Dynamics of an upland stream fish community over 40 years: trajectories and support for the loose equilibrium concept

Previous theoretical models and empirical studies suggested that communities can exist in a "stochastic" or "loose" equilibrium, diverging transiently but eventually returning toward earlier or average structure, in what we call here the "loose equilibrium concept" (LEC). We sampled the fish communities at 12 local stream reaches spaced broadly throughout a relatively undisturbed watershed in the Ozark Mountains of northern Arkansas, USA, in 11 surveys from 1972 to 2012 at a scale of decades, and at a subset of five of these local sites in a total of 16 surveys, allowing tests of the LEC at different spatial and temporal scales. Multivariate analyses of the dynamics of communities over the 40-year period provided support for the LEC at both "global" and "local" scales within the watershed. At the broadest spatial scale, core species numerically dominated the community, and most common species remained so across all decades. In spite of two extraordinary floods, and interannual variation in abundance of some species, the 12-site and five-site global communities and eight of 12 local communities repeatedly returned toward average positions in multivariate space. Trajectories of the global and local fish communities varied relative to model hypothetical trajectories that were based on gradual vs. saltatory changes, and prevalence of returns toward average community structure. Beta diversity among sites was variable across time, but beta partitioning consistently showed that pure spatial turnover dominated over nestedness, because many common species were consistently distributed either upstream or downstream. This study suggests that vertebrate communities in relatively undisturbed environments may display dynamics consistent with the LEC. The LEC, combined with quantification of community trajectory patterns, can help to clarify whether systems are moving about within ranges of conditions that reflect expected noise, or, conversely, have moved so far out of previous bounds, as a result of climate change or human intervention, that they are permanently changed or "novel."