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138-520: The River Gilpin is a river in the English county of Cumbria . Formerly, the river was in the county of Westmorland . The river rises near Gilpin Lodge in the vicinity of Bowness-on-Windermere and flows in a general south-by-south-easterly direction through Gilpinpark Plantation. At Crosthwaite , the direction of the river changes from south to south-east, continuing past the hamlet of Row into

276-497: A caddisfly ), Plecoptera (also known as a stonefly , Diptera (also known as a true fly ), some types of Coleoptera (also known as a beetle ), Odonata (the group that includes the dragonfly and the damselfly ), and some types of Hemiptera (also known as true bugs). Additional invertebrate taxa common to flowing waters include mollusks such as snails , limpets , clams , mussels , as well as crustaceans like crayfish , amphipoda and crabs . Fish are probably

414-582: A lake , an ocean , or another river. A stream refers to water that flows in a natural channel , a geographic feature that can contain flowing water. A stream may also be referred to as a watercourse. The study of the movement of water as it occurs on Earth is called hydrology , and their effect on the landscape is covered by geomorphology . Rivers are part of the water cycle , the continuous processes by which water moves about Earth. This means that all water that flows in rivers must ultimately come from precipitation . The sides of rivers have land that

552-547: A trip hammer , and grind grains with a millstone . In the Middle Ages , water mills began to automate many aspects of manual labor , and spread rapidly. By 1300, there were at least 10,000 mills in England alone. A medieval watermill could do the work of 30–60 human workers. Water mills were often used in conjunction with dams to focus and increase the speed of the water. Water wheels continued to be used up to and through

690-740: A boat along certain stretches. In these religions, such as that of the Altai in Russia , the river is considered a living being that must be afforded respect. Rivers are some of the most sacred places in Hinduism. There is archeological evidence that mass ritual bathing in rivers at least 5,000 years ago in the Indus river valley . While most rivers in India are revered, the Ganges is most sacred. The river has

828-644: A broad spectrum of tolerances to conditions ranging, from oligotrophic to eutrophic. Algae, consisting of phytoplankton and periphyton , are the most significant sources of primary production in most streams and rivers. Phytoplankton float freely in the water column and thus are unable to maintain populations in fast flowing streams. They can, however, develop sizeable populations in slow moving rivers and backwaters. Periphyton are typically filamentous and tufted algae that can attach themselves to objects to avoid being washed away by fast currents. In places where flow rates are negligible or absent, periphyton may form

966-515: A central role in various Hindu myths, and its water is said to have properties of healing as well as absolution from sins. Hindus believe that when the cremated remains of a person is released into the Ganges, their soul is released from the mortal world. Freshwater fish make up 40% of the world's fish species, but 20% of these species are known to have gone extinct in recent years. Human uses of rivers make these species especially vulnerable. Dams and other engineered changes to rivers can block

1104-406: A combination of factors such as historical rates of speciation and extinction , type of substrate , microhabitat availability, water chemistry, temperature, and disturbance such as flooding seem to be important. Although many alternate theories have been postulated for the ability of guild-mates to coexist (see Morin 1999), resource partitioning has been well documented in lotic systems as

1242-413: A common outlet. Rivers have a great effect on the landscape around them. They may regularly overflow their banks and flood the surrounding area, spreading nutrients to the surrounding area. Sediment or alluvium carried by rivers shapes the landscape around it, forming deltas and islands where the flow slows down. Rivers rarely run in a straight line, instead, they bend or meander ; the locations of

1380-408: A continuous flow of water throughout the year. This may be because an arid climate is too dry depending on the season to support a stream, or because a river is seasonally frozen in the winter (such as in an area with substantial permafrost ), or in the headwaters of rivers in mountains, where snowmelt is required to fuel the river. These rivers can appear in a variety of climates, and still provide

1518-413: A flying stage and spend their entire life cycle in the river. Like most of the primary consumers, lotic invertebrates often rely heavily on the current to bring them food and oxygen. Invertebrates are important as both consumers and prey items in lotic systems. The common orders of insects that are found in river ecosystems include Ephemeroptera (also known as a mayfly ), Trichoptera (also known as

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1656-770: A gelatinous, unanchored floating mat. Plants exhibit limited adaptations to fast flow and are most successful in reduced currents. More primitive plants, such as mosses and liverworts attach themselves to solid objects. This typically occurs in colder headwaters where the mostly rocky substrate offers attachment sites. Some plants are free floating at the water's surface in dense mats like duckweed or water hyacinth . Others are rooted and may be classified as submerged or emergent. Rooted plants usually occur in areas of slackened current where fine-grained soils are found. These rooted plants are flexible, with elongated leaves that offer minimal resistance to current. Living in flowing water can be beneficial to plants and algae because

1794-564: A habitat for aquatic life and perform other ecological functions. Subterranean rivers may flow underground through flooded caves. This can happen in karst systems, where rock dissolves to form caves. These rivers provide a habitat for diverse microorganisms and have become an important target of study by microbiologists . Other rivers and streams have been covered over or converted to run in tunnels due to human development. These rivers do not typically host any life, and are often used only for stormwater or flood control. One such example

1932-495: A large scale. This has been attributed to unusually large floods destroying infrastructure; however, there is evidence that permanent changes to climate causing higher aridity and lower river flow may have been the determining factor in what river civilizations succeeded or dissolved. Water wheels began to be used at least 2,000 years ago to harness the energy of rivers. Water wheels turn an axle that can supply rotational energy to move water into aqueducts , work metal using

2070-510: A means of reducing competition. The three main types of resource partitioning include habitat, dietary, and temporal segregation. Habitat segregation was found to be the most common type of resource partitioning in natural systems (Schoener, 1974). In lotic systems, microhabitats provide a level of physical complexity that can support a diverse array of organisms (Vincin and Hawknis, 1998). The separation of species by substrate preferences has been well documented for invertebrates. Ward (1992)

