Spatial Variation in Fish Assemblages across a Beaver-Influenced Successional Landscape

Beavers are increasingly viewed as "ecological engineers," having broad effects on physical, chemical, and biological attributes of north-temperate landscapes. We examine the influence of both local successional processes associated with beaver activity and regional geomorphic boundaries on spatial variation in fish assemblages along the Kabetogama Peninsula in Voyageurs National Park, northern Minnesota, USA. Fish abundance and species richness exhibited considerable variation among drainages along the peninsula. Geological barriers to fish dispersal at outlets of some drainages has reduced fish abundance and species richness. Fish abundance and species richness also varied within drainages among local environments associated with beaver pond succession. Fish abundance was higher in upland ponds than in lowland ponds, collapsed ponds, or streams, whereas species richness was highest in collapsed ponds and streams. Cluster analyses based on fish abundance at sites classified according to successional environment indicated that four species (northern redbelly dace, Phoxinus eos; brook stickleback, Culaea inconstans; finescale dace, P. neogaeus; and fathead minnow, Pimephales promelas), were predominant in all successional environments. Several less abundant species were added in collapsed ponds and streams, with smaller size classes of large lake species (e.g., black crappie, Pomoxis nigromaculatus; smallmouth bass, Micropertus dolomieui; yellow perch, Perca flavescens; and burbot, Lota lota) being a component of these less abundant species. The addition of smaller size classes of large lake species indicates that dispersal of early life-history stages from Kabetogama Lake played a role in determining the species richness and composition of less abundant species in successional environments on the peninsula. Furthermore, collapsed-pond and stream environments closer to Kabetogama Lake had higher species richness than similar successional sites located farther from the lake. Cluster analyses based on fish abundance at sites classified according to drainage indicated that species composition among drainages was influenced both by the presence or absence of geological barriers to fish dispersal and the nonrandom distribution of collapsed ponds and streams. Based on these results, we present a hierarchical conceptual model suggesting how geomorphic boundaries and beaver pond succession interact to influence fish assemblage attributes. The presence of a productive and diverse fish assemblage in headwater streams of north-temperate areas requires the entire spatial and temporal mosaic of successional habitats associated with beaver activity, including those due to the creation and abandonment of beaver ponds. The ultimate impact of the local successional mosaic on fishes, however, will be strongly influenced by the regional geomorphic context in which the mosaic occurs.