River ecosystems are flowing waters that drain the landscape, and include the biotic (living) interactions amongst plants, animals and micro-organisms, as well as abiotic (nonliving) physical and chemical interactions of its many parts.Angelier, E. 2003. Ecology of Streams and Rivers. Science Publishers, Inc., Enfield. Pp. 215.”Biology Concepts & Connections Sixth Edition”, Campbell, Neil A. (2009), page 2, 3 and G-9. Retrieved 2010-06-14. River ecosystems are part of larger

watershed Watershed is a hydrological term, which has been adopted in other fields in a more or less figurative sense. It may refer to: Hydrology * Drainage divide, the line that separates neighbouring drainage basins * Drainage basin, called a "watershe ...

networks or catchments, where smaller

headwater The headwaters of a river or stream is the farthest place in that river or stream from its estuary or downstream confluence with another river, as measured along the course of the river. It is also known as a river's source. Definition The ...

streams drain into mid-size streams, which progressively drain into larger river networks. The major zones in river ecosystems are determined by the river bed's gradient or by the velocity of the current. Faster moving turbulent water typically contains greater concentrations of

dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...

, which supports greater biodiversity than the slow-moving water of pools. These distinctions form the basis for the division of rivers into

upland and lowland Upland and lowland are conditional descriptions of a plain based on elevation above sea level. In studies of the ecology of freshwater rivers, habitats are classified as upland or lowland. Definitions Upland and lowland are portions of p ...

rivers. The food base of streams within riparian forests 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 An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...

. A dam produces negative effects that continue down the watershed. The most important negative effects are 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 Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...

''lotus'', meaning washed. Lotic waters range from

spring Spring(s) may refer to: Common uses * Spring (season), a season of the year * Spring (device), a mechanical device that stores energy * Spring (hydrology), a natural source of water * Spring (mathematics), a geometric surface in the shape of a h ...

s only a few centimeters wide to major

river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of ...

s kilometers in width.Allan, J.D. 1995. Stream Ecology: structure and function of running waters. Chapman and Hall, London. Pp. 388. Much of this article applies to lotic ecosystems in general, including related lotic systems such as

stream A stream is a continuous body of surface water flowing within the bed and banks of a channel. Depending on its location or certain characteristics, a stream may be referred to by a variety of local or regional names. Long large streams ...

s and springs. Lotic ecosystems can be contrasted with lentic ecosystems, which involve relatively still terrestrial waters such as lakes, ponds, and

wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...

s. Together, these two ecosystems form 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 the biota is specialized to live with flow conditions.Giller, S. and B. Malmqvist. 1998. The Biology of Streams and Rivers. Oxford University Press, Oxford. Pp. 296.

Abiotic components (non-living)

The non-living components of an ecosystem are called abiotic components. E.g. stone, air, soil, etc.

Water flow

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 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 the channel, sinuosity, obstructions, and the incline gradient. In addition, the amount of water input into the system from direct precipitation,

snowmelt In hydrology, snowmelt is surface runoff produced from melting snow. It can also be used to describe the period or season during which such runoff is produced. Water produced by snowmelt is an important part of the annual water cycle in many p ...

, and/or groundwater can affect the flow rate. The amount of water in a stream is measured as

discharge Discharge may refer to Expel or let go * Discharge, the act of firing a gun * Discharge, or termination of employment, the end of an employee's duration with an employer * Military discharge, the release of a member of the armed forces from ser ...

(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 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 Geomorphology (from Ancient Greek: , ', "earth"; , ', "form"; and , ', "study") is the scientific study of the origin and evolution of topographic and bathymetric features created by physical, chemical or biological processes operating at or ...

. 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 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.Cushing, C.E. and J.D. Allan. 2001. Streams: their ecology and life. Academic Press, San Diego. Pp. 366.

Light

Light is important to lotic systems, because it provides the energy necessary to drive

primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through ...

via

photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...

, 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

Temperature

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.

Chemistry

Water chemistry in river ecosystems varies depending on which dissolved solutes and gases are present in the

water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...

of the stream. Specifically river water can include, apart from the water itself, * dissolved inorganic matter and major ions (calcium, sodium, magnesium, potassium, bicarbonate, sulphide, chloride) * dissolved inorganic nutrients (nitrogen, phosphorus, silica) * suspended and

dissolved organic matter Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called parti ...

