![Stream in the redwoods](https://upload.wikimedia.org/wikipedia/commons/e/e6/Stream_in_the_redwoods.jpg)
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
In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
(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
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 ...
are part of larger
watershed networks or catchments, where smaller
headwater 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
Canopy may refer to:
Plants
* Canopy (biology), aboveground portion of plant community or crop (including forests)
* Canopy (grape), aboveground portion of grapes
Religion and ceremonies
* Baldachin or canopy of state, typically placed over an ...
derive the majority of their food base from algae.
Anadromous fish
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 thousan ...
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 ther ...
.
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 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 the power of the ...
''lotus'', meaning washed. Lotic waters range from
spring
Spring(s) may refer to:
Common uses
* Spring (season)
Spring, also known as springtime, is one of the four temperate seasons, succeeding winter and preceding summer. There are various technical definitions of spring, but local usage of ...
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 w ...
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
streams and
springs. Lotic ecosystems can be contrasted with
lentic ecosystems
A lake ecosystem or lacustrine ecosystem includes biotic (living) plants, animals and micro-organisms, as well as abiotic (non-living) physical and chemical interactions. Lake ecosystems are a prime example of lentic ecosystems (''lentic'' ref ...
, 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) processes prevailing, especially in the soils. The p ...
s. Together, these two ecosystems form the more general study area of freshwater or
aquatic ecology
An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms that are dependent on each other and on their environment. The tw ...
.
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 (
microhabitat
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s), 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
![Overlander Falls](https://upload.wikimedia.org/wikipedia/commons/4/43/Overlander_Falls.JPG)
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
Sinuosity, sinuosity index, or sinuosity coefficient of a continuously differentiable curve having at least one inflection point is the ratio of the curvilinear length (along the curve) and the Euclidean distance (straight line) between the en ...
, 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 par ...
, 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 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 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 riffle
A riffle is a shallow landform in a flowing channel. Colloquially, it is a shallow place in a river where water flows quickly past rocks. However, in geology a riffle has specific characteristics.
Topographic, sedimentary and hydraulic indica ...
s, 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 i ...
, 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 poikilotherm
A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
s 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 ion
An ion () is an atom or molecule with a net electrical charge.
The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conve ...
s (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 part ...
* 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
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 ...
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 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 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 as ...
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 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
Particle size is a notion introduced for comparing dimensions of solid particles ('' flecks''), liquid particles ('' droplets''), or gaseous particles ('' bubbles''). The notion of particle size applies to particles in colloids, in ecology, in ...
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
Large woody debris (LWD) are the logs, sticks, branches, and other wood that falls into streams and rivers. This debris can influence the flow and the shape of the stream channel. Large woody debris, grains, and the shape of the bed of the stream ...
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
A 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 ...
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 olde ...
s that exist in a film along the streambed 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
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 ...](_b ...<br></span></div>. Biofilm assemblages themselves are complex, and add to the complexity of a streambed.
The different biofilm components (algae and bacteria are the principal components) are embedded in an <div class=)
matrix (EPS), and are net receptors of inorganic and organic elements and remain submitted to the influences of the different environmental factors.[ ]
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 river
Intermittent, temporary or seasonal rivers or streams cease to flow every year or at least twice every five years.(Tzoraki et al., 2007) Such rivers drain large arid and semi-arid areas, covering approximately a third of the earth's surface. ...
s, where the importance of 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 ...
is reduced during extended low-activity periods of the hydrological cycle. Biofilms can be understood as microbial consortia of 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 ...
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 substances (EPS). These two main biological components are respectively mainly algae and cyanobacteria on one side, and bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
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 from ...
on the other. Micro- and meiofauna
Meiobenthos, also called meiofauna, are small benthic invertebrates that live in both marine and fresh water environments. The term ''meiofauna'' loosely defines a group of organisms by their size, larger than microfauna but smaller than macrof ...
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 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
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 radiation i ...
, 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.
Microorganisms
Bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
are present in large numbers in lotic waters. Free-living forms are associated with decomposing organic material, 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 ...
on the surfaces of rocks and vegetation, in between particles that compose the substrate, and suspended 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 ...
. Other forms are also associated with the guts of lotic organisms as parasites or in commensal
Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
relationships.[ Bacteria play a large role in energy recycling (see below).][
Diatoms are one of the main dominant groups of ]periphytic
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 ...
