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Metacommunity patterns of highly diverse stream midges: gradients, chequerboards, and nestedness, or is there only randomness?


*Jani Heino, Finnish Environment Institute, Research Programme for Biodiversity, PO Box 413, FIN-90014 University of Oulu, Finland. E-mail:


Abstract.  1. Several non-random patterns in the distribution of species have been observed, including Clementsian gradients, Gleasonian gradients, nestedness, chequerboards, and evenly spaced gradients. Few studies have examined these patterns simultaneously, although they have often been studied in isolation and contrasted with random distribution of species across sites.

2. This study examined whether assemblages of chironomid midges exhibit any of the idealised distribution patterns as opposed to random distribution of species across sites within the metacommunity context in a boreal drainage system. Analyses were based on stream surveys conducted during three consecutive years. Analytical approaches included ordinations, cluster analysis, null models, and associated randomisation methods.

3. Midge assemblages did not conform to Clementsian gradients, which was evidenced by the absence of clearly definable assemblage types with numerous species exclusive to each assemblage type. Rather, there were signs of continuous Gleasonian variability of assemblage composition, as well as significant nested subset patterns of species distribution.

4. Midge assemblages showed only weak relationships with any of the measured environmental variables, and even these weak environmental relationships varied among years.

5. Midge assemblages did not appear to be structured by competition. This finding was somewhat problematic, however, because the two indices measuring co-occurrence provided rather different signs of distribution patterns. This was probably a consequence of how they actually measure co-occurrence.

6. Although midge assemblages did not show a perfect match with any of the idealised distribution patterns, they nevertheless showed a resemblance to the empirical patterns found previously for several plant and animal groups.