* gases (nitrogen, nitrous oxide, carbon dioxide, oxygen) * trace metals and pollutants

Dissolved ions and nutrients

Dissolved stream solutes can be considered either ''reactive'' or ''conservative''. Reactive solutes are readily biologically assimilated by the

autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...

ic and heterotrophic biota of 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 is foremost determined by inputs from the geology of its

, 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 the river's source.

Dissolved gases

In terms of dissolved gases,

oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...

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 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.

Suspended matter

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.

Substrate

The inorganic substrate of lotic systems is composed of the

geologic Geology () is a branch of natural science concerned with Earth and other astronomical objects, the features or rocks of which it is composed, and the processes by which they change over time. Modern geology significantly overlaps all other E ...

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 Grain size (or particle size) is the diameter of individual grains of sediment, or the lithified particles in clastic rocks. The term may also be applied to other granular materials. This is different from the crystallite size, which refer ...

, 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 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.

Biotic components (living)

The living components of an ecosystem are called 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.

Biofilm

biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular po ...

is a combination of algae (diatoms etc.), fungi, bacteria, and other small

microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...

s that exist in a film along the

streambed A stream bed or streambed is the bottom of a stream or river (bathymetry) or the physical confine of the normal water flow ( channel). The lateral confines or channel margins are known as the stream banks or river banks, during all but flood ...

or the

benthos Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.exopolysaccharide matrix (EPS), and are net receptors of inorganic and organic elements and remain submitted to the influences of the different environmental factors. CC-BY icon

Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
Biofilms are one of the main biological interphases in river ecosystems, and probably the most important in intermittent rivers, where the importance of the

is reduced during extended low-activity periods of the

hydrological cycle The water cycle, also known as the hydrologic cycle or the hydrological cycle, is a biogeochemical cycle that describes the continuous movement of water on, above and below the surface of the Earth. The mass of water on Earth remains fairly const ...

. Biofilms can be understood as microbial consortia of

s and

heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...

s, coexisting in a matrix of hydrated

extracellular polymeric substance Extracellular polymeric substances (EPSs) are natural polymers of high molecular weight secreted by microorganisms into their environment. EPSs establish the functional and structural integrity of biofilms, and are considered the fundamental comp ...

s (EPS). These two main biological components are respectively mainly

algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...

and

cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...

on one side, and

bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...

and

fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...

on the other.

Micro Micro may refer to: Measurement * micro- (μ), a metric prefix denoting a factor of 10−6 Places * Micro, North Carolina, town in U.S. People * DJ Micro, (born Michael Marsicano) an American trance DJ and producer * Chii Tomiya (都宮ち ...

- 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 the formation of synergistic consortia. The EPS is able to retain

extracellular enzyme Extracellular enzymes or exoenzymes are synthesized inside the cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a relig ...

s 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.,

biocide A biocide is defined in the European legislation as a chemical substance or microorganism intended to destroy, deter, render harmless, or exert a controlling effect on any harmful organism. The US Environmental Protection Agency (EPA) uses a sli ...

UV radiation Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30 PHz) to 400 nm (750 THz), shorter than that of visible light, but longer than X-rays. UV radiatio ...

, etc.) from 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. Co-occurrence networks of bacterial communities in a stream

Co-occurrence networks of bacterial communities in a stream

Microorganisms

Bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...

are present in large numbers in lotic waters. Free-living forms are associated with decomposing organic material,

on the surfaces of rocks and vegetation, in between particles that compose the substrate, and suspended in the

. 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 Below may refer to: *Earth * Ground (disambiguation) *Soil *Floor * Bottom (disambiguation) *Less than *Temperatures below freezing *Hell or underworld People with the surname *Ernst von Below (1863–1955), German World War I general *Fred Below ...

Diatom A diatom ( Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising se ...

s 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 a broad spectrum of tolerances to conditions ranging, from oligotrophic to eutrophic.

Primary producers

Algae, consisting of

phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. ...

and

periphyton Periphyton is a complex mixture of algae, cyanobacteria, heterotrophic microbes, and detritus that is attached to submerged surfaces in most aquatic ecosystems. The related term Aufwuchs (German "surface growth" or "overgrowth") refers to the col ...

