Rivers with a course that drops in altitude very slowly will have slower water flow and lower force. This in turn produces the other characteristics of a lowland river - a meandering course lacking rapids, a river bed dominated by fine sediments and higher water temperatures. Lowland rivers tend to carry more suspended sediment and organic matter as well, but some lowland rivers have periods of high water clarity in seasonal low flow periods. The generally clear, cool, fast-flowing waters and bedrock and coarse sediment beds of upland rivers encourage fish species with limited temperature tolerances, high oxygen needs, strong swimming ability and specialized reproductive strategies to prevent eggs or larvae being swept away. These characteristics also encourage invertebrate species with limited temperature tolerances, high oxygen needs and ecologies revolving around coarse sediments and interstices or "gaps" between those coarse sediments.