River ecosystems are prime examples of lotic ecosystems. Lotic refers to flowing water, from the Latin lotus, washed. Lotic waters range from springs only a few centimeters wide to major rivers kilometers in width. Much of this article applies to lotic ecosystems in general, including related lotic systems such as streams and springs. Lotic ecosystems can be contrasted with lentic ecosystems, which involve relatively still terrestrial waters such as lakes and ponds. Together, these two fields form the more general study area of freshwater oraquatic ecology.
The following unifying characteristics make the ecology of running waters unique from that of other aquatic habitats.
- Flow is unidirectional.
- There is a state of continuous physical change.
- There is a high degree of spatial and temporal heterogeneity at all scales (microhabitats).
- Variability between lotic systems is quite high.
- The biota is specialized to live with flow conditions.
Water flow is the key factor in lotic systems influencing their ecology. The strength of water flow can vary between systems, ranging from torrential rapids to slow backwaters that almost seem like lentic systems. The speed of the water flow can also vary within a system and is subject to chaotic turbulence. This turbulence results in divergences of flow from the mean downslope flow vector as typified by eddy currents. The mean flow rate vector is based on variability of friction with the bottom or sides of the channel, sinuosity, obstructions, and the incline gradient. In addition, the amount of water input into the system from direct precipitation, snowmelt, and/or groundwater can affect flow rate. Flowing waters can alter the shape of the streambed through erosion and deposition, creating a variety of habitats, including riffles, glides, and pools.
Light is important to lotic systems, because it provides the energy necessary to drive primary production via photosynthesis, and can also provide refuge for prey species in shadows it casts. The amount of light that a system receives can be related to a combination of internal and external stream variables. The area surrounding a small stream, for example, might be shaded by surrounding forests or by valley walls. Larger river systems tend to be wide so the influence of external variables is minimized, and the sun reaches the surface. These rivers also tend to be more turbulent, however, and particles in the water increasingly attenuate light as depth increases. Seasonal and diurnal factors might also play a role in light availability because the angle of incidence, the angle at which light strikes water can lead to light lost from reflection. Known as Beer’s Law, the shallower the angle, the more light is reflected and the amount of solar radiation received declines logarithmically with depth. Additional influences on light availability include cloud cover, altitude, and geographic position (Brown 1987).
Most lotic species are poikilotherms whose internal temperature varies with their environment, thus temperature is a key abiotic factor for them. Water can be heated or cooled through radiation at the surface and conduction to or from the air and surrounding substrate. Shallow streams are typically well mixed and maintain a relatively uniform temperature within an area. In deeper, slower moving water systems, however, a strong difference between the bottom and surface temperatures may develop. Spring fed systems have little variation as springs are typically from groundwater sources, which are often very close to ambient temperature. Many systems show strong diurnal fluctuations and seasonal variations are most extreme in arctic, desert and temperate systems. The amount of shading, climate and elevation can also influence the temperature of lotic systems.
Water chemistry between systems varies tremendously. The chemistry is foremost determined by inputs from the geology of its watershed, or catchment area, but can also be influenced by precipitation and the addition of pollutants from human sources. Large differences in chemistry do not usually exist within small lotic systems due to a high rate of mixing. In larger river systems, however, the concentrations of most nutrients, dissolved salts, and pH decrease as distance increases from the river’s source.
Oxygen is likely the most important chemical constituent of lotic systems, as all aerobic organisms require it for survival. It enters the water mostly via diffusion at the water-air interface. Oxygen’s solubility in water decreases as water temperature increases. Fast, turbulent streams expose more of the water’s surface area to the air and tend to have low temperatures and thus more oxygen than slow, backwaters. Oxygen is a byproduct of photosynthesis, so systems with a high abundance of aquatic algae and plants may also have high concentrations of oxygen during the day. These levels can decrease significantly during the night when primary producers switch to respiration. Oxygen can be limiting if circulation between the surface and deeper layers is poor, if the activity of lotic animals is very high, or if there is a large amount of organic decay occurring.
The inorganic substrate of lotic systems is composed of the geologic material present in the catchment that is eroded, transported, sorted, and deposited by the current. Inorganic substrates are classified by size on the Wentworth scale, which ranges from boulders, to pebbles, to gravel, to sand, and to silt. Typically, particle size decreases downstream with larger boulders and stones in more mountainous areas and sandy bottoms in lowland rivers. This is because the higher gradients of mountain streams facilitate a faster flow, moving smaller substrate materials further downstream for deposition. Substrate can also be organic and may include fine particles, autumn shed leaves, submerged wood, moss, and more evolved plants. Substrate deposition is not necessarily a permanent event, as it can be subject to large modifications during flooding events.