2208-556: A peak in the 1970s, when between two or three dams were completed every day, and has since begun to decline. New dam projects are primarily focused in China , India , and other areas in Asia . The first civilizations of Earth were born on floodplains between 5,500 and 3,500 years ago. The freshwater, fertile soil, and transportation provided by rivers helped create the conditions for complex societies to emerge. Three such civilizations were

2346-636: A predator population increases. This limits the available prey population, which limits the availability of energy for lower trophic levels within the food chain. Many biotic and abiotic factors can influence top-down and bottom-up interactions. Another example of food web interactions are trophic cascades . Understanding trophic cascades has allowed ecologists to better understand the structure and dynamics of food webs within an ecosystem. The phenomenon of trophic cascades allows keystone predators to structure entire food web in terms of how they interact with their prey. Trophic cascades can cause drastic changes in

2484-891: A ritualistic sense has been compared to the Christian ritual of baptism , famously the Baptism of Jesus in the Jordan River . Floods also appear in Norse mythology , where the world is said to emerge from a void that eleven rivers flowed into. Aboriginal Australian religion and Mesoamerican mythology also have stories of floods, some of which contain no survivors, unlike the Abrahamic flood. Along with mythological rivers, religions have also cared for specific rivers as sacred rivers. The Ancient Celtic religion saw rivers as goddesses. The Nile had many gods attached to it. The tears of

2622-425: A river can take several forms. Tidal rivers (often part of an estuary ) have their levels rise and fall with the tide . Since the levels of these rivers are often already at or near sea level, the flow of alluvium and the brackish water that flows in these rivers may be either upriver or downriver depending on the time of day. Rivers that are not tidal may form deltas that continuously deposit alluvium into

2760-420: A river in England is a stub . You can help Misplaced Pages by expanding it . River A river is a natural freshwater stream that flows on land or inside caves towards another body of water at a lower elevation , such as an ocean , lake , or another river. A river may run dry before reaching the end of its course if it runs out of water, or only flow during certain seasons. Rivers are regulated by

2898-484: A river or stream includes the general patterns of discharge over annual or decadal time scales, and may capture seasonal changes in flow. While water flow is strongly determined by slope, flowing waters can alter the general shape or direction of the stream bed, a characteristic also known as geomorphology . The profile of the river water column is made up of three primary actions: erosion, transport, and deposition. Rivers have been described as "the gutters down which run

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3036-1026: A river's banks can change frequently. Rivers get their alluvium from erosion , which carves rock into canyons and valleys . Rivers have sustained human and animal life for millennia, including the first human civilizations . The organisms that live around or in a river such as fish , aquatic plants , and insects have different roles, including processing organic matter and predation . Rivers have produced abundant resources for humans, including food , transportation , drinking water , and recreation. Humans have engineered rivers to prevent flooding, irrigate crops, perform work with water wheels , and produce hydroelectricity from dams. People associate rivers with life and fertility and have strong religious, political, social, and mythological attachments to them. Rivers and river ecosystems are threatened by water pollution , climate change , and human activity. The construction of dams, canals , levees , and other engineered structures has eliminated habitats, has caused

3174-514: 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. Most lotic species are poikilotherms whose internal temperature varies with their environment, thus temperature

3312-460: A section of the river behind them into a lake or reservoir. This can provide nearby cities with a predictable supply of drinking water. Hydroelectricity is desirable as a form of renewable energy that does not require any inputs beyond the river itself. Dams are very common worldwide, with at least 75,000 higher than 6 feet (1.8 m) in the U.S. Globally, reservoirs created by dams cover 193,500 square miles (501,000 km ). Dam-building reached

3450-410: A species is added or removed from an ecosystem it will have an effect on the remaining food web, the intensity of this effect is related to species connectedness and food web robustness. When a new species is added to a river ecosystem the intensity of the effect is related to the robustness or resistance to change of the current food web. When a species is removed from a river ecosystem the intensity of

3588-504: 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

3726-438: A water body is that body's riparian zone . Plants in the riparian zone of a river help stabilize its banks to prevent erosion and filter alluvium deposited by the river on the shore, including processing the nitrogen and other nutrients it contains. Forests in a riparian zone also provide important animal habitats . River ecosystems have also been categorized based on the variety of aquatic life they can sustain, also known as

3864-535: A water cycle that involved precipitation. The term flumen , in planetary geology , refers to channels on Saturn 's moon Titan that may carry liquid. Titan's rivers flow with liquid methane and ethane . There are river valleys that exhibit wave erosion , seas, and oceans. Scientists hope to study these systems to see how coasts erode without the influence of human activity, something that isn't possible when studying terrestrial rivers. River ecosystems The food base of streams within riparian forests

4002-474: A wide range of prey. These can be floral , faunal , and/or detrital in nature. Finally, parasites live off of host species, typically other fishes. Fish are flexible in their feeding roles, capturing different prey with regard to seasonal availability and their own developmental stage. Thus, they may occupy multiple feeding guilds in their lifetime. The number of species in each guild can vary greatly between systems, with temperate warm water streams having

4140-445: Is a tributary , and the place they meet is a confluence . Rivers must flow to lower altitudes due to gravity . The bed of a river is typically within a river valley between hills or mountains . Rivers flowing through an impermeable section of land such as rocks will erode the slopes on the sides of the river. When a river carves a plateau or a similar high-elevation area, a canyon can form, with cliffs on either side of

4278-403: 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

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4416-729: 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

4554-762: Is a large amount of organic decay occurring. Rivers can also transport suspended inorganic and organic matter. These materials can include sediment or terrestrially-derived organic matter that falls into the stream channel. Often, organic matter is processed within the stream via mechanical fragmentation, consumption and grazing by invertebrates, and microbial decomposition. Leaves and woody debris recognizable coarse particulate organic matter (CPOM) into particulate organic matter (POM), down to fine particulate organic matter. Woody and non-woody plants have different instream breakdown rates, with leafy plants or plant parts (e.g., flower petals) breaking down faster than woody logs or branches. The inorganic substrate of lotic systems

