Theoretical habitat templets, species traits, and species richness: 548 plant and animal species in the Upper Rhône River and its floodplain

Abstract

SUMMARY For practical reasons, conceptual developments in community ecology are usually based on studies of a restricted systematic group. The cooperation of thirty or so specialists in the synthesis of long‐term ecological research on the Upper Rhône River, France, provided a unique occasion to investigate relationships among species traits, the habitat utilization by species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the framework of spatial‐temporal habitat variability for 548 species of plants (Hyphomycetes, aquatic macrophytes, floodplain vegetation) and animals (Tricladida, Oligochaeta, several groups of Crustacea, Insecta and Vertebrata). Using correspondence analysis, 100 modalities of eighteen species traits were examined; the resulting typology demonstrates that systematic groups are the most important elements for separating species traits such as size, fecundity of individuals, parental care, mobility, body form, and food type. Small species have an intermediate number of descendants per reproductive cycle and few reproductive cycles both per year and per individual; in contrast, large species have a high number of descendants per reproductive cycle and few reproductive cycles per year but many potential reproductive cycles per individual. The analysis of habitat utilization in the Upper Rhône River and its floodplain by the 548 species demonstrated a vertical gradient separating interstitial from superficial habitats; a transverse gradient for superficial habitats from the main channel towards more terrestrial ones is also evident. Because of a significant (P < 0.01) relationship between species traits and habitat utilization, traits such as size, fecundity of individuals, parental care, tolerance to variation in humidity, and respiration are arranged along the vertical and transverse habitat gradient. Size, the number of reproductive cycles per individual, and the tolerance to variation of humidity increases from permanent waters to temporary waters, aggrading habitats, and terrestrial habitats. Species traits showed significant (P < 0.01) trends in the framework of spatial‐temporal habitat variability and were compared with predictions based on the river habitat templet. Although each habitat showed a mixture of species traits at low temporal and spatial variability, and at high variability sites, trends corresponded to predictions for three traits (number of descendants per reproductive cycle, number of reproductive cycles per individual, attachment to soil or substrate) along a gradient of increasing temporal habitat variability. The species richness of each habitat within the Upper Rhône River and its floodplain significantly (P = 0.03) increased as the spatial variability of habitats increased but there is no statistical correlation between spedes richness and temporal variability. An altemative hypothesis predicting that fewer spedes per resource occur in temporally stable habitats is also not supported.