THEORETICAL HABITAT TEMPLETS, SPECIES TRAITS, AND SPECIES RICHNESS - AQUATIC OLIGOCHAETES IN THE UPPER RHONE RIVER AND ITS FLOODPLAIN

1. This paper summarizes twenty years of ecological research on aquatic oligochaetes of the Upper Rhone River and its alluvial floodplain (France). Species traits of fifty species of the ninety taxa recorded from two areas (Jons and Bregnier-Cordon) were used to examine the relationships among species traits, habitat utilization of these species, whether a relationship exists between species traits and habitat utilization, and the applicability of predictions from the river habitat templet and the patch dynamics concept in the framework of spatial and temporal habitat variability. We used fourteen habitat types and sixteen species traits in this analysis. 2. When examined by correspondence analysis, species traits separate the Naididae (with a higher potential for reproduction, small size, high mobility, and opportunistic diet) from all other families. 3. Habitat utilization by oligochaetes demonstrates two gradients: a vertical gradient that arranges species by their affinity for interstitial habitats (stygophily) and a transversal gradient that arranges them by their affinity for main channel habitats (rheophily). 4. No significant relationship was found between species traits and habitat utilization in a co-inertia analysis. 5. Trends observed for species traits within the framework of spatial-temporal habitat variability show only minor agreement with predictions of the river habitat templet. 6. Species richness is generally higher in superficial and interstitial habitats that are permanently connected with the main channel, and peaks in the superficial parapotamons (backwaters that are permanently connected with the main channel) characterized by intermediate levels of spatial as well as temporal variability; this pattern