4692-633: Is a linear system of links that is part of a food web, and represents the order in which organisms are consumed from one trophic level to the next. Each link in a food chain is associated with a trophic level in the ecosystem. The numbered steps it takes for the initial source of energy starting from the bottom to reach the top of the food web is called the food chain length. While food chain lengths can fluctuate, aquatic ecosystems start with primary producers that are consumed by primary consumers which are consumed by secondary consumers, and those in turn can be consumed by tertiary consumers so on and so forth until

4830-504: Is also important for the lumber industry , as logs can be shipped via river. Countries with dense forests and networks of rivers like Sweden have historically benefited the most from this method of trade. The rise of highways and the automobile has made this practice less common. One of the first large canals was the Canal du Midi , connecting rivers within France to create a path from

4968-564: Is an ancient dam built on the Nile 4,500 years ago. The Ancient Roman civilization used aqueducts to transport water to urban areas . Spanish Muslims used mills and water wheels beginning in the seventh century. Between 130 and 1492, larger dams were built in Japan, Afghanistan, and India, including 20 dams higher than 15 metres (49 ft). Canals began to be cut in Egypt as early as 3000 BC, and

5106-448: Is at a higher elevation than the river itself, and in these areas, water flows downhill into the river. The headwaters of a river are the smaller streams that feed a river, and make up the river's source. These streams may be small and flow rapidly down the sides of mountains . All of the land uphill of a river that feeds it with water in this way is in that river's drainage basin or watershed. A ridge of higher elevation land

5244-405: Is because any natural impediment to the flow of the river may cause the current to deflect in a different direction. When this happens, the alluvium carried by the river can build up against this impediment, redirecting the course of the river. The flow is then directed against the opposite bank of the river, which will erode into a more concave shape to accommodate the flow. The bank will still block

5382-491: 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, large woody debris such as submerged tree logs, moss, and semi-aquatic plants. Substrate deposition is not necessarily a permanent event, as it can be subject to large modifications during flooding events. The living components of an ecosystem are called

5520-490: 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, substrate particle size decreases downstream with larger boulders and stones in more mountainous areas and sandy bottoms in lowland rivers. This

5658-453: Is correlated with and thus can be used to predict certain data points related to rivers, such as the size of the drainage basin (drainage area), and the length of the channel. The ecosystem of a river includes the life that lives in its water, on its banks, and in the surrounding land. The width of the channel of a river, its velocity, and how shaded it is by nearby trees. Creatures in a river ecosystem may be divided into many roles based on

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5796-741: Is ever changing with the seasons and differing habitats within the river ecosystem. Another highly variable component to river ecosystems is nutrient input from wetland and terrestrial detritus . Food and nutrient supply variability is important for the succession , robustness and connectedness of river ecosystem organisms. Energy sources can be autochthonous or allochthonous. Invertebrates can be organized into many feeding guilds in lotic systems. Some species are shredders, which use large and powerful mouth parts to feed on non-woody CPOM and their associated microorganisms. Others are suspension feeders , which use their setae , filtering aparati, nets, or even secretions to collect FPOM and microbes from

5934-448: Is foremost determined by inputs from the geology of its watershed , or catchment area. Stream water chemistry 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

6072-497: Is in part because of a projected loss of snowpack in mountains, meaning that melting snow can't replenish rivers during warm summer months, leading to lower water levels. Lower-level rivers also have warmer temperatures, threatening species like salmon that prefer colder upstream temperatures. Attempts have been made to regulate the exploitation of rivers to preserve their ecological functions. Many wetland areas have become protected from development. Water restrictions can prevent

6210-407: Is mostly derived from the trees, but wider streams and those that lack a canopy derive the majority of their food base from algae. Anadromous fish are also an important source of nutrients. Environmental threats to rivers include loss of water, dams, chemical pollution and introduced species . A dam produces negative effects that continue down the watershed. The most important negative effects are

6348-402: Is part of permafrost ice caps, or trace amounts of water vapor in the atmosphere. However, there is evidence that rivers flowed on Mars for at least 100,000 years. The Hellas Planitia is a crater left behind by an impact from an asteroid. It has sedimentary rock that was formed 3.7 billion years ago, and lava fields that are 3.3 billion years old. High resolution images of the surface of

6486-633: Is rarely static, the exact location of a river border may be called into question by countries. The Rio Grande between the United States and Mexico is regulated by the International Boundary and Water Commission to manage the right to fresh water from the river, as well as mark the exact location of the border. Up to 60% of fresh water used by countries comes from rivers that cross international borders. This can cause disputes between countries that live upstream and downstream of

6624-437: Is related to the ability of lotic systems to return to the original community configuration relatively quickly after a disturbance (Townsend et al. 1987). This is one example of temporal succession, a site-specific change in a community involving changes in species composition over time. Another form of temporal succession might occur when a new habitat is opened up for colonization . In these cases, an entirely new community that

6762-701: Is that of the Mississippi River , whose drainage basin covers 40% of the contiguous United States . The river was then used for shipping crops from the American Midwest and cotton from the American South to other states as well as the Atlantic Ocean. The role of urban rivers has evolved from when they were a center of trade, food, and transportation to modern times when these uses are less necessary. Rivers remain central to

6900-553: Is the Sunswick Creek in New York City, which was covered in the 1800s and now exists only as a sewer-like pipe. While rivers may flow into lakes or man-made features such as reservoirs , the water they contain will always tend to flow down toward the ocean . However, if human activity siphons too much water away from a river for other uses, the riverbed may run dry before reaching the sea. The outlets mouth of

7038-413: Is used by producers (plants) to turn inorganic substances into organic substances which can be used as food by consumers (animals). Plants release portions of this energy back into the ecosystem through a catabolic process. Animals then consume the potential energy that is being released from the producers. This system is followed by the death of the consumer organism which then returns nutrients back into

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7176-401: Is well adapted to the conditions found in this new area can establish itself. The River continuum concept (RCC) was an attempt to construct a single framework to describe the function of temperate lotic ecosystems from the headwaters to larger rivers and relate key characteristics to changes in the biotic community (Vannote et al. 1980). The physical basis for RCC is size and location along

