The consequences of scale: assessing the distribution of benthic populations in a complex estuarine fjord

Evidence suggests that patterns of benthic community structure are functionally linked to estuarine processes and physical characteristics of the benthos. To assess these linkages for coarse-sediment shorelines, we used a spatially nested sampling design to quantify patterns of distribution and abundance of both macroinfauna and macroepibiota. We examined replicate beach segments within a site (∼1 km), sites within areas of relatively uniform salinity and temperature (∼10 km), and areas (∼100 km) in the two major basins of Puget Sound, Washington. Because slight variations in physical characteristics of a beach can lead to significant alterations in biota, we minimized confounding physical influences by working only in the predominant shoreline habitat type in Puget Sound, a mixture of sand, pebbles and cobbles. Species richness decreased steadily from north to south along gradients of declining wave energy, increasing temperature and decreasing salinity. A few taxa were confined to the South Basin, but many more were found in the North. Most of the variability in population abundance was captured at the smaller spatial scales. Physical conditions tend to become increasingly different with distance among sites. Communities became more different from north to south as species intolerant of more estuarine conditions dropped out. There was significant spatial autocorrelation among populations on neighboring beach segments for 73 of the 172 species sampled. Populations of these benthic species may be connected via dispersal on scales of at least km in Puget Sound. Our results strengthen prior conclusions about the strong linkages between the biota and physical patterns and processes in estuaries. It is important for monitoring and impact-detection studies to account for natural variation of physical gradients across the sampling scales used. Nested, replicated sampling designs can facilitate the detection of environmental change at spatial scales ranging from global (e.g., warming or El Niño), to regional (e.g., estuary-wide changes in salinity patterns), to local (e.g., from development at a site).