, 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 a gelatinous, unanchored floating mat. water hyacinth

Plants exhibit limited adaptations to fast flow and are most successful in reduced currents. More primitive plants, such as

moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta ('' sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and ...

es and

liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ...

s 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 Lemnoideae is a subfamily of flowering aquatic plants, known as duckweeds, water lentils, or water lenses. They float on or just beneath the surface of still or slow-moving bodies of fresh water and wetlands. Also known as bayroot, they arose f ...

water hyacinth ''Pontederia crassipes'' (formerly ''Eichhornia crassipes''), commonly known as common water hyacinth is an aquatic plant native to South America, naturalized throughout the world, and often invasive outside its native range.invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...

s in some lotic systems are

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

s. These species exhibit tremendous diversity and can be found occupying almost every available habitat, including the surfaces of stones, deep below the substratum in the

hyporheic zone The hyporheic zone is the region of sediment and porous space beneath and alongside a stream bed, where there is mixing of shallow groundwater and surface water. The flow dynamics and behavior in this zone (termed hyporheic flow or underflow) is re ...

, 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 the giant water bug (

Belostomatidae Belostomatidae is a family of freshwater hemipteran insects known as giant water bugs or colloquially as toe-biters, Indian toe-biters, electric-light bugs, alligator ticks, or alligator fleas (in Florida). They are the largest insects in the or ...

), avoid flood events by leaving the stream when they sense rainfall. In addition to these behaviors and body shapes, insects have different life history

adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...

s 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 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 Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are aquatic insects belonging to the orde ...

), Trichoptera (also known as a caddisfly),

Plecoptera Plecoptera is an order of insects, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the mos ...

(also known as a

stonefly Plecoptera is an order of insects, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the mos ...

Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...

(also known as a true

fly Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...

), some types of

Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describe ...

(also known as a

beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...

Odonata Odonata is an order of flying insects that includes the dragonflies and damselflies. Members of the group first appeared during the Triassic, though members of their total group, Odonatoptera, first appeared in Late Carboniferous. The two com ...

(the group that includes the

dragonfly A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat thre ...

and the

damselfly Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies, which constitute the other odonatan suborder, Anisoptera, but are smaller and have slimmer bodies. Most species fold the wings along ...

), and some types of

Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to arou ...

(also known as true bugs). Additional invertebrate

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

common to flowing waters include

mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is e ...

s such as

snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...

limpet Limpets are a group of aquatic snails that exhibit a conical gastropod shell, shell shape (patelliform) and a strong, muscular foot. Limpets are members of the class Gastropoda, but are polyphyletic, meaning the various groups called "limpets" ...

s, clams,

mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other edible clams, which ...

s, as well as

crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...

s like

crayfish Crayfish are freshwater crustaceans belonging to the clade Astacidea, which also contains lobsters. In some locations, they are also known as crawfish, craydids, crawdaddies, crawdads, freshwater lobsters, mountain lobsters, rock lobsters, m ...

amphipoda Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far descri ...

and

crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all th ...

Fish and other vertebrates

Fish are probably 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 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 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 Salmon () is the common name for several commercially important species of euryhaline ray-finned fish from the family Salmonidae, which are native to tributaries of the North Atlantic (genus '' Salmo'') and North Pacific (genus '' Onco ...

, for example, are

anadromous Fish migration is mass relocation by fish from one area or body of water to another. Many types of fish migrate on a regular basis, on time scales ranging from daily to annually or longer, and over distances ranging from a few metres to thousa ...

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 the opposite, living in freshwater as adults but migrating to the ocean to spawn. Other vertebrate taxa that inhabit lotic systems include

amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...

s, such as

salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...

s (e.g. snakes, turtles, crocodiles and alligators) various bird species, and mammals (e.g.,

otter Otters are carnivorous mammals in the subfamily Lutrinae. The 13 extant otter species are all semiaquatic, aquatic, or marine, with diets based on fish and invertebrates. Lutrinae is a branch of the Mustelidae family, which also includes we ...

beaver Beavers are large, semiaquatic rodents in the genus ''Castor'' native to the temperate Northern Hemisphere. There are two extant species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers a ...

s, 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 the larger vertebrates mentioned above.