7314-842: Is what typically separates drainage basins; water on one side of a ridge will flow into one set of rivers, and water on the other side will flow into another. One example of this is the Continental Divide of the Americas in the Rocky Mountains . Water on the western side of the divide flows into the Pacific Ocean , whereas water on the other side flows into the Atlantic Ocean . Not all precipitation flows directly into rivers; some water seeps into underground aquifers . These, in turn, can still feed rivers via

7452-599: The 2024 Summer Olympics . Another example is the restoration of the Isar in Munich from being a fully canalized channel with hard embankments to being wider with naturally sloped banks and vegetation. This has improved wildlife habitat in the Isar, and provided more opportunities for recreation in the river. As a natural barrier , rivers are often used as a border between countries , cities, and other territories . For example,

7590-487: The Atlantic Ocean to the Mediterranean Sea . The nineteenth century saw canal-building become more common, with the U.S. building 4,400 miles (7,100 km) of canals by 1830. Rivers began to be used by cargo ships at a larger scale, and these canals were used in conjunction with river engineering projects like dredging and straightening to ensure the efficient flow of goods. One of the largest such projects

7728-501: The Industrial Revolution as a source of power for textile mills and other factories, but were eventually supplanted by steam power . Rivers became more industrialized with the growth of technology and the human population . As fish and water could be brought from elsewhere, and goods and people could be transported via railways , pre-industrial river uses diminished in favor of more complex uses. This meant that

7866-791: The Lamari River in New Guinea separates the Angu and the Fore people in New Guinea. The two cultures speak different languages and rarely mix. 23% of international borders are large rivers (defined as those over 30 meters wide). The traditional northern border of the Roman Empire was the Danube , a river that today forms the border of Hungary and Slovakia . Since the flow of a river

8004-612: The Lyth Valley , where it is swelled by the much larger River Pool . From there, the river continues moving south to Sampool, where it meets the River Kent . The name of the river is in honour of the Gilpin family. 54°15′N 2°48′W  /  54.250°N 2.800°W  / 54.250; -2.800 This Cumbria location article is a stub . You can help Misplaced Pages by expanding it . This article related to

8142-525: The Nile and the Ganges . The Quran describes these four rivers as flowing with water, milk, wine, and honey, respectively. The book of Genesis also contains a story of a great flood . Similar myths are present in the Epic of Gilgamesh , Sumerian mythology, and in other cultures. In Genesis, the flood's role was to cleanse Earth of the wrongdoing of humanity. The act of water working to cleanse humans in

8280-568: The River Continuum Concept . "Shredders" are organisms that consume this organic material. The role of a "grazer" or "scraper" organism is to feed on the algae that collects on rocks and plants. "Collectors" consume the detritus of dead organisms. Lastly, predators feed on living things to survive. The river can then be modeled by the availability of resources for each creature's role. A shady area with deciduous trees might experience frequent deposits of organic matter in

8418-627: The River Lethe to forget their previous life. Rivers also appear in descriptions of paradise in Abrahamic religions , beginning with the story of Genesis . A river beginning in the Garden of Eden waters the garden and then splits into four rivers that flow to provide water to the world. These rivers include the Tigris and Euphrates , and two rivers that are possibly apocryphal but may refer to

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8556-787: The Sumerians in the Tigris–Euphrates river system , the Ancient Egyptian civilization in the Nile, and the Indus Valley Civilization on the Indus River . The desert climates of the surrounding areas made these societies especially reliant on rivers for survival, leading to people clustering in these areas to form the first cities . It is also thought that these civilizations were the first to organize

8694-485: The climate . The alluvium carried by rivers, laden with minerals, is deposited into the floodplain when the banks spill over, providing new nutrients to the soil, allowing them to support human activity like farming as well as a host of plant and animal life. Deposited sediment from rivers can form temporary or long-lasting fluvial islands . These islands exist in almost every river. About half of all waterways on Earth are intermittent rivers , which do not always have

8832-685: The cultural identity of cities and nations. Famous examples include the River Thames 's relationship to London , the Seine to Paris , and the Hudson River to New York City . The restoration of water quality and recreation to urban rivers has been a goal of modern administrations. For example, swimming was banned in the Seine for over 100 years due to concerns about pollution and the spread of E. coli , until cleanup efforts to allow its use in

8970-484: The discharge of a river, the amount of water passing through it at a particular time. The flow of a river can act as a means of transportation for plant and animal species, as well as a barrier. For example, the Amazon River is so wide in parts that the variety of species on either side of its basin are distinct. Some fish may swim upstream to spawn as part of a seasonal migration . Species that travel from

9108-465: The extinction of some species, and lowered the amount of alluvium flowing through rivers. Decreased snowfall from climate change has resulted in less water available for rivers during the summer. Regulation of pollution, dam removal , and sewage treatment have helped to improve water quality and restore river habitats. A river is a natural flow of freshwater that flows on or through land towards another body of water downhill. This flow can be into

9246-436: The sea . The sediment yield of a river is the quantity of sand per unit area within a watershed that is removed over a period of time. The monitoring of the sediment yield of a river is important for ecologists to understand the health of its ecosystems, the rate of erosion of the river's environment, and the effects of human activity. Rivers rarely run in a straight direction, instead preferring to bend or meander . This

9384-429: The water column , and gatherers who feed on FPOM found on the substrate of the river or stream. The secondary consumers in a river ecosystem are the predators of the primary consumers. This includes mainly insectivorous fish. Consumption by invertebrate insects and macro-invertebrates is another step of energy flow up the food chain. Depending on their abundance, these predatory consumers can shape an ecosystem by

9522-475: The water cycle , the processes by which water moves around the Earth. Water first enters rivers through precipitation , whether from rainfall, the runoff of water down a slope, the melting of glaciers or snow , or seepage from aquifers beneath the surface of the Earth. Rivers flow in channeled watercourses and merge in confluences to form drainage basins , areas where surface water eventually flows to

9660-521: The water table , the groundwater beneath the surface of the land stored in the soil . Water flows into rivers in places where the river's elevation is lower than that of the water table. This phenomenon is why rivers can still flow even during times of drought . Rivers are also fed by the melting of snow glaciers present in higher elevation regions. In summer months, higher temperatures melt snow and ice, causing additional water to flow into rivers. Glacier melt can supplement snow melt in times like