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 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.][
]
Plants exhibit limited adaptations to fast flow and are most successful in reduced currents. More primitive plants, such as mosses 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 ...
or 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 chordate ...]
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 ...
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 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 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
The caddisflies, or order Trichoptera, are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the b ...
(also known as a caddisfly
The caddisflies, or order Trichoptera, are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the ...
), Plecoptera (also known as a stonefly, Diptera (also known as a true fly), some types of Coleoptera (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 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 (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 G ...
s, limpets, clam
Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shel ...
s, mussels, as well as crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s like crayfish, amphipoda and crabs.[
]
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 '' Oncorhy ...
, 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. Eel
Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
s are catadromous
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 thousan ...
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 amphibians, 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 ...
s, reptiles (e.g. snakes, turtles, crocodiles and alligators) various bird species, and mammals (e.g., otters, beavers, hippos
A hippo or hippopotamus is either of two species of large African mammal which live mainly in and near water:
* Hippopotamus
* Pygmy hippopotamus
Hippo or Hippos may also refer to:
Toponymy
* The ancient city of Hippo Regius (modern Annaba, Al ...
, and river dolphin
River dolphins are a polyphyletic group of fully aquatic mammals that reside exclusively in freshwater or brackish water. They are an informal grouping of dolphins, which itself is a paraphyletic group within the infraorder Cetacea. Extant riv ...
s). 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 web
A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one o ...
s 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
A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
.
# Level one: Producers
Producer or producers may refer to:
Occupations
*Producer (agriculture), a farm operator
*A stakeholder of economic production
*Film producer, supervises the making of films
**Executive producer, contributes to a film's budget and usually does not ...
, plant-like organisms that generate their own food using solar radiation, including algae, phytoplankton, mosses and lichens.
# Level two: 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 crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s.
# Level three: Decomposer
Decomposers are organisms that break down dead or decaying organisms; they carry out decomposition, a process possible by only certain kingdoms, such as fungi. Like herbivores and predators, decomposers are heterotrophic, meaning that they use o ...
s, organisms that break down the dead matter of consumers and producers and return the nutrients back to the system. Example are bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
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 from ...
.
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 syste ...
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, lipid ...
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 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 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 cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce t ...
s. 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 maintaini ...
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
A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
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 ...
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 producer
Primary or primaries may refer to:
Arts, entertainment, and media Music Groups and labels
* Primary (band), from Australia
* Primary (musician), hip hop musician and record producer from South Korea
* Primary Music, Israeli record label
Works
* ...
s start every food chain. Their production of energy and nutrients comes from the sun through 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 i ...
. 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
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 w ...
. 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 chordate ...
s 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
Primary consumers 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 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 ...
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 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 ...
, and gatherers who feed on FPOM found on the substrate of the river or stream.
Secondary consumers
The secondary consumers in a river ecosystem are the predators
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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 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
Diversity, diversify, or diverse may refer to:
Business
*Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce
*Diversity marketing, marketing communication targeting diverse customers
* ...
, 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
Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative ab ...
, composition
Composition or Compositions may refer to:
Arts and literature
*Composition (dance), practice and teaching of choreography
*Composition (language), in literature and rhetoric, producing a work in spoken tradition and written discourse, to include v ...
and stability
Stability may refer to:
Mathematics
*Stability theory, the study of the stability of solutions to differential equations and dynamical systems
** Asymptotic stability
** Linear stability
** Lyapunov stability
** Orbital stability
** Structural sta ...
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'') l ...
. 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
Stability may refer to:
Mathematics
*Stability theory, the study of the stability of solutions to differential equations and dynamical systems
** Asymptotic stability
** Linear stability
** Lyapunov stability
** Orbital stability
** Structural sta ...
/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 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 Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce t ...
. One highly variable component to river ecosystems is food supply ( biomass of primary producer
Primary or primaries may refer to:
Arts, entertainment, and media Music Groups and labels
* Primary (band), from Australia
* Primary (musician), hip hop musician and record producer from South Korea
* Primary Music, Israeli record label
Works
* ...
s). 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. Food and nutrient supply variability is important for the succession, 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
Autochthon, autochthons or autochthonous may refer to:
Fiction
* Autochthon (Atlantis), a character in Plato's myth of Atlantis
* Autochthons, characters in the novel ''The Divine Invasion'' by Philip K. Dick
* Autochthon, a Primordial in the ...
or allochthonous.
* Autochthonous (from the Latin "auto" = "self) energy sources are those derived from within the lotic system. During 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 i ...