Trophic level dynamics

The concept of

trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it ...

s 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. # Level one: Producers, plant-like organisms that generate their own food using solar radiation, including

es and

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.Consumer A consumer is a person or a group who intends to order, or uses purchased goods, products, or services primarily for personal, social, family, household and similar needs, who is not directly related to entrepreneurial or business activities. ...

s, animal-like organism that get their energy from eating producers, such as

zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...

, small fish, and

s. # Level three: Decomposers, organisms that break down the dead matter of consumers and producers and return the nutrients back to the system. Example are

and

. All energy transactions within an

ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...

derive from a single external source of energy, the sun. Some of this solar radiation 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 Catabolism () is the set of metabolic pathways that breaks down molecules into smaller units that are either oxidized to release energy or used in other anabolic reactions. Catabolism breaks down large molecules (such as polysaccharides, lip ...

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 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 Ecological succession is the process of change in the species structure of an ecological community over time. The time scale can be decades (for example, after a wildfire) or more or less. Bacteria allows for the cycling of nutrients such as car ...

when observing the transfer of energy within a system.

Top-down and bottom-up affect

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 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 Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...

availability to higher trophic organism. Top-down regulations occur when 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.

Trophic cascade

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 predator A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintain ...

s to structure entire food web in terms of how they interact with their prey. Trophic cascades can cause drastic changes in 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.

Food chain

A food chain is a linear system of links that is part of a food web, and represents the order in which

organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...

s 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 the top of the food chain has been reached.

Primary producers

Primary producers start every food chain. Their production of energy and nutrients comes from the sun through

. 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

. Production of organic compounds like carbon is what gets transferred up the food chain. Primary producers are consumed by herbivorous

invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...

s that act as the

primary consumer A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s. 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

Primary consumer A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s are the invertebrates and macro-invertebrates that feed upon the primary producers. They play an important role in initiating the transfer of energy from the base trophic level to the next. They are regulatory organisms which facilitate and control rates of

nutrient cycling A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cycli ...

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

that collects on submerged objects, shredders that feed on large leaves and detritus and help break down large material. Also

filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...

s, macro-invertebrates that rely on stream flow to deliver them fine particulate organic matter (FPOM) suspended in the

, and gatherers who feed on FPOM found on the substrate of the river or stream.

Secondary consumers

The

secondary consumer A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other ...

s in a river ecosystem are the predators of the primary consumers. This includes mainly

insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...

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 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 the food chain and depending on the ecosystem, may end with these predatory fish.

Food web complexity

Diversity,

productivity Productivity is the efficiency of production of goods or services expressed by some measure. Measurements of productivity are often expressed as a ratio of an aggregate output to a single input or an aggregate input used in a production proces ...

, 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 Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...

. 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 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 the effect is related to the connectedness of the species to the food web. An

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...

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 (

of primary producers). Food supply or type of producers 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 In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts comm ...

. Food and nutrient supply variability is important for the

succession Succession is the act or process of following in order or sequence. Governance and politics *Order of succession, in politics, the ascension to power by one ruler, official, or monarch after the death, resignation, or removal from office of ...

robustness Robustness is the property of being strong and healthy in constitution. When it is transposed into a system, it refers to the ability of tolerating perturbations that might affect the system’s functional body. In the same line ''robustness'' ca ...

and connectedness of river ecosystem organisms.

Trophic relationships

Energy inputs

Energy sources can be autochthonous or allochthonous. * Autochthonous (from the Latin "auto" = "self) energy sources are those derived from within the lotic system. During

, for example, primary producers form organic carbon compounds out of carbon dioxide and inorganic matter. The energy they produce is important for the community because it may be transferred to higher

s via consumption. Additionally, high rates of

can introduce

(DOM) to the waters. Another form of autochthonous energy comes from the decomposition of dead organisms and feces that originate within the lotic system. In this case, bacteria decompose the

or coarse particulate organic material (CPOM; >1 mm pieces) into fine particulate organic matter (FPOM; <1 mm pieces) and then further into inorganic compounds that are required for photosynthesis. This process is discussed in more detail below. Stream at the Walhalla ravine

* Allochthonous energy sources are those derived from outside the lotic system, that is, from the terrestrial environment. Leaves, twigs, fruits, etc. are typical forms of terrestrial CPOM that have entered the water by direct litter fall or lateral leaf blow. In addition, terrestrial animal-derived materials, such as feces or carcasses that have been added to the system are examples of allochthonous CPOM. The CPOM undergoes a specific process of degradation. Allan gives the example of a leaf fallen into a stream. First, the soluble chemicals are dissolved and leached from the leaf upon its saturation with water. This adds to the DOM load in the system. Next

microbe A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...