9798-457: The best-known inhabitants of lotic systems. The ability of a fish species to live in flowing waters depends upon the speed at which it can swim and the duration that its speed can be maintained. This ability can vary greatly between species and is tied to the habitat in which it can survive. Continuous swimming expends a tremendous amount of energy and, therefore, fishes spend only short periods in full current. Instead, individuals remain close to

9936-553: The biota is specialized to live with flow conditions. The non-living components of an ecosystem are called abiotic components. E.g. stone, air, soil, etc. Unidirectional water flow is the key factor in lotic systems influencing their ecology. Streamflow can be continuous or intermittent, though. Streamflow is the result of the summative inputs from groundwater, precipitation, and overland flow. Water flow can vary between systems, ranging from torrential rapids to slow backwaters that almost seem like lentic systems. The speed or velocity of

10074-424: The biotic components. Streams have numerous types of biotic organisms that live in them, including bacteria, primary producers, insects and other invertebrates, as well as fish and other vertebrates. A biofilm is a combination of algae (diatoms etc.), fungi, bacteria, and other small microorganisms that exist in a film along the streambed or the benthos . Biofilm assemblages themselves are complex, and add to

10212-695: The bodies of humans and animals worldwide, as well as in the soil, with potentially negative health effects. Research into how to remove it from the environment, and how harmful exposure is, is ongoing. Fertilizer from farms can lead to a proliferation of algae on the surface of rivers and oceans, which prevents oxygen and light from dissolving into water, making it impossible for underwater life to survive in these so-called dead zones . Urban rivers are typically surrounded by impermeable surfaces like stone, asphalt , and concrete. Cities often have storm drains that direct this water to rivers. This can cause flooding risk as large amounts of water are directed into

10350-442: The bottom or the banks, behind obstacles, and sheltered from the current, swimming in the current only to feed or change locations. Some species have adapted to living only on the system bottom, never venturing into the open water flow. These fishes are dorso-ventrally flattened to reduce flow resistance and often have eyes on top of their heads to observe what is happening above them. Some also have sensory barrels positioned under

10488-527: 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 the flow rate. The amount of water in a stream is measured as discharge (volume per unit time). As water flows downstream, streams and rivers most often gain water volume, so at base flow (i.e., no storm input), smaller headwater streams have very low discharge, while larger rivers have much higher discharge. The "flow regime" of

10626-405: The complete draining of rivers. Limits on the construction of dams, as well as dam removal , can restore the natural habitats of river species. Regulators can also ensure regular releases of water from dams to keep animal habitats supplied with water. Limits on pollutants like pesticides can help improve water quality. Today, the surface of Mars does not have liquid water. All water on Mars

10764-445: The complexity of a streambed. The different biofilm components (algae and bacteria are the principal components) are embedded in an exopolysaccharide matrix (EPS), and are net receptors of inorganic and organic elements and remain submitted to the influences of the different environmental factors. Biofilms are one of the main biological interphases in river ecosystems, and probably the most important in intermittent rivers , where

10902-541: The current is usually well aerated and it provides a continuous supply of nutrients. These organisms are limited by flow, light, water chemistry, substrate, and grazing pressure. Algae and plants are important to lotic systems as sources of energy, for forming microhabitats that shelter other fauna from predators and the current, and as a food resource. Up to 90% of invertebrates in some lotic systems are insects . These species exhibit tremendous diversity and can be found occupying almost every available habitat, including

11040-399: The ecosystem. This allow further growth for the plants, and the cycle continues. Breaking cycles down into levels makes it easier for ecologists to understand ecological succession when observing the transfer of energy within a system. A common issue with trophic level dynamics is how resources and production are regulated. The usage and interaction between resources have a large impact on

11178-403: The effect is related to the connectedness of the species to the food web. An invasive species could be removed with little to no effect, but if important and native primary producers, prey or predatory fish are removed you could have a negative trophic cascade . One highly variable component to river ecosystems is food supply ( biomass of primary producers ). Food supply or type of producers

11316-436: The effect of normalizing the effects of rivers; the greatest floods are smaller and more predictable, and larger sections are open for navigation by boats and other watercraft. A major effect of river engineering has been a reduced sediment output of large rivers. For example, the Mississippi River produced 400 million tons of sediment per year. Due to the construction of reservoirs , sediment buildup in man-made levees , and

11454-449: The energy flow within a food web. For example, when a top or keystone predator consumes organisms below them in the food web, the density and behavior of the prey will change. This, in turn, affects the abundance of organisms consumed further down the chain, resulting in a cascade down the trophic levels. However, empirical evidence shows trophic cascades are much more prevalent in terrestrial food webs than aquatic food webs. A food chain

11592-451: The fish zonation concept. Smaller rivers can only sustain smaller fish that can comfortably fit in its waters, whereas larger rivers can contain both small fish and large fish. This means that larger rivers can host a larger variety of species. This is analogous to the species-area relationship , the concept of larger habitats being host to more species. In this case, it is known as the species-discharge relationship, referring specifically to

11730-673: The floating of wood on rivers to transport it, was especially important. Rivers also were an important source of drinking water . For civilizations built around rivers, fish were an important part of the diet of humans. Some rivers supported fishing activities, but were ill-suited to farming, such as those in the Pacific Northwest . Other animals that live in or near rivers like frogs , mussels , and beavers could provide food and valuable goods such as fur . Humans have been building infrastructure to use rivers for thousands of years. The Sadd el-Kafara dam near Cairo , Egypt,

11868-412: The flow of the river beneath its surface. These help rivers flow straighter by increasing the speed of the water at the middle of the channel, helping to control floods. Levees are also used for this purpose. They can be thought of as dams constructed on the sides of rivers, meant to hold back water from flooding the surrounding area during periods of high rainfall. They are often constructed by building up

