Environmental heterogeneity and species diversity of forest sedges


  • Graham Bell,

    1. Biology Department, McGill University, 1205 Ave Dr Penfield, Montreal, Quebec, Canada H3A 1B1;
    2. Redpath Museum, McGill University, 859 Sherbrooke St West, Montreal, Quebec, Canada H3A 2K6; and
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  • Martin J. Lechowicz,

    1. Biology Department, McGill University, 1205 Ave Dr Penfield, Montreal, Quebec, Canada H3A 1B1;
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  • Marcia J. Waterway

    1. Department of Plant Science, Macdonald Campus, McGill University, 21111 Lakeshore Road, Ste-Anne de Bellevue, Quebec, Canada H9X 3V9
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Graham Bell, Biology Department, McGill University, 1205 Ave Dr Penfield, Montreal, Quebec, Canada H3A 1B1 (fax 1 514 398 3185; e-mail gbell2@maclan.mcgill.ca).


1 A field experiment was designed to investigate the relationship between environmental heterogeneity and species diversity in a group of sedges (Cyperaceae: Carex) growing in old-growth forest.

2 A measure of environmental quality, as perceived by the sedges, was obtained from the survival of clonal ramets of 11 species of Carex planted at 10-m intervals along each of three 1-km transect lines.

3 The resident assemblage of sedges was censused along the same three transect lines and along a further 24 km of survey lines in the same forest.

4 The general state of a site was represented by the overall survival of the experimental implants at that site. The general environmental variance between sites provided a measure of environmental heterogeneity. This could be partitioned into a specific variance (mean environmental variance of species) and an environmental covariance. The rate of increase of the general and specific variances with distance between sites reflected environmental structure.

5 The three transects differed in scale. The species diversity of the resident Carex assemblage was correlated with general environmental quality both among and within transects.

6 The three transects differed in structure. The number of resident species, relative to the number expected from the number of individuals sampled, was greatest on the most coarse-grained transect (steepest increase in general environmental variance with distance).

7 Within each transect, species diversity increased with general environmental variance because the specific correlation of performance (correlation among species of survival in pair-wise combinations of sites) decreased as the general environmental variance increased.

8 The effect of specific environmental variance was weaker. Overall survival of a species on the transects was not correlated with its abundance in the forest. Neither the transects nor a targeted implant experiment provided evidence for a close relationship between the distribution of species and the state of the environment.

9 As a general explanation of our results, we propose a ‘marginal-specialist’ model in which the species that dominate the most productive sites also have the broadest ranges, whereas other species are superior in a more restricted range of less productive sites.