s such as

and

colonize Colonization, or colonisation, constitutes large-scale population movements wherein migrants maintain strong links with their, or their ancestors', former country – by such links, gain advantage over other inhabitants of the territory. When ...

the leaf, softening it as the

mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates ...

of the fungus grows into it. The composition of the microbial community is influenced by the species of tree from which the leaves are shed (Rubbo and Kiesecker 2004). This combination of bacteria, fungi, and leaf are a food source for shredding invertebrates, which leave only FPOM after consumption. These fine particles may be colonized by microbes again or serve as a food source for animals that consume FPOM. Organic matter can also enter the lotic system already in the FPOM stage by wind,

surface runoff Surface runoff (also known as overland flow) is the flow of water occurring on the ground surface when excess rainwater, stormwater, meltwater, or other sources, can no longer sufficiently rapidly infiltrate in the soil. This can occur when t ...

bank A bank is a financial institution that accepts Deposit account, deposits from the public and creates a demand deposit while simultaneously making loans. Lending activities can be directly performed by the bank or indirectly through capital m ...

erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is d ...

, or

groundwater Groundwater is the water present beneath Earth's surface in rock and Pore space in soil, soil pore spaces and in the fractures of stratum, rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit ...

. Similarly, DOM can be introduced through canopy drip from rain or from surface flows.

Invertebrates

Invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...

s 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 feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...

s, which use their

setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...

, filtering aparati, nets, or even secretions to collect FPOM and microbes from 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

and

. Finally, several families are predatory, capturing and consuming animal prey. Both 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

Fish can also be placed into feeding guilds. Planktivores pick

plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cruc ...

out of the

Herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...

detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrate ...

s are bottom-feeding species that ingest both

and

indiscriminately. Surface and water column feeders capture surface prey (mainly terrestrial and emerging insects) and drift (

benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning " ...

invertebrates floating downstream). Benthic invertebrate feeders prey primarily on immature insects, but will also consume other benthic invertebrates.

Top predators An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic le ...

consume fishes and/or large invertebrates.

Omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nu ...

s ingest a wide range of prey. These can be floral,

fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is '' flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. ...

l, and/or detrital in nature. Finally,

parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...

s 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 the most benthic invertebrate feeders, and tropical systems having large numbers of detritus feeders due to high rates of allochthonous input.

Community patterns and diversity

Local species richness

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, a combination of factors such as historical rates of

speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution withi ...

and

extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...

, type of substrate, microhabitat availability, water chemistry, temperature, and disturbance such as flooding seem to be important.

Resource partitioning

Although many alternate theories have been postulated for the ability of guild-mates to coexist (see Morin 1999),

resource partitioning In ecology, niche differentiation (also known as niche segregation, niche separation and niche partitioning) refers to the process by which competing species use the environment differently in a way that helps them to coexist. The competitive exclu ...

has been well documented in lotic systems as 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) 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 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

s, for example, can filter varying particle size of FPOM from 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 the timing of maximum growth among guild mates. Tropical fishes in

Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and e ...

, for example, have shifted to shorter life spans in response to the

ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...

reduction felt with increasing levels of species richness in their ecosystem (Watson and Balon 1984).

Persistence and succession

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 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 Colonization, or colonisation, constitutes large-scale population movements wherein migrants maintain strong links with their, or their ancestors', former country – by such links, gain advantage over other inhabitants of the territory. When ...

. In these cases, an entirely new community that is well adapted to the conditions found in this new area can establish itself.

River continuum concept

The

River continuum concept The River Continuum Concept (RCC) is a model for classifying and describing flowing water, in addition to the classification of individual sections of waters after the occurrence of indicator organisms. The theory is based on the concept of dyna ...