12006-399: The flow, causing it to reflect in the other direction. Thus, a bend in the river is created. Rivers may run through low, flat regions on their way to the sea. These places may have floodplains that are periodically flooded when there is a high level of water running through the river. These events may be referred to as "wet seasons' and "dry seasons" when the flooding is predictable due to

12144-538: The food chain and depending on the ecosystem, may end with these predatory fish. Diversity , productivity , species richness , composition and stability are all interconnected by a series of feedback loops. Communities can have a series of complex, direct and/or indirect, responses to major changes in biodiversity . Food webs can include a wide array of variables, the three main variables ecologists look at regarding ecosystems include species richness, biomass of productivity and stability /resistant to change. When

12282-399: The food chain. Primary producers are consumed by herbivorous invertebrates that act as the primary consumers . Productivity of these producers and the function of the ecosystem as a whole are influenced by the organism above it in the food chain. Primary consumers are the invertebrates and macro-invertebrates that feed upon the primary producers. They play an important role in initiating

12420-650: The form of leaves. In this type of ecosystem, collectors and shredders will be most active. As the river becomes deeper and wider, it may move slower and receive more sunlight . This supports invertebrates and a variety of fish , as well as scrapers feeding on algae. Further downstream, the river may get most of its energy from organic matter that was already processed upstream by collectors and shredders. Predators may be more active here, including fish that feed on plants, plankton , and other fish. The flood pulse concept focuses on habitats that flood seasonally, including lakes and marshes . The land that interfaces with

12558-461: The formation of synergistic consortia. The EPS is able to retain extracellular enzymes and therefore allows the utilization of materials from the environment and the transformation of these materials into dissolved nutrients for the use by algae and bacteria. At the same time, the EPS contributes to protect the cells from desiccation as well from other hazards (e.g., biocides , UV radiation , etc.) from

12696-582: The giant water bug ( Belostomatidae ), avoid flood events by leaving the stream when they sense rainfall. In addition to these behaviors and body shapes, insects have different life history adaptations to cope with the naturally-occurring physical harshness of stream environments. Some insects time their life events based on when floods and droughts occur. For example, some mayflies synchronize when they emerge as flying adults with when snowmelt flooding usually occurs in Colorado streams. Other insects do not have

12834-508: The goddess Isis were said to be the cause of the river's yearly flooding, itself personified by the goddess Hapi . Many African religions regard certain rivers as the originator of life. In Yoruba religion , Yemọja rules over the Ogun River in modern-day Nigeria and is responsible for creating all children and fish. Some sacred rivers have religious prohibitions attached to them, such as not being allowed to drink from them or ride in

12972-484: The gravel, while others reside in the crevices between one piece of gravel and the next, while still others live on the bottom of this gravel piece. Dietary segregation is the second-most common type of resource partitioning. High degrees of morphological specializations or behavioral differences allow organisms to use specific resources. The size of nets built by some species of invertebrate suspension feeders , for example, can filter varying particle size of FPOM from

13110-452: The head to assist in the testing of substratum. Lotic systems typically connect to each other, forming a path to the ocean (spring → stream → river → ocean), and many fishes have life cycles that require stages in both fresh and salt water. Salmon , for example, are anadromous species that are born in freshwater but spend most of their adult life in the ocean, returning to fresh water only to spawn. Eels are catadromous species that do

13248-413: The importance of the water column is reduced during extended low-activity periods of the hydrological cycle . Biofilms can be understood as microbial consortia of autotrophs and heterotrophs , coexisting in a matrix of hydrated extracellular polymeric substances (EPS). These two main biological components are respectively mainly algae and cyanobacteria on one side, and bacteria and fungi on

13386-428: The irrigation of desert environments for growing food. Growing food at scale allowed people to specialize in other roles, form hierarchies, and organize themselves in new ways, leading to the birth of civilization. In pre-industrial society , rivers were a source of transportation and abundant resources. Many civilizations depended on what resources were local to them to survive. Shipping of commodities, especially

13524-429: The larger vertebrates mentioned above. The concept of trophic levels are used in food webs to visualise the manner in which energy is transferred from one part of an ecosystem to another. Trophic levels can be assigned numbers determining how far an organism is along the food chain . All energy transactions within an ecosystem derive from a single external source of energy, the sun. Some of this solar radiation

13662-434: The late summer, when there may be less snow left to melt, helping to ensure that the rivers downstream of the glaciers have a continuous supply of water. Rivers flow downhill, with their direction determined by gravity . A common misconception holds that all or most rivers flow from North to South, but this is not true. As rivers flow downstream, they eventually merge to form larger rivers. A river that feeds into another

13800-474: The level of river branching in a drainage basin. Several systems of stream order exist, one of which is the Strahler number . In this system, the first tributaries of a river are 1st order rivers. When two 1st order rivers merge, the resulting river is 2nd order. If a river of a higher order and a lower order merge, the order is incremented from whichever of the previous rivers had the higher order. Stream order

13938-561: The local ecosystems of rivers needed less protection as humans became less reliant on them for their continued flourishing. River engineering began to develop projects that enabled industrial hydropower , canals for the more efficient movement of goods, as well as projects for flood prevention . River transportation has historically been significantly cheaper and faster than transportation by land. Rivers helped fuel urbanization as goods such as grain and fuel could be floated downriver to supply cities with resources. River transportation

14076-405: The manner in which they affect the trophic levels below them. When fish are at high abundance and eat lots of invertebrates, then algal biomass and primary production in the stream is greater, and when secondary consumers are not present, then algal biomass may decrease due to the high abundance of primary consumers. Energy and nutrients that starts with primary producers continues to make its way up

14214-407: The mechanical shadoof began to be used to raise the elevation of water. Drought years harmed crop yields, and leaders of society were incentivized to ensure regular water and food availability to remain in power. Engineering projects like the shadoof and canals could help prevent these crises. Despite this, there is evidence that floodplain-based civilizations may have been abandoned occasionally at

14352-400: The migration routes of fish and destroy habitats. Rivers that flow freely from headwaters to the sea have better water quality, and also retain their ability to transport nutrient-rich alluvium and other organic material downstream, keeping the ecosystem healthy. The creation of a lake changes the habitat of that portion of water, and blocks the transportation of sediment, as well as preventing