, for example, primary producer
Primary or primaries may refer to:
Arts, entertainment, and media Music Groups and labels
* Primary (band), from Australia
* Primary (musician), hip hop musician and record producer from South Korea
* Primary Music, Israeli record label
Works
* ...
s 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 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 via consumption. Additionally, high rates of 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 ...
can introduce 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 part ...
(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 detritus or coarse ]particulate
Particulates – also known as atmospheric aerosol particles, atmospheric particulate matter, particulate matter (PM) or suspended particulate matter (SPM) – are microscopic particles of solid or liquid matter suspended in the air. The te ...
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.
]
* 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 microbes such as ]bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
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 from ...
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, bank
A bank is a financial institution that accepts 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 markets.
Because ...
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 dis ...
, or groundwater
Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidated ...
. Similarly, DOM can be introduced through canopy
Canopy may refer to:
Plants
* Canopy (biology), aboveground portion of plant community or crop (including forests)
* Canopy (grape), aboveground portion of grapes
Religion and ceremonies
* Baldachin or canopy of state, typically placed over an ...
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 chordate ...
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 feeders, 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 ]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 ...
and detritus. 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 crucia ...
out of 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 ...
. Herbivore- detritivores are bottom-feeding species that ingest both 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 ...
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
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. Omnivores ingest a wide range of prey. These can be floral
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
, 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 (ecology ...
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 has ...
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
Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative ab ...
. In these cases, a combination of factors such as historical rates of speciation 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 last individual of the species, although the capacity to breed and ...
, type of substrate, microhabitat
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
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 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 suspension feeders, 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 ea ...
, 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. 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 (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 s ...
, 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
Humans exert a geomorphic
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 ...
force that now rivals that of the natural Earth. The period of human dominance has been termed the Anthropocene
The Anthropocene ( ) is a proposed geological epoch dating from the commencement of significant human impact on Earth's geology and ecosystems, including, but not limited to, anthropogenic climate change.
, neither the International Commissio ...
, 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.[ ]
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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 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 catego ...
, 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, 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, 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 dam
A dam is a barrier that stops or restricts the flow of surface water or underground streams. Reservoirs created by dams not only suppress floods but also provide water for activities such as irrigation, human consumption, industrial use ...
s, water regulation and extraction, channel modification, and the destruction of the river floodplain and adjacent riparian zones.
Dam
A dam is a barrier that stops or restricts the flow of surface water or underground streams. Reservoirs created by dams not only suppress floods but also provide water for activities such as irrigation, human consumption, industrial use ...
s 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 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 ...
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 anadromous and catadromous
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 thousan ...
species.
Invasive species
Invasive species 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.
See also
* Betty's Brain software that "learns" about river ecosystems
* Flood pulse concept
* Lake ecosystem
A lake ecosystem or lacustrine ecosystem includes biotic (living) plants, animals and micro-organisms, as well as abiotic (non-living) physical and chemical interactions. Lake ecosystems are a prime example of lentic ecosystems (''lentic'' ref ...
* Rheophile
A rheophile is an animal that prefers to live in fast-moving water.
Examples of rheophilic animals
Insects
*Many aquatic insects living in riffles require current to survive.
*'' Epeorus sylvicola'', a rheophilic mayfly species ( Ephemeroptera ...
* Riparian zone
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 ...
* River continuum concept
* RIVPACS
* '' The Riverkeepers''
* Upland and lowland rivers
References
Further reading
* Brown, A.L. 1987. Freshwater Ecology. Heinimann Educational Books, London. Pp. 163.
* Carlisle, D.M. and M.D. Woodside. 2013.
Ecological health in the nation's streams
United States Geological Survey
The United States Geological Survey (USGS), formerly simply known as the Geological Survey, is a scientific agency of the United States government. The scientists of the USGS study the landscape of the United States, its natural resources, ...
. Pp. 6.
* Edington, J.M., Edington, M.A., and J.A. Dorman. 1984. Habitat partitioning amongst hydrophyschid larvae of a Malaysian stream. Entomologica 30: 123–129.
* Hynes, H.B.N. 1970. Ecology of Running Waters. Originally published in Toronto by University of Toronto Press, 555p
* Morin, P.J. 1999. Community Ecology. Blackwell Science, Oxford. Pp. 424.
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* Ward, J.V. 1992. Aquatic Insect Ecology: biology and habitat. Wiley, New York. Pp. 456.
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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