(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 the gradient from a small stream eventually linked to a large river. Stream order (see characteristics of streams) is used as the physical measure of the position along the RCC. According to the RCC, low ordered sites are small shaded streams where allochthonous inputs of CPOM are a necessary resource for consumers. As the river widens at mid-ordered sites, energy inputs should change. Ample sunlight should reach the bottom in these systems to support significant periphyton production. Additionally, the biological processing of CPOM (Coarse Particulate Organic Matter - larger than 1 mm) inputs at upstream sites is expected to result in the transport of large amounts of FPOM (Fine Particulate Organic Matter - smaller than 1 mm) to these downstream ecosystems. Plants should become more abundant at edges of the river with increasing river size, especially in lowland rivers where finer sediments have been deposited and facilitate rooting. The main channels likely have too much current and turbidity and a lack of substrate to support plants or periphyton. Phytoplankton should produce the only autochthonous inputs here, but photosynthetic rates will be limited due to turbidity and mixing. Thus, allochthonous inputs are expected to be the primary energy source for large rivers. This FPOM will come from both upstream sites via the decomposition process and through lateral inputs from floodplains. Biota should change with this change in energy from the headwaters to the mouth of these systems. Namely, shredders should prosper in low-ordered systems and grazers in mid-ordered sites. Microbial decomposition should play the largest role in energy production for low-ordered sites and large rivers, while photosynthesis, in addition to degraded allochthonous inputs from upstream will be essential in mid-ordered systems. As mid-ordered sites will theoretically receive the largest variety of energy inputs, they might be expected to host the most biological diversity (Vannote et al. 1980). Just how well the RCC actually reflects patterns in natural systems is uncertain and its generality can be a handicap when applied to diverse and specific situations. The most noted criticisms of the RCC are: 1. It focuses mostly on

macroinvertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...

, disregarding that plankton and fish diversity is highest in high orders; 2. It relies heavily on the fact that low ordered sites have high CPOM inputs, even though many streams lack

riparian A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks a ...

habitats; 3. It is based on pristine systems, which rarely exist today; and 4. It is centered around the functioning of temperate streams. Despite its shortcomings, the RCC remains a useful idea for describing how the patterns of ecological functions in a lotic system can vary from the source to the mouth. Disturbances such as congestion by dams or natural events such as shore flooding are not included in the RCC model.Junk J.W., P.B. Bayley, R.E. Sparks: “The flood pulse concept in river flood plain systems”. Canadian Special Publications of Fisheries and Aquatic Sciences. 106. 1989. Various researchers have since expanded the model to account for such irregularities. For example, J.V. Ward and J.A. Stanford came up with the Serial Discontinuity Concept in 1983, which addresses the impact of geomorphologic disorders such as congestion and integrated inflows. The same authors presented the Hyporheic Corridor concept in 1993, in which the vertical (in depth) and lateral (from shore to shore) structural complexity of the river were connected. The flood pulse concept, developed by W.J. Junk in 1989, further modified by P.B. Bayley in 1990 and K. Tockner in 2000, takes into account the large amount of nutrients and organic material that makes its way into a river from the sediment of surrounding flooded land.

Human impacts

Human impact factors on river–floodplain systems

Humans exert a geomorphic force that now rivals that of the natural Earth. The period of human dominance has been termed the Anthropocene, and several dates have been proposed for its onset. Many researchers have emphasised the dramatic changes associated with the

Industrial Revolution The Industrial Revolution was the transition to new manufacturing processes in Great Britain, continental Europe, and the United States, that occurred during the period from around 1760 to about 1820–1840. This transition included going f ...

in Europe after about 1750 CE (Common Era) and the Great Acceleration in technology at about 1950 CE. CC-BY icon

Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
However, a detectable human imprint on the environment extends back for thousands of years, and an emphasis on recent changes minimises the enormous landscape transformation caused by humans in antiquity. Important earlier human effects with significant environmental consequences include megafaunal extinctions between 14,000 and 10,500 cal yr BP;

domestication of plants Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...

and

animals Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...

close to the start of the

Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...

at 11,700 cal yr BP; agricultural practices and deforestation at 10,000 to 5000 cal yr BP; and widespread generation of anthropogenic soils at about 2000 cal yr BP. Key evidence of early anthropogenic activity is encoded in early fluvial successions, long predating

anthropogenic effects Human impact on the environment (or anthropogenic impact) refers to changes to biophysical environments and to ecosystems, biodiversity, and natural resources caused directly or indirectly by humans. Modifying the environment to fit the needs ...

that have intensified over the past centuries and led to the modern worldwide river crisis.