14490-400: The more general study area of freshwater or aquatic ecology . The following unifying characteristics make the ecology of running waters unique among aquatic habitats: the flow is unidirectional, there is a state of continuous physical change, and there is a high degree of spatial and temporal heterogeneity at all scales ( microhabitats ), the variability between lotic systems is quite high and

14628-432: The most benthic invertebrate feeders, and tropical systems having large numbers of detritus feeders due to high rates of allochthonous input. Large rivers have comparatively more species than small streams. Many relate this pattern to the greater area and volume of larger systems, as well as an increase in habitat diversity. Some systems, however, show a poor fit between system size and species richness . In these cases,

14766-399: The natural meandering of the river. Dams block the migration of fish such as salmon for which fish ladder and other bypass systems have been attempted, but these are not always effective. Pollution from factories and urban areas can also damage water quality. " Per- and polyfluoroalkyl substances (PFAS) is a widely used chemical that breaks down at a slow rate. It has been found in

14904-429: The natural terrain with soil or clay. Some levees are supplemented with floodways, channels used to redirect floodwater away from farms and populated areas. Dams restrict the flow of water through a river. They can be built for navigational purposes, providing a higher level of water upstream for boats to travel in. They may also be used for hydroelectricity , or power generation from rivers. Dams typically transform

15042-755: The number of species and the abundance of individuals within each guild is largely dependent upon food availability. Thus, these values may vary across both seasons and systems. Fish can also be placed into feeding guilds . Planktivores pick plankton out of the water column . Herbivore - detritivores are bottom-feeding species that ingest both periphyton and detritus indiscriminately. Surface and water column feeders capture surface prey (mainly terrestrial and emerging insects) and drift ( benthic invertebrates floating downstream). Benthic invertebrate feeders prey primarily on immature insects, but will also consume other benthic invertebrates. Top predators consume fishes and/or large invertebrates. Omnivores ingest

15180-551: The opposite , living in freshwater as adults but migrating to the ocean to spawn. Other vertebrate taxa that inhabit lotic systems include amphibians , such as salamanders , reptiles (e.g. snakes, turtles, crocodiles and alligators) various bird species, and mammals (e.g., otters , beavers , hippos , and river dolphins ). With the exception of a few species, these vertebrates are not tied to water as fishes are, and spend part of their time in terrestrial habitats. Many fish species are important as consumers and as prey species to

15318-481: The other. Micro - and meiofauna also inhabit the biofilm, predating on the organisms and organic particles and contributing to its evolution and dispersal. Biofilms therefore form a highly active biological consortium, ready to use organic and inorganic materials from the water phase, and also ready to use light or chemical energy sources. The EPS immobilize the cells and keep them in close proximity allowing for intense interactions including cell-cell communication and

15456-593: The outer world. On the other hand, the packing and the EPS protection layer limits the diffusion of gases and nutrients, especially for the cells far from the biofilm surface, and this limits their survival and creates strong gradients within the biofilm. Both the biofilm physical structure, and the plasticity of the organisms that live within it, ensure and support their survival in harsh environments or under changing environmental conditions. Bacteria are present in large numbers in lotic waters. Free-living forms are associated with decomposing organic material, biofilm on

15594-425: The plain show evidence of a river network, and even river deltas. These images reveal channels formed in the rock, recognized by geologists who study rivers on Earth as being formed by rivers, as well as "bench and slope" landforms, outcroppings of rock that show evidence of river erosion. Not only do these formations suggest that rivers once existed, but that they flowed for extensive time periods, and were part of

15732-735: The reduction of spring flooding, which damages wetlands, and the retention of sediment, which leads to the loss of deltaic wetlands. River ecosystems are prime examples of lotic ecosystems. Lotic refers to flowing water, from the Latin lotus , meaning 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, ponds, and wetlands . Together, these two ecosystems form

15870-448: The removal of natural banks replaced with revetments , this sediment output has been reduced by 60%. The most basic river projects involve the clearing of obstructions like fallen trees. This can scale up to dredging , the excavation of sediment buildup in a channel, to provide a deeper area for navigation. These activities require regular maintenance as the location of the river banks changes over time, floods bring foreign objects into

16008-536: The reverse, death and destruction, especially through floods . This power has caused rivers to have a central role in religion , ritual , and mythology . In Greek mythology , the underworld is bordered by several rivers. Ancient Greeks believed that the souls of those who perished had to be borne across the River Styx on a boat by Charon in exchange for money. Souls that were judged to be good were admitted to Elysium and permitted to drink water from

16146-488: The river's source. In terms of dissolved gases, 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 pH and 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

16284-468: The river, and natural sediment buildup continues. Artificial channels are often constructed to "cut off" winding sections of a river with a shorter path, or to direct the flow of a river in a straighter direction. This effect, known as channelization, has made the distance required to traverse the Missouri River in 116 kilometres (72 mi) shorter. Dikes are channels built perpendicular to

16422-552: The river. Areas of a river with softer rock weather faster than areas with harder rock, causing a difference in elevation between two points of a river. This can cause the formation of a waterfall as the river's flow falls down a vertical drop. A river in a permeable area does not exhibit this behavior and may even have raised banks due to sediment. Rivers also change their landscape through their transportation of sediment , often known as alluvium when applied specifically to rivers. This debris comes from erosion performed by

16560-625: The river. A country that is downstream of another may object to the upstream country diverting too much water for agricultural uses, pollution, as well as the creation of dams that change the river's flow characteristics. For example, Egypt has an agreement with Sudan requiring a specific minimum volume of water to pass into the Nile yearly over the Aswan Dam , to maintain both countries access to water. The importance of rivers throughout human history has given them an association with life and fertility . They have also become associated with

16698-457: The rivers themselves, debris swept into rivers by rainfall, as well as erosion caused by the slow movement of glaciers. The sand in deserts and the sediment that forms bar islands is from rivers. The particle size of the debris is gradually sorted by the river, with heavier particles like rocks sinking to the bottom, and finer particles like sand or silt carried further downriver . This sediment may be deposited in river valleys or carried to

16836-412: The rivers. Due to these impermeable surfaces, these rivers often have very little alluvium carried in them, causing more erosion once the river exits the impermeable area. It has historically been common for sewage to be directed directly to rivers via sewer systems without being treated, along with pollution from industry. This has resulted in a loss of animal and plant life in urban rivers, as well as

16974-493: The ruins of continents". Rivers are continuously eroding , transporting, and depositing substrate, sediment, and organic material. The continuous movement of water and entrained material creates 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

17112-409: The sea from their mouths. Depending on the activity of waves, the strength of the river, and the strength of the tidal current, the sediment can accumulate to form new land. When viewed from above, a delta can appear to take the form of several triangular shapes as the river mouth appears to fan out from the original coastline . In hydrology , a stream order is a positive integer used to describe

17250-414: The sea to breed in freshwater rivers are anadromous. Salmon are an anadromous fish that may die in the river after spawning, contributing nutrients back to the river ecosystem. Modern river engineering involves a large-scale collection of independent river engineering structures that have the goal of flood control , improved navigation, recreation, and ecosystem management. Many of these projects have

17388-519: The spread of waterborne diseases such as cholera . In modern times, sewage treatment and controls on pollution from factories have improved the water quality of urban rivers. Climate change can change the flooding cycles and water supply available to rivers. Floods can be larger and more destructive than expected, causing damage to the surrounding areas. Floods can also wash unhealthy chemicals and sediment into rivers. Droughts can be deeper and longer, causing rivers to run dangerously low. This

17526-477: The stream; examples can include inorganic nitrogen species such as nitrate or ammonium, some forms of phosphorus (e.g., soluble reactive phosphorus), and silica. Other solutes can be considered conservative, which indicates that the solute is not taken up and used biologically; chloride is often considered a conservative solute. Conservative solutes are often used as hydrologic tracers for water movement and transport. Both reactive and conservative stream water chemistry

17664-423: The structure of food webs as a whole. Temperature plays a role in food web interactions including top-down and bottom-up forces within ecological communities. Bottom-up regulations within a food web occur when a resource available at the base or bottom of the food web increases productivity, which then climbs the chain and influence the biomass availability to higher trophic organism. Top-down regulations occur when

17802-564: The surfaces of rocks and vegetation, in between particles that compose the substrate, and suspended in the water column . Other forms are also associated with the guts of lotic organisms as parasites or in commensal relationships. Bacteria play a large role in energy recycling (see below ). Diatoms are one of the main dominant groups of periphytic algae in lotic systems and have been widely used as efficient indicators of water quality, because they respond quickly to environmental changes, especially organic pollution and eutrophication, with

17940-423: The surfaces of stones, deep below the substratum in the hyporheic zone , adrift in the current, and in the surface film. Insects have developed several strategies for living in the diverse flows of lotic systems. Some avoid high current areas, inhabiting the substratum or the sheltered side of rocks. Others have flat bodies to reduce the drag forces they experience from living in running water. Some insects, like

18078-429: The temperature of lotic systems. Water chemistry in river ecosystems varies depending on which dissolved solutes and gases are present in the water column of the stream. Specifically river water can include, apart from the water itself, Dissolved stream solutes can be considered either reactive or conservative . Reactive solutes are readily biologically assimilated by the autotrophic and heterotrophic biota of

18216-724: The timing of maximum growth among guild mates. Tropical fishes in Borneo , for example, have shifted to shorter life spans in response to the ecological niche reduction felt with increasing levels of species richness in their ecosystem (Watson and Balon 1984). Over long time scales, there is a tendency for species composition in pristine systems to remain in a stable state. This has been found for both invertebrate and fish species. On shorter time scales, however, flow variability and unusual precipitation patterns decrease habitat stability and can all lead to declines in persistence levels. The ability to maintain this persistence over long time scales

18354-409: The top of the food chain has been reached. Primary producers start every food chain. Their production of energy and nutrients comes from the sun through photosynthesis . Algae contributes to a lot of the energy and nutrients at the base of the food chain along with terrestrial litter-fall that enters the stream or river . Production of organic compounds like carbon is what gets transferred up

18492-664: The transfer of energy from the base trophic level to the next. They are regulatory organisms which facilitate and control rates of nutrient cycling and the mixing of aquatic and terrestrial plant materials. They also transport and retain some of those nutrients and materials. There are many different functional groups of these invertebrate, including grazers, organisms that feed on algal biofilm that collects on submerged objects, shredders that feed on large leaves and detritus and help break down large material. Also filter feeders , macro-invertebrates that rely on stream flow to deliver them fine particulate organic matter (FPOM) suspended in

18630-490: The water (Edington et al. 1984). Similarly, members in the grazing guild can specialize in the harvesting of algae or detritus depending upon the morphology of their scraping apparatus. In addition, certain species seem to show a preference for specific algal species. Temporal segregation is a less common form of resource partitioning, but it is nonetheless an observed phenomenon. Typically, it accounts for coexistence by relating it to differences in life history patterns and

18768-419: The water flow of the water column can also vary within a system and is subject to chaotic turbulence, though water velocity tends to be highest in the middle part of the stream channel (known as the thalveg ). 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 the variability of friction with the bottom or sides of

18906-585: The water. These species may be passive collectors, utilizing the natural flow of the system, or they may generate their own current to draw water, and also, FPOM in Allan. Members of the gatherer-collector guild actively search for FPOM under rocks and in other places where the stream flow has slackened enough to allow deposition. Grazing invertebrates utilize scraping, rasping, and browsing adaptations to feed on periphyton and detritus . Finally, several families are predatory, capturing and consuming animal prey. Both

19044-403: Was able to divide substrate dwellers into six broad assemblages, including those that live in: coarse substrate, gravel, sand, mud, woody debris, and those associated with plants, showing one layer of segregation. On a smaller scale, further habitat partitioning can occur on or around a single substrate, such as a piece of gravel. Some invertebrates prefer the high flow areas on the exposed top of

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