Jack A. Stanford

Flathead Lake Biological Station, University of Montana

I. INTRODUCTION

Streams, rivers, and groundwater flow pathways are the plumbing of the continents. Water coalesces and flows downhill in surface channels and subsurface pathways in response to precipitation patterns and the dynamic form of river basins (catchments). Uplift of mountain ranges, caused by continental drift and volcanism, is continually countered by erosion and deposition (sedimentation) mediated by the forces of wind and water. Catchment landscapes are formed by the long geologic and biological history of the region as well as recent events such as floods, fires, and human-caused environmental disturbances (e.g., deforestation, dams, pollution, exotic species).

The term landscape is used extensively, referring generally to the collective attributes of local geography. An expansive view of a stream or river and its catchment, including natural and cultural attributes and interactions, is the “riverscape.” For a stream ecologist, a riverscape view of a catchment (river) basin encompasses the entire stream network, including interconnection with groundwater flow pathways, embedded in its terrestrial setting and flowing from the highest elevation in the catchment to the ocean, with considerable animal and human modifications of flow paths likely along the way (Fausch et al. 2002). For example, the earth’s largest catchment, the Amazon River basin, occupies over half of the South American continent. Headwaters flow from small catchments containing glaciers and snowfields over 4300 m above sea level on the spine of the Andes Mountains to feed the major tributaries. The tributary rivers converge to form the mainstem Amazon, which flows from the base of the Andes across a virtually flat plate covered by equatorial tropical forest to the Atlantic Ocean. The altitude change is less than 200 m over the nearly 3000 km length of the mainstem river from the base of the Andes to the ocean. Because of the enormous transport power of the massive water volume of the Amazon River, some channels are >100 m deep. In other places along the river corridor the channel is >5km wide, relatively shallow, and filled by sediment deposition (alluviation). Flood waters spread out over huge and heavily vegetated floodplains that support a myriad of fishes and other animals (Day and Davies 1986).

The riverscape of the Amazon River, as among all rivers, was molded over time with the river cutting steep canyons through mountain ranges while building (alluviating) expansive floodplains where the slope of the river valley decreased. Rivers drain the continents; transport sediments, nutrients, and other materials from the highlands to the lowlands and oceans; and constantly modify the biophysical character of their catchment basins. These processes occur in direct relation to a particular catchment’s global position, climate, orography, and biotic character, coupled with spatial variations in bedrock and other geomorphic features of the riverscape.

Within a catchment basin, stream channels usually grow in size and complexity in a downstream direction (Figure 1.1). The smallest or first-order stream channels in the network often begin as outflows from snowfields or springs below porous substrata forming ridges dividing one catchment from another. Two first-order streams coalesce to form a second-order channel and so on to create the network (Strahler 1963). A very large river, like the Amazon, often has several large tributaries, and each of those river tributaries may be fed by several to many smaller streams (Figure 1.1). Thus, each large catchment basin has many subcatchments.

Erosive power generally increases with stream size. Boulders, gravel, sand, and silt are transported from one reach of the stream network to the next in relation to discharge and valley geomorphometry (e.g., slope and relative resistance of substrata to erosion). Expansive deposition zones (floodplains) form between steep canyons, where downcutting predominates.

All rivers feature this basic theme of alternating cut and fill alluviation. Floodplains occur like beads on a string between gradient breaks or transitions in the altitudinal profile of the flow pathway (Leopold et al. 1964). Rivers of very old geologic age have exhausted much of their erosive power; mountains are rounded, valleys are broadly U-shaped, and river channels are single threads in the valley bottom with ancient, abandoned floodplains called terraces rising on either side. Whereas, in geologically young, recently uplifted catchments, stream power and associated erosive influence on valley form is great; mountains are steep-sided, valleys narrowly V-shaped, and the river spills out of many interconnected channels on alluvial floodplains in aggraded areas during flooding. Of course, no two rivers are exactly alike, but a general longitudinal (upstream to downstream) pattern of cut and fill alluviation usually exists (Figure 1.1A). Many small and usually erosive streams coalesce in the headwaters to form a main channel that grows in size and power with each primary tributary. The main channel alternately cuts through reaches constrained by bedrock canyons and spills water and sediments onto aggraded floodplain reaches where the river may be quite erosive (cutting) in one place and time and building sedimentary structures (filling) in another; thus, creating a suite of dynamic habitats for biota.

The riverscape at any point within the stream network is four-dimensional (Figure 1.1B). The river continuum or corridor from headwaters to ocean is the longitudinal (upstream to downstream) dimension. The second dimension is the transitional area from the river channel laterally into the terrestrial environment of the valley uplands (aquatic to terrestrial dimension). Except where rivers flow over impervious bedrock, some amount of porous alluvium is present within the channel owing to erosion at points upstream. Hence, water from the river may penetrate deeply into the substrata of the river bottom. Moreover, substrata of floodplains are composed of alluvial gravels and/or sands and silts, which allow lateral flow of river water. Hence, interstitial flow pathways constitute a vertical dimension in the river channel and on the floodplains. All of the physical dimensions change in size over time (the fourth dimension), as floods and droughts alter hydrology, sediment transport, and distribution of vegetation and other biota (Ward 1989, Stanford et al. 2005b).

Plants and animals are distributed in relation to biophysical gradients expressed by the four-dimensional nature of the stream network within catchment basins. For example, certain species of aquatic insects reside only in the cold, rocky environs of cascading headwater streams in the high mountains (rhithron environments), whereas other species are found only in the much warmer waters of the often sandy, turbid, and meandering reaches of the lowlands near the ocean (potamon environments) (Ward 1989). Thus, riverine biota have distinct preferences for specific environmental conditions that are optimal only at certain locations within longitudinal (upstream-downstream), lateral (aquatic-terrestrial), vertical (surface-ground water), and temporal (certain time) gradients that characterize lotic ecosystems (Figure 1.1). Andrewartha and Birch (1954) observed that the essence of ecology is understanding the distribution and abundance of biota. Because environmental conditions at any point in a stream are continuously influenced by conditions at points upstream, biophysical controls on distribution and abundance of riverine biota must be examined in the context...