Pollution

River pollution can include but is not limited to: increasing sediment export, excess nutrients from fertilizer or urban runoff, sewage and septic inputs,

plastic pollution Plastic pollution is the accumulation of plastic objects and particles (e.g. plastic bottles, bags and microbeads) in the Earth's environment that adversely affects humans, wildlife and their habitat. Plastics that act as pollutants are cate ...

, nano-particles, pharmaceuticals and personal care products, synthetic chemicals, road salt, inorganic contaminants (e.g., heavy metals), and even heat via thermal pollutions. The effects of pollution often depend on the context and material, but can reduce

ecosystem function An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...

ing, limit

ecosystem services Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...

, reduce stream biodiversity, and impact human health. Pollutant sources of lotic systems are hard to control because they can derive, often in small amounts, over a very wide area and enter the system at many locations along its length. While direct pollution of lotic systems has been greatly reduced in the United States under the government's

Clean Water Act The Clean Water Act (CWA) is the primary federal law in the United States governing water pollution. Its objective is to restore and maintain the chemical, physical, and biological integrity of the nation's waters; recognizing the responsibiliti ...

, contaminants from diffuse non-point sources remain a large problem. Agricultural fields often deliver large quantities of sediments, nutrients, and chemicals to nearby streams and rivers. Urban and residential areas can also add to this pollution when contaminants are accumulated on impervious surfaces such as roads and parking lots that then drain into the system. Elevated nutrient concentrations, especially nitrogen and phosphorus which are key components of fertilizers, can increase periphyton growth, which can be particularly dangerous in slow-moving streams. Another pollutant,

acid rain Acid rain is rain or any other form of precipitation that is unusually acidic, meaning that it has elevated levels of hydrogen ions (low pH). Most water, including drinking water, has a neutral pH that exists between 6.5 and 8.5, but ac ...

, forms from sulfur dioxide and nitrous oxide emitted from factories and power stations. These substances readily dissolve in atmospheric moisture and enter lotic systems through precipitation. This can lower the pH of these sites, affecting all trophic levels from algae to vertebrates. Mean species richness and total species numbers within a system decrease with decreasing pH.

Flow modification

Flow modification can occur as a result of dams, water regulation and extraction, channel modification, and the destruction of the river floodplain and adjacent riparian zones. Dams alter the flow, temperature, and sediment regime of lotic systems. Additionally, many rivers are dammed at multiple locations, amplifying the impact. Dams can cause enhanced clarity and reduced variability in stream flow, which in turn cause an increase in

abundance. \Invertebrates immediately below a dam can show reductions in species richness due to an overall reduction in habitat heterogeneity. Also, thermal changes can affect insect development, with abnormally warm winter temperatures obscuring cues to break egg

diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...

and overly cool summer temperatures leaving too few acceptable days to complete growth. Finally, dams fragment river systems, isolating previously continuous populations, and preventing the migrations of

and catadromous species.

Invasive species

Invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...

have been introduced to lotic systems through both purposeful events (e.g. stocking game and food species) as well as unintentional events (e.g. hitchhikers on boats or fishing waders). These organisms can affect natives via competition for prey or habitat, predation, habitat alteration, hybridization, or the introduction of harmful diseases and parasites. Once established, these species can be difficult to control or eradicate, particularly because of the connectivity of lotic systems. Invasive species can be especially harmful in areas that have

endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and in ...

biota, such as mussels in the Southeast United States, or those that have localized

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...

species, like lotic systems west of the Rocky Mountains, where many species evolved in isolation.

References

External links

USGS real time stream flow data for gauged systems nationwide
{{modelling ecosystems, expanded=none Aquatic ecology Ecosystems Freshwater ecology Limnology Riparian zone Rivers Water streams

Abiotic components (non-living)

Water flow

Light

Temperature

Chemistry

Dissolved ions and nutrients

Dissolved gases

Suspended matter

Substrate

Biotic components (living)

Biofilm

Microorganisms

Primary producers

Fish and other vertebrates

Trophic level dynamics

Top-down and bottom-up affect

Trophic cascade

Food chain

Primary producers

Primary consumers

Secondary consumers

Food web complexity

Trophic relationships

Energy inputs

Invertebrates

Fish

Community patterns and diversity

Local species richness

Resource partitioning

Persistence and succession

River continuum concept

Human impacts

Pollution

Flow modification

Invasive species

See also

References

Further reading

External links