Riparian plant species richness along lateral and longitudinal gradients of water stress and flood disturbance, San Pedro River, Arizona, USA

Diversity theory predicts that species numbers should be highest at intermediate levels of both disturbance and environmental stress. We examined woody and herbaceous plant species richness and cover in the San Pedro River flood plain, along lateral gradients of water availability (ground-water depth), flood disturbance (inundation frequency), and distance from and elevation above the channel, and along longitudinal gradients of water availability (ground-water depth, surface flow permanence, and rainfall) and flood disturbance (total stream power). Herbaceous species were recorded during four sampling periods, and spatial patterns for this group were time-dependent, reflecting temporal variation in limiting factors. During the summer dry season of a dry year, when overall richness was low, richness and cover of herbaceous species declined laterally from the stream channel with increasing ground-water depth, consistent with the idea that low resource levels can limit species richness. Following the summer monsoon rains and floods, when water was less limiting and annuals were seasonally abundant, lateral patterns shifted such that herbaceous species richness and cover increased with increasing plot location above or from the channel. The relationship of herbaceous species richness with tree canopy cover also varied seasonally, shifting from positive (greater richness under canopy) in dry seasons to negative (lesser richness under canopy) in wet seasons. Longitudinally, herbaceous species richness and cover were limited primarily by stream flow and/or ground-water availability during the summer dry season of a dry year. Following the summer monsoon rains and floods, patterns were weighted by the seasonally abundant annuals, and richness increased among sites primarily with distance upstream (and related rainfall gradients). Richness and cover patterns also varied between years with different flood conditions. During the two sampling seasons in the year following a large flood, herbaceous species richness increased with flood disturbance intensity but declined at the few most intensely disturbed sites, consistent with intermediate disturbance theory. For woody species, richness within plant functional groups varied in opposing ways along the lateral gradients: hydromesic pioneer species decreased and hydromesic and xeric competitors increased with distance from or above the channel, with no overall change in species richness. Among sites, woody species richness patterns were related to water availability, but not to flood disturbance. However, richness of woody hydromesic pioneer species increased with both increasing site moisture and flood disturbance. Woody and herbaceous species richness both increased among sites as a function of increasing flood-plain width, likely due to species-area effects. Overall, results indicate that flood disturbance and water availability both influence species richness of riparian plants in the flood plain of this semi-arid region river, with the relative influence of each factor varying among plant groups and over time.