P. & Lauenroth, W.K. (1998) Effects of soil texture and precipitation on aboveground net primary production across the central grassland region. Journal of Vegetation Science, 9, 239–250.
- Leibold, M.A., Holyoak, M., Mouquet, N., Amarasekare, P., Chase, J.M., Hoopes, M.F., Holt, R.D., Shurin, J.B., Law, R., Tilman, D., Loreau, M. & Gonzales, A. (2004) The metacommunity concept: a framework for multi-scale community ecology. Ecology Letters, 7, 601–613.
- Levenson, J.B. (1981) Woodlots as biogeographic islands in southeastern Wisconsin. Forest island dynamics in man-dominated landscapes (ed. by R.L. Burgess and D.M. Sharpe), pp. 13–49. Springer Verlag, New York.
-
MacArthur, R.H. &
Wilson, E.O. (1967) The theory of island biogeography. Princeton University Press, Princeton, NJ.
10.1111/j.1463-6409.2007.00280.x Google Scholar
-
Magurran, A.E. (1988) Ecological diversity and its measurement. Princeton University Press, Princeton, NJ.
10.1007/978-94-015-7358-0 Google Scholar
- Matlack, G.R. (1994) Plant species migration in a mixed history forest landscape in eastern North America. Ecology, 75, 1491–1502.
- McAuliffe, J.R. (1994) Landscape evolution, soil formation, and ecological patterns and processes in Sonoran Desert bajadas. Ecological Monographs, 64, 111–148.
- McAuliffe, J.R. (1995) The aftermath of wildfire in the Sonoran Desert. The Sonoran Quarterly, 4, 4–8.
- McAuliffe, J.R. (1999) The Sonoran Desert: landscape complexity and ecological diversity. Ecology of Sonoran Desert plants and plant communities (ed. by R.H. Robichaux), pp. 68–114. University of Arizona Press, Tucson, AZ.
- Mittelbach, G.G., Steiner, C.F., Scheiner, S.M., Gross, K.L., Reynolds, H.L., Waide, R.B., Willig, M.R., Dodson, S.I. & Gough, L. (2001) What is the observed relationship between species richness and productivity? Ecology, 82, 2381–2396.
- Palmer, M.W. (1990) The estimation of species richness by extrapolation. Ecology, 71, 1195–1198.
- Palmer, M.W. (1991) Estimating species richness – the second order jackknife reconsidered. Ecology, 72, 1512–1513.
- Palmer, M.W. & White, P.S. (1994) Scale dependence and the species–area relationship. The American Naturalist, 144, 717–740.
- Pautasso, M. (2007) Scale dependence of the correlation between human population presence and vertebrate and plant species richness. Ecology Letters, 10, 16–24.
- Peterken, G.F. & Game, M. (1984) Historical factors affecting the number and distribution of vascular plant species in the woodlands of central Lincolnshire. Journal of Ecology, 72, 155–182.
-
Phillips, D.L. &
MacMahon, J.A. (1978) Gradient analysis of a Sonoran Desert bajada.
The Southwestern Naturalist, 23, 669–680.
10.2307/3671189 Google Scholar
- Pierson, E.A. & Turner, R.M. (1998) An 85-year study of saguaro (Carnegia gigantea) demography. Ecology, 79, 2676–2693.
- Preston, F.W. (1962) The canonical distribution of commonness and rarity: Part I. Ecology, 43, 185–215.
- Rebele, F. (1994) Urban ecology and special features of urban ecosystems. Global Ecology and Biogeography Letters, 4, 173–187.
- Ricklefs, R.E. (1987) Community diversity – relative roles of local and regional processes. Science, 235, 167–171.
-
Rosenzweig, M.L. (1995) Species diversity in space and time. Cambridge University Press, Cambridge.
10.1111/j.2006.0906-7590.04272.x Google Scholar
-
Scanlan, M.J. (1981) Biogeography of forest plants in the prairie forest ecotone in western Minnesota.
Forest island dynamics in man-dominated landscapes (ed. by
R.L. Burgess and
D.M. Sharpe), pp. 97–124. Springer Verlag, New York.
10.1007/978-1-4612-5936-7_7 Google Scholar
- Scheiner, S.M. (2003) Six types of species–area curves. Global Ecology and Biogeography, 12, 441–447.
- Scheiner, S.M. & Jones, S. (2002) Diversity, productivity, and scale in Wisconsin vegetation. Evolutionary Ecology Research, 4, 1097–1117.
- Scheiner, S.M. & Willig, M.R. (2005) Developing unified theories in ecology as exemplified with diversity gradients. The American Naturalist, 166, 458–469.
- Scheiner, S.M., Cox, S.B., Willig, M., Mittelbach, G.G., Osenberg, O. & Kaspari, M. (2000) Species richness, species–area curves, and Simpson’s paradox. Evolutionary Ecology Research, 2, 791–802.
- Schmid, M.K. & Rogers, G.F. (1988) Trends in fire occurrence in the Arizona Upland subdivision of the Sonoran Desert, 1955 to 1983. Southwestern Naturalist, 33, 437–444.
- Schneider, D.C. (2001) The rise of the concept of scale in ecology. BioScience, 51, 545–553.
- Shmida, A. & Wilson, M.V. (1985) Biological determinants of species diversity. Journal of Biogeography, 12, 1–20.
- Shreve, F. & Wiggins, I.L. (1964) Vegetation and flora of the Sonoran Desert. Stanford University Press, Stanford, CA.
- Simberloff, D.S. & Gotelli, N. (1984) Effects of insularization on plant species richness in the prairie–forest ecotone. Biological Conservation, 29, 27–46.
- Simmerling, D., Waldhardt, R. & Otte, A. (2006) Quantifying determinants contributing to plant species richness in mosaic landscapes: a single- and multi-patch perspective. Landscape Ecology, 21, 1233–1251.
- Soil Survey Geographic Database (2002) Maricopa county. National Cartographic and Geospatial Center, Fort Worth, TX.
- Soulé, M.E., Alberts, A.C. & Bolger, D.T. (1992) The effects of habitat fragmentation on chaparral plants and vertebrates. Oikos, 63, 39–47.
- SPSS (1994) Systat 6.1 for Windows. SPSS Inc., Chicago, IL.
- Stiles, A. & Scheiner, S.M. (2008) Nestedness of remnant Sonoran Desert plant communities in metropolitan Phoenix, Arizona. Ecology, 89, 2473–2481.
- Turner, R.M. & Brown, D.E. (1982) Sonoran desertscrub. Biotic communities of the American Southwest-United States and Mexico (ed. by D.E. Brown), pp. 181–221. University of Arizona Press, Tucson, AZ.
- Williams, M.R. (1995) An extreme-value function model of the species incidence and species–area relations. Ecology, 76, 2607–2616.
- Yang, T.W. & Lowe, C.H. (1956) Correlation of major vegetation climaxes with soil characteristics in the Sonoran Desert. Science